Corded Ware—Uralic (II): Finno-Permic and the expansion of N-L392/Siberian ancestry

finno-ugric-samoyedic

This is the second of three posts on the Corded Ware—Uralic identification. See Corded Ware—Uralic(I): Differences and similarities with Yamna.

I read from time to time that “we have not sampled Uralic speakers yet”, and “we are waiting to see when Uralic-speaking peoples are sampled”. Are we, though?

Proto-language homelands are based on linguistic data, such as guesstimates for dialectal evolution, loanwords and phonetic changes for language contacts, toponymy for ancient territories, etc. depending on the available information. The trace is then followed back, using available archaeological data, from the known historic speakers and territory to the appropriate potential prehistoric cultures. Only then can genetic analyses help us clarify the precise prehistoric population movements that better fit the models.

uralic-language-family
The traditional family tree of the Uralic branches. Kallio (2014)

The linguistic homeland

We thought – using linguistic guesstimates and fitting prehistoric cultures and their expansion – that Yamna was the Late Proto-Indo-European culture, so when Yamna was sampled, we had Late Proto-Indo-Europeans sampled. Simple deduction.

We thought that north-eastern Europe was a Uralic-speaking area during the Neolithic:

  • For those supporting a western continuity (and assuming CWC was Indo-European), the language was present at least since the Comb Ware culture, potentially since the Mesolithic.
  • For those supporting a late introduction into Finland, Uralic expanded the latest with Abashevo-related movements after its incorporation of Volosovo and related hunter-gatherers.

The expansion to the east must have happened through progressive infiltrations with Seima-Turbino / Andronovo-related expansions.

uralic-time-space
Some datings for the traditional proto-stages from Uralic to Finnic. Kallio (2014).

Finding the linguistic homeland going backwards can be described today as follows:

I. Proto-Fennic homeland

Based on the number of Baltic loanwords, not attested in the more eastern Uralic branches (and reaching only partially Mordvinic), the following can be said about western Finno-Permic languages (Junttila 2014):

The Volga-Kama Basin lies still too far east to be included in a list of possible contact locations. Instead, we could look for the contact area somewhere between Estonia in the west and the surroundings of Moscow in the east, a zone with evidence of Uralic settlement in the north and Baltic on the south side.

The only linguistically well-grounded version of the Stone Age continuation theory was presented by Mikko Korhonen in 1976. Its validity, however, became heavily threatened when Koivulehto 1983a-b proved the existence of a Late Proto-Indo-European or Pre-Baltic loanword layer in Saami, Finnic, and Mordvinic. Since this layer must precede the Baltic one and it was presumably acquired in the Baltic Sea region, Koivulehto posited it on the horizon of the Battle Axe period. This forces a later dating for the Baltic–Finnic contacts.

Today the Battle Axe culture is dated at 3200 to 3000 BC, a period far too remote to correspond linguistically with Proto-Baltic (Kallio 1998a).

Since the Baltic contacts began at a very initial phase of Proto-Finnic, the language must have been relatively uniform at that time. Hence, if we consider that the layer of Baltic loanwords may have spread over the Gulf of Finland at that time, we could also insist that the whole of the Proto-Finnic language did so.

migration-theory
Prehistoric Balts as the southern neighbours of Proto-Finnic speakers. 1 = The approximated area of Proto-Uralic. 2 = The approximated area of Finnic during the Iron Age. 3 = The area of ancient Baltic hydronyms. 4 = The area of Baltic languages in about 1200 AD. 5 = The problem: When did Uralic expand westwards and when did it meet Baltic? Junntila (2012).

II. Proto-Finno-Saamic homeland

The evidence of continued Palaeo-Germanic loanwords (from Pre- to Proto-Germanic stages) is certainly the most important data to locate the Finno-Saamic homeland, and from there backwards into the true Uralic homeland. Following Kallio (2017):

(…) the loanword evidence furthermore suggests that the ancestors of Finnic and Saamic had at least phonologically remained very close to Proto-Uralic as late as the Bronze Age (ca. 1700–500 BC). In particular, certain loanwords, whose Baltic and Germanic sources point to the first millennium BC, after all go back to the Finno-Saamic proto-stage, which is phonologically almost identical to the Uralic proto-stage (see especially the table in Sammallahti 1998: 198–202). This being the case, Dahl’s wave model could perhaps have some use in Uralic linguistics, too.

The presence of Pre-Germanic loanwords points rather to the centuries around the turn of the 2nd – 1st millennium BC or earlier. Proto-Germanic words must have been borrowed before the end of Germanic influence in the eastern Baltic at the beginning of the Iron Age, which sets a clear terminus ante quem ca. 800 BC.

The arrival of Bell Beaker peoples in Scandinavia ca. 2350 BC, heralding the formation of the Dagger Period, as well as the development of Pre-Germanic in common with Finnic-like populations point to the late 3rd / early 2nd millennium BC as the first time of close interaction through the Baltic region.

III. Proto-Uralic homeland

(…) the earliest Indo-European loanwords in the Uralic languages (…) show that Proto-Uralic cannot have been spoken much earlier than Proto-Indo-European dated about 3500 BC (Koivulehto 2001: 235, 257). As the same loanword evidence naturally also shows that the Uralic and Indo-European homelands were not located far from one another, the Uralic homeland can most likely be located in the Middle and Upper Volga region, right north of the Indo-European homeland*. From the beginning of the Subneolithic period about 5900 BC onwards, this region was an important innovation centre, from where several cultural waves spread to the Finnish Gulf area, such as the Sperrings Ware wave about 4900 BC, the Combed Ware wave about 3900 BC, and the Netted Ware wave about 1900 BC (Carpelan & Parpola 2001: 78–90).

The mainstream position is nowadays trying to hold together the traditional views of Corded Ware as Indo-European, and a Uralic Fennoscandia during the Bronze Age.

The following is an example of how this “Volosovo/Forest Zone hunter-gatherer theory” of Uralic origins looks like, as a ‘mixture’ of cultures and languages that benefits from the lack of genetic data for certain regions and periods (taken from Parpola 2018):

asbestos-ware
The extent of Typical Comb Ware (TCW), Asbestos- and Organic-tempered Wares (AOW) and Volosovo and Garino-Bor cultures; areas with deposits of native copper in Karelia and copperbearing sandstone in Volga-Kama-area are marked dark gray (after Zhuravlev 1977; Krajnov 1987; Nagovitsyn 1987; Chernykh 1992; Carpelan 1999; Zhul´nikov 1999). From Nordqvist et al. (2012).

The Corded Ware (or Battle Axe) culture intruded into the Eastern Baltic and coastal Finland already around 3100 BCE. The continuity hypothesis maintains that the early Proto-Finnic speakers of the coastal regions, who had come to Finland in the 4th millennium BCE with the Comb-Pitted Ware, coexisted with the Corded Ware newcomers, gradually adopting their pastoral culture and with it a number of NW-IE loanwords, but assimilating the immigrants linguistically.

The fusion of the Corded Ware and the local Comb-Pitted Ware culture resulted into the formation of the Kiukais culture (c. 2300–1500) of southwestern Finland, which around 2300 received some cultural impulses from Estonia, manifested in the appearance of the Western Textile Ceramic (which is different from the more easterly Textile Ceramic or Netted Ware, and which is first attested in Estonia c. 2700 BCE, cf. Kriiska & Tvauri 2007: 88), and supposed to have been accompanied by an influx of loanwords coming from Proto-Baltic. At the same time, the Kiukais culture is supposed to have spread the custom of burying chiefs in stone cairns to Estonia.

The coming of the Corded Ware people and their assimilation created a cultural and supposedly also a linguistic split in Finland, which the continuity hypothesis has interpreted to mean dividing Proto-Saami-Finnic unity into its two branches. Baltic Finnic, or simply Finnic, would have emerged in the coastal regions of Finland and in the northern East Baltic, while preforms of Saami would have been spoken in the inland parts of Finland.

The Nordic Bronze Age culture, correlated above with early Proto-Germanic, exerted a strong influence upon coastal Finland and Estonia 1600–700 BCE. Due to this, the Kiukais culture was transformed into the culture of Paimio ceramics (c. 1600–700 BCE), later continued by Morby ceramics (c. 700 BCE – 200 CE). The assumption is that clear cultural continuity was accompanied by linguistic continuity. Having assimilated the language of the Germanic traders and relatively few settlers of the Bronze Age, the language of coastal Finland is assumed to have reached the stage of Proto-Finnish at the beginning of the Christian era. In Estonia, the Paimio ceramics have a close counterpart in the contemporaneous Asva ceramics.

Eastern homelands?

I will not comment on Siberian or Central Asian homeland proposals, because they are obviously not mainstream, still less today when we know that Uralic was certainly in contact with Proto-Indo-European, and then with Pre- and Proto-Indo-Iranian, as supported even by the Copenhagen group in Damgaard et al. (2018).

This is what Kallio (2017) has to say about the agendas behind such proposals:

Interestingly, the only Uralicists who generally reject the Central Russian homeland are the Russian ones who prefer the Siberian homeland instead. Some Russians even advocate that the Central Russian homeland is only due to Finnish nationalism or, as one of them put it a bit more tactfully, “the political and ideological situation in Finland in the first decades of the 20th century” (Napolskikh 1995: 4).

Still, some Finns (and especially those who also belong to the “school who wants it large and wants it early”) simultaneously advocate that exactly the same Central Russian homeland is due to Finnlandisierung (Wiik 2001: 466).

Hence, for those of you willing to learn about fringe theories not related to North-Eastern Europe, you also have then the large and early version of the Uralic homeland, with Wiik’s Palaeolithic continuity of Uralic peoples spread over all of eastern and central Europe (hence EHG and R1a included):

atlantic-finnic-theory
Palaeolithic boat peoples and Finno-Ugric. Source

These fringe Finnish theories look a lot like the Corded Ware expansion… Better not go the Russian or Finnish nationalist ways? Agreed then, let’s discuss only rational proposals based on current data.

The archaeological homeland

For a detailed account of the Corded Ware expansion with Battle Axe, Fatyanovo-Balanovo, and Abashevo groups into the area, you can read my recent post on the origin of R1a-Z645.

1. Textile ceramics

During the 2nd millennium BC, textile impressions appear in pottery as a feature across a wide region, from the Baltic area through the Volga to the Urals, in communities that evolve from late Corded Ware groups without much external influence.

While it has been held that this style represents a north-west expansion from the Volga region (with the “Netted Ware” expansion), there are actually at least two original textile styles, one (earlier) in the Gulf of Finland, common in the Kiukainen pottery, which evolves into the Textile ware culture proper, and another which seems to have an origin in the Middle Volga region to the south-east.

The Netted ware culture is the one that apparently expands into inner Finland – a region not densely occupied by Corded Ware groups until then. There are, however, no clear boundaries between groups of both styles; textile impressions can be easily copied without much interaction or population movement; and the oldest textile ornamentation appeared on the Gulf of Finland. Hence the tradition of naming all as groups of Textile ceramics.

textile-ware-cultures
Maximum distribution of Textile ceramics during the Bronze Age (ca. 2000-800 BC). Asbestos-tempered ware lies to the north (and is also continued in western Fennoscandia).

The fact that different adjacent groups from the Gulf of Finland and Forest Zone share similar patterns making it very difficult to differentiate between ‘Netted Ware’ or ‘Textile Ware’ groups points to:

  • close cultural connections that are maintained through the Gulf of Finland and the Forest Zone after the evolution of late Corded Ware groups; and
  • no gross population movements in the original Battle Axe / Fatyanovo regions, except for the expansion of Netted Ware to inner Finland, Karelia, and the east, where the scattered Battle Axe finds and worsening climatic conditions suggest most CWC settlements disappeared at the end of the 3rd millennium BC and recovered only later.

NOTE. This lack of population movement – or at least significant replacement by external, non-CWC groups – is confirmed in genetic investigation by continuity of CWC-related lineages (see below).

The technology present in Textile ceramics is in clear contrast to local traditions of sub-Neolithic Lovozero and Pasvik cultures of asbestos-tempered pottery to the north and east, which point to a different tradition of knowledge and learning network – showing partial continuity with previous asbestos ware, since these territories host the main sources of asbestos. We have to assume that these cultures of northern and eastern Fennoscandia represent Palaeo-European (eventually also Palaeo-Siberian) groups clearly differentiated from the south.

The Chirkovo culture (ca. 1800-700 BC) forms on the middle Volga – at roughly the same time as Netted Ware formed to the west – from the fusion of Abashevo and Balanovo elites on Volosovo territory, and is also related (like Abashevo) to materials of the Seima-Turbino phenomenon.

Bronze Age ethnolinguistic groups

In the Gulf of Finland, Kiukainen evolves into the Paimio ceramics (in Finland) — Asva Ware (in Estonia) culture, which lasts from ca. 1600 to ca. 700 BC, probably representing an evolving Finno-Saamic community, while the Netted Ware from inner Finland (the Sarsa and Tomitsa groups) and the groups from the Forest Zone possibly represent a Volga-Finnic community.

NOTE. Nevertheless, the boundaries between Textile ceramic groups are far from clear, and inner Finland Netted Ware groups seem to follow a history different from Netted Ware groups from the Middle and Upper Volga, hence they could possibly be identified as an evolving Pre-Saamic community.

Based on language contacts, with Early Baltic – Early Finnic contacts starting during the Iron Age (ca. 500 BC onwards), this is a potential picture of the situation at the end of this period, when Germanic influence on the coast starts to fade, and Lusatian culture influence is stronger:

aikio-finnic-saamic
The linguistic situation in Lapland and the northern Baltic Sea Area in the Early Iron Age prior to the expansion of Saami languages; the locations of the language groups are schematic. The black line indicates the distribution of Saami languages in the 19th century, and the gray line their approximate maximal distribution before the expansion of Finnic. Aikio (2012)

The whole Finno-Permic community remains thus in close contact, allowing for the complicated picture that Kallio mentions as potentially showing Dahl’s wave model for Uralic languages.

Genetic data shows a uniform picture of these communities, with exclusively CWC-derived ancestry and haplogroups. So in Mittnik et al. (2018) all Baltic samples show R1a-Z645 subclades, while the recent session on Estonian populations in ISBA 8 (see programme in PDF) clearly states that:

[Of the 24 Bronze Age samples from stone-cist graves] all 18 Bronze Age males belong to R1a.

Regarding non-Uralic substrates found in Saami, supposedly absorbed during the expansion to the north (and thus representing languages spoken in northern Fennoscandia during the Bronze Age) this is what Aikio (2012) has to say:

The Saami substrate in the Finnish dialects thus reveals that also Lakeland Saami languages had a large number of vocabulary items of obscure origin. Most likely many of these words were substrate in Lakeland Saami, too, and ultimately derive from languages spoken in the region before Saami. In some cases the loan origin of these words is obvious due to their secondary Proto-Saami vowel combinations such as *ā–ë in *kāvë ‘bend; small bay’ and *šāpšë ‘whitefish’. This substrate can be called ‘Palaeo-Lakelandic’, in contrast to the ‘Palaeo-Laplandic’ substrate that is prominent in the lexicon of Lapland Saami. As the Lakeland Saami languages became extinct and only fragments of their lexicon can be reconstructed via elements preserved in Finnish place-names and dialectal vocabulary, we are not in a position to actually study the features of this Palaeo-Lakelandic substrate. Its existence, however, appears evident from the material above.

If we wanted to speculate further, based on the data we have now, it is very likely that two opposing groups will be found in the region:

A) The central Finnish group, in this hypothesis the Palaeo-Lakelandic group, made up of the descendants of the Mesolithic pioneers of the Komsa and Suomusjärvi cultures, and thus mainly Baltic HG / Scandinavian HG ancestry and haplogroups I / R1b(xM269) (see more on Scandinavian HG).

siberian-ancestry-map
Frequency map of the so-called ‘Siberian’ component. From Tambets et al. (2018).

B) Lapland and Kola were probably also inhabited by similar Mesolithic populations, until it was eventually assimilated by expanding Siberian groups (of Siberian ancestry and N1c-L392 lineages) from the east – entering the region likely through the Kola peninsula – , forming the Palaeo-Laplandic group, which was in turn later replaced by expanding Proto-Saamic groups.

Siberian ancestry appears first in Fennoscandia at Bolshoy Oleni Ostrov ca. 1520 BC, with haplogroup N1c-L392 (2 samples, BOO002 and BOO004), and with Siberian ancestry. This is their likely movement in north-eastern Europe, from Lamnidis et al (2018):

The large Siberian component in the Bolshoy individuals from the Kola Peninsula provides the earliest direct genetic evidence for an eastern migration into this region. Such contact is well documented in archaeology, with the introduction of asbestos-mixed Lovozero ceramics during the second millenium BC, and the spread of even-based arrowheads in Lapland from 1,900 BCE. Additionally, the nearest counterparts of Vardøy ceramics, appearing in the area around 1,600-1,300 BCE, can be found on the Taymyr peninsula, much further to the east. Finally, the Imiyakhtakhskaya culture from Yakutia spread to the Kola Peninsula during the same period.

saamic-lovozero-pca
PCA plot of 113 Modern Eurasian populations, with individuals from this study projected on the principal components. Uralic speakers are highlighted in light purple. Image modified from Lamnidis et al. (2018)

Obviously, these groups of asbestos-tempered ware are not connected to the Uralic expansion. From the same paper:

The fact that the Siberian genetic component is consistently shared among Uralic-speaking populations, with the exceptions of Hungarians and the non-Uralic speaking Russians, would make it tempting to equate this component with the spread of Uralic languages in the area. However, such a model may be overly simplistic. First, the presence of the Siberian component on the Kola Peninsula at ca. 4000 yBP predates most linguistic estimates of the spread of Uralic languages to the area. Second, as shown in our analyses, the admixture patterns found in historic and modern Uralic speakers are complex and in fact inconsistent with a single admixture event. Therefore, even if the Siberian genetic component partly spread alongside Uralic languages, it likely presented only an addition to populations carrying this component from earlier.

2. The Early Iron Age

The Ananino culture appears in the Vyatka-Kama area, famed for its metallurgy, with traditions similar to the North Pontic area, by this time developing Pre-Sauromatian traditions. It expanded to the north in the first half of the first millennium BC, remaining in contact with the steppes, as shown by the ‘Scythian’ nature of its material culture.

NOTE. The Ananino culture can be later followed through its zoomorphic styles into Iron Age Pjanoborskoi and Gljadenovskoi cultures, later to Ural-Siberian Middle Age cultures – Itkuska, Ust’-Poluiska, Kulaiska cultures –, which in turn can be related as prototypes of medieval Permian styles.

ananino-culture-homeland
Territory of (early and maximum) Ananino material culture. Vasilyev (2002).

At the same time as the Ananino culture begins to expand ca. 1000 BC, the Netted Ware tradition from the middle Oka expanded eastwards into the Oka-Vyatka interfluve of the middle Volga region, until then occupied by the Chirkovo culture. Eventually the Akozino or Akhmylovo group (ca. 800-300 BC) emerged from the area, showing a strong cultural influence from the Ananino culture, by that time already expanding into the Cis-Urals region.

The Akozino culture remains nevertheless linked to the western Forest Zone traditions, with long-ranging influences from as far as the Lusatian culture in Poland (in metallurgical techniques), which at this point is also closely related with cultures from Scandinavia (read more on genetics of the Tollense Valley).

malar-celts-ananino
Mälar celts and molds for casting (a) and the main distribution area (в) of Mälar-type celts of the Mälar type in the Volga-Kama region (according to Kuzminykh 1983: figure 92) and Scandinavia (according to Baudou 1960: Karte 10); Ananino celts and molds for casting (б) and the main distribution area (г) of the distribution of the celts of the Ananino type in the Volga-Kama area (according to Kuzminykh 1983: figure 9); dagger of Ananino type (д).Map from (Yushkova 2010)

Different materials from Akozino reach Fennoscandia late, at the end of the Bronze Age and beginning of the Early Iron Age, precisely when the influence of the Nordic Bronze Age culture on the Gulf of Finland was declining.

This is a period when Textile ceramic cultures in north-eastern Europe evolve into well-armed chiefdom-based groups, with each chiefdom including thousands or tens of thousands, with the main settlements being hill forts, and those in Fennoscandia starting ca. 1000-400 BC.

Mälar-type celts and Ananino-type celts appear simultaneously in Fennoscandia and the Forest Zone, with higher concentrations in south-eastern Sweden (Mälaren) and the Volga-Kama region, supporting the existence of a revived international trade network.

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

The Paimio—Asva Ware culture evolves (ca. 700-200 BC) into the Morby (in Finland) — Ilmandu syle (in Estonia, Latvia, and Mälaren) culture. The old Paimio—Asva tradition continues side by side with the new one, showing a clear technical continuity with it, but with ornamentation compared to the Early Iron Age cultures of the Upper Volga area. This new south-eastern influence is seen especially in:

  • Akozino-Mälar axes (ca. 800-500 BC): introduced into the Baltic area in so great numbers – especially south-western Finland, the Åland islands, and the Mälaren area of eastern Sweden – that it is believed to be accompanied by a movement of warrior-traders of the Akozino-Akhmylovo culture, following the waterways that Vikings used more than a thousand years later. Rather than imports, they represent a copy made with local iron sources.
  • Tarand graves (ca. 500 BC – AD 400): these ‘mortuary houses’ appear in the coastal areas of northern and western Estonia and the islands, at the same time as similar graves in south-western Finland, eastern Sweden, northern Latvia and Courland. Similar burials are found in Akozino-Akhmylovo, with grave goods also from the upper and middle Volga region, while grave goods show continuity with Textile ware.

The use of asbestos increases in mainland Finnish wares with Kjelmøy Ware (ca. 700 BC – AD 300), which replaced the Lovozero Ware; and in the east in inner Finland and Karelia with the Luukonsaari and Sirnihta wares (ca. 700-500 BC – AD 200), where they replaced the previous Sarsa-Tomitsa ceramics.

The Gorodets culture appears during the Scythian period in the forest-steppe zone north and west of the Volga, shows fortified settlements, and there are documented incursions of Gorodets iron makers into the Samara valley, evidenced by deposits of their typical pottery and a bloom or iron in the region.

Iron Age ethnolinguistic groups

According to (Koryakova and Epimakhov 2007):

It is commonly accepted by archaeology, ethnography, and linguistics that the ancestors of the Permian peoples (the Udmurts, Komi-Permians, and Komi-Zyryans) left the sites of Ananyino cultural intercommunity.

NOTE. For more information on the Late Metal Ages and Early Medieval situation of Finno-Ugric languages, see e.g. South-eastern contact area of Finnic languages in the light of onomastics (Rahkonen 2013).

finno-saamic-mordvin
Yakhr-, -khra, yedr-, -dra and yer-/yar, -er(o), -or(o) names of lakes in Central and North Russia and the possible boundary of the proto-language words *jäkra/ä and *järka/ä. Rahkonen (2011)

Certain innovations shared between Proto-Fennic (identified with the Gulf of Finland) and Proto-Mordvinic (from the Gorodets culture) point to their close contact before the Proto-Fennic expansion, and thus to the identification of Gorodets as Proto-Mordvinic, hence Akozino as Volgaic (Parpola 2018):

  • the noun paradigms and the form and function of individual cases,
  • the geminate *mm (foreign to Proto-Uralic before the development of Fennic under Germanic influence) and other non-Uralic consonant clusters.
  • the change of numeral *luka ‘ten’ with *kümmen.
  • The presence of loanwords of non-Uralic origin, related to farming and trees, potentially Palaeo-European in nature (hence possibly from Siberian influence in north-eastern Europe).
ananino-textile-ware-cultures
Map of archaeological cultures in north-eastern Europe ca. 8th-3rd centuries BC. [The Mid-Volga Akozino group not depicted] Shaded area represents the Ananino cultural-historical society. Purple area show likely zones of predominant Siberian ancestry and N1c-L392 lineages. Blue areas likely zones of predominant CWC ancestry and R1a-Z645 lineages. Fading purple arrows represent likely stepped movements of haplogroup N1c-L392 for centuries (Siberian → Ananino → Akozino → Fennoscandia), found eventually in tarand graves. Blue arrows represent eventual expansions of Fennic and (partially displaced) Saamic. Modified image from Vasilyev (2002).

The introduction of a strongly hierarchical chiefdom system can quickly change the pre-existing social order and lead to a major genetic shift within generations, without a radical change in languages, as shown in Sintashta-Potapovka compared to the preceding Poltavka society (read more about Sintashta).

Fortified settlements in the region represented in part visiting warrior-traders settled through matrimonial relationships with local chiefs, eager to get access to coveted goods and become members of a distribution network that could guarantee them even military assistance. Such a system is also seen synchronously in other cultures of the region, like the Nordic Bronze Age and Lusatian cultures (Parpola 2013).

The most likely situation is that N1c subclades were incorporated from the Circum-Artic region during the Anonino (Permic) expansion to the north, later emerged during the formation of the Akozino group (Volgaic, under Anonino influence), and these subclades in turn infiltrated among the warrior traders that spread all over Fennoscandia and the eastern Baltic (mainly among Fennic, Saamic, Germanic, and Balto-Slavic peoples), during the age of hill forts, creating alliances partially based on exogamy strategies (Parpola 2013).

Over the course of these events, no language change is necessary in any of the cultures involved, since the centre of gravity is on the expanding culture incorporating new lineages:

  • first on the Middle Volga, when Ananino expands to the north, incorporatinig N1c lineages from the Circum-Artic region.
  • then with the expansion of the Akozino-Akhmylovo culture into Ananino territory, admixing with part of its population;
  • then on the Baltic region, when materials are imported from Akozino into Fennoscandia and the eastern Baltic (and vice versa), with local cultures being infiltrated by foreign (Akozino) warrior-traders and their materials;
  • and later with the different population movements that led eventually to a greater or lesser relevance of N1c in modern Finno-Permic populations.

To argue that this infiltration and later expansion of lineages changed the language in one culture in one of these events seems unlikely. To use this argument of “opposite movement of ethnic and language change” for different successive events, and only on selected regions and cultures (and not those where the greatest genetic and cultural impact is seen, like e.g. Sweden for Akozino materials) is illogical.

NOTE. Notice how I write here about “infiltration” and “lineages”, not “migration” or “populations”. To understand that, see below the next section on autosomal studies to compare Bronze Age, Iron Age, Medieval and Modern Estonians, and see how little the population of Estonia (homeland of Proto-Fennic and partially of Proto-Finno-Saamic) has changed since the Corded Ware migrations, suggesting genetic continuity and thus mostly close inter-regional and intra-regional contacts in the Forest Zone, hence a very limited impact of the absorbed N1c lineages (originally at some point incorporated from the Circum-Artic region). You can also check on the most recent assessment of R1a vs. N1c in modern Uralic populations.

Iron Age and later populations

From the session on Estonian samples on ISBA 8, by Tambets et al.:

[Of the 13 samples from the Iron Age tarand-graves] We found that the Iron Age individuals do in fact carry chrY hg N3 (…) Furthermore, based on their autosomal data, all of the studied individuals appear closer to hunter-gatherers and modern Estonians than Estonian CWC individuals do.

estonian-pca
PCA of Estonian samples from the Bronze Age, Iron Age and Medieval times. Tambets et al. (2018, upcoming).

Looking at the plot, the genetic inflow marking the change from the Bronze Age to the Iron Age looks like an obvious expansion of nearby peoples with CWC-related ancestry, i.e. likely from the south-east, near the Middle Volga, where influence of steppe peoples is greater (hence likely Akozino) into a Proto-Fennic population already admixed (since the arrival of Corded Ware groups) with Comb Ware-like populations.

All of these groups were probably R1a-Z645 (likely R1a-Z283) since the expansion of Corded Ware peoples, with an introduction of some N1c lineages precisely during this Iron Age period. This infiltration of N1c-L392 with Akozino is obviously not directly related to Siberian cultures, given what we know about the autosomal description of Estonian samples.

Rather, N1c-L392 lineages were likely part of the incoming (Volgaic) Akozino warrior-traders, who settled among developing chiefdoms based on hill fort settlements of cultures all over the Baltic area, and began to appear thus in some of the new tarand graves associated with the Iron Age in north-eastern Europe.f

A good way to look at this is to realize that no new cluster appears compared to the data we already have from Baltic LN and BA samples from Mittnik et al. (2018), so the Estonian BA and IA clusters must be located (in a proper PCA) in the cline from Pit-Comb Ware culture through Baltic BA to Corded Ware groups:

baltic-samples
PCA and ADMIXTURE analysis reflecting three time periods in Northern European prehistory. a Principal components analysis of 1012 present-day West Eurasians (grey points, modern Baltic populations in dark grey) with 294 projected published ancient and 38 ancient North European samples introduced in this study (marked with a red outline). Population labels of modern West Eurasians are given in Supplementary Fig. 7 and a zoomed-in version of the European Late Neolithic and Bronze Age samples is provided in Supplementary Fig. 8. b Ancestral components in ancient individuals estimated by ADMIXTURE (k = 11)

This genetic continuity from Corded Ware (the most likely Proto-Uralic homeland) to the Proto-Fennic and Proto-Saamic communities in the Gulf of Finland correlates very well with the known conservatism of Finno-Saamic phonology, quite similar to Finno-Ugric, and both to Proto-Uralic (Kallio 2017): The most isolated region after the expansion of Corded Ware peoples, the Gulf of Finland, shielded against migrations for almost 1,500 years, is then the most conservative – until the arrival of Akozino influence.

NOTE. This has its parallel in the phonetic conservatism of Celtic or Italic compared to Finno-Ugric-influenced Germanic, Balto-Slavic, or Indo-Iranian.

Only later would certain regions (like Finland or Lappland) suffer Y-DNA bottlenecks and further admixture events associated with population displacements and expansions, such as the spread of Fennic peoples from their Estonian homeland (evidenced by the earlier separation of South Estonian) to the north and east:

diversification-finnic
The Finnic family tree. Kallio (2014).

The initial Proto-Fennic expansion was probably coupled with the expansion of Proto-Saami to the north, with the Kjelmøy Ware absorbing the Siberian population of Lovozero Ware, and potentially in inner Finland and Karelia with the Luukonsaari and Sirnihta wares (Carpelan and Parpola 2017).

This Proto-Saami population expansion from the mainland to the north, admixing with Lovozero-related peoples, is clearly reflected in the late Iron Age Saamic samples from Levänluhta (ca. 400-800 AD), as a shift (of 2 out of 3 samples) to Siberian-like ancestry from their original CWC_Baltic-like situation (see PCA from Lamnidis et al. 2018 above).

Also, Volgaic and Permic populations from inner Finland and the Forest Zone to the Cis-Urals and Circum-Artic regions probably incorporate Siberian ancestry and N1c-L392 lineages during these and later population movements, while the westernmost populations – Estonian, Mordvinic – remain less admixed (see PCA from Tambets et al. 2018 below).

We also have data of N1c-L392 in Nordic territory in the Middle Ages, proving its likely strong presence in the Mälaren area since the Iron Age, with the arrival of Akozino warrior traders. Similarly, it is found among Balto-Slavic groups along the eastern Baltic area. Obviously, no language change is seen in Nordic Bronze Age and Lusatian territory, and none is expected in Estonian or Finnish territory, either.

Therefore, no “N1c-L392 + Siberian ancestry” can be seen expanding Finno-Ugric dialects, but rather different infiltrations and population movements with limited effects on ancestry and Y-DNA composition, depending on the specific period and region.

estonians-hungarians-mordvinian
Selection of the PCA, with the group of Estonians, Mordovians, and Hungarians selected. See Tambets et al. (2018) for more information.

An issue never resolved

Because N1c-L392 subclades & Siberian ancestry, which appear in different proportions and with different origins among some modern Uralic peoples, do not appear in cultures supposed to host Uralic-speaking populations until the Iron Age, people keep looking into any direction to find the ‘true’ homeland of those ‘Uralic N1c peoples’? Kind of a full circular reasoning, anyone? The same is valid for R1a & steppe ancestry being followed for ‘Indo-Europeans’, or R1b-P312 & Neolithic farmer ancestry being traced for ‘Basques’, because of their distribution in modern populations.

I understand the caution of many pointing to the need to wait and see how samples after 2000 BC are like, in every single period, from the middle and upper Volga, Kama, southern Finland, and the Forest Zone between Fennoscandia and the steppe. It’s like waiting to see how people from Western Yamna and the Carpathian Basin after 3000 BC look like, to fill in what is lacking between East Yamna and Bell Beakers, and then between them and every single Late PIE dialect.

But the answer for Yamna-Bell Beaker-Poltavka peoples during the Late PIE expansion is always going to be “R1b-L23, but with R1a-Z645 nearby” (we already have a pretty good idea about that); and the answer for the Forest Zone and northern Cis- and Trans-Urals area – during the time when Uralic languages are known to have already been spoken there – is always going to be “R1a-Z645, but with haplogroup N nearby”, as is already clear from the data on the eastern Baltic region.

So, without a previously proposed model as to where those amateurs expressing concern about ‘not having enough data’ expect to find those ‘Uralic peoples’, all this waiting for the right data looks more like a waiting for N1c and Siberian ancestry to pop up somewhere in the historic Uralic-speaking area, to be able to say “There! A Uralic-speaking male!”. Not a very reasonable framework to deal with prehistoric peoples and their languages, I should think.

But, for those who want to do that, let me break the news to you already:

ananino-culture-balto-slavic
First N1c – Finno-Ugric person arrives in Estonia to teach Finno-Saamic to Balto-Slavic peoples.

And here it is, an appropriate fantasy description of the ethnolinguistic groups from the region. You are welcome:

  • During the Bronze Age, late Corded Ware groups evolve as the western Textile ware Fennic Balto-Slavic group in the Gulf of Finland; the Netted Ware Saamic Balto-Slavic group of inner Finland; the south Netted Ware / Akozino Volgaic Balto-Slavic groups of the Middle Volga; and the Anonino Permic Balto-Slavic group in the north-eastern Forest Zone; all developing still in close contact with each other, allowing for common traits to permeate dialects.
  • These Balto-Slavic groups would then incorporate west of the Urals during and after the Iron Age (ca. 800-500 BC first, and also later during their expansion to the north) limited ancestry and lineages from eastern European hunter-gatherer groups of Palaeo-European Fennic and Palaeo-Siberian Volgaic and Permic languages from the Circum-Artic region, but they adopted nevertheless the language of the newcomers in every single infiltration of N1c lineages and/or admixture with Siberian ancestry. Oh and don’t forget the Saamic peoples from central Sweden, of course, the famous N1c-L392 ‘Rurikid’ lineages expanding Saamic to the north and replacing Proto-Germanic…

The current model for those obsessed with modern Y-DNA is, therefore, that expanding Neolithic, Bronze Age and Iron Age cultures from north-eastern Europe adopted the languages of certain lineages originally from sub-Neolithic (Scandinavian and Siberian) hunter-gatherer populations of the Circum-Artic region; lineages that these cultures incorporated unevenly during their expansions. Hmmmm… Sounds like an inverse Western movie, where expanding Americans end up speaking Apache, and the eastern coast speaks Spanish until Italian migrants arrive and make everyone speak English… or something. A logic, no-nonsense approach to ethnolinguistic identification.

I kid you not, this is the kind of models we are going to see very soon. In 2018 and 2019, with ancient DNA able to confirm or reject archaeological hypotheses based on linguistic data, people will keep instead creating new pet theories to support preconceived ideas based on the Y-DNA prevalent among modern populations. That is, information available in the 2000s.

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

[Next post on the subject: Corded Ware—Uralic (III): Seima-Turbino and the Ugric and Samoyedic expansion]

Related

Corded Ware—Uralic (I): Differences and similarities with Yamna

indo-european-uralic-migrations-corded-ware

I was reading The Bronze Age Landscape in the Russian Steppes: The Samara Valley Project (2016), and I was really surprised to find the following excerpt by David W. Anthony:

The Samara Valley links the central steppes with the western steppes and is a north-south ecotone between the pastoral steppes to the south and the forest-steppe zone to the north [see figure below]. The economic contrast between pastoral steppe subsistence, with its associated social organizations, and forest-zone hunting and fishing economies probably explains the shifting but persistent linguistic border between forest-zone Uralic languages to the north (today largely displaced by Russian) and a sequence of steppe languages to the south, recently Turkic, before that Iranian, and before that probably an eastern dialect of Proto-Indo-European (Anthony 2007). The Samara Valley represents several kinds of borders, linguistic, cultural, and ecological, and it is centrally located in the Eurasian steppes, making it a critical place to examine the development of Eurasian steppe pastoralism.

uralic-languages-forest-zone-volga
Language map of the middle Volga-Ural region. After “Geographical Distribution of the Uralic Languages” by Finno-Ugrian Society, Helsinki, 1993.

Khokhlov (translated by Anthony) further insists on the racial and ethnic divide between both populations, Abashevo to the north, and Poltavka to the south, during the formation of the Abashevo – Sintashta-Potapovka community that gave rise to Proto-Indo-Iranians:

Among all cranial series in the Volga-Ural region, the Potapovka population represents the clearest example of race mixing and probably ethnic mixing as well. The cultural advancements seen in this period might perhaps have been the result of the mixing of heterogeneous groups. Such a craniometric observation is to some extent consistent with the view of some archaeologists that the Sintashta monuments represent a combination of various cultures (principally Abashevo and Poltavka, but with other influences) and therefore do not correspond to the basic concept of an archaeological culture (Kuzmina 2003:76). Under this option, the Potapovka-Sintashta burial rite may be considered, first, a combination of traits to guarantee the afterlife of a selected part of a heterogeneous population. Second, it reflected a kind of social “caste” rather than a single population. In our view, the decisive element in shaping the ethnic structure of the Potapovka-Sintashta monuments was their extensive mobility over a fairly large geographic area. They obtained knowledge of various cultures from the populations with whom they interacted.

steppe-lmba-sintashta-potapovka-filatovka
Late Middle Bronze Age cultures with the Proto-Indo-Iranian Sintashta-Potapovka-Filatovka group (shaded). After Anthony (2007 Figure 15.5), from Anthony (2016).

Interesting is also this excerpt about the predominant population in the Abashevo – Sintashta-Potapovka admixture (which supports what Chetan said recently, although this does not seemed backed by Y-DNA haplogroups found in the richest burials), coupled with the sign of incoming “Uraloid” peoples from the east, found in both Sintashta and eastern Abashevo:

The socially dominant anthropological component was Europeoid, possibly the descendants of Yamnaya. The association of craniofacial types with archaeological cultures in this period is difficult, primarily because of the small amount of published anthropological material of the cultures of steppe and forest belt (Balanbash, Vol’sko-Lbishche) and the eastern and southern steppes (Botai-Tersek). The crania associated with late MBA western Abashevo groups in the Don-Volga forest zone were different from eastern Abashevo in the Urals, where the expression of the Old Uraloid craniological complex was increased. Old Uraloid is found also on a single skull of Vol’sko-Lbishche culture (Tamar Utkul VII, Kurgan 4). Potentially related variants, including Mongoloid features, could be found among the Seima-Turbino tribes of the forest-steppe zone, who mixed with Sintashta and Abashevo. In the Sintashta Bulanova cemetery from the western Urals, some individuals were buried with implements of Seima-Turbino type (Khalyapin 2001; Khokhlov 2009; Khokhlov and Kitov 2009). Previously, similarities were noted between some individual skulls from Potapovka I and burials of the much older Botai culture in northern Kazakhstan (Khokhlov 2000a). Botai-Tersek is, in fact, a growing contender for the source of some “eastern” cranial features.

khvalynsk-yamna-srubna-facial-reconstruction
Facial reconstructions based on skulls from (a) Khvalynsk II Grave 24, a young adult male; (b) Poludin Grave 6, Yamnaya culture, a mature male (both by A. I. Nechvaloda); and (c) Luzanovsky cemetery, Srubnaya culture (by L. T. Yablonsky). In Khokhlov (2016).

The wave of peoples associated with “eastern” features can be seen in genetics in the Sintashta outliers from Narasimhan et al. (2018), and it probably will be eventually seen in Abashevo, too. These may be related to the Seima-Turbino international network – but most likely it is directly connected to Sintashta through the starting Andronovo and Seima-Turbino horizons, by admixing of prospective groups and small-scale back-migrations.

Corded Ware – Yamna similarities?

So, if peoples of north-eastern Europe have been assumed for a long time to be Uralic speakers, what is happening with the Corded Ware = IE obsession? Is it Gimbutas’ ghost possessing old archaeologists? Probably not.

It is about certain cultural similarities evident at first sight, which have been traditionally interpreted as a sign of cultural diffusion or migration. Not dissimilar to the many Bell Beaker models available, where each archaeologist is pushing certain differences, mixing what seemed reasonable, what still might seem reasonable, and what certainly isn’t anymore after the latest ancient DNA data.

kurgan-expansion
“European dialect” expansion of Proto-Indo-European according to Gimbutas (1963)

The initial models of Gimbutas, Kristiansen, or Anthony – which are known to many today – were enunciated in the infancy of archaeological studies in the regions, during and just after the fall of the USSR, and before many radiocarbon dates that we have today were published (with radiocarbon dating being still today in need of refinement), so it is only logical that gross mistakes were made.

We have similar gross mistakes related to the origins of Bell Beakers, and studying them was certainly easier than studying eastern data.

  • Gimbutas believed – based mainly on Kurgan-like burials – that Bell Beaker formed from a combination of Yamna settlers with the Vučedol culture, so she was not that far from the truth.
  • The expansion of Corded Ware from peoples of the North Pontic forest-steppe area, proposed by Gimbutas and later supported also by Kristiansen (1989) as the main Indo-European expansion – , is probably also right about the approximate origins of the culture. Only its ‘Indo-European’ nature is in question, given the differences with Khvalynsk and Yamna evolution.
  • Anthony only claimed that Yamna migrants settled in the Balkans and along the Danube into the Hungarian steppes. He never said that Corded Ware was a Yamna offshoot until after the first genetic papers of 2015 (read about his newest proposal). He initially claimed that only certain neighbouring Corded Ware groups “adopted” Indo-European (through cultural diffusion) because of ‘patron-client’ relationships, and was never preoccupied with the fate of Corded Ware and related cultures in the east European forest zone and Finland.

So none of them was really that far from the true picture; we might say a lot people are more way off the real picture today than the picture these three researchers helped create in the 1990s and 2000s. Genetics is just putting the last nail in the coffin of Corded Ware as a Yamna offshoot, instead of – as we believed in the 2000s – to Vučedol and Bell Beaker.

So let’s revise some of these traditional links between Corded Ware and Yamna with today’s data:

Archaeology

Even more than genetics – at least until we have an adequate regional and temporary sampling – , archaeological findings lead what we have to know about both cultures.

It is essential to remember that Corded Ware, starting ca. 3000/2900 BC in east-central Europe, has been proposed to be derived from Early Yamna, which appeared suddenly in the Pontic-Caspian steppes ca. 3300 BC (probably from the late Repin expansion), and expanded to the west ca. 3000.

Early Yamna is in turn identified as the expanding Late Proto-Indo-European community, which has been confirmed with the recent data on Afanasevo, Bell Beaker, and Sintashta-Potapovka and derived cultures.

The question at hand, therefore, is if Corded Ware can be considered an offshoot of the Late PIE community, and thus whether the CWC ethnolinguistic community – proven in genetics to be quite homogeneous – spoke a Late PIE dialect, or if – alternatively – it is derived from other neighbouring cultures of the North Pontic region.

NOTE. The interpretation of an Indo-Slavonic group represented by a previous branching off of the group is untenable with today’s data, since Indo-Slavonic – for those who support it – would itself be a branch of Graeco-Aryan, and Palaeo-Balkan languages expanded most likely with West Yamna (i.e. R1b-L23, mainly R1b-Z2103) to the south.

The convoluted alternative explanation would be that Corded Ware represents an earlier, Middle PIE branch (somehow carrying R1a??) which influences expanding Late PIE dialects; this has been recently supported by Kortlandt, although this simplistic picture also fails to explain the Uralic problem.

Kurgans: The Yamna tradition was inherited from late Repin, in turn inherited from Khvalynsk-Novodanilovka proto-Kurgans. As for the CWC tradition, it is unclear if the tumuli were built as a tradition inherited from North and West Pontic cultures (in turn inherited or copied from Khvalynsk-Novodanilovka), such as late Trypillia, late Kvityana, late Dereivka, late Sredni Stog; or if they were built because of the spread of the ‘Transformation of Europe’, set in motion by the Early Yamna expansion ca. 3300-3000 BC (as found in east-central European cultures like Coţofeni, Lizevile, Șoimuș, or the Adriatic Vučedol). My guess is that it inherits an older tradition than Yamna, with an origin in east-central Europe, because of the mound-building distribution in the North Pontic area before the Yamna expansion, but we may never really know.

pit-graves-central-europe-cwc
Distribution of Pit-Grave burials west of the Black Sea likely dating to the 2nd half of the IVth millennium BC (triangles: side-crouched burials; filled circles: supine extended burials; open circles: suspected). Frînculeasa, Preda, and Heyd (2015)

Burial rite: Yamna features (with regional differences) single burials with body on its back, flexed upright knees, poor grave goods, common orientation east-west (heads to the west) inherited from Repin, in turn inherited from Khvalynsk-Novodanilovka. CWC tradition – partially connected to Złota and surrounding east-central European territories (in turn from the Khvalynsk-Novodanilovka expansion) – features single graves, body in fetal position, strict gender differentiation – men on the right, women on the left -, looking to the south, graves with standardized assemblages (objects representing affirmation of battle, hunting, and feasting). The burial rites clearly represent different ideologies.

pit-grave-burial-schemes
Left: Pit-Grave burial types expanded with Khvalynsk-Novodanilovka. Right: Pit-Grave burial types associated with the Yamna expansion and influence. Frînculeasa, Preda, and Heyd (2015)

Corded decoration: Corded ware decoration appears in the Balkans during the 5th millennium, and represents a simple technique whereby a cord is twisted, or wrapped around a stick, and then pressed directly onto the fresh surface of a vessel leaving a characteristic decoration. It appears in many groups of the 5th and 4th millennium BC, but it was Globular Amphorae the culture which popularized the drinking vessels and their corded ornamentation. It appears thus in some regional groups of Yamna, but it becomes the standard pottery only in Corded Ware (especially with the A-horizon), which shows continuity with GAC pottery.

corded-ware-first-horizon
Origins of the first Corded Ware horizon (5th millennium BC) after the Khvalynsk-Novodanilovka expansion. Corded Ware (circles) and horse-head scepters (rectangles) and other steppe elements (triangles). Image from Bulatović (2014).

Economy: Yamna expands from Repin (and Repin from Khvalynsk-Novodanilovka) as a nomadic or semi-nomadic purely pastoralist society (with occasional gathering of wild seeds), which naturally thrives in the grasslands of the Pontic-Caspian, lower Danube and Hungarian steppes. Corded Ware shows agropastoralism (as late Eneolithic forest-steppe and steppe groups of eastern Europe, such as late Trypillian, TRB, and GAC groups), inhabits territories north of the loess line, with heavy reliance of hunter-gathering depending on the specific region.

Cattle herding: Interestingly, both west Yamna and Corded Ware show more reliance on cattle herding than other pastoralist groups, which – contrasted with the previous Eneolithic herding traditions of the Pontic-Caspian steppe, where sheep-goats predominate – make them look alike. However, the cattle-herding economy of Yamna is essential for its development from late Repin and its expansion through the steppes (over western territories practising more hunter-gathering and sheep-goat herding economy), and it does not reach equally the Volga-Ural region, whose groups keep some of the old subsistence economy (read more about the late Repin expansion). Corded Ware, on the other hand, inherits its economic strategy from east European groups like TRB, GAC, and especially late Trypillian communities, showing a predominance of cattle herding within an agropastoral community in the forest-steppe and forest zones of Volhynia, Podolia, and surrounding forest-steppe and forest regions.

yamna-scheme
Scheme of interlinked socio-economic-ideological innovations forming the Yamnaya. Frînculeasa, Preda, and Heyd (2015)

Horse riding: Horse riding and horse transport is proven in Yamna (and succeeding Bell Beaker and Sintashta), assumed for late Repin (essential for cattle herding in the seas of grasslands that are the steppes, without nearby water sources), quite likely during the Khvalynsk expansion (read more here), and potentially also for Samara, where the predominant horse symbolism of early Khvalynsk starts. Corded Ware – like the north Pontic forest-steppe and forest areas during the Eneolithic – , on the other hand, does not show a strong reliance on horse riding. The high mobility and short-term settlements characteristic of Corded Ware, that are often associated with horse riding by association with Yamna, may or may not be correct, but there is no need for horses to explain their herding economy or their mobility, and the north-eastern European areas – the one which survived after Bell Beaker expansion – did certainly not rely on horses as an essential part of their economy.

NOTE: I cannot think of more supposed similarities right now. If you have more ideas, please share in the comments and I will add them here.

Genetic similarities

EHG: This is the clearest link between both communities. We thought it was related to the expansion of ANE-related ancestry to the west into WHG territory, but now it seems that it will be rather WHG expanding into ANE territory from the Pontic-Caspian region to the east (read more on recent Caucasus Neolithic, on , and on Caucasus HG).

NOTE. Given how much each paper changes what we know about the Palaeolithic, the origin and expansion of the (always developing) known ancestral components and specific subclades (see below) is not clear at all.

CHG: This is the key link between both cultures, which will delimit their interaction in terms of time and space. CHG is intermediate between EHG and Iran N (ca. 8000 BC). The ancestry is thus linked to the Caucasus south of the steppe before the emergence of North Pontic (western) and Don-Volga-Ural (eastern) communities during the Mesolithic. The real question is: when we have more samples from the steppe and the Caucasus during the Neolithic, how many CHG groups are we going to find? Will the new specific ancestral components (say CHG1, CHG2, CHG3, etc.) found in Yamna (from Khvalynsk, in the east) and Corded Ware (probably from the North Pontic forest-steppe) be the same? My guess is, most likely not, unless they are mediated by the Khvalynsk-Novodanilovka expansion (read more on CHG in the Caucasus).

yamnaya-chg-ancestry
Formation of Yamna and CHG contribution, in Damgaard et al. (Science 2018). A 10-leaf model based on combining the models in Fig. S16 and Fig. S19 and re-estimating the model parameters.

WHG/EEF: This is the obvious major difference – known today – in the formation of both communities in the steppe, and shows the different contacts that both groups had at least since the Eneolithic, i.e. since the expansion of Repin with its renewed Y-DNA bottleneck, and probably since before the early Khvalynsk expansion (read more on Yamna-Corded Ware differences contrasting with Yamna-Afanasevo, Yamna-Bell Beaker, and Yamna-Sintashta similarities).

NOTE 1. Some similarities between groups can be seen depending on the sampled region; e.g. Baltic groups show more similarities with southern Pontic-Caspian steppe populations, probably due to exogamy.

yamna-corded-ware-diff-qpgraph
Tested qpGraph model in Tambets et al. (2018). The qpGraph model fitting the data for the tested populations. “Colour codes for the terminal nodes: pink—modern populations (‘Population X’ refers to test population) and yellow—ancient populations (aDNA samples and their pools). Nodes coloured other than pink or yellow are hypothetical intermediate populations. We putatively named nodes which we used as admixture sources using the main recipient among known populations. The colours of intermediate nodes on the qpGraph model match those on the admixture proportions panel.”

NOTE 2. We have this information on the differences in “steppe ancestry” between Yamna and Corded Ware, compared to previous studies, because now we have more samples of neighbouring, roughly contemporaneous Eneolithic groups, to analyse the real admixture processes. This kind of fine scale studies is what is going to show more and more differences between Khvalynsk-Yamna and Sredni Stog-Corded Ware as more data pours in. The evolution of both communities in archaeology and in PCA (see below) is probably witness to those differences yet to be published.

R1: Even though some people try very hard to think in terms of “R1” vs. (Caucasus) J or G or any other upper clade, this is plainly wrong. It is possible, given what we know now, that Q1a2-M242 expanded ANE ancestry to the west ca. 13000 BC, while R1b-P279 expanded WHG ancestry to the east with the expansion of post-Swiderian cultures, creating EHG as a WHG:ANE cline. The role of R1a-M459 is unknown, but it might be related to any of these migrations, or others (plural) along northern Eurasia (read more on the expansion of R1b-P279, on Palaeolithic Q1a2, and on R1a-M417).

NOTE. I am inclined to believe in a speculative Mesolithic-Early Neolithic community involving Eurasiatic movements accross North Eurasia, and Indo-Uralic movements in its western part, with the last intense early Uralic-PIE contacts represented by the forming west (Mariupol culture) and east (Don-Volga-Ural cultures, including Samara) communities developing side by side. Before their known Eneolithic expansions, no large-scale Y-DNA bottleneck is going to be seen in the Pontic-Caspian steppe, with different (especially R1a and R1b subclades) mixed among them, as shown in North Pontic Neolithic, Samara HG, and Khvalynsk samples.

PCA-trypillia-greece-neolithic-outlier-anatolian
Image modified from Wang et al. (2018). Samples projected in PCA of 84 modern-day West Eurasian populations (open symbols). Previously known clusters have been marked and referenced. Marked and labelled are the Balkan samples referenced in this text An EHG and a Caucasus ‘clouds’ have been drawn, leaving Pontic-Caspian steppe and derived groups between them. See the original file here.

Corded Ware and ‘steppe ancestry’

If we take a look at the evolution of Corded Ware cultures, the expansion of Bell Beakers – dominated over most previous European cultures from west to east Europe – influenced the development of the whole European Bronze Age, up to Mierzanowice and Trzciniec in the east.

The only relevant unscathed CWC-derived groups, after the expansion of Sintashta-Potapovka as the Srubna-Andronovo horizon in the Eurasian steppes, were those of the north-eastern European forest zone: between Belarus to the west, Finland to the north, the Urals to the east, and the forest-steppe region to the south. That is, precisely the region supposed to represent Uralic speakers during the Bronze Age.

This inconsistency of steppe ancestry and its relation with Uralic (and Balto-Slavic) peoples was observed shortly after the publication of the first famous 2015 papers by Paul Heggarty, of the Max-Planck Institute for Evolutionary Anthropology (read more):

Haak et al. (2015) make much of the high Yamnaya ancestry scores for (only some!) Indo-European languages. What they do not mention is that those same results also include speakers of other languages among those with the highest of all scores for Yamnaya ancestry. Only these are languages of the Uralic family, not Indo-European at all; and their Yamnaya-ancestry signals are far higher than in many branches of Indo-European in (southern) Europe. Estonian ranks very high, while speakers of the very closely related Finnish are curiously not shown, and nor are the Saami. Hungarian is relevant less directly since this language arrived only c. 900 AD, but also high.

uralic-steppe-ancestry

These data imply that Uralic-speakers too would have been part of the Yamnaya > Corded Ware movement, which was thus not exclusively Indo-European in any case. And as well as the genetics, the geography, chronology and language contact evidence also all fit with a Yamnaya > Corded Ware movement including Uralic as well as Balto-Slavic.

Both papers fail to address properly the question of the Uralic languages. And this despite — or because? — the only Uralic speakers they report rank so high among modern populations with Yamnaya ancestry. Their linguistic ancestors also have a good claim to have been involved in the Corded Ware and Yamnaya cultures, and of course the other members of the Uralic family are scattered across European Russia up to the Urals.

NOTE. Although the author was trying to support the Anatolian hypothesis – proper of glottochronological studies often published from the Max Planck Institute – , the question remains equally valid: “if Proto-Indo-European expands with Corded Ware and steppe ancestry, what is happening with Uralic peoples?”

For my part, I claimed in my draft that ancestral components were not the only relevant data to take into account, and that Y-DNA haplogroups R1a and R1b (appearing separately in CWC and Yamna-Bell Beaker-Afanasevo), together with their calculated timeframes of formation – and therefore likely expansion – did not fit with the archaeological and linguistic description of the spread of Proto-Indo-European and its dialects.

In fact, it seemed that only one haplogroup (R1b-M269) was constantly and consistenly associated with the proposed routes of Late PIE dialectal expansions – like Anthony’s second (Afanasevo) and third (Lower Danube, Balkan) waves. What genetics shows fits seamlessly with Mallory’s association of the North-West Indo-European expansion with Bell Beakers (read here how archaeologists were right).

balanovksy-yamnaya-ancestry
Map of the much beloved steppe (or “Yamnaya”) ancestry in modern populations, by Balanovsky. Modified from Klejn (2017).

More precise inconsistencies were observed after the publication of Olalde et al. (2017) and Mathieson et al. (2017), by Volker Heyd in Kossinna’s smile (2017). Letting aside the many details enumerated (you can read a summary in my latest draft), this interesting excerpt is from the conclusion:

NOTE. An open access ealier draft version of the paper is offered for download by the author.

Simple solutions to complex problems are never the best choice, even when favoured by politicians and the media. Kossinna also offered a simple solution to a complex prehistoric problem, and failed therein. Prehistoric archaeology has been aware of this for a century, and has responded by becoming more differentiated and nuanced, working anthropologically, scientifically and across disciplines (cf. Müller 2013; Kristiansen 2014), and rejecting monocausal explanations. The two aDNA papers in Nature, powerful and promising as they are for our future understanding, also offer rather straightforward messages, heavily pulled by culture-history and the equation of people with culture. This admittedly is due partly to the restrictions of the medium that conveys them (and despite the often relevant additional detail given as supplementary information, which is unfortunately not always given full consideration).

While I have no doubt that both papers are essentially right, they do not reflect the complexity of the past. It is here that archaeology and archaeologists contributing to aDNA studies find their role; rather than simply handing over samples and advising on chronology, and instead of letting the geneticists determine the agenda and set the messages, we should teach them about complexity in past human actions and interactions. If accepted, this could be the beginning of a marriage made in heaven, with the blessing smile of Gustaf Kossinna, and no doubt Vere Gordon Childe, were they still alive, in a reconciliation of twentieth- and twenty-first-century approaches. For us as archaeologists, it could also be the starting point for the next level of a new archaeology.

heyd-yamnaya-expansion
Main distribution of Yamnaya kurgans in the Pontic-Caspian steppe of modern day Russia, Ukraine, and Kazakhstan, and its western branch in modern south-east European countries of Romania, Bulgaria, Serbia, and Hungary, with numbers of excavated kurgans and graves given. Picture: Volker Heyd (2018).

The question was made painfully clear with the publication of Olalde et al. (2018) & Mathieson et al. (2018), where the real route of Yamna expansion into Europe was now clearly set through the steppes into the Carpathian basin, later expanded as Bell Beakers.

This has been further confirmed in more recent papers, such as Narasimhan et al. (2018), Damgaard et al. (2018), or Wang et al. (2018), among others.

However, the discussion is still dominated by political agendas based on prevalent Y-DNA haplogroups in modern countries and ethnic groups.

Related

Haplogroup R1a and CWC ancestry predominate in Fennic, Ugric, and Samoyedic groups

uralic-languages

Open access Genes reveal traces of common recent demographic history for most of the Uralic-speaking populations, by Tambets et al. Genome Biology (2018).

Interesting excerpts (emphasis mine):

Methods

A total of 286 samples of Uralic-speaking individuals, of those 121 genotyped in this study, were analysed in the context of 1514 Eurasian samples (including 14 samples published for the first time) based on whole genome single nucleotide polymorphisms (SNPs) (Additional file 1: Table S1). All these samples, together with the larger sample set of Uralic speakers, were characterized for mtDNA and chrY markers.

The question as which material cultures may have co-spread together with proto-Uralic and Uralic languages depends on the time estimates of the splits in the Uralic language tree. Deeper age estimates (6,000 BP) of the Uralic language tree suggest a connection between the spread of FU languages from the Volga River basin towards the Baltic Sea either with the expansion of the Neolithic culture of Combed Ware, e.g. [6, 7, 17, 26] or with the Neolithic Volosovo culture [7]. Younger age estimates support a link between the westward dispersion of Proto-Finno-Saamic and eastward dispersion of Proto-Samoyedic with a BA Sejma-Turbino (ST) cultural complex [14, 18, 27, 28] that mediated the diffusion of specific metal tools and weapons from the Altai Mountains over the Urals to Northern Europe or with the Netted Ware culture [23], which succeeded Volosovo culture in the west. It has been suggested that Proto-Uralic may have even served as the lingua franca of the merchants involved in the ST phenomenon [18]. All these scenarios imply that material culture of the Baltic Sea area in Europe was influenced by cultures spreading westward from the periphery of Europe and/or Siberia. Whether these dispersals involved the spread of both languages and people remains so far largely unknown.

The population structure of Uralic speakers

To contextualize the autosomal genetic diversity of Uralic speakers among other Eurasian populations (Additional file 1: Table S1), we first ran the principal component (PC) analysis (Fig. 2a, Additional file 3: Figure S1). The first two PCs (Fig. 2a, Additional file 3: Figure S1A) sketch the geography of the Eurasian populations along the East-West and North-South axes, respectively. The Uralic speakers, along with other populations speaking Slavic and Turkic languages, are scattered along the first PC axis in agreement with their geographic distribution (Figs. 1 and 2a) suggesting that geography is the main predictor of genetic affinity among the groups in the given area. Secondly, in support of this, we find that FST-distances between populations (Additional file 3: Figure S2) decay in correlation with geographical distance (Pearson’s r = 0.77, p < 0.0001). On the UPGMA tree based on these FST-distances (Fig. 2b), the Uralic speakers cluster into several different groups close to their geographic neighbours.

uralic-pca
Principal component analysis (PCA) and genetic distances of Uralic-speaking populations. a PCA (PC1 vs PC2) of the Uralic-speaking populations.

We next used ADMIXTURE [48], which presents the individuals as composed of inferred genetic components in proportions that maximize Hardy-Weinberg and linkage equilibrium in the overall sample (see the ‘Methods’ section for choice of presented K). Overall, and specifically at lower values of K, the genetic makeup of Uralic speakers resembles that of their geographic neighbours. The Saami and (a subset of) the Mansi serve as exceptions to that pattern being more similar to geographically more distant populations (Fig. 3a, Additional file 3: S3). However, starting from K = 9, ADMIXTURE identifies a genetic component (k9, magenta in Fig. 3a, Additional file 3: S3), which is predominantly, although not exclusively, found in Uralic speakers. This component is also well visible on K = 10, which has the best cross-validation index among all tests (Additional file 3: S3B). The spatial distribution of this component (Fig. 3b) shows a frequency peak among Ob-Ugric and Samoyed speakers as well as among neighbouring Kets (Fig. 3a). The proportion of k9 decreases rapidly from West Siberia towards east, south and west, constituting on average 40% of the genetic ancestry of FU speakers in Volga-Ural region (VUR) and 20% in their Turkic-speaking neighbours (Bashkirs, Tatars, Chuvashes; Fig. 3a). The proportion of this component among the Saami in Northern Scandinavia is again similar to that of the VUR FU speakers, which is exceptional in the geographic context. It is also notable that North Russians, sampled from near the White Sea, differ from other Russians by sporting higher proportions of k9 (10–15%), which is similar to the values we observe in their Finnic-speaking neighbours. Notably, Estonians and Hungarians, who are geographically the westernmost Uralic speakers, virtually lack the k9 cluster membership.

siberian-ancestry
Population structure of Uralic-speaking populations inferred from ADMIXTURE analysis on autosomal SNPs in Eurasian context. a Individual ancestry estimates for populations of interest for selected number of assumed ancestral populations (K3, K6, K9, K11). Ancestry components discussed in a main text (k2, k3, k5, k6, k9, k11) are indicated and have the same colours throughout. The names of the Uralic-speaking populations are indicated with blue (Finno-Ugric) or orange (Samoyedic). The full bar plot is presented in Additional file 3: Figure S3. b Frequency map of component k9

We also tested the different demographic histories of female and male lineages by comparing outgroup f3 results for autosomal and X chromosome (chrX) data for pairs of populations (Estonians, Udmurts or Khanty vs others) with high versus low probability to share their patrilineal ancestry in chrY hg N (see the ‘Methods’ section, Additional file 3: Figure S13). We found a minor but significant excess of autosomal affinity relative to chrX for pairs of populations that showed a higher than 10% chance of two randomly sampled males across the two groups sharing their chrY ancestry in hg N3-M178, compared to pairs of populations where such probability is lower than 5% (Additional file 3: Figure S13).

In sum, these results suggest that most of the Uralic speakers may indeed share some level of genetic continuity via k9, which, however, also extends to the geographically close Turkic speakers.

uralic-modern-europe

Identity-by-descent

We found that it is the admixture with the Siberians that makes the Western Uralic speakers different from the tested European populations (Additional file 3: Figure S4A-F, H, J, L). Differentiating between Estonians and Finns, the Siberians share more derived alleles with Finns, while the geographic neighbours of Estonians (and Finns) share more alleles with Estonians (Additional file 3: Figure S4M). Importantly, Estonians do not share more derived alleles with other Finnic, Saami, VUR FU or Ob-Ugric-speaking populations than Latvians (Additional file 3: Figure S4O). The difference between Estonians and Latvians is instead manifested through significantly higher levels of shared drift between Estonians and Siberians on the one hand and Latvians and their immediate geographic neighbours on the other hand. None of the Uralic speakers, including linguistically close Khanty and Mansi, show significantly closer affinities to the Hungarians than any non-FU population from NE Europe (Additional file 3: Figure S4R).

ibd-uralic-genetics
Share of ~ 1–2 cM identity-by-descent (IBD) segments within and between regional groups of Uralic speakers. For each Uralic-speaking population representing lines in this matrix, we performed permutation test to estimate if it shows higher IBD segment sharing with other population (listed in columns) as compared to their geographic control group. Empty rectangles indicate no excess IBD sharing, rectangles filled in blue indicate comparisons when statistically significant excess IBD sharing was detected between one Uralic-speaking population with another Uralic-speaking population (listed in columns), rectangles filled in green mark the comparisons when a Uralic-speaking population shows excess IBD sharing with a non-Uralic-speaking population. For each tested Uralic speaker (matrix rows) populations in the control group that were used to generate permuted samples are indicated using small circles. For example, the rectangle filled in blue for Vepsians and Komis (A) implies that the Uralic-speaking Vepsians share more IBD segments with the Uralic-speaking Komis than the geographic control group for Vepsians, i.e. populations indicated with small circles (Central and North Russians, Swedes, Latvians and Lithuanians). The rectangle filled in green for Vepsians and Dolgans shows that the Uralic-speaking Vepsians share more IBD segments with the non-Uralic-speaking Dolgans than the geographic control group

Time of Siberian admixture

The time depth of the Globetrotter (Fig. 5b) inferred admixture events is relatively recent—500–1900 AD (see also complementary ALDER results, in Additional file 13: Table S12 and Additional file 3: Figure S7)—and agrees broadly with the results reported in Busby et al. [55]. A more detailed examination of the ALDER dates, however, reveals an interesting pattern. The admixture events detected in the Baltic Sea region and VUR Uralic speakers are the oldest (800–900 AD or older) followed by those in VUR Turkic speakers (∼1200–1300 AD), while the admixture dates for most of the Siberian populations (>1500 AD) are the most recent (Additional file 3: Figure S7). The West Eurasian influx into West Siberia seen in modern genomes was thus very recent, while the East Eurasian influx into NE Europe seems to have taken place within the first millennium AD (Fig. 5b, Additional file 3: Figure S7).

Affinities of the Uralic speakers with ancient Eurasians

We next calculated outgroup f3-statistics [48] to estimate the extent of shared genetic drift between modern and ancient Eurasians (Additional file 14: Table S13, Additional file 3: Figures S8-S9). Consistent with previous reports [45, 50], we find that the NE European populations including the Uralic speakers share more drift with any European Mesolithic hunter-gatherer group than Central or Western Europeans (Additional file 3: Figure S9A-C). Contrasting the genetic contribution of western hunter-gatherers (WHG) and eastern hunter-gatherers (EHG), we find that VUR Uralic speakers and the Saami share more drift with EHG. Conversely, WHG shares more drift with the Finnic and West European populations (Additional file 3: Figure S9A). Interestingly, we see a similar pattern of excess of shared drift between VUR and EHG if we substitute WHG with the aDNA sample from the Yamnaya culture (Additional file 3: Figure S9D). As reported before [2, 45], the genetic contribution of European early farmers decreases along an axis from Southern Europe towards the Ural Mountains (Fig. 6, Additional file 3: Figure S9E-F).

yamna-cwc-qpgraph-admixture-uralic
Proportions of ancestral components in studied European and Siberian populations and the tested qpGraph model. a The qpGraph model fitting the data for the tested populations. Colour codes for the terminal nodes: pink—modern populations (‘Population X’ refers to test population) and yellow—ancient populations (aDNA samples and their pools). Nodes coloured other than pink or yellow are hypothetical intermediate populations. We putatively named nodes which we used as admixture sources using the main recipient among known populations. The colours of intermediate nodes on the qpGraph model match those on the admixture proportions panel. b Admixture proportions (%) of ancestral components. We calculated the admixture proportions summing up the relative shares of a set of intermediate populations to explain the full spectrum of admixture components in the test population. We further did the same for the intermediate node CWC’ and present the proportions of the mixing three components in the stacked column bar of CWC’. Colour codes for ancestral components are as follows: dark green—Western hunter gatherer (WHG’); light green—Eastern hunter gatherer (EHG’); grey—European early farmer (LBK’); dark blue—carriers of Corded Ware culture (CWC’); and dark grey—Siberian. CWC’ consists of three sub-components: blue—Caucasian hunter-gatherer in Yamnaya (CHGinY’); light blue—Eastern hunter-gatherer in Yamnaya (EHGinY’); and light grey—Neolithic Levant (NeolL’)

We then used the qpGraph software [48] to test alternative demographic scenarios by trying to fit the genetic diversity observed in a range of the extant Finno-Ugric populations through a model involving the four basic European ancestral components: WHG, EHG, early farmers (LBK), steppe people of Yamnaya/Corded Ware culture (CWC) and a Siberian component (Fig. 6, Additional file 3: Figure S10). We chose the modern Nganasans to serve as a proxy for the latter component because we see least evidence for Western Eurasian admixture (Additional file 3: Figure S3) among them. We also tested the Khantys for that proxy but the model did not fit (yielding f2-statistics, Z-score > 3). The only Uralic-speaking population that did not fit into the tested model with five ancestral components were Hungarians. The qpGraph estimates of the contributions from the Siberian component show that it is the main ancestry component in the West Siberian Uralic speakers and constitutes up to one third of the genomes of modern VUR and the Saami (Fig. 6). It drops, however, to less than 10% in most of NE Europe, to 5% in Estonians and close to zero in Latvians and Lithuanians.

Discussion

uralic-groups-haplogroup-r1a
Additional file 6: Table S5. Y chromosome haplogroup frequencies in Eurasia. Modified by me: in bold haplogroup N1c and R1a from Uralic-speaking populations, with those in red showing where R1a is the major haplogroup. Observe that all Uralic subgroups – Finno-Permic, Ugric, and Samoyedic – have some populations with a majority of R1a lineages.

One of the notable observations that stands out in the fineSTRUCTURE analysis is that neither Hungarians nor Estonians or Mordovians form genetic clusters with other Uralic speakers but instead do so with a broad spectrum of geographically adjacent samples. Despite the documented history of the migration of Magyars [63] and their linguistic affinity to Khantys and Mansis, who today live east of the Ural Mountains, there is nothing in the present-day gene pool of the sampled Hungarians that we could tie specifically to other Uralic speakers.

Perhaps even more surprisingly, we found that Estonians, who show close affinities in IBD analysis to neighbouring Finnic speakers and Saami, do not share an excess of IBD segments with the VUR or Siberian Uralic speakers. This is eIn this context, it is important to remind that the limited (5%, Fig. 6) East Eurasian impact in the autosomal gene pool of modern Estonians contrasts with the fact that more than 30% of Estonian (but not Hungarian) men carry chrY N3 that has an East Eurasian origin and is very frequent among NE European Uralic speakers [36]. However, the spread of chrY hg N3 is not language group specific as it shows similar frequencies in Baltic-speaking Latvians and Lithuanians, and in North Russians, who in all our analyses are very similar to Finnic-speakers. The latter, however, are believed to have either significantly admixed with their Uralic-speaking neighbours or have undergone a language shift from Uralic to Indo-European [38].ven more striking considering that the immediate neighbours—Finns, Vepsians and Karelians—do.

With some exceptions such as Estonians, Hungarians and Mordovians, both IBD sharing and Globetrotter results suggest that there are detectable inter-regional haplotype sharing ties between Uralic speakers from West Siberia and VUR, and between NE European Uralic speakers and VUR. In other words, there is a fragmented pattern of haplotype sharing between populations but no unifying signal of sharing that unite all the studied Uralic speakers.

Comments

The paper is obviously trying to find a “N1c/Siberian ancestry = Uralic” link, but it shows (as previous papers using ancient DNA) that this identification is impossible, because it is not possible to identify “N1c=Siberian ancestry”, “N1c=Uralic”, or “Siberian ancestry = Uralic”. In fact, the arrival of N subclades and Siberian ancestry are late, both events (probably multiple stepped events) are unrelated to each other, and represent east-west demic diffusion waves (as well as founder effects) that probably coincide in part with the Scythian and Turkic (or associated) expansions, i.e. too late for any model of Proto-Uralic or Proto-Finno-Ugric expansion.

On the other hand, it shows interesting data regarding ancestry of populations that show increased Siberian influence, such as those easternmost groups admixed with Yeniseian-like populations (Samoyedic), those showing strong founder effects (Finnic), or those isolated in the Circum-Artic region with neighbouring Siberian peoples in Kola (Saami). All in all, Hungarians, Estonians and Mordovians seem to show the original situation better than the other groups, which is also reflected in part in Y-DNA, conserved as a majority of R1a lineages precisely in these groups. Just another reminder that CWC-related ancestry is found in every single Uralic group, and that it represents the main ancestral component in all non-Samoyedic groups.

estonians-hungarians-mordvinian
Selection of the PCA, with the group of Estonians, Mordovians, and Hungarians selected.

The qpGraph shows the ancestor of Yamna (likely Khvalynsk) and Corded Ware stemming as different populations from a common (likely Neolithic) node – whose difference is based on the proportion of Anatolian-related ancestry – , that is, probably before the Indo-Hittite expansion; and ends with CWC groups forming the base for all Uralic peoples. Below is a detail of the qpGraph on the left, and my old guess (2017) on the right, for comparison:

yamna-corded-ware-qpgraph

#EDIT (22 sep 2018): I enjoyed re-reading it, and found this particular paragraph funny:

Despite the documented history of the migration of Magyars [63] and their linguistic affinity to Khantys and Mansis, who today live east of the Ural Mountains, there is nothing in the present-day gene pool of the sampled Hungarians that we could tie specifically to other Uralic speakers.

They are so obsessed with finding a link to Siberian ancestry and N1c, and so convinced of Kristiansen’s idea of CWC=Indo-European, that they forgot to examine their own data from a critical point of view, and see the clear link between all Uralic peoples with Corded Ware ancestry and R1a-Z645 subclades… Here is a reminder about Hungarians and R1a-Z282, and about the expansion of R1a-Z645 with Uralic peoples.

Related

The Iron Age expansion of Southern Siberian groups and ancestry with Scythians

iron_age-sarmatians

Maternal genetic features of the Iron Age Tagar population from Southern Siberia (1st millennium BC), by Pilipenko et al. (2018).

Interesting excerpts (emphasis mine):

The positions of non-Tagar Iron Age groups in the MDS plot were correlated with their geographic position within the Eurasian steppe belt and with frequencies of Western and Eastern Eurasian mtDNA lineages in their gene pools. Series from chronological Tagar stages (similar to the overall Tagar series) were located within the genetic variability (in terms of mtDNA) of Scythian World nomadic groups (Figs 5 and 6; S4 and S6 Tables). Specifically, the Early Tagar series was more similar to western nomads (North Pontic Scythians), while the Middle Tagar was more similar to the Southern Siberian populations of the Scythian period. The Late Tagar group (Tes`culture) belonging to the Early Xiongnu period had the “western-most” location on the MDS plot with the maximal genetic difference from Xiongnu and other eastern nomadic groups (but see Discussion concerning the low sample size for the Tes`series).

In a comparison of our Tagar series with modern populations in Eurasia, we detected similarity between the Tagar group and some modern Turkic-speaking populations (with the exception of the Indo-Iranian Tajik population) (Fig 7; S2 Table). Among the modern Turkic-speaking groups, populations from the western part of the Eurasian steppe belt, such as Bashkirs from the Volga-Ural region and Siberian Tatars from the West Siberian forest-steppe zone, were more similar to the Tagar group than modern Turkic-speaking populations of the Altay-Sayan mountain system (including the Khakassians from the Minusinsk basin) (Fig 7).

tagar-archaeology
Location of Tagar archaeological sites from which samples for this study were obtained. Burial grounds: 1—Novaya Chernaya-1; 2—Podgornoe Ozero, Barsuchiha-1, Barsuchiha-6, Barsuchiha-7; 3—Perevozinskiy; 4—Ulug-Kyuzyur, Kichik-Kyuzyur, Sovetskaya Khakassiya; 5—Tepsey-3, Tepsey-8, Tepsey-9; 6—Dolgiy Kurgan. https://doi.org/10.1371/journal.pone.0204062.g001

Mitochondrial DNA diversity and genetic relationships of the Tagar population

Our results are not inconsistent with the assumption of a probable role of gene flow due to the migration from Western Eurasia to the Minusinsk basin in the Bronze Age in the formation of the genetic composition of the Tagar population. Particularly, we detected many mtDNA lineages/clusters with probable West Eurasian origin that were dominant in modern populations of different parts of Europe, Caucasus, and the Near East (such as K and HV6) in our Tagar series based on a phylogeographic analysis.

We detected relatively low genetic distances between our Tagar population and two Bronze Age populations from the Minusinsk basin—the Okunevo culture population (pre-Andronovo Bronze Age) and Andronovo culture population, followed by Afanasievo population from the Minusinsk Basin and Middle Bronze Age population from the Mongolian Altai Mountains (the region adjacent to the Minusinsk basin) (Figs 3 and 6; S3 and S5 Tables). Among West Eurasian part of our Tagar series we also observed haplogroups/sub-haplogroups and haplotypes shared with Early and Middle Bronze Age populations from Minusinsk Basin and western part of Eurasian steppe belt (Fig 4; S5 Table). Thus, our results suggested a potentially significant role of the genetic components, introduced by migrants from Western Eurasia during the Bronze Age, in the formation of the genetic composition of the Tagar population. It is necessary to note the relatively small size of available mtDNA samples from the Bronze Age populations of Minusinsk basin; accordingly, additional mtDNA data for these populations are required to further confirm our inference.

tagar-mtdna-tree
Phylogenetic tree of mtDNA lineages from the Tagar population. Color coding of the Tagar stages: orange—the Early Tagar stage; blue—the Middle Tagar Stage; green—the Late Tagar stage. Color of haplogroup labels: yellow—for Western Eurasian haplogroups; red—for Eastern Eurasian haplogroups. https://doi.org/10.1371/journal.pone.0204062.g002

Another substantial part of the mtDNA pool of the Tagar and other eastern populations of the Scythian World is typical of populations in Southern Siberia and adjacent regions of Central Asia (autochthonous Central Asian mtDNA clusters). Most of these components belong to the East Eurasian cluster of mtDNA haplogroups. Moreover, the role of each of these components in the formation of the genetic composition of subsequent (to the present) populations in South Siberia and Central Asia could be very different. In this regard, cluster C4a2a (and its subcluster C4a2a1), and haplogroup A8 are of particular interest.

Genetic features of successive Tagar groups

We compared successive Tagar groups (Early, Middle, and Late Tagar) with each other and with other Iron Age nomadic populations to evaluate changes in the mtDNA pool structure. Despite the genetic similarity between the Early and Middle Tagar series and Scythian World nomadic groups (Figs 5 and 6; S4 and S6 Tables), there were some peculiarities. For example, the Early Tagar series was more similar to North Pontic Classic Scythians, while the Middle Tagar samples were more similar to the Southern Siberian populations of the Scythian period (i.e., completely synchronous populations of regions neighboring the Minusinsk basin, such as the Pazyryk population from the Altay Mountains and Aldy-Bel population from Tuva).

We observed differences in the mtDNA pool structure between the Early and the Middle chronological stages of the Tagar culture population, as evidenced by the change in the ratio of Western to Eastern Eurasian mtDNA components. The contribution of Eastern Eurasian lineages increased from about one-third (34.8%) in the Early Tagar group to almost one-half (45.8%) in the Middle Tagar group.

tagar-mtdna-fst
Results of multidimensional scaling based on matrix of Slatkin population differentiation (FST) according to frequencies of mtDNA haplogroup in Tagar populations and modern populations of Eurasia. Populations: Tagar (red pentagon) (this study); Mongolian-speaking populations: Khamnigans (Buryat Republic, Russia) [43]; Barghuts (Inner Mongolia, China) [44]; Buryats (Buryat Republic, Southern Siberia, Russia) [43]; Mongols (Mongolia) [45]. Turkic-speaking populations: Tuvinians (Tuva Republic, Russia) [43]; Tofalars (Irkutsk region, Russia) [46]; Altai-Kizhi ((Altai Republic, Russia) [43, 47]; Telenghits (Altai Republic, Russia) [43,47]; Tubalars (Altai Republic) [48]; Shors (Kemerovo region, Russia) [43, 47]; Khakassians (Khakassian Rupublic, Russia) [43, 46]; Altaian Kazakhs (Altai Republic) [49]; Kazakhs (Kazakhstan, Uzbekistan) [50, 51]; Kirghiz (Kyrgyzstan) [50, 51]; Uighurs (Kazakhstan and Xinjiang) [50, 52]; Siberian Tatars (Tyumen and Omsk regions, Russia) [53]; Tatars (Volga-Ural rigion, Russia) [54]; Bashkirs (Volga-Ural region, Russia) [55]; Uzbeks (Uzbekistan) [51, 56]; Turkmens (Turkmenistan) [51, 56]; Nogays [57]; Turkeys [58]; other populations: Evenks [43, 46]; Ulchi [59]; Koreans (South Korea) [43]; Han Chinese [60]; Zhuang (Guangxi, China) [61]; Tadjiks (Tadjikistan) [43, 51]; Iranians [60]; Russians [62]. https://doi.org/10.1371/journal.pone.0204062.g007

At the level of mtDNA haplogroups, we detected a decrease in the diversity of phylogenetic clusters during the transition from the Early Tagar to the Middle Tagar. This decline in diversity equally affected the West Eurasian and East Eurasian components of the Tagar mtDNA pool. It should be noted that this decrease can be partially explained by the smaller number of Middle Tagar than Early Tagar samples. Under a simple binomial approximation the mtDNA clusters, observed at frequencies of 6.3% and 11.7%, could be lost by chance in our Early (N = 46) and Middle (N = 24) Tagar samples, respectively. However, the simultaneous lack of several such clusters, with a total frequency in the gene pool of the Early group of 34.8%, is unlikely.

The observed reduction in the genetic distance between the Middle Tagar population and other Scythian-like populations of Southern Siberia(Fig 5; S4 Table), in our opinion, is primarily associated with an increase in the role of East Eurasian mtDNA lineages in the gene pool (up to nearly half of the gene pool) and a substantial increase in the joint frequency of haplogroups C and D (from 8.7% in the Early Tagar series to 37.5% in the Middle Tagar series). These features are characteristic of many ancient and modern populations of Southern Siberia and adjacent regions of Central Asia, including the Pazyryk population of the Altai Mountains. We did not obtain strong evidence for an intensification of genetic contact between the population of the Minusinsk basin and the Altai Mountains in the Middle Tagar period compared with the Early Tagar period. Although, several archaeologists have found evidence for the intensification of contact at the level of material culture, namely, a cultural influence of the population of the Altai Mountains (represented by the Pazyryk population) on the population of the Minusinsk basin (the Saragash Tagar group) [6, 71, 72].

Another important issue is the change in the genetic structure of the Tagar population during the transition from the Middle (Saragash) to the Late (Tes`) stage. The Late Tagar stage refers to the Xiongnu period. Many archaeologists suggest that the formation of the Tes`stage involved the direct cultural influence of the Xiongnu and/or related groups of nomads from more eastern regions of Central Asia [71, 73]. Some archaeologists have even suggested renaming the Tes`stage in the Tes`culture [71], emphasizing the role of new eastern cultural elements. If this influence also existed at the genetic level, then we would expect to observe new genetic elements in the Tes`gene pool, particularly those of East Eurasian origin.

Siberian ancestry

Just a reminder of the recent session in ISBA 8 on expanding Scythians (and also Mongolians and Turks) spreading Siberian ancestry, usually (wrongly) identified as “Uralic-Yeniseian” based on modern populations (similar to how steppe ancestry is wrongly identified as “Indo-European”), see the following graphic including the Tagar population:

siberian-genetic-component-chronology
Very important observation with implication of population turnover is that pre-Turkic Inner Eurasian populations’ Siberian ancestry appears predominantly “Uralic-Yeniseian” in contrast to later dominance of “Tungusic-Mongolic” sort (which does sporadically occur earlier). Alexander M. Kim

And also the poster by Alexander M. Kim et al. Yeniseian hypotheses in light of genome-wide ancient DNA from historical Siberia:

The relevance of ancient DNA data to debates in historical linguistics is an emphatic strand in much recent work on the archaeogenetics of Eurasia, where the discussion has focused heavily on Indo-European (Haak et al. 2015; Narasimhan et al. 2018; de Barros Damgaard et al. 2018a,b). We present new genome-wide ancient DNA data from a historical Siberian individual in relation to Yeniseian, an isolated language “microfamily” (Vajda 2014) that nonetheless sits at the center of numerous controversial proposals in historical linguistics and cultural interaction. Yeniseian’s sole surviving representative is Ket, a critically endangered language fluently spoken by only a few dozen individuals near the Middle Yenisei River of Central Siberia.

In strong contrast to the present-day picture, river names and argued substrate influences and loanwords in languages outside the current range of Yeniseian, as well as direct records from the Russian colonial period, indicate that speakers of extinct Yeniseian languages had a formerly much broader presence in the taiga of Central Siberia as well as further south in the mountainous Altai-Sayan region – and perhaps even further afield in Inner Asia (Vajda 2010; Gorbachov 2017; Blažek 2016). The consilience of these proposals with genetic data is not straightforward (Flegontov et al. 2015, 2017) and faces a major obstacle in the lack of genetic information from verifiable speakers of Yeniseian languages other than the Kets, who have had complex ongoing interactions with speakers of non-Yeniseian languages such as the Samoyedic Selkups. We attempt to remedy this with new historical Siberian aDNA data, orienting our search for common denominators and systematic difference in a broader landscape of concordance, discordance, and uncertainty at the interface of diachronic linguistics and genetics.

Related

Evolution of Steppe, Neolithic, and Siberian ancestry in Eurasia (ISBA 8, 19th Sep)

jena-isba8

Some information is already available from ISBA 8 (see programme in PDF), thanks to the tweets from Alexander M. Kim.

Official abstracts are listed first (emphasis mine), then reports and images with link to Kim’s tweets. Here is the list for quick access:

Updates (17:00 CET):

Turkic and Hunnic expansions

Tracing the origin and expansion of the Turkic and Hunnic confederations, by Flegontov et al.

Turkic-speaking populations, now spread over a vast area in Asia, are highly heterogeneous genetically. The first confederation unequivocally attributed to them was established by the Göktürks in the 6th c. CE. Notwithstanding written resources from neighboring sedentary societies such as Chinese, Persian, Indian and Eastern Roman, earlier history of the Turkic speakers remains debatable, including their potential connections to the Xiongnu and Huns, which dominated the Eurasian steppe in the first half of the 1st millennium CE. To answer these questions, we co-analyzed newly generated human genome-wide data from Central Asia (the 1240K panel), spanning the period from ca. 3000 to 500 YBP, and the data published by de Barros Damgaard et al. (137 ancient human genomes from across the Eurasian steppes, Nature, 2018). Firstly, we generated a PCA projection to understand genetic affinities of ancient individuals with respect to present-day Tungusic, Mongolic, Turkic, Uralic, and Yeniseian-speaking groups. Secondly, we modeled hundreds of present-day and few ancient Turkic individuals using the qpAdm tool, testing various modern/ancient Siberian and ancient West Eurasian proxies for ancestry sources.

A majority of Turkic speakers in Central Asia, Siberia and further to the west share the same ancestry profile, being a mixture of Tungusic or Mongolic speakers and genetically West Eurasian populations of Central Asia in the early 1st millennium CE. The latter are themselves modelled as a mixture of Iron Age nomads (western Scythians or Sarmatians) and ancient Caucasians or Iranian farmers. For some Turkic groups in the Urals and the Altai regions and in the Volga basin, a different admixture model fits the data: the same West Eurasian source + Uralic- or Yeniseian-speaking Siberians. Thus, we have revealed an admixture cline between Scythians and the Iranian farmer genetic cluster, and two further clines connecting the former cline to distinct ancestry sources in Siberia. Interestingly, few Wusun-period individuals harbor substantial Uralic/Yeniseian-related Siberian ancestry, in contrast to preceding Scythians and later Turkic groups characterized by the Tungusic/Mongolic-related ancestry. It remains to be elucidated whether this genetic influx reflects contacts with the Xiongnu confederacy. We are currently assembling a collection of samples across the Eurasian steppe for a detailed genetic investigation of the Hunnic confederacies.

jeong-population-clines
Three distinct East/West Eurasian clines across the continent with some interesting linguistic correlates, as earlier reported by Jeong et al. (2018). Alexander M. Kim.
siberian-genetic-component-chronology
Very important observation with implication of population turnover is that pre-Turkic Inner Eurasian populations’ Siberian ancestry appears predominantly “Uralic-Yeniseian” in contrast to later dominance of “Tungusic-Mongolic” sort (which does sporadically occur earlier). Alexander M. Kim

New interesting information on the gradual arrival of the “Uralic-Yeniseian” (Siberian) ancestry in eastern Europe with Iranian and Turkic-speaking peoples. We already knew that Siberian ancestry shows no original relationship with Uralic-speaking peoples, so to keep finding groups who expanded this ancestry eastwards in North Eurasia should be no surprise for anyone at this point.

Central Asia and Indo-Iranian

The session The Genomic Formation of South and Central Asia, by David Reich, on the recent paper by Narasimhan et al. (2018).

bmac-reich
One important upside of dense genomic sampling at single localities – greater visibility of outliers and better constraints on particular incoming ancestries’ arrival times. Gonur Tepe as a great case study of this. Alexander M. Kim
ani-asi-steppe-cline
– Tale of three clines, with clear indication that “Indus Periphery” samples drawn from an already-cosmopolitan and heterogeneous world of variable ASI & Iranian ancestry. (I know how some people like to pore over these pictures – so note red dots = just dummy data for illustration.)
– Some more certainty about primary window of steppe ancestry injection into S. Asia: 2000-1500 BC
Alexander M. Kim

British Isles

Ancient DNA and the peopling of the British Isles – pattern and process of the Neolithic transition, by Brace et al.

Over recent years, DNA projects on ancient humans have flourished and large genomic-scale datasets have been generated from across the globe. Here, the focus will be on the British Isles and applying aDNA to address the relative roles of migration, admixture and acculturation, with a specific focus on the transition from a Mesolithic hunter-gatherer society to the Neolithic and farming. Neolithic cultures first appear in Britain ca. 6000 years ago (kBP), a millennium after they appear in adjacent areas of northwestern continental Europe. However, in Britain, at the margins of the expansion the pattern and process of the British Neolithic transition remains unclear. To examine this we present genome-wide data from British Mesolithic and Neolithic individuals spanning the Neolithic transition. These data indicate population continuity through the British Mesolithic but discontinuity after the Neolithic transition, c.6000 BP. These results provide overwhelming support for agriculture being introduced to Britain primarily by incoming continental farmers, with surprisingly little evidence for local admixture. We find genetic affinity between British and Iberian Neolithic populations indicating that British Neolithic people derived much of their ancestry from Anatolian farmers who originally followed the Mediterranean route of dispersal and likely entered Britain from northwestern mainland Europe.

british-isles
Millennium of lag between farming establishment in NW mainland Europe & British Isles. Only 25 Mesolithic human finds from Britain. Alexander M. Kim.
british-admixture
– Evidently no resurgence of hunter-gatherer ancestry across Neolithic
– Argument for at least two geographically distinct entries of Neolithic farmers
Alexander M. Kim.

MN Atlantic / Megalithic cultures

Genomics of Middle Neolithic farmers at the fringe of Europe, by Sánchez Quinto et al.

Agriculture emerged in the Fertile Crescent around 11,000 years before present (BP) and then spread, reaching central Europe some 7,500 years ago (ya.) and eventually Scandinavia by 6,000 ya. Recent paleogenomic studies have shown that the spread of agriculture from the Fertile Crescent into Europe was due mainly to a demic process. Such event reshaped the genetic makeup of European populations since incoming farmers displaced and admixed with local hunter-gatherers. The Middle Neolithic period in Europe is characterized by such interaction, and this is a time where a resurgence of hunter-gatherer ancestry has been documented. While most research has been focused on the genetic origin and admixture dynamics with hunter-gatherers of farmers from Central Europe, the Iberian Peninsula, and Anatolia, data from farmers at the North-Western edges of Europe remains scarce. Here, we investigate genetic data from the Middle Neolithic from Ireland, Scotland, and Scandinavia and compare it to genomic data from hunter-gatherers, Early and Middle Neolithic farmers across Europe. We note affinities between the British Isles and Iberia, confirming previous reports. However, we add on to this subject by suggesting a regional origin for the Iberian farmers that putatively migrated to the British Isles. Moreover, we note some indications of particular interactions between Middle Neolithic Farmers of the British Isles and Scandinavia. Finally, our data together with that of previous publications allow us to achieve a better understanding of the interactions between farmers and hunter-gatherers at the northwestern fringe of Europe.

megalithic-europe
-Novel genomic data from 21 individuals from 6 sites.
– “Megalithic” individuals not systematically diff. from geographically proximate “non-megalithic” burials
– Mild evidence for over-representation of males in some British Isles megalithic tombs
– Megalithic tombs in W & N Neolithic Europe may have link to kindred structures
Alexander M. Kim

Central European Bronze Age

Ancient genomes from the Lech Valley, Bavaria, suggest socially stratified households in the European Bronze Age, by Mittnik et al.

Archaeogenetic research has so far focused on supra-regional and long-term genetic developments in Central Europe, especially during the third millennium BC. However, detailed high-resolution studies of population dynamics in a microregional context can provide valuable insights into the social structure of prehistoric societies and the modes of cultural transition.

Here, we present the genomic analysis of 102 individuals from the Lech valley in southern Bavaria, Germany, which offers ideal conditions for such a study. Several burial sites containing rich archaeological material were directly dated to the second half of the 3rd and first half of the 2nd millennium BCE and were associated with the Final Neolithic Bell Beaker Complex and the Early and Middle Bronze Age. Strontium isotope data show that the inhabitants followed a strictly patrilocal residential system. We demonstrate the impact of the population movement that originated in the Pontic-Caspian steppe in the 3rd millennium BCE and subsequent local developments. Utilising relatedness inference methods developed for low-coverage modern DNA we reconstruct farmstead related pedigrees and find a strong association between relatedness and grave goods suggesting that social status is passed down within families. The co-presence of biologically related and unrelated individuals in every farmstead implies a socially stratified complex household in the Central European Bronze Age.

lech-bavaria
Diminishing steppe ancestry and resurgent Neolithic ancestry over time. Alexander M. Kim

Notice how the arrival of Bell Beakers, obviously derived from Yamna settlers in Hungary, and thus clearly identified as expanding North-West Indo-Europeans all over Europe, marks a decrease in steppe ancestry compared to Corded Ware groups, in a site quite close to the most likely East BBC homeland. Copenhagen’s steppe ancestry = Indo-European going down the toilet, step by step…

UPDATES

Russian Far East populations

Gene geography of the Russian Far East populations – faces, genome-wide profiles, and Y-chromosomes, by Balanovsky et al.

Russian Far East is not only a remote area of Eurasia but also a link of the chain of Pacific coast regions, spanning from East Asia to Americas, and many prehistoric migrations are known along this chain. The Russian Far East is populated by numerous indigenous groups, speaking Tungusic, Turkic, Chukotko-Kamchatka, Eskimo-Aleut, and isolated languages. This linguistic and geographic variation opens question about the patterns of genetic variation in the region, which was significantly undersampled and received minor attention in the genetic literature to date. To fill in this gap we sampled Aleuts, Evenks, Evens, Itelmens, Kamchadals, Koryaks, Nanais, Negidals, Nivkhs, Orochi, Udegeis, Ulchi, and Yakuts. We also collected the demographic information of local populations, took physical anthropological photos, and measured the skin color. The photos resulted in the “synthetic portraits” of many studied groups, visualizing the main features of their faces.

north-eurasia

far-east-pca
Impressive North Eurasian biobank including 30,500 individual samples with broad consent, some genealogical info, phenotypic data. Alexander M. Kim

Finland AD 5th-8th c.

Sadly, no information will be shared on the session A 1400-year transect of ancient DNA reveals recent genetic changes in the Finnish population, by Salmela et al. We will have to stick to the abstract:

Objectives: Our objective was to use aDNA to study the population history of Finland. For this aim, we sampled and sequenced 35 individuals from ten archaeological sites across southern Finland, representing a time transect from 5th to 18th century.

Methods: Following genomic DNA extraction and preparation of indexed libraries, the samples were enriched for 1,2 million genomewide SNPs using in-solution capture and sequenced on an Illumina HighSeq 4000 instrument. The sequence data were then compared to other ancient populations as well as modern Finns, their geographical neighbors and worldwide populations. Authenticity testing of the data as well as population history inference were based on standard computational methods for aDNA, such as principal component analysis and F statistics.

Results: Despite the relatively limited temporal depth of our sample set, we are able to see major genetic changes in the area, from the earliest sampled individuals – who closely resemble the present-day Saami population residing markedly further north – to the more recent ancient individuals who show increased affinity to the neighboring Circum-Baltic populations. Furthermore, the transition to the present-day population seems to involve yet another perturbation of the gene pool.

So, most likely then, in my opinion – although possibly Y-DNA will not be reported – Finns were in the Classical Antiquity period mostly R1a with secondary N1c in the Circum-Baltic region (similar to modern Estonians, as I wrote recently), while Saami were probably mostly a mix of R1a-Z282 and I1 in southern Finland. That’s what the first transition after the 5th c. probably reflects, the spread of Finns (with mainly N1c lineages) to the north, while the more recent transition shows probably the introduction of North Germanic ancestry (and thus also R1b-U106, R1a-Z284, and I1 lineages) in the west.

Dairying in ancient Mongolia

The History of Dairying in ancient Mongolia, by Wilkin et al.

The use of mass spectrometry based proteomics presents a novel method for investigating human dietary intake and subsistence strategies from archaeological materials. Studies of ancient proteins extracted from dental calculus, as well as other archaeological material, have robustly identified both animal and plant-based dietary components. Here we present a recent case study using shotgun proteomics to explore the range and diversity of dairying in the ancient eastern Eurasian steppe. Contemporary and prehistoric Mongolian populations are highly mobile and the ephemerality of temporarily occupied sites, combined with the severe wind deflation common across the steppes, means detecting evidence of subsistence can be challenging. To examine the time depth and geographic range of dairy use in Mongolia, proteins were extracted from ancient dental calculus from 32 individuals spanning burial sites across the country between the Neolithic and Mongol Empire. Our results provide direct evidence of early ruminant milk consumption across multiple time periods, as well as a dramatic increase in the consumption of horse milk in the late Bronze Age. These data provide evidence that dairy foods from multiple species were a key part of subsistence strategies in prehistoric Mongolia and add to our understanding of the importance of early pastoralism across the steppe.

The confirmation of the date 3000-2700 BC for dairying in the eastern steppe further supports what was already known thanks to archaeological remains, that the pastoralist subsistence economy was brought for the first time to the Altai region by expanding late Khvalynsk/Repin – Early Yamna pastoralists that gave rise to the Afanasevo culture.

Neolithic transition in Northeast Asia

Genomic insight into the Neolithic transition peopling of Northeast Asia, by C. Ning

East Asian representing a large geographic region where around one fifth of the world populations live, has been an interesting place for population genetic studies. In contrast to Western Eurasia, East Asia has so far received little attention despite agriculture here evolved differently from elsewhere around the globe. To date, only very limited genomic studies from East Asia had been published, the genetic history of East Asia is still largely unknown. In this study, we shotgun sequenced six hunter-gatherer individuals from Houtaomuga site in Jilin, Northeast China, dated from 12000 to 2300 BP and, 3 farming individuals from Banlashan site in Liaoning, Northeast China, dated around 5300 BP. We find a high level of genetic continuity within northeast Asia Amur River Basin as far back to 12000 BP, a region where populations are speaking Tungusic languages. We also find our Compared with Houtaomuga hunter-gatherers, the Neolithic farming population harbors a larger proportion of ancestry from Houtaomuga related hunter-gathers as well as genetic ancestry from central or perhaps southern China. Our finding further suggests that the introduction of farming technology into Northeast Asia was probably introduced through demic diffusion.

A detail of the reported haplogroups of the Houtaomuga site:

houtaomuga-site-y-dna-mtdna

Y-DNA in Northeast Asia shows thus haplogroup N1b1 ~5000 BC, probably representative of the Baikal region, with a change to C2b-448del lineages before the Xiongnu period, which were later expanded by Mongols.

Mitogenomes from Avar nomadic elite show Inner Asian origin

ring-pommel-swords

Inner Asian maternal genetic origin of the Avar period nomadic elite in the 7th century AD Carpathian Basin, by Csáky et al. bioRxiv (2018).

Abstract (emphasis mine):

After 568 AD the nomadic Avars settled in the Carpathian Basin and founded their empire, which was an important force in Central Europe until the beginning of the 9th century AD. The Avar elite was probably of Inner Asian origin; its identification with the Rourans (who ruled the region of today’s Mongolia and North China in the 4th-6th centuries AD) is widely accepted in the historical research.

Here, we study the whole mitochondrial genomes of twenty-three 7th century and two 8th century AD individuals from a well-characterised Avar elite group of burials excavated in Hungary. Most of them were buried with high value prestige artefacts and their skulls showed Mongoloid morphological traits.

The majority (64%) of the studied samples’ mitochondrial DNA variability belongs to Asian haplogroups (C, D, F, M, R, Y and Z). This Avar elite group shows affinities to several ancient and modern Inner Asian populations.

The genetic results verify the historical thesis on the Inner Asian origin of the Avar elite, as not only a military retinue consisting of armed men, but an endogamous group of families migrated. This correlates well with records on historical nomadic societies where maternal lineages were as important as paternal descent.

mds-ancient-avar-elite
MDS with 23 ancient populations. The Multidimensional Scaling plot is based on linearised Slatkin FST values that were calculated based on whole mitochondrial sequences (stress value is 0.1581). The MDS plot shows the connection of the Avars (AVAR) to the Central-Asian populations of the Late Iron Age (C-ASIA_LIAge) and Medieval period (C-ASIA_Medieval) along coordinate 1 and coordinate 2, which is caused by non-significant genetic distances between these populations. The European ancient populations are situated on the left part of the plot, where the Iberian (IB_EBRAge), Central-European (C-EU_BRAge) and British (BRIT_BRAge) populations from Early Bronze Age and Bronze Age are clustered along coordinate 2, while the Neolithic populations from Germany (GER_Neo), Hungary (HUN_Neo), Near-East (TUR_ _Neo) and Baltic region (BALT_Neo) are located on the skirt of the plot along coordinate 1. The linearised Slatkin FST values, abbreviations and references are presented in Table S4.

Interesting excerpts:

The mitochondrial genome sequences can be assigned to a wide range of the Eurasian haplogroups with dominance of the Asian lineages, which represent 64% of the variability: four samples belong to Asian macrohaplogroup C (two C4a1a4, one C4a1a4a and one C4b6); five samples to macrohaplogroup D (one by one D4i2, D4j, D4j12, D4j5a, D5b1), and three individuals to F (two F1b1b and one F1b1f). Each haplogroup M7c1b2b, R2, Y1a1 and Z1a1 is represented by one individual. One further haplogroup, M7 (probably M7c1b2b), was detected (sample AC20); however, the poor quality of its sequence data (2.19x average coverage) did not allow further analysis of this sample.

European lineages (occurring mainly among females) are represented by the following haplogroups: H (one H5a2 and one H8a1), one J1b1a1, three T1a (two T1a1 and one T1a1b), one U5a1 and one U5b1b (Table S1).

We detected two identical F1b1f haplotypes (AC11 female and AC12 male) and two identical C4a1a4 haplotypes (AC13 and AC15 males) from the same cemetery of Kunszállás; these matches indicate the maternal kinship of these individuals. There is no chronological difference between the female and the male from Grave 30 and 32 (AC11 and AC12), but the two males buried in Grave 28 and 52 (AC13 and AC15) are not contemporaries; they lived at least 2-3 generations apart.

ward-clustering-ancient-populations
Ward type clustering of 44 ancient populations. The Ward type clustering shows separation of Asian and European populations. The Avar elite group (AVAR) is situated on an Asian branch and clustered together with Central Asian populations from Late Iron Age (C-ASIA_LIAge) and Medieval period (C-ASIA_Medieval), furthermore with Xiongnu period population from Mongolia (MON_Xiongnu) and Scythians from the Altai region (E-EU_IAge_Scyth). P values are given in percent as red numbers on the dendogram, where red rectangles indicate clusters with significant p values. The abbreviations and references are presented in Table S2.

The Avar period elite shows the lowest and non-significant genetic distances to ancient Central Asian populations dated to the Late Iron Age (Hunnic) and to the Medieval period, which is displayed on the ancient MDS plot (Fig. 4); these connections are also reflected on the haplogroup based Ward-type clustering tree (Fig. 3). Building of these large Central Asian sample pools is enabled by the small number of samples per cultural/ethnic group. Further mitogenomic data from Inner Asia are needed to specify the ancient genetic connections; however, genomic analyses are also set back by the state of archaeological research, i.e. the lack of human remains from the 4th-5th century Mongolia, which would be a particularly important region in the study of the Avar elite’s origin.

The investigated elite group from the Avar period elite also shows low genetic distances and phylogenetic connections to several Central and Inner Asian modern populations. Our results indicate that the source population of the elite group of the Avar Qaganate might have existed in Inner Asia (region of today’s Mongolia and North China) and the studied stratum of the Avars moved from there westwards towards Europe. Further genetic connections of the Avars to modern populations living to East and North of Inner Asia (Yakuts, Buryats, Tungus) probably indicate common source populations.

mds-eurasian-avar-elite-group
MDS with the 44 modern populations and the Avar elite group. The Multidimensional Scaling plot is displayed based on linearised Slatkin FST values calculated based on whole mitochondrial sequences (stress value is 0.0677). The MDS plot shows differentiation of European, Near-Eastern, Central- and East-Asian populations along coordinates 1 and 2. The Avar elite (AVAR) is located on the Asian part of plot and clustered with Uyghurs from Northwest-China (NW-CHIN_UYG) and Han Chinese (CHIN), as well as with Burusho and Hazara populations from the Central-Asian Highland (Pakistan). The linearised Slatkin FST values, abbreviations and references are presented in Table S5.

Sadly, no Y-DNA is available from this paper, although haplogroups Q, C2, or R1b (xM269) are probably to be expected, given the reported mtDNA. A replacement of the male population with subsequent migrations is obvious from the current distribution of Y-DNA haplogroups in the Carpathian Basin.

Hungarians and Corded Ware

Ancient Hungarians are important to understand the evolution, not only of Ugric, but also of Finno-Ugric peoples and their origin, since they show a genetic picture before more recent population expansions, genetic drift, and bottlenecks in eastern Europe.

By now it is evident that the migration of Magyar clans from their homeland in the Cis-Urals region (from the 4th century AD on) happened after the first waves of late and gradual expansion of N1c subclades among Finno-Ugric peoples, but before the bottlenecks seen in modern populations of eastern Europe.

In Ob-Ugric peoples, from the scarce data found in Pimenoff et al. (2018), we can see how Siberian N subclades expanded further after the separation of Magyars, evidenced by the inverted proportion of haplogroups R1a and N in modern Khantys and Mansis compared to Hungarians, and the diversity of N subclades compared to modern Fennic peoples.

Similarly to Hungarians, the situation of modern Estonians (where R1a and N subclades show approximately the same proportion, ca. 33%) is probably closer to Fennic peoples in Antiquity, not having undergone the latest strong founder effect evident in modern Finns after their expansion to the north.

middle-age-hungarian
Hungarian expansion from the 4th to the 10th century AD.

Modern Hungary

This is data from recent papers, summed up in Wikipedia:

  • In Semino et al. (2001) they found among 45 Palóc from Budapest and northern Hungary: 60% R1a, 13% R1b, 11% I, 9% E, 2% G, 2% J2.
  • In Csányi et al. (2008) Among 100 Hungarian men, 90 of whom from the Great Hungarian Plain: 30% R1a, 15% R1b, 13% I2a1, 13% J2, 9% E1b1b1a, 8% I1, 3% G2, 3% J1, 3% I*, 1% E*, 1% F*, 1% K*. Among 97 Székelys, in Romania: 20% R1b, 19% R1a, 17% I1, 11% J2, 10% J1, 8% E1b1b1a, 5% I2a1, 5% G2, 3% P*, 1% E*, 1% N.
  • In Pamjav et al. (2011), among 230 samples expected to include 6-8% Gypsy peoples: 26% R1a, 20% I2a, 19% R1b, 7% I, 6% J2, 5% H, 5% G2a, 5% E1b1b1a1, 3% J1, <1% N, <1% R2.
  • In Pamjav et al. (2017), from the Bodrogköz population: R1a-M458 (20.4%), I2a1-P37 (19%), R1b-M343 (15%), R1a-Z280 (14.3%), E1b-M78 (10.2%), and N1c-Tat (6.2%).

NOTE. The N1c-Tat found in Bodrogköz belongs to the N1c-VL29 subgroup, more frequent among Balto-Slavic peoples, which may suggest (yet again) an initial stage of the expansion of N subclades among Finno-Ugric peoples by the time of the Hungarian migration.

This is the data from FTDNA group on Hungary (copied from a Wikipedia summary of 2017 data):

  • 26.1% R1a (15% Z280, 6.5% M458, 0.9% Z93=>S23201, 3.7% unknown)
  • 19.2% R1b (6% L11-P312/U106, 5.3% P312, 4.2% L23/Z2103, 3.7% U106)
  • 16.9% I2 (15.2% CTS10228, 1.4% M223, 0.5% L38)
  • 8.3% I1
  • 8.1% J2 (5.3% M410, 2.8% M102)
  • 6.9% E1b1b1 (6% V13, 0.3% V22, 0.3% M123, 0.3% M81)
  • 6.9% G2a
  • 3.2% N (1.4% Z9136, 0.5% M2019/VL67, 0.5% Y7310, 0.9% Z16981)- note: only unrelated males are sampled
  • 2.3% Q (1.2% YP789, 0.9% M346, 0.2% M242)
  • 0.9% T
  • 0.5% J1
  • 0.2% L
  • 0.2% C

R1a-Z280 stands out in FDNA (which we have to assume has no geographic preference among modern Hungarians), while R1a-M458 is prevalent in the north, which probably points to its relationship with (at least West) Slavic populations.

Ancient Hungarians

We already knew that Hungarians show similarities with Srubna and Hunnic peoples, and this paper shows a good reason for the similarities with the Huns.

Also, recent population movements in the region (before the Avars) probably increased the proportion of R1b-L23 and I1 subclades (related to Roman and Germanic peoples) as well as possibly R1a-Z283 (mainly M458, related to the expansion of Slavs). From Understanding 6th-century barbarian social organization and migration through paleogenomics, by Amorim et al. (2018):

szolad-collegno
Y-chromosome haplogroup attribution for 37 medieval and 1 Bronze age individuals.

NOTE. The sample SZ15, of haplogroup R1a1a1b1a3a (S200), belongs to the Germanic branch Z284, which has a completely different history with its integration into the Nordic Bronze Age community.

Interesting is the Szólád Bronze Age sample of R1a1a1b2a2a (Z2123) subclade (ca. 2100-1700 BC), which is possibly the same haplogroup found in King Béla III [Z93+ (80.6%), Z2123+ (10.8%)]*. Nevertheless, Z2123 refers to an upper clade, found also in East Andronovo sites in Narasimhan et al. (2018), as well as in the modern population of the Tarim Basin.

NOTE. For more on the analysis of probability of the actual subclade, see here.

Bronze Age R1a-Z93 samples of central-east Europe – like the Balkans BA sample (ca. 1750-1625 BC) from Merichleri, of R1a1a1b2 subclade – correspond most likely to the expansion of Iranian-speaking peoples in the early 2nd millennium BC, probably to the westward expansion of the Srubna culture.

The specific subclade of King Béla III, on the other hand, probably corresponds to the more recent expansion of Magyar tribes settled in the region during the 9th century AD, so the specific subclade must have separated from those found in central-east Europe and in Andronovo during the Corded Ware expansion.

r1a-z282-z93-distribution
Modified image, from Underhill et al. (2015). Spatial frequency distributions of Z282 (green) and Z93 (blue) affiliated haplogroups. Notice the potential Finno-Ugric-associated distribution of Z282 (including M558, a Z280 subclade) according to ancient maps; the northern Eurasian finds of Z2125 (upper clade of Z2123); and the potential of M458 subclades representing a west-east expansion of Balto-Slavic as a western outgroup of an original Fenno-Ugric population, equivalent to Z284 in Scandinavia.

The study by Csányi et al. (2008), where the Tat C allele was found in 2 of 4 ancient samples, showed thus a potential 50:50 relationship of N1c in ancient Magyars, which is striking given the modern 1-3% a mere 1,000 years later, without any relevant population movement in between. This result remains to be reproduced with the current technology.

In fact, recent studies of ancient Magyars, from the 10th to the 12th century, have not shown any N1c sample, and have confirmed instead the ancient presence of R1a (two other samples, interred near Béla III), R1b (four samples), I2a (two samples) J1, and E1b, a mixed genetic picture which is more in line with what is expected.

So the question that I recently posed about east Corded Ware groups remains open: were Proto-Ugric peoples mainly of R1a-Z282 or R1a-Z93 subclades? Without ancient DNA from Middle Dnieper, Fatyanovo, Afanasevo, and the succeeding cultures (like Netted Ware) in north-eastern Europe, it is difficult to say.

It is very likely that they are going to show mainly a mixture of both R1a-Z282 and R1a-Z93 lineages, with later populations showing a higher proportion of R1a-Z280 subclades. Whether this mixture happened already during the Corded Ware period, or is the result of later developments, is still unknown. What is certain is that Hungarian N1a1a1a-L708 subclades belong to more recent additions of Siberian haplogroups to the Ugric stock, probably during the Iron Age, just centuries before the Magyar expansion.

Related

Viking Age town shows higher genetic diversity than Neolithic and Bronze Age

sigtuna-vikings

Open access Genomic and Strontium Isotope Variation Reveal Immigration Patterns in a Viking Age Town, by Krzewińska et al., Current Biology (2018).

Interesting excerpts (emphasis mine, some references deleted for clarity):

The town of Sigtuna in eastern central Sweden was one of the pioneer urban hubs in the vast and complex communicative network of the Viking world. The town that is thought to have been royally founded was planned and organized as a formal administrative center and was an important focal point for the establishment of Christianity [19]. The material culture in Sigtuna indicates that the town had intense international contacts and hosted several cemeteries with a Christian character. Some of them may have been used by kin-based groups or by people sharing the same sociocultural background. In order to explore the character and magnitude of mobility and migration in a late Viking Age town, we generated and analyzed genomic (n = 23) and strontium isotope (n = 31) data from individuals excavated in Sigtuna.

y-dna-vikings

The mitochondrial genomes were sequenced at 1.5× to 367× coverage. Most of the individuals were assigned to haplogroups commonly found in current-day Europeans, such as H, J, and U [14, 26, 27]. All of these haplotypes are present in Scandinavia today.

The Y chromosome haplogroups were assigned in seven males. The Y haplogroups include I1a, I2a, N1a, G2a, and R1b. Two identified lineages (I2a and N1a) have not been found in modern-day Sweden or Norway [28, 29]. Haplogroups I and N are associated with eastern and central Europe, as well as Finno-Ugric groups [30]. Interestingly, I2a was previously identified in a middle Neolithic Swedish hunter-gatherer dating to ca. 3,000 years BCE [31].

In Sigtuna, the genetic diversity in the late Viking Age was greater than the genetic diversity in late Neolithic and Bronze Age cultures (Unetice and Yamnaya as examples) and modern East Asians; it was on par with Roman soldiers in England but lower than in modern-day European groups (GBR and FIN; Figure 2B). Within the town, the group excavated at church 1 has somewhat greater diversity than that at cemetery 1. Interestingly, the diversity at church 1 is nearly as high as that observed in Roman soldiers in England, which is remarkable, since the latter was considered to be an exceptionally heterogeneous group in contemporary Europe [39].

pca-vikings
A PCA plot visualising all 23 individuals from Sigtuna used in ancient DNA analyses (m – males, f – females).

Different sex-related mobility patterns for Sigtuna inhabitants have been suggested based on material culture, especially ceramics. Building on design and clay analyses, some female potters in Sigtuna are thought to have grown up in Novgorod in Rus’ [40]. Moreover, historical sources mention female mobility in connection to marriage, especially among the elite from Rus’ and West Slavonic regions [41, 42]. Male mobility is also known from historical sources, often in connection to clergymen moving to the town [43].

Interestingly, we found a number of individuals from Sigtuna to be genetically similar to the modern-day human variation of eastern Europeans, and most harbor close genetic affinities to Lithuanians (Figure 2A). The strontium isotope ratios in 28 adult individuals with assigned biological sex and strontium values obtained from teeth (23 M1 and five M2) show that 70% of the females and 44% of the males from Sigtuna were non-locals (STAR Methods). The difference in migrant ratios between females and male mobility patterns was not statistically significant (Fisher’s exact test, p = 0.254 for 28 individuals and p = 0.376 for 16 individuals). Hence, no evidence of a sex-specific mobility pattern was found.

(…) As these social groups are not mirrored by our genetic or strontium data, this suggests that the inclusion in them was not based on kinship. Therefore, it appears as if socio-cultural factors, not biological bonds, governed where people were interred (i.e., the choice of cemetery).

diversity-yamna
Average pairwise genetic diversity measured in complete Sigtuna, St. Gertrud (church 1) and cemetery 1 (the Nunnan block) compared to both ancient and modern populations ranked by time period (Yamnaya, Unetice, and GBR-Roman, Roman Age individuals from Great Britain; GBR-AS, Anglo-Saxon individuals from Great Britain; GBR-IA, Iron Age individuals from Great Britain; JPT-Modern, presentday Japanese from Tokyo; FIN-Modern, present-day Finnish; GBR-Modern, present-day British; GIHModern, present-day Gujarati Indian from Houston, Texas). Error bars show ±2 SEs.

Interesting from this paper is the higher genetic (especially Y-DNA) diversity found in more recent periods (see e.g. here) compared to Neolithic and Bronze Age cultures, which is probably the reason behind some obviously wrong interpretations, e.g. regarding links between Yamna and Corded Ware populations.

The sample 84001, a “first-generation short-distance migrant” of haplogroup N1c-L392 (N1a in the new nomenclature) brings yet more proof of how:

  • Admixture changes completely within a certain number of generations. In this case, the N1c-L392 sample clusters within the genetic variation of modern Norwegians, near to the Skane Iron Age sample, and not with its eastern origin (likely many generations before).
  • This haplogroup appeared quite late in Fennoscandia but still managed to integrate and expand into different ethnolinguistic groups; in this case, this individual was probably a Viking of Nordic language, given its genetic admixture and its non-local (but neighbouring Scandinavian) strontium values.

Related

On the origin and spread of haplogroup R1a-Z645 from eastern Europe

indo-european-uralic-migrations-corded-ware

In my recent post about the origin and expansion of haplogroup R1b-L51, Chetan made an interesting comment on the origin and expansion of R1a-Z645. Since this haplogroup is also relevant for European history and dialectal North-West Indo-European and Indo-Iranian expansion, I feel compelled to do a similar post, although the picture right now is more blurry than that of R1b-L51.

I find it interesting that many geneticists would question the simplistic approach to the Out of Africa model as it is often enunciated, but they would at the same time consider the current simplistic model of Yamna expansion essentially right; a model – if anyone is lost here – based on proportions of the so-called Yamnaya™ ancestral component, as found in a small number of samples, from four or five Eneolithic–Chalcolithic cultures spanning more than a thousand years.

The “75% Yamnaya ancestry of Corded Ware”, which has been given so much publicity since 2015, made geneticists propose a “Yamna → Corded Ware → Únětice / Bell Beaker” migration model, in order of decreasing Yamnaya proportions. Y-DNA and solid archaeological models suggested that this model was wrong, and recent findings have proven it was. In fact, the CWC sample closest to Yamna was a late outlier of Esperstedt in Central Europe, whose ancestry is most likely directly related to Yamna settlers from Hungary.

These wrong interpretations have been now substituted by data from two new early samples from the Baltic, which cluster closely to Yamna, and which – based on the Y-DNA and PCA cluster formed by all Corded Ware samples – are likely the product of female exogamy with Yamna peoples from the neighbouring North Pontic region (as we are seeing, e.g. in the recent Nikitin et al. 2018).

NOTE. There is also another paper from Nikitin et al. (2017), with more ancient mtDNA, “Subdivisions of haplogroups U and C encompass mitochondrial DNA lineages of Eneolithic-Early Bronze Age Kurgan populations of western North Pontic steppe”. Link to paper (behind paywall). Most interesting data is summarized in the following table:

yamna-corded-ware-mtdna

Even after the publication of Olalde et al. (2018) and Wang et al. (2018) – where expanding Yamna settlers and Bell Beakers are clearly seen highly admixed within a few generations, and are found spread across a wide Eurasian cline (sharing one common invariable trait, the paternally inherited haplogroup, as supported by David Reich) – fine-scale studies of population structure and social dynamics is still not a thing for many, even though it receives more and more advocates among geneticists (e.g. Lazaridis, or Veeramah).

NOTE. I have tried to explain, more than once, that the nature and origin of the so-called “Yamnaya ancestry” (then “steppe ancestry”, and now subdivided further as Steppe_EMBA and Steppe_MLBA) is not known with precision before Yamna samples of ca. 3000 BC, and especially that it is not necessarily a marker of Indo-European speakers. Why some people are adamant that steppe ancestry and thus R1a must be Indo-European is mostly related to a combination of grandaddy’s haplogroup, the own modern ethnolinguistic attribution, and an aversion to sharing grandpa with other peoples and cultures.

In the meantime, we are seeing the “Yamnaya proportion” question often reversed: “how do we make Corded Ware stem from Yamna, now that we believed it?”. This is a funny circular reasoning, akin to the one used by proponents of the Franco-Cantabrian origin of R1b, when they look now at EEF proportions in Iberian R1b-L23 samples. It seems too comic to be true.

R1a and steppe ancestry

The most likely origin of haplogroup R1a-Z645 is to be found in eastern Europe. Samples published in the last year support this region as a sort of cradle of R1a expansions:

  • I1819, Y-DNA R1a1-M459, mtDNA U5b2, Ukraine Mesolithic ca. 8825-8561 calBCE, from Vasilievka.
  • I5876, Y-DNA R1a, mtDNA U5a2a, Ukraine Mesolithic 7040-6703 calBCE, from Dereivka.
  • I0061, hg R1a1-M459 (xR1a1a-M17), mtDNA C1, ca. 6773-6000 calBCE (with variable dates), from Yuzhnyy Oleni Ostrov in Karelia.
  • Samples LOK_1980.006 and LOK_1981.024.01, of hg MR1a1a-M17, mtDNA F, Baikalic cultures, dated ca. 5500-5000 BC.
  • Sample I0433, hg R1a1-M459(xM198), mtDNA U5a1i, from Samara Eneolithic, ca. 5200-4000 BCE
  • Samples A3, A8, A9, of hg R1a1-M459, mtDNA H, from sub-Neolithic cultures (Comb Ware and Zhizhitskaya) at Serteyea, although dates (ca. 5th-3rd millennium BC) need possibly a revision (from Chekunova 2014).

NOTE. The fact that Europe is better sampled than North Asia, coupled with the finding of R1a-M17 in Baikalic cultures, poses some problems as to the precise origin of this haplogroup and its subclades. While the first (Palaeolithic or Mesolithic) expansion was almost certainly from Northern Eurasia to the west – due to the Mal’ta sample – , it is still unknown if the different subclades of R1a in Europe are the result of local developments, or rather different east—west migrations through North Eurasia.

Y-Full average estimates pointed to R1a-M417 formation ca. 6500 BC, TMRCA ca. 3500 BC, and R1a-Z645 formation ca. 3300 BC, TMRCA ca. 2900 BC, so the most likely explanation was that R1a-Z645 and its subclades – similar to R1b-L23 subclades, but slightly later) expanded quickly with the expansion of Corded Ware groups.

The presence of steppe ancestry in Ukraine Eneolithic sample I6561, of haplogroup R1a-M417, from Alexandria, dated ca. 4045-3974 calBCE, pointed to the forest steppe area and late Sredni Stog as the most likely territory from where the haplogroup related to the Corded Ware culture expanded.

However, the more recent Y-SNP call showing R1a-Z93 (L657) subclade rendered Y-Full’s (at least formation) estimates too young, so we have to rethink the actual origin of both subclades, R1a-Z93 (formation ca. 2900 BC, TMRCA ca 2700 BC), and R1a-Z283 (formation ca. 2900 BC, TMRCA ca. 2800 BC).

Contrary to what we thought before this, then, it is possible that the expansion of Khvalynsk-Novodanilovka chieftains through the steppes, around the mid-5th millennium BC, had something to do with the expansion of R1a-Z645 to the north, in the forest steppe.

We could think that the finding of Z93 in Alexandria after the expansion of Khvalynsk-Novodanilovka chiefs would make it more likely that R1a-Z645 will be found in the North Pontic area. However, given that Lower Mikhailovka and Kvitjana seem to follow a steppe-related cultural tradition, different to forest steppe cultures (like Dereivka and Alexandria), and that forest steppe cultures show connections to neighbouring northern and western forest regions, the rest of the expanding R1a-Z645 community may not be related directly to the steppe at all.

Adding a hypothetical split and expansion of Z645 subclades to the mid-/late-5th millennium could place the expansion of this haplogroup to the north and west, pushed by expanding Middle PIE-speaking steppe peoples from the east:

distribution-horse-scepters
Schematic depiction of the spread of horse-head scepters in the Middle Eneolithic, representing expanding Khvalynsk-Novodanilovka chieftains. See a full version with notes here.

The Złota culture

I have already written about the Podolia-Volhynia region: about the North Pontic steppe cultures in contact with this area, and about the chaotic period of migrations when Corded Ware seem to have first emerged there among multi-directional and multi-ethnic migrants.

This is what Włodarczak (2017) says about the emergence of Corded Ware with ‘steppe features’ after the previous expansion of such features in Central Europe with Globular Amphorae peoples. He refers here to the Złota culture (appearing ca. 2900-2800 BC) in Lesser Poland, believed to be the (or a) transitional stage between GAC and Corded Ware, before the emergence of the full-fledged “Corded Ware package”.

So far, to the north of the Carpathian Mountains, including Polish lands, no graves indicating their relationship with communities of the steppe zone have been found. On the contrary, the funeral rites always display a local, central European nature. However, individual elements typical of steppe communities do appear, such as the “frog-like” arrangement of the body (Fig. 20), or items associated with Pit Grave milieux (cf. Klochko, Kośko 2009; Włodarczak 2014). A spectacular example of the latter is the pointed-base vessel of Pit Grave culture found at the cemetery in Święte, site 11 near Jarosław (Kośko et al. 2012). These finds constitute a confirmation of the importance of the relationships between communities of Pit Grave culture and Corded Ware culture. They are chronologically diverse, although most of them are dated to 2600-2400 BC – that is, to the “classic” period of Corded Ware culture.

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

However, when discussing the relationships with the steppe communities, Polish lands deserve particular attention since part of the groups inhabiting it belonged to the eastern province of Corded Ware culture (cf. Häusler 2014), which neighboured Pit Grave culture both from the east and south. In addition, there was a tradition of varied relationships with the north Pontic zone, which began to intensify from the second half of the 4th millennium BC (Kośko, Szmyt, 2009; Kośko, Klochko, 2009). These connections are especially readable in Małopolska and Kujawy (Kośko 2014; Włodarczak 2014). The emergence of the community of Globular Amphora culture in the north Pontic zone at the end of the 4th and the beginnings of the 3rd millennium BC (Szmyt 1999) became a harbinger of a cultural closening between the worlds of central Europe and the steppe.

The second important factor taking place at that time was the expansion of the people of Pit Grave culture in a westerly direction, along the Danube thoroughfare. As a result of this, also to the south of the Carpathian Mountains, e.g., along the upper Tisza River, a new “kurgan” cultural system was formed. As one outcome, the areas of central Europe, above all Małopolska, found themselves in the vicinity of areas inhabited by communities characterized by new principles of social organization and a new funeral rite. Around 2800 BC these changes became evident in different regions of Poland, with the most numerous examples being documented in south-eastern Poland and Kujawy. The nature of the funeral rite and the features of the material culture perceptible at that time do not have straight forward analogies in the world of north Pontic communities. In this respect, the “A-horizon” is a phenomenon of local, central European origin. The events preceding the emergence of the said horizon (that is, the expansion of the people of Pit Grave culture into the area north of the arc of the Carpathians) are nowadays completely unidentifiable and remain merely an interesting theoretical matter (cf. e.g., Kośko 2000). Therefore, analysis of the archaeological sources cannot confirm the first archaeogenetic analysis suggesting a bond between the communities of the Pit Grave culture and Corded Ware culture (e.g., Haak et al. 2015).

Artefacts of the “A-horizon”, i.e., shaft-hole axes, amphorae (Fig. 21), beakers, and pots with a plastic wavy strip (Fig. 7) are found in different funerary and settlement contexts, sometimes jointly with finds having characteristics of various cultures (e.g., in graves of Złota culture, or at settlements of Rzucewo culture). Hence, they primarily represent a chronological phase (c. 2800-2600 BC), one obviously related to the expansion of a new ideology.

Eastern CWC expansion

Before continuing tracing the Corded Ware culture’s main features, it is worth it to trace first their movement forward in time, as Corded Ware settlers, from Poland to the east.

Circum-Baltic CWC

According to Klochko and Kośko (1998):

The colonizing Neolithic waves are continued by the Circum-Baltic Corded Ware culture, closely related to the traditions of the Single Grave culture and traditions of the Northern European Lowlands. After ca. 2900 BC, certain cultural systems with ‘corded’ traits –genetically related to the catchment area of the south-western Baltic – appear in the drainages of the Nemen, Dvina, Upper Dnieper, and even the Volga. These communities are considered the vector of Neolithisation in the Forest Zone.

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

The picture in the Baltic (Pamariu / Rzucewo) and Finland (Battle Axe) is thus more or less clearly connected with early dates ca. 2900-2800 BC:

There is a clear interaction sphere between the eastern Gulf of Finland area – reaching from Estonia to the areas of present-day Finland and the Karelian Isthmus in Russia –, evidenced e.g. by the sharp-butted axes, derived from the Estonian Karlova axe.

Interesting in this regard is the expansion of the Corded Ware culture in Finland, into a far greater territory than previously thought, that is poorly represented in most maps depicting the extent of the culture in Europe. Here is summary of CWC findings in Finland, using images from Nordqvist and Häkäla (2014):

finland-cwc
Corded Ware culture remains in Finland, excluding the so-called ‘imitations’. [Notice in the top left image the often depicted border of the culture]. Combination of maps from Nordqvist & Häkälä (2014)

Middle Dnieper and Fatyanovo

The earliest Middle Dnieper remains are related to CWC graves between the Upper Vistula and the Bug, containing pottery with Middle Dnieper traits, dated probably ca. 2700 BC, which links it with the expansion of the A-horizon. In fact, during the period ca. 2800-2400 BC, the area of Lesser Poland (with its numerous kurgans and catacomb burials) is considered the western fringe of an area spreading to the east, to the middle Dniester and middle Dnieper river basins, i.e. regions bordering the steppe oecumene. This ‘eastern connection’ of funeral ritual, raw materials, and stylistic traits of artefacts is also identified in some graves of the Polish Lowlands (Włodarczak 2017).

gac-cwc-baltic-dnieper
Cultural situation in Eastern Europe in approximately the middle of the III mill. BC. Key: 1 – areas settled by Globular Amphora culture populations; 2 – areas penetrated by Globular Amphora culture populations; 3 – border between central and eastern group; 4 – Pamariu/Rzucewo culture area; 5 – zone of Pamariu/Rzucewo culture influences; 6 – directions of Comb Pottery culture influence; 7 – Zhizhitskaya culture; 8 – eastern border of “pure” Corded Ware site; 9 – North Belarussian culture; 10 – Middle Dnieper culture; 11 – Fatyanovo culture; 12 – Yamnaya culture; 13 – eastern border of Dniester group; 14 – Kemi-Oba culture and influences; 15 – Foltesti culture; 16 – syncretic sites with evidence of Globular Amphora culture traits (1 – Nida; 2 – Butinge; 3 – Palanga; 4 – Juodkrante; 5 – Azyarnoye; 6 – Mali Rogi; 7 – Prorva; 8 – Strumen/Losha; 9 – Syabrovichi; 10 – Luchin-Zavale; 11 – Lunevo (?); 12 – Belynets; 13 – Losiatyn; 14 – Corpaci; 15 – Ocnita; 16-17 – Camenca; 18 – Marculesti; 19 – Orhei; 20 – Efimovka; 21 – Tatarbunary; 22 – Novoselitsa; 23 – Primorskoye; 24 – Sanzhiyka; 25 – Akkermen; 26 – Maydanetskoye; 27 – Grigorevka; 28 – Kholmskoye; 29 – Purcari; 30 – Roscani; 31 – Semenovka; 32 – Grishevka; 33 – Durna Skela; 34 – Iskovshchina; 35 – Primorskoye); 17 – borders of ecological zones. From Szmyt (2010)

The Fatyanovo (or Fatyanovo-Balanovo) culture was the easternmost group of the Corded Ware culture, and occupied the centre of the Russian Plain, from Lake Ilmen and the Upper Dnieper drainage to the Wiatka River and the middle course of the Volga. From the few available dates, the oldest ones from the plains of the Moskva river, and from the late Volosovo culture containing also Fatyanovo materials, and in combination they show a date of ca. 2700 BC for its appearance in the region. The Volosovo culture of foragers eventually disappeared when the Fatyanovo culture expanded into the Upper and Middle Volga basin.

The origin of the culture is complicated, because it involves at its earliest stage different Corded Ware influences in neighbouring sites, at least on the Moskva river plains (Krenke et al. 2013): some materials (possibly earlier) show Circum-Baltic and Polish features; other sites show a connection to western materials, in turn a bridge to the Middle Dnieper culture. This suggests that groups belonging to different groups of the corded ware tradition penetrated the Moscow region.

The split of subclades Z93 – Z283

If we take into account that the split between R1-Z93 and R1a-Z283 must have happened during the 5th millennium BC, we have R1a-Z93 likely around the middle Dnieper area (as supported by the Alexandria sample), and R1a-Z283 possibly to the north(-west), so that it could have expanded easily into Central Europe, and – through the northern, Baltic region – to the east.

Where exactly lies the division is unclear, but for the moment all reported Circum-Baltic samples with Z645 subclades seem to belong to Z282, while R1a samples from Sintashta/Potapovka (including the Poltavka outlier) point to Abashevo being dominated by R1a-Z93 subclades.

We have to assume, then, that an original east-west split betwen R1a-Z283 and R1a-Z93 turned, in the eastern migrations, into a north-south split between Z282 and Z93, where Finland and Battle Axe in general is going to show Z282, and Middle Dnieper – Abashevo Z93 subclades.

copper-age-early-cwc
Early Copper Age migrations ca. 3100-2600 BC.

I can think of two reasons why this is important:

  1. Depending on how Proto-Corded Ware peoples expanded, we may be talking about one community overcoming the other and imposing its language. Because either
    • clans of both Z93 and Z283 were quite close and kept intense cultural contacts around Dnieper-Dniester area; or
    • if the split is as early as the 5th millennium BC, and both communities separated then without contact, we are probably going to see a difference in the language spoken by both of them.
  2. In any case, the main north-south division of eastern Corded Ware groups is pointing to an important linguistic division within the Uralic-speaking communities, specifically between a Pre-Finno-Ugric and a Pre-Samoyedic one, and potentially between Pre-Finno-Permic and Pre-Ugric.

These may seem irrelevant questions – especially for people interested only in Indo-European migrations. However, for those interested in the history of Eurasian peoples and languages as a whole, they are relevant: even those who support an ‘eastern’ origin of Proto-Uralic, like Häkkinen, or Parpola (who are, by the way, in the minority, because most Uralicists would point to eastern Europe well before the Yamna expansion), place the Finno-Ugric expansion with the Netted Ware culture as the latest possible Finno-Ugric immigrants in Fennoscandia.

The Netted Ware culture

The image below shows the approximate expansion of Corded Ware peoples of Battle Axe traditions in Finland, as well as neighbouring Fennoscandian territories, from ca. 2800 BC until the end of the 3rd millennium. A controversial 2nd (late) wave of the so-called Estonian Corded Ware is popular in texts about this region, but has not been substantiated, and it seems to be a regional development, rather than the product of migrations.

finland-corded-ware
Left: Corded Ware remains in Finland from ca. 2800 BC, according to Nordqvist & Häkälä (2014), combined in a single image. Right: Distribution of the Corded Ware culture within Finland. Mapped (black dots) are finds of typical stone battle axes, used as a proxy (data from [8]). The red isolines indicate average permanent snow cover period from 1981 to 2010 (data from [9]). A recent study estimates the snow cover period ca 4500 years ago would have been 40–50 days less than today [10]. Overlying coloration refers to the lactose persistance (LP) allele gradient in modern northeastern Europe (see the electronic supplementary material, appendix B: Material and methods and table 1, for details); lozenge dots specify the dataset mean points for the triangulation. From Cramp et al. (2014).

As we have seen, Fatyanovo represents the most likely cultural border zone between Circum-Baltic peoples reaching from the Russian Battle Axe to the south, and Middle Dnieper peoples reaching from Abashevo to the north. In that sense, it also represents the most likely border culture between north-western (mainly R1a-Z282) and south-eastern (R1a-Z93) subclades.

With worsening climatic conditions (cooler seasons) at the end of the 3rd millennium, less settlements are apparent in the archaeological record in Finland. After ca. 2000 BC, two CWC-related cultures remain: in the coast, the Kiukainen culture, derived from the original Circum-Baltic Corded Ware settlers, reverts to a subsistence economy which includes hunting and fishing, and keeps mainly settlements (from the best territories) along the coast. In the inland, Netted Ware immigrants eventually appear from the south.

cultures-western-finland
Image modified from Cramp et al. (2014) “The timeline shows the archaeological cultures
discussed here alongside actual sherds sampled and typical vessel forms (after [26–28]) (latter not shown to scale). Distribution maps show the geographical range of (f) Typical Comb Ware, (g) Corded Ware, (h) Kiukainen Ware and (i) Bronze Age cultures in the region (after [10,20,29]).”

The Netted Ware culture emerged in the Upper Volga–Oka region, derived from the Abashevo culture and its interaction with the Seima-Turbino network, and spread ca. 1900-1800 BC to the north into Finland, spreading into eastern regions previously occupied by cultures producing asbestos and organic-tempered wares (Parpola 2018).

NOTE. Those ‘contaminated’ by the Copenhagen fantasy map series may think that Volosovo hunter-gatherers somehow survived the expansion of Fatyanovo-Balanovo and Abashevo, hidden for hundreds of years in the forest, and then reappeared and expanded the Netted Ware culture. Well, they didn’t. At least not in archaeological terms, and certainly not with the genetic data we have.

If we combine all this information, and we think about these peoples in terms of Pre-Finno-Permic and Pre-Ugric languages developing side by side, we get a really interesting picture (see here for Proto-Fennic estimates):

  • The Battle Axe around the Baltic Sea – including the Gulf of Finland and Scandinavia – would be the area of expansion of Pre-Finno-Permic peoples, of R1a-Z283 subclades, which became later concentrated mainly on coastal regions;
  • the southern areas may correspond to Pre-Ugric peoples, which expanded later to the north with Netted Ware (see image below) – their precise subclades may be dependent on what will be found in Fatyanovo;
  • and Pre-Samoyedic peoples (of R1a-Z93 subclades) would have become isolated somewhere in the Cis- or (more likely) Trans-Urals region after 2000 BC, possibly from the interaction of the latest Balanovo stages and the Seima-Turbino phenomenon.
netted-ware-parpola
Distribution of the Netted Ware according to Carpelan (2002: 198). A: Emergence of the Netted Ware on the Upper Volga c. 1900 calBC. B: Spread of Netted Ware by c. 1800 calBC. C: Early Iron Age spread of Netted Ware. (After Carpelan 2002: 198 > Parpola 2012a: 151.)

These communities in contact would have allowed for:

  • the known Indo-Iranian loanwords in Finno-Ugric to spread through a continuum of early dialects formed by Abashevo – Fatyanovo – Battle Axe groups;
  • the Finno-Saamic substrate of Germanic to be associated with Battle Axe groups in Scandinavia;
  • the important Palaeo-Germanic loanwords in Finno-Saamic spreading with long-term contacts (from Pre-Germanic to the Proto-Germanic, and later North Germanic period) through the Baltic Sea, between Scandinavia and the Gulf of Finland;
  • and Tocharian contacts with Samoyedic (although limited, and in part controversial), which point to its early expansion to the east of the Ural Mountains.

On the other hand, if one is inclined to believe that R1a and steppe ancestry do represent Indo-European speakers… which language was spoken from the Gulf of Finland well into the north, the inland, and Karelia, and in Northern Russia, by Corded Ware peoples and their cultural heirs (like Kiukainen or Netted Ware) for almost three thousand years?

Because we know that no other peoples of different haplogroups dominated over eastern Fennoscandia until the Iron Age, and N1c and Siberian ancestry expanded separately, and probably due to late bottlenecks, especially with Fennic peoples expanding recently to the north at the expense of the Saami population.

After the expansion of Bell Beaker peoples, the geographic distribution of late Corded Ware groups in the second half of the 3rd millennium, just before their demise – and before the expansion of Netted Ware to the north – , can be depicted thus as follows:

bronze_age_early_cut
Early Bronze Age Europe.

Territories in cyan must then represent, for some people who believe in an archaic Indo-Slavonic of sorts, the famous Fennoscandian Balto-Slavic to the north (before they were displaced by incoming Finno-Saamic peoples of hg N1c during the Iron Age and up to the Middle Ages); and the also famous Tundra-Forest Indo-Iranian in the Upper Volga area, a great environment for the development of the two-wheeled chariot…

But let’s leave the discussion on imaginary IE dialects for another post, and continue with the real question at hand.

A steppe funerary connection?

Back to Złota as a transitional culture, we have already seen how the corded ware vessels characteristic of the Classic CWC are related to Globular Amphora tradition, and show no break with this culture. It is usually believed that the funerary rites were adopted from steppe influence, too. That is probably right; but it does not mean that it came from Yamna or other coeval (or previous) steppe culture; at least not directly.

NOTE. A similar problem is seen when we read that Mierzanowice or Trzciniec show “Corded Ware” traits from a neighbouring CWC group, when CWC groups disappeared long before these cultures emerged. For cultural groups that are separated centuries from each other, an assertion as to their relationship needs specifics in terms of dates and material connection, or it is plainly wrong.

These are the funerary ritual features from Złota (later specialized in Corded Ware), as described by Włodarczak (2017):

  • Single burial graves; along with the habit of interring the deceased in multiple burial graves, but emphasizing their individual character by careful deposition of the body and personal nature of the grave goods.
  • Grave goods with materials and stylistiscs belonging to an older system (e.g. amber products); and others correlated to the ‘new world’ of the CWC, such as flint products made of the raw materials tipical of Lesser Poland’s CWC, copper ornaments, stone shaft-hole axes, bone and shell ornaments, and characteristic forms of vessels like beakers and amphoras.
  • Military goods, which would become prevalent in later periods, are present in a moderate number, compatible with their lesser importance.
  • There are also cases of the characteristic catacomb (“niche”) graves – with an entrance pit, a more extensive niche, and a narrow corridor leading to a vault – , as well as some individual cases of application of ochre and deformation of skulls.
catacomb-grave-ksiaznice
Catacomb grave no. 2a/06 from Książnice, Złota culture (acc. to Wilk 2013). Image from Włodarczak (2017)

It seems that the Złota funerary tradition was also “transitional”, like corded ware vessels, into the classical Corded Ware ideology. But “transitional” from what exactly? Yamna? Probably not.

The Lublin-Volhynia culture

One needs not look for a too distant culture to find similarities. Włodarczak (2017) points to CWC in south-eastern Poland and Kuyavia showing, by the time of the Yamna expansion, a funeral rite and features of the material culture without straightforward analogies in the world of north Pontic communities, and thus suggests that the “A-horizon” is a local phenomenon of central European origin.

This assertion is interesting, in so far as most Corded Ware samples investigated to date seem to come precisely from an East-Central territory near the Ukraine forest steppe, with a cluster already established by the end of the 5th millennium:

PCA-caucasus-lola-ane-chg
Image modified from Wang et al. (2018). Samples projected in PCA of 84 modern-day West Eurasian populations (open symbols). Previously known clusters have been marked and referenced. An EHG and a Caucasus ‘clouds’ have been drawn, leaving Pontic-Caspian steppe and derived groups between them.See the original file here.

The following text is from Stanisław Wilk (2018), about the Lublin-Volhynian (and related) cemeteries at Wyciąże and Książnice:

lublin-volhynia-culture
A reach of the Wyciąże-Złotniki group and Lublin-Volhynian culture in the south-eastern Poland and western Ukraine: 1. Area of the Wyciąże-Złotniki group; 2. Area of the Lublin-Volhynian culture. A. Cemetery of the Lublin-Volhynian culture at site 2 in Książnice; B. Cemetery of the Wyciąże-Złotniki group at site 5 in Kraków Nowa Huta-Wyciąże (drawing by S. Wilk based on Zakościelna 2006 and Nowak 2014, on a background downloaded from https://maps-for-free.com/).

Regardless of the differences between the two necropolises (such as the number of burials, the area which has been explored, the orientation and layout of burials), it seems that they have several key elements in common:

  • concentration of graves in separate cemeteries;
  • differentiation of burials with regard to sex (the principle of the ‘left ̶ right’ side, different burial goods for males and females);
  • stratification of graves with regard to the richness of their inventories (this mainly applied to copper artefacts);
  • occurrence of indicators of the richest male burials (a copper dagger in Wyciąże, a copper battle axe, a small axe and a chisel in Książnice);
  • allocation of a separate area for elite burials (the eastern burial area in Książnice, and the southeastern and north-central part of the necropolis in Wyciąże), as well as one for egalitarian burials (the western area in Książnice, and the south-central and western part of the cemetery in Wyciąże).
lublin-volhynian-eneolithic-cemetery
Plan of the Lublin-Volhynian culture cemetery at site 2 in Książnice: 1. female graves; 2. man graves; 3. copper traces; 4. cenothap; 5. cremation grave; 6. partial grave; 7. estimated area of the L-VC cemetery; 8. estimated area of an elite and poor burial fields; 9. area of burials containing copper artefacts (drawing by S. Wilk).

The above-mentioned characteristics prove that the patterns of social and religious behaviours from areas lying beyond the Carpathian Mountains exerted a strong influence on the two societies living in Lesser Poland.

Anna Zakościelna, while describing the similarities between the burial ritual of the late Polgár groups and cultures from areas on the Tisza river and the Lublin-Volhynia culture, claimed that:

a characteristic feature of the burial ritual of both cultures was practicing various group norms, which required different treatment of the deceased depending on their sex, age and social rank. As in the Lublin-Volhynia culture, the opposition ‘male – female’ can the most clearly be observed ̶ particularly, in the consistent positioning of males on the right, and females, on the left side. And, there is much indication that this ritual norm divided the deceased from early childhood (Sofaer Derevensky 1997: 877, Tab. 1; Lichter 2001: 276- 280, 322-323) (Zakościelna 2010: 227-228).

It seems that these observations can also be extended to the Wyciąże-Złotniki group.

Another question is whether the evidence of the influences of the copper civilization observed in both cemeteries emerged as a result of the literal copying of patterns from the south, or whether the latter were only a source of inspiration for local solutions.

Looking at this problem form the perspective of the details of burial ritual, between the Carpathian Basin and Lesser Poland, we can observe clear differences, among others, in the size of cemeteries and orientation of burials. While, in the Carpathian Basin there were large necropolises, consisting of several dozen burials located in rows, with the dominant orientation along the SE-NW and E-W axis (Lichter 2001: Abb. 123, 143; Kadrow 2008: 87); in Lesser Poland there were small cemeteries of several to a dozen or so burials, mostly oriented along the S-N axis (in the Lublin-Volhynia culture; Zakościelna 2010: 66), as well as S-E and NE-SW (in the Wyciąże-Złotniki group; Kaczanowska 2009: 77). Similarly, there are differences in the details of the burial goods. North of the Carpathians, there is a much smaller frequency of copper artefacts, particularly in the group of prestigious, heavy items (battle axes, axes and daggers), as well as a complete lack of objects made of gold. Want is more, the pottery found in the graves has a distinct local character, only supplemented by imitating or imports from areas beyond the Carpathians (Zakościelna 2006: 85; Nowak 2014: 273; a different opinion Kozłowski 2006: 57). Therefore, the suggestion made by Nowak seems right ̶ namely, that these influences were not caused by migrations of groups of the population living on the Tisza river to Lesser Poland, but were rather due to processes of selective cultural transmission (Nowak 2014: 273).

Therefore, the sharing of a similar funerary rite (as happened later between Lublin-Volhynia and Złota), although it shows a strong cultural connection with autochthonous cultures, is obviously not the same as sharing ancestors; and even if it were so, they would not need to be paternal ancestors. But it shows that important Corded Ware cultural traits are local developments, and it disconnects thus still more supposed CWC ‘steppe traits’ from steppe cultures, and connects them with the first steppe-related cultural wave that reached central Europe in the 5th millennium BC.

Prehistoric Pontic—Caspian links

How would a Lublin-Volhynia culture be related to the North Pontic area ca. 4500-3000 BC? We can enjoy the map series of Baltic—Pontic migrations by Viktor Klochko (2009), and make a wild guess:

baltic-pontic-routes
Pontic—Baltic routes of migrations during the Eneolithic. Top left: Linear Pottery expansion. Top right: Funnel Beaker expansion. Bottom left: late Trypillia expansion. Bottom right: GAC expansion.

And then read the account of Sławomir Kadrow, in Exchange of People, Ideas and Things between Cucuteni-Trypillian Complex and Areas of South-Eastern Poland (2016):

In the second half of the 5th millennium BC (horizon 1), communities of the Tripolye culture, phases BI-BII, had contacts with the population of the late (IIa) phase of the Malice culture. The areas settled by both cultural complexes were located at a great distance from each other. The communities of the Tripolye culture adopted selected features of Malice ceramic production (fig 2). This seems to have resulted from marital exchange: on a moderate scale, Tripolye men sought out their wives in the area of the Malice culture and, according to patrilocal marriage customs, the women then moved to the Tripolye settlements, sporadically transferring ready-made ceramic products, so-called imports, to the Tripolye culture. Thus, the wives were responsible for the considerably more numerous imitations of the Malice ceramics and the long-lasting, though selective, traditions of Malice pottery passed down in their new environment. The patrilocal marriage customs involving the Malice women and the Tripolye men (never the other way round), and the fact that pottery was women’s domain, led to the unidirectional transfer of vessels, technology and norms of ceramic production from the Malice culture to the Tripolye culture.

The turn of the 5th and the 4th millennia and the early 4th millennium BC (horizon 2) witnessed the deepening interaction between the populations of the youngest (IIb) phase of the Malice culture and the classic (II) phase of the Lublin-Volhynia culture on the one hand and the communities of phase BII of the Tripolye culture on the other. The Danube and the Tripolye settlement complexes came into contact on the upper Dniester and between the Styr and the Horyn rivers in Volhynia. This helped to continue the previous forms of marital exchange, which resulted in further popularisation of the ceramics and the traditions of ceramic production typical of the Danube cultures, i.e. the Malice and the Lublin-Volhynia cultures, and also the Polgár culture, in the areas settled by the Tripolye cultural complex.

As the civilizational norms of the Eneolithic (Copper) Age became widespread in that period, the forms of interaction described above acquired new elements. The deepening internal diversification of the early Eneolithic communities of the Lublin-Volhynia culture led to a growing demand for prestige objects, which was met with import or imitation of copper artefacts, mainly those from the Carpathian Basin, and with flint tools produced from long blades. That type of flint production depended largely on new technologies derived from the Tripolye culture, as proven by such borrowings as troughlike retouch or the very idea and technology for the production of long flint blades in the Lublin-Volhynia culture. It seems that the influx of Tripolye settlers into flintbearing areas in Volhynia and on the upper Dniester, adjacent to the settlement centres of the late phase of the Malice culture and the Lublin-Volhynia culture, created sufficient conditions for the expanding influence of the Tripolye flint working on the communities of the Eneolithic Lublin-Volhynia culture.

In the mid-4th millennium BC (horizon 3), those forms of interaction between the Danube communities (the late phase of the Lublin-Volhynian culture) and the Tripolye communities (phase CI)were continued. Elements of the Danube pottery still grew in popularity in the Tripolye population, while selected features of the Tripolye flint working were adopted by the Lublin-Volhynia culture.

In that period, the population of the Funnel Beaker culture of the pre-classic and early classic phases (the beginnings of Gródek 1 and Bronocice III), until then absent from those areas, quite quickly drove out and replaced the Danube population in western Volhynia and the upper Dniester basin. This caused significant changes in the forms and intensity of the intercultural interaction, which became fully apparent already in the 2nd half of the 4th millennium BC.

In the following period (horizon 4), the population of the classic phase of the Funnel Beaker culture (Gródek 1, Bronocice III) settled more and more intensively the upper Dniester basin, up to the Hnyla Lypa river, and western Volhynia, up to the Styr river. East of those rivers, the Funnel Beaker settlers created considerable areas where they mixed with settlers from early phase CII of the Tripolye culture. Their coexistence, lasting there for many generations, resulted in deepening the interactions between members of both cultural complexes and in developing entirely new forms of relationships.

(…)

The intensifying interaction between the communities of the Funnel Beaker culture and the Tripolye culture, early phase CII, in the 2nd half of the 4th millennium BC (horizon 4) was an introduction to, and perhaps a condition for, even more frequent contacts in the next period, the first centuries of the 3rd millennium BC (horizon 5). In that case, the interaction was mainly triggered by multidirectional migrations of larger human groups, involving a significant part of the population of all cultures from the areas discussed here. The Tripolye communities of younger phase CII settled Volhynia, its eastern areas in particular, from the south and the south-east, while groups representing the younger phases of the Funnel Beaker culture (Gródek 2), often with Baden features (Bronocice IV and V), moved increasingly into the western part of that region. The Yamna communities expanded along the lower and central Danube to the west, whereas the populations of the late phase of the Baden culture took the opposite direction, reaching as far as Kiev in the northeast, and contributed to the cultural character of the Sofievka group.

The communities of the Globular Amphora culture migrated from the north-west, from eastern Poland, towards the Danube Delta and as far as the Dnieper in the east, while the multicultural population from the areas around the mouth of the Danube moved in the opposite direction, carrying with them cultural elements from Thrace, or even from Anatolia. Some of them returned to the starting point (to south-eastern Poland), bringing with them a new form of pottery, so-called Thuringian amphora, borrowed from the late Trypillian Usatovo group. This resulted in origins of the Złota culture, a cultural phenomenon that gave beginnings to the oldest Corded Ware culture. Inventories of both cultures contained the already mentioned Thuringian amphorae.

lublin-volhynia-alexandria
Graves and cemeteries with gender differentiated burial rites in Europe; A — Hamangia and Varna cultures; B — Tiszapolgar and Bodrogkeresztur cultures; C — Lublin-Volhynia culture; D — Brześć Kujawski culture. Added star symbol with approximate location of the Alexandria site. Modified image from Sławomir Kadrow (2016)

Here is a more recent assessment (2017) of the latest radiocarbon analyses of the available settlements of cultures in the area, published by Marek Novak (announced in a previous post), which gives the following data on Wyciąże-Złotniki, Lublin-Volhynia, and Wyciąże/Niedźwiedź:

This scheme unambiguously suggests both the overlapping and contiguous nature of cultural development in western Lesser Poland within the Middle Neolithic. The basic elements of this development are: 1) the Wyciąże-Złotniki group and the Lublin-Volhynian culture, until c. 3650–3550 cal BC; 2) the Funnel Beaker culture proper, which appeared c. 3750–3700c al BC, and existed until c. 3300–3250 cal BC, perhaps accompanied by the Wyciąże/Niedźwiedź materials from c. 3650–3550 cal BC; and 3) the Baden culture and the Funnel Beaker/Baden assemblages from 3100 and 3300–3100 cal BC, respectively, until 2850–2750 and 2850 cal BC, with – possibly – later Funnel Beaker culture and Wyciąże/ Niedźwiedź materials, existing until c. 3100 cal BC.

The final scheme shows that the Lublin-Volhynian culture could have coincided with the Wyciąże-Złotniki group. In view of the territorial relationship between them, relations from the point of view of material culture, primarily in the field of pottery, become particularly interesting. It is relatively easy to see clear similarities between these units. However, the most evident similarities apply only to some categories of ceramics, including, for example, vessels with Scheibenhenkel handles. What is more, in the period between the late 38th and early 36th centuries BC, the early Funnel Beaker and possibly early Baden influences are superimposed on this Lublin-Volhynian/Wyciąże-Złotniki ‘mix’.

[About Corded Ware: The] development of this unit in central Europe, including western Lesser Poland, [] usually point to c. 2800 cal BC (Włodarczak 2006a). (…) the calibration curve makes it possible to alternatively refer several dates earlier than c. 3100 to c. 2850–2800 cal BC.

Conclusion

There is no direct archaeological link of Lublin-Volhynia-related groups with Corded Ware, beyond the fact that they shared homeland and Central European (‘steppe-related’) traits, as found in the Złota culture. But there is no direct link of Yamna with Corded Ware, either, whether in terms of culture or population.

So, given the evident link of R1a-Z93 and steppe ancestry with the forest steppe ca. 4000 BC, the surrounding North Pontic areas in contact along the Dniester, Dnieper, Bug, and Prut are the best candidates for the appearance of R1a-Z283: steppe cultures to the south and south-west; sub-Neolithic (Comb Ware) groups to the north in the forest zone; and Eneolithic groups to the west and north-west.

Seeing how ‘ancestral components’ and PCA cluster can change within a few generations, the question of the spread of R1a-Z645 subclades is still not settled by a single sample in Alexandria. However, based on the explosive expansions we are seeing from small territories, it would not be surprising to find R1a-Z93 and R1a-Z283 side by side in the same small area within the forest steppe.

NOTE. An archaeological link may not mean anything relevant in genetics, especially – as in this case – when no clear migration event has been traced to date. We have seen exactly that with Kristiansen’s proposal of a long-term genetic admixture of Yamna with Trypillia and GAC to form Corded Ware, which didn’t happen. The cultural and ideological connection of CWC peoples with Lublin-Volhynian tradition may be similar to the already known connection with GAC, and not mean anything in genetic finds; at least in terms of Y-DNA haplogroup.

We believed in the 2000s that Corded Ware represented the expansion of Late Proto-Indo-European, because the modern map of haplogroup R1a showed a distribution similar to how we thought the European and Indo-Iranian languages could have expanded. This has been proven wrong, and that’s what ancient DNA is for; not to confirm the own ideas or models, or to support modern ideologies.

It is impossible to know if R1a-Z645 comes from the steppe, forest steppe, or forest zone, until more samples are published. I don’t think there will be any big surprise, no matter where it is eventually found. By now, adding linguistic reconstruction to archaeological traits, and to the genetic data from Yamna and Corded Ware settlers, the only clear pattern is that patrilineal clans expanded, during the Final Eneolithic / Chalcolithic:

  • Late Proto-Indo-European with Yamna and R1b-L23 subclades, given the known genomic data from Khvalynsk, Yamna, Afanasevo, Bell Beaker, Catacomb, and Poltavka—Sintashta/Potapovka.
  • Uralic with Corded Ware and R1a-Z645 subclades, given the known genomic data from Fennoscandia and the Forest Zone.

Everything else is just wishful thinking at this moment.

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