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

steppe-forest-steppe-biomes

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

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

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

A Persistent ecological and cultural frontier

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

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

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

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

A barrier to steppe migrations into northern Europe

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

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

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

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

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

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

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

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

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

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

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

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

Related

ASoSaH Reread (II): Y-DNA haplogroups among Uralians (apart from R1a-M417)

corded-ware-yamna-ancestry

This is mainly a reread of from Book Two: A Game of Clans of the series A Song of Sheep and Horses: chapters iii.5. Early Indo-Europeans and Uralians, iv.3. Early Uralians, v.6. Late Uralians and vi.3. Disintegrating Uralians.

“Sredni Stog”

While the true source of R1a-M417 – the main haplogroup eventually associated with Corded Ware, and thus Uralic speakers – is still not known with precision, due to the lack of R1a-M198 in ancient samples, we already know that the Pontic-Caspian steppes were probably not it.

We have many samples from the north Pontic area since the Mesolithic compared to the Volga-Ural territory, and there is a clear prevalence of I2a-M223 lineages in the forest-steppe area, mixed with R1b-V88 (possibly a back-migration from south-eastern Europe).

R1a-M459 (xR1a-M198) lineages appear from the Mesolithic to the Chalcolithic scattered from the Baltic to the Caucasus, from the Dniester to Samara, in a situation similar to haplogroups Q1a-M25 and R1b-L754, which supports the idea that R1a, Q1a, and R1b expanded with ANE ancestry, possibly in different waves since the Epipalaeolithic, and formed the known ANE:EHG:WHG cline.

y-dna-khvalynsk
Y-DNA samples from Khvalynsk and neighbouring cultures. See full version.

The first confirmed R1a-M417 sample comes from Alexandria, roughly coinciding with the so-called steppe hiatus. Its emergence in the area of the previous “early Sredni Stog” groups (see the mess of the traditional interpretation of the north Pontic groups as “Sredni Stog”) and its later expansion with Corded Ware supports Kristiansen’s interpretation that Corded Ware emerged from the Dnieper-Dniester corridor, although samples from the area up to ca. 4000 BC, including the few Middle Eneolithic samples available, show continuity of hg. I2a-M223 and typical Ukraine Neolithic ancestry.

NOTE. The further subclade R1a-Z93 (Y26) reported for the sample from Alexandria seems too early, given the confidence interval for its formation (ca. 3500-2500 BC); even R1a-Z645 could be too early. Like the attribution of the R1b-L754 from Khvalynsk to R1b-V1636 (after being previously classifed as of Pre-V88 and M73 subclade), it seems reasonable to take these SNP calls with a pinch of salt: especially because Yleaf (designed to look for the furthest subclade possible) does not confirm for them any subclade beyond R1a-M417 and R1b-L754, respectively.

The sudden appearance of “steppe ancestry” in the region, with the high variability shown by Ukraine_Eneolithic samples, suggests that this is due to recent admixture of incoming foreign peoples (of Ukraine Neolithic / Comb Ware ancestry) with Novodanilovka settlers.

The most likely origin of this population, taking into account the most common population movements in the area since the Neolithic, is the infiltration of (mainly) hunter-gatherers from the forest areas. That would confirm the traditional interpretation of the origin of Uralic speakers in the forest zone, although the nature of Pontic-Caspian settlers as hunter-gatherers rather than herders make this identification today fully unnecessary (see here).

EDIT (3 FEB 2019): As for the most common guesstimates for Proto-Uralic, roughly coinciding with the expansion of this late Sredni Stog community (ca. 4000 BC), you can read the recent post by J. Pystynen in Freelance Reconstruction, Probing the roots of Samoyedic.

eneolithic-ukraine-corded-ware
Late Sredni Stog admixture shows variability proper of recent admixture of forest-steppe peoples with steppe-like population. See full version here.

NOTE. Although my initial simplistic interpretation (of early 2017) of Comb Ware peoples – traditionally identified as Uralic speakers – potentially showing steppe ancestry was probably wrong, it seems that peoples from the forest zone – related to Comb Ware or neighbouring groups like Lublyn-Volhynia – reached forest-steppe areas to the south and eventually expanded steppe ancestry into east-central Europe through the Volhynian Upland to the Polish Upland, during the late Trypillian disintegration (see a full account of the complex interactions of the Final Eneolithic).

The most interesting aspect of ascertaining the origin of R1a-M417, given its prevalence among Uralic speakers, is to precisely locate the origin of contacts between Late Proto-Indo-European and Proto-Uralic. Traditionally considered as the consequence of contacts between Middle and Upper Volga regions, the most recent archaeological research and data from ancient DNA samples has made it clear that it is Corded Ware the most likely vector of expansion of Uralic languages, hence these contacts of Indo-Europeans of the Volga-Ural region with Uralians have to be looked for in neighbours of the north Pontic area.

sredni-stog-repin-contacts
Sredni Stog – Repin contacts representing Uralic – Late Indo-European contacts were probably concentrated around the Don River.

My bet – rather obvious today – is that the Don River area is the source of the earliest borrowings of Late Uralic from Late Indo-European (i.e. post-Indo-Anatolian). The borrowing of the Late PIE word for ‘horse’ is particularly interesting in this regard. Later contacts (after the loss of the initial laryngeal) may be attributed to the traditionally depicted Corded Ware – Yamna contact zone in the Dnieper-Dniester area.

NOTE. While the finding of R1a-M417 populations neighbouring R1b-L23 in the Don-Volga interfluve would be great to confirm these contacts, I don’t know if the current pace of more and more published samples will continue. The information we have right now, in my opinion, suffices to support close contacts of neighbouring Indo-Europeans and Uralians in the Pontic-Caspian area during the Late Eneolithic.

Classical Corded Ware

After some complex movements of TRB, late Trypillia and GAC peoples, Corded Ware apparently emerged in central-east Europe, under the influence of different cultures and from a population that probably (at least partially) stemmed from the north Pontic forest-steppe area.

Single Grave and central Corded Ware groups – showing some of the earliest available dates (emerging likely ca. 3000/2900 BC) – are as varied in their haplogroups as it is expected from a sink (which does not in the least resemble the Volga-Ural population):

Interesting is the presence of R1b-L754 in Obłaczkowo, potentially of R1b-V88 subclade, as previously found in two Central European individuals from Blätterhole MN (ca. 3650 and 3200 BC), and in the Iron Gates and north Pontic areas.

Haplogroups I2a and G have also been reported in early samples, all potentially related to the supposed Corded Ware central-east European homeland, likely in southern Poland, a region naturally connected to the north Pontic forest-steppe area and to the expansion of Neolithic groups.

corded-ware-haplogroups
Y-DNA samples from early Corded Ware groups and neighbouring cultures. See full version.

The true bottlenecks under haplogroup R1a-Z645 seem to have happened only during the migration of Corded Ware to the east: to the north into the Battle Axe culture, mainly under R1a-Z282, and to the south into Middle Dnieper – Fatyanovo-Balanovo – Abashevo, probably eventually under R1a-Z93.

This separation is in line with their reported TMRCA, and supports the split of Finno-Permic from an eastern Uralic group (Ugric and Samoyedic), although still in contact through the Russian forest zone to allow for the spread of Indo-Iranian loans.

This bottleneck also supports in archaeology the expansion of a sort of unifying “Corded Ware A-horizon” spreading with people (disputed by Furholt), the disintegrating Uralians, and thus a source of further loanwords shared by all surviving Uralic languages.

Confirming this ‘concentrated’ Uralic expansion to the east is the presence of R1a-M417 (xR1a-Z645) lineages among early and late Single Grave groups in the west – which essentially disappeared after the Bell Beaker expansion – , as well as the presence of these subclades in modern Central and Western Europeans. Central European groups became thus integrated in post-Bell Beaker European EBA cultures, and their Uralic dialect likely disappeared without a trace.

NOTE. The fate of R1b-L51 lineages – linked to North-West Indo-Europeans undergoing a bottleneck in the Yamna Hungary -> Bell Beaker migration to the west – is thus similar to haplogroup R1a-Z645 – linked to the expansion of Late Uralians to the east – , hence proving the traditional interpretation of the language expansions as male-driven migrations. These are two of the most interesting genetic data we have to date to confirm previous language expansions and dialectal classifications.

It will be also interesting to see if known GAC and Corded Ware I2a-Y6098 subclades formed eventually part of the ancient Uralic groups in the east, apart from lineages which will no doubt appear among asbestos ware groups and probably hunter-gatherers from north-eastern Europe (see the recent study by Tambets et al. 2018).

Corded Ware ancestry marked the expansion of Uralians

Sadly, some brilliant minds decided in 2015 that the so-called “Yamnaya ancestry” (now more appropriately called “steppe ancestry”) should be associated to ‘Indo-Europeans’. This is causing the development of various new pet theories on the go, as more and more data contradicts this interpretation.

There is a clear long-lasting cultural, populational, and natural barrier between Yamna and Corded Ware: they are derived from different ancestral populations, which show clearly different ancestry and ancestry evolution (although they did converge to some extent), as well as different Y-DNA bottlenecks; they show different cultures, including those of preceding and succeeding groups, and evolved in different ecological niches. The only true steppe pastoralists who managed to dominate over grasslands extending from the Upper Danube to the Altai were Yamna peoples and their cultural successors.

corded-ware-yamna-pca
Corded Ware admixture proper of expanding late Sredni Stog-like populations from the forest-steppe. See full version here.

NOTE. You can also read two recent posts by FrankN in the blog aDNA era, with detailed information on the Pontic-Caspian cultures and the formation of “steppe ancestry” during the Palaeolithic, Mesolithic and Neolithic: How did CHG get into Steppe_EMBA? Part 1: LGM to Early Holocene and How did CHG get into Steppe_EMBA? Part 2: The Pottery Neolithic. Unlike your typical amateur blogger on genetics using few statistical comparisons coupled with ‘archaeolinguoracial mumbo jumbo’ to reach unscientific conclusions, these are obviously carefully redacted texts which deserve to be read.

I will not enter into the discussion of “steppe ancestry” and the mythical “Siberian ancestry” for this post, though. I will just repost the opinion of Volker Heyd – an archaeologist specialized in Yamna Hungary and Bell Beakers who is working with actual geneticists – on the early conclusions based on “steppe ancestry”:

[A]rchaeologist Volker Heyd at the University of Bristol, UK, disagreed, not with the conclusion that people moved west from the steppe, but with how their genetic signatures were conflated with complex cultural expressions. Corded Ware and Yamnaya burials are more different than they are similar, and there is evidence of cultural exchange, at least, between the Russian steppe and regions west that predate Yamnaya culture, he says. None of these facts negates the conclusions of the genetics papers, but they underscore the insufficiency of the articles in addressing the questions that archaeologists are interested in, he argued. “While I have no doubt they are basically right, it is the complexity of the past that is not reflected,” Heyd wrote, before issuing a call to arms. “Instead of letting geneticists determine the agenda and set the message, we should teach them about complexity in past human actions.

Related

“Steppe ancestry” step by step: Khvalynsk, Sredni Stog, Repin, Yamna, Corded Ware

dzudzuana_pca-large

Wang et al. (2018) is obviously a game changer in many aspects. I have already written about the upcoming Yamna Hungary samples, about the new Steppe_Eneolithic and Caucasus Eneolithic keystones, and about the upcoming Greece Neolithic samples with steppe ancestry.

An interesting aspect of the paper, hidden among so many relevant details, is a clearer picture of how the so-called Yamnaya or steppe ancestry evolved from Samara hunter-gatherers to Yamna nomadic pastoralists, and how this ancestry appeared among Proto-Corded Ware populations.

anatolia-neolithic-steppe-eneolithic
Image modified from Wang et al. (2018). Marked are in orange: equivalent Steppe_Maykop ADMIXTURE; in red, approximate limit of Anatolia_Neolithic ancestry found in Yamna populations; in blue, Corded Ware-related groups. “Modelling results for the Steppe and Caucasus cluster. Admixture proportions based on (temporally and geographically) distal and proximal models, showing additional Anatolian farmer-related ancestry in Steppe groups as well as additional gene flow from the south in some of the Steppe groups as well as the Caucasus groups.”

Please note: arrows of “ancestry movement” in the following PCAs do not necessarily represent physical population movements, or even ethnolinguistic change. To avoid misinterpretations, I have depicted arrows with Y-DNA haplogroup migrations to represent the most likely true ethnolinguistic movements. Admixture graphics shown are from Wang et al. (2018), and also (the K12) from Mathieson et al. (2018).

1. Samara to Early Khvalynsk

The so-called steppe ancestry was born during the Khvalynsk expansion through the steppes, probably through exogamy of expanding elite clans (eventually all R1b-M269 lineages) originally of Samara_HG ancestry. The nearest group to the ANE-like ghost population with which Samara hunter-gatherers admixed is represented by the Steppe_Eneolithic / Steppe_Maykop cluster (from the Northern Caucasus Piedmont).

Steppe_Eneolithic samples, of R1b1 lineages, are probably expanded Khvalynsk peoples, showing thus a proximate ancestry of an Early Eneolithic ghost population of the Northern Caucasus. Steppe_Maykop samples represent a later replacement of this Steppe_Eneolithic population – and/or a similar population with further contribution of ANE-like ancestry – in the area some 1,000 years later.

PCA-caucasus-steppe-samara

This is what Steppe_Maykop looks like, different from Steppe_Eneolithic:

steppe-maykop-admixture

NOTE. This admixture shows how different Steppe_Maykop is from Steppe_Eneolithic, but in the different supervised ADMIXTURE graphics below Maykop_Eneolithic is roughly equivalent to Eneolithic_Steppe (see orange arrow in ADMIXTURE graphic above). This is useful for a simplified analysis, but actual differences between Khvalynsk, Sredni Stog, Afanasevo, Yamna and Corded Ware are probably underestimated in the analyses below, and will become clearer in the future when more ancestral hunter-gatherer populations are added to the analysis.

2. Early Khvalynsk expansion

We have direct data of Khvalynsk-Novodanilovka-like populations thanks to Khvalynsk and Steppe_Eneolithic samples (although I’ve used the latter above to represent the ghost Caucasus population with which Samara_HG admixed).

We also have indirect data. First, there is the PCA with outliers:

PCA-khvalynsk-steppe

Second, we have data from north Pontic Ukraine_Eneolithic samples (see next section).

Third, there is the continuity of late Repin / Afanasevo with Steppe_Eneolithic (see below).

3. Proto-Corded Ware expansion

It is unclear if R1a-M459 subclades were continuously in the steppe and resurged after the Khvalynsk expansion, or (the most likely option) they came from the forested region of the Upper Dnieper area, possibly from previous expansions there with hunter-gatherer pottery.

Supporting the latter is the millennia-long continuity of R1b-V88 and I2a2 subclades in the north Pontic Mesolithic, Neolithic, and Early Eneolithic Sredni Stog culture, until ca. 4500 BC (and even later, during the second half).

Only at the end of the Early Eneolithic with the disappearance of Novodanilovka (and beginning of the steppe ‘hiatus’ of Rassamakin) is R1a to be found in Ukraine again (after disappearing from the record some 2,000 years earlier), related to complex population movements in the north Pontic area.

NOTE. In the PCA, a tentative position of Novodanilovka closer to Anatolia_Neolithic / Dzudzuana ancestry is selected, based on the apparent cline formed by Ukraine_Eneolithic samples, and on the position and ancestry of Sredni Stog, Yamna, and Corded Ware later. A good alternative would be to place Novodanilovka still closer to the Balkan outliers (i.e. Suvorovo), and a source closer to EHG as the ancestry driven by the migration of R1a-M417.

PCA-sredni-stog-steppe

The first sample with steppe ancestry appears only after 4250 BC in the forest-steppe, centuries after the samples with steppe ancestry from the Northern Caucasus and the Balkans, which points to exogamy of expanding R1a-M417 lineages with the remnants of the Novodanilovka population.

steppe-ancestry-admixture-sredni-stog

4. Repin / Early Yamna expansion

We don’t have direct data on early Repin settlers. But we do have a very close representative: Afanasevo, a population we know comes directly from the Repin/late Khvalynsk expansion ca. 3500/3300 BC (just before the emergence of Early Yamna), and which shows fully Steppe_Eneolithic-like ancestry.

afanasevo-admixture

Compared to this eastern Repin expansion that gave Afanasevo, the late Repin expansion to the west ca. 3300 BC that gave rise to the Yamna culture was one of colonization, evidenced by the admixture with north Pontic (Sredni Stog-like) populations, no doubt through exogamy:

PCA-repin-yamna

This admixture is also found (in lesser proportion) in east Yamna groups, which supports the high mobility and exogamy practices among western and eastern Yamna clans, not only with locals:

yamnaya-admixture

5. Corded Ware

Corded Ware represents a quite homogeneous expansion of a late Sredni Stog population, compatible with the traditional location of Proto-Corded Ware peoples in the steppe-forest/forest zone of the Dnieper-Dniester region.

PCA-latvia-ln-steppe

We don’t have a comparison with Ukraine_Eneolithic or Corded Ware samples in Wang et al. (2018), but we do have proximate sources for Abashevo, when compared to the Poltavka population (with which it admixed in the Volga-Ural steppes): Sintashta, Potapovka, Srubna (with further Abashevo contribution), and Andronovo:

sintashta-poltavka-andronovo-admixture

The two CWC outliers from the Baltic show what I thought was an admixture with Yamna. However, given the previous mixture of Eneolithic_Steppe in north Pontic steppe-forest populations, this elevated “steppe ancestry” found in Baltic_LN (similar to west Yamna) seems rather an admixture of Baltic sub-Neolithic peoples with a north Pontic Eneolithic_Steppe-like population. Late Repin settlers also admixed with a similar population during its colonization of the north Pontic area, hence the Baltic_LN – west Yamna similarities.

NOTE. A direct admixture with west Yamna populations through exogamy by the ancestors of this Baltic population cannot be ruled out yet (without direct access to more samples), though, because of the contacts of Corded Ware with west Yamna settlers in the forest-steppe regions.

steppe-ancestry-admixture-latvia

A similar case is found in the Yamna outlier from Mednikarovo south of the Danube. It would be absurd to think that Yamna from the Balkans comes from Corded Ware (or vice versa), just because the former is closer in the PCA to the latter than other Yamna samples. The same error is also found e.g. in the Corded Ware → Bell Beaker theory, because of their proximity in the PCA and their shared “steppe ancestry”. All those theories have been proven already wrong.

NOTE. A similar fallacy is found in potential Sintashta→Mycenaean connections, where we should distinguish statistically that result from an East/West Yamna + Balkans_BA admixture. In fact, genetic links of Mycenaeans with west Yamna settlers prove this (there are some related analyses in Anthrogenica, but the site is down at this moment). To try to relate these two populations (separated more than 1,000 years before Sintashta) is like comparing ancient populations to modern ones, without the intermediate samples to trace the real anthropological trail of what is found…Pure numbers and wishful thinking.

Conclusion

Yamna and Corded Ware show a similar “steppe ancestry” due to convergence. I have said so many times (see e.g. here). This was clear long ago, just by looking at the Y-chromosome bottlenecks that differentiate them – and Tomenable noticed this difference in ADMIXTURE from the supplementary materials in Mathieson et al. (2017), well before Wang et al. (2018).

This different stock stems from (1) completely different ancestral populations + (2) different, long-lasting Y-chromosome bottlenecks. Their similarities come from the two neighbouring cultures admixing with similar populations.

If all this does not mean anything, and each lab was going to support some pre-selected archaeological theories from the 1960s or the 1980s, coupled with outdated linguistic models no matter what – Anthony’s model + Ringe’s glottochronological tree of the early 2000s in the Reich Lab; and worse, Kristiansen’s CWC-IE + Germano-Slavonic models of the 1940s in the Copenhagen group – , I have to repeat my question again:

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

Related

Early Iranian steppe nomadic pastoralists also show Y-DNA bottlenecks and R1b-L23

New paper (behind paywall) Ancient genomes suggest the eastern Pontic-Caspian steppe as the source of western Iron Age nomads, by Krzewińska et al. Science (2018) 4(10):eaat4457.

Interesting excerpts (emphasis mine, some links to images and tables deleted for clarity):

Late Bronze Age (LBA) Srubnaya-Alakulskaya individuals carried mtDNA haplogroups associated with Europeans or West Eurasians (17) including H, J1, K1, T2, U2, U4, and U5 (table S3). In contrast, the Iron Age nomads (Cimmerians, Scythians, and Sarmatians) additionally carried mtDNA haplogroups associated with Central Asia and the Far East (A, C, D, and M). The absence of East Asian mitochondrial lineages in the more eastern and older Srubnaya-Alakulskaya population suggests that the appearance of East Asian haplogroups in the steppe populations might be associated with the Iron Age nomads, starting with the Cimmerians.

scythian-cimmerian-sarmatian-y-dna-mtdna

#UPDATE (5 OCT 2018): Some Y-SNP calls have been published in a Molgen thread, with:

  • Srubna samples have possibly two R1a-Z280, three R1a-Z93.
  • Cimmerians may not have R1b: cim357 is reported as R1a.
  • Some Scythians have low coverage to the point where it is difficult to assign even a reliable haplogroup (they report hg I2 for scy301, or E for scy197, probably based on some shared SNPs?), but those which can be reliably assigned seem R1b-Z2103 [hence probably the use of question marks and asterisks in the table, and the assumption of the paper that all Scythians are R1b-L23]:
    • The most recent subclade is found in scy305: R1b-Z2103>Z2106 (Z2106+, Y12538/Z8131+)
    • scy304: R1b-Z2103 (M12149/Y4371/Z8128+).
    • scy009: R1b-P312>U152>L2 (P312+, U152?, L2+)?
  • Sarmatians are apparently all R1a-Z93 (including tem002 and tem003);
  • You can read here the Excel file with (some probably as speculative as the paper’s own) results.

    About the PCA

    1. Srubnaya-Alakulskaya individuals exhibited genetic affinity to northern and northeastern present-day Europeans, and these results were also consistent with outgroup f3 statistics.
    2. The Cimmerian individuals, representing the time period of transition from Bronze to Iron Age, were not homogeneous regarding their genetic similarities to present-day populations according to the PCA. F3 statistics confirmed the heterogeneity of these individuals in comparison with present-day populations
    3. The Scythians reported in this study, from the core Scythian territory in the North Pontic steppe, showed high intragroup diversity. In the PCA, they are positioned as four visually distinct groups compared to the gradient of present-day populations:
      1. A group of three individuals (scy009, scy010, and scy303) showed genetic affinity to north European populations (…).
      2. A group of four individuals (scy192, scy197, scy300, and scy305) showed genetic similarities to southern European populations (…).
      3. A group of three individuals (scy006, scy011, and scy193) located between the genetic variation of Mordovians and populations of the North Caucasus (…). In addition, one Srubnaya-Alakulskaya individual (kzb004), the most recent Cimmerian (cim357), and all Sarmatians fell within this cluster. In contrast to the Scythians, and despite being from opposite ends of the Pontic-Caspian steppe, the five Sarmatians grouped close together in this cluster.
      4. A group of three Scythians (scy301, scy304, and scy311) formed a discrete group between the SC and SE and had genetic affinities to present-day Bulgarian, Greek, Croatian, and Turkish populations (…).
      5. Finally, one individual from a Scythian cultural context (scy332) is positioned outside of the modern West Eurasian genetic variation (Fig. 1C) but shared genetic drift with East Asian populations.
    scythian-cimmerian-pca
    Radiocarbon ages and geographical locations of the ancient samples used in this study. Figure panels presented (Left) Bar plot visualizing approximate timeline of presented and previously published individuals. (Right) Principal component analysis (PCA) plot visualizing 35 Bronze Age and Iron Age individuals presented in this study and in published ancient individuals (table S5) in relation to modern reference panel from the Human Origins data set (41).

    Cimmerians

    The presence of an SA component (as well as finding of metals imported from Tien Shan Mountains in Muradym 8) could therefore reflect a connection to the complex networks of the nomadic transmigration patterns characteristic of seasonal steppe population movements. These movements, although dictated by the needs of the nomads and their animals, shaped the economic and social networks linking the outskirts of the steppe and facilitated the flow of goods between settled, semi-nomadic, and nomadic peoples. In contrast, all Cimmerians carried the Siberian genetic component. Both the PCA and f4 statistics supported their closer affinities to the Bronze Age western Siberian populations (including Karasuk) than to Srubnaya. It is noteworthy that the oldest of the Cimmerians studied here (cim357) carried almost equal proportions of Asian and West Eurasian components, resembling the Pazyryks, Aldy-Bel, and Iron Age individuals from Russia and Kazakhstan (12). The second oldest Cimmerian (cim358) was also the only one with both uniparental markers pointing toward East Asia. The Q1* Y chromosome sublineage of Q-M242 is widespread among Asians and Native Americans and is thought to have originated in the Altai Mountains (24)

    Scythians

    In contrast to the eastern steppe Scythians (Pazyryks and Aldy-Bel) that were closely related to Yamnaya, the western North Pontic Scythians were instead more closely related to individuals from Afanasievo and Andronovo groups. Some of the Scythians of the western Pontic-Caspian steppe lacked the SA and the East Eurasian components altogether and instead were more similar to a Montenegro Iron Age individual (3), possibly indicating assimilation of the earlier local groups by the Scythians.

    Toward the end of the Scythian period (fourth century CE), a possible direct influx from the southern Ural steppe zone took place, as indicated by scy332. However, it is possible that this individual might have originated in a different nomadic group despite being found in a Scythian cultural context.

    scythian-alakul-variation
    Genetic diversity and ancestral components of Srubnaya-Alakulskaya population.(here called “Srubnaya”): (Left) Mean f3 statistics for Srubnaya and other Bronze Age populations. Srubnaya group was color-coded the same as with PCA. (Right) Pairwise mismatch estimates for Bronze Age populations.

    Comments

    I am surprised to find this new R1b-L23-based bottleneck in Eastern Iranian expansions so late, but admittedly – based on data from later times in the Pontic-Caspian steppe near the Caucasus – it was always a possibility. The fact that pockets of R1b-L23 lineages remained somehow ‘hidden’ in early Indo-Iranian communities was clear already since Narasimhan et al. (2018), as I predicted could happen, and is compatible with the limited archaeological data on Sintashta-Potapovka populations outside fortified settlements. I already said that Corded Ware was out of Indo-European migrations then, this further supports it.

    Even with all these data coming just from a north-west Pontic steppe region (west of the Dnieper), these ‘Cimmerians’ – or rather the ‘Proto-Scythian’ nomadic cultures appearing before ca. 800 BC in the Pontic-Caspian steppes – are shown to be probably formed by diverse peoples from Central Asia who brought about the first waves of Siberian ancestry (and Asian lineages) seen in the western steppes. You can read about a Cimmerian-related culture, Anonino, key for the evolution of Finno-Permic peoples.

    Also interesting about the Y-DNA bottleneck seen here is the rejection of the supposed continuous western expansions of R1a-Z645 subclades with steppe tribes since the Bronze Age, and thus a clearest link of the Hungarian Árpád dynasty (of R1a-Z2123 lineage) to either the early Srubna-related expansions or – much more likely – to the actual expansions of Hungarian tribes near the Urals in historic times.

    NOTE. I will add the information of this paper to the upcoming post on Ugric and Samoyedic expansions, and the late introduction of Siberian ancestry to these peoples.

    A few interesting lessons to be learned:

    • Remember the fantasy story about that supposed steppe nomadic pastoralist society sharing different Y-DNA lineages? You know, that Yamna culture expanding with R1b from Khvalynsk-Repin into the whole Pontic-Caspian steppes and beyond, developing R1b-dominated Afanasevo, Bell Beaker, and Poltavka, but suddenly appearing (in the middle of those expansions through the steppes) as a different culture, Corded Ware, to the north (in the east-central European forest zone) and dominated by R1a? Well, it hasn’t happened with any other steppe migration, so…maybe Proto-Indo-Europeans were that kind of especially friendly language-teaching neighbours?
    • Remember that ‘pure-R1a’ Indo-Slavonic society emerged from Sintashta ca. 2100 BC? (or even Graeco-Aryan??) Hmmmm… Another good fantasy story that didn’t happen; just like a central-east European Bronze Age Balto-Slavic R1a continuity didn’t happen, either. So, given that cultures from around Estonia are those showing the closest thing to R1a continuity in Europe until the Iron Age, I assume we have to get ready for the Gulf of Finland Balto-Slavic soon.
    • Remember that ‘pure-R1a’ expansion of Indo-Europeans based on the Tarim Basin samples? This paper means ipso facto an end to the Tarim Basin – Tocharian artificial controversy. The Pre-Tocharian expansion is represented by Afanasevo, and whether or not (Andronovo-related) groups of R1a-Z645 lineages replaced part or eventually all of its population before, during, or after the Tocharian expansion into the Tarim Basin, this does not change the origin of the language split and expansion from Yamna to Central Asia; just like this paper does not change the fact that these steppe groups were Proto-Iranian (Srubna) and Eastern Iranian (Scythian) speakers, regardless of their dominant haplogroup.
    • And, best of all, remember the Copenhagen group’s recent R1a-based “Indo-Germanic” dialect revival vs. the R1b-Tocharo-Italo-Celtic? Yep, they made that proposal, in 2018, based on the obvious Yamna—R1b-L23 association, and the desire to support Kristiansen’s model of Corded Ware – Indo-European expansion. Pepperidge Farm remembers. This new data on Early Iranians means another big NO to that imaginary R1a-based PIE society. But good try to go back to Gimbutas’ times, though.
    olander-classificatoin
    Olander’s (2018) tree of Indo-European languages. Presented at Languages and migrations in pre-historic Europe (7-12 Aug 2018)

    Do you smell that fresher air? It’s the Central and East European post-Communist populist and ethnonationalist bullshit (viz. pure blond R1a-based Pan-Nordicism / pro-Russian Pan-Slavism / Pan-Eurasianism, as well as Pan-Turanism and similar crap from the 19th century) going down the toilet with each new paper.

    #EDIT (5 OCT 2018): It seems I was too quick to rant about the consequences of the paper without taking into account the complexity of the data presented. Not the first time this impulsivity happens, I guess it depends on my mood and on the time I have to write a post on the specific work day…

    While the data on Srubna, Cimmerians, and Sarmatians shows clearer Y-DNA bottlenecks (of R1a-Z645 subclades) with the new data, the Scythian samples remain controversial, because of the many doubts about the haplogroups (although the most certain cases are R1b-Z2103), their actual date, and cultural attribution. However, I doubt they belong to other peoples, given the expansionist trends of steppe nomads before, during, and after Scythians (as shown in statistical analyses), so most likely they are Scythian or ‘Para-Scythian’ nomadic groups that probably came from the east, whether or not they incorporated Balkan populations. This is further supported by the remaining R1b-P312 and R1b-Z2103 populations in and around the modern Eurasian steppe region.

    scythian-peoples-balkans
    Early Iron Age cultures of the Carpathian basin ca. 7-6th century BC, including steppe groups Basarabi and Scythians. Ďurkovič et al. (2018).

    You can find an interesting and detailed take on the data published (in Russian) at Vol-Vlad’s LiveJournal (you can read an automatic translation from Google). I think that post is maybe too detailed in debunking all information associated to the supposed Scythians – to the point where just a single sample seems to be an actual Scythian (?!) -, but is nevertheless interesting to read the potential pitfalls of the study.

    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.

    Related

    Mitogenomes show likely origin of elevated steppe ancestry in neighbouring Corded Ware groups

    west-yamna-corded-ware

    Open Access Mitochondrial genomes reveal an east to west cline of steppe ancestry in Corded Ware populations, by Juras et al., Scientific Reports (2018) 8:11603.

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

    Ancient DNA was extracted from the Corded Ware culture individuals excavated in southeastern Poland (N = 12) and Moravia (N = 3). Late Eneolithic (N = 5) and Bronze Age human remains (N = 25) originated from western Ukraine and came from the Yampil barrow cemetery complex located in the north–western region of the Black Sea. Bronze Age individuals were associated with different archaeological cultures, including Yamnaya (N = 14), Catacomb (N = 2), Babyno (N = 4) and Noua (N = 5).

    The PCA results described 50.62% of the variability and were combined with the k-means clustering (with the k value of 5 as the best representation of the data, at the average silhouette of 0.2608). Based on these results individuals associated with the western and eastern Yamnaya horizon (YAE and YAW in Fig. 2) were grouped within a cluster consisting of populations from central Eurasia and Europe (blue cluster) including people associated with eastern Corded Ware culture (CWPlM) and Baltic Corded Ware culture (CWBal). This cluster did not contain any populations linked with early Neolithic farmers (red), or hunter-gatherers (green and yellow). On the other hand, k-means clustering linked the western Corded Ware culture-associated population (CWW) with Near East and Neolithic farmer ancestry groups from western and central Europe.

    pca-cwc-yamna
    Modified image, from the paper. PCA based on mitochondrial DNA haplogroup frequencies with k-means clustering. The two principal components explained 50.62% of the total variance. Loading vectors, representing mitochondrial haplogroup contributions, are highlighted as grey arrows. Populations are grouped into four clusters according to k-means. Population abbreviations are as follows: BABA – Bronze Age Balkans; CAT – Catacomb Culture; CWPlM – Corded Ware Culture from Poland and Moravia; CWBal – Baltic Corded Ware Culture; IAK – Iron Age Kazakchstan; IASI – Iron Age Syberia – Aldy Bel Culture; SCA – Scytho-Siberian Pazyryk (Altai); SCR – Rostov-Scythians, Samara; SCU – Scythians from Moldova and Ukraine; TAG – Tagar Culture; GAC – Globular Amphora Culture; YAW – western Yamnaya horizon population from Ukraine and Bulgaria; YAE – eastern Yamnaya horizon population; BAC – Baalberge Culture; BANE – Bronze Age Near East; BEC – Bernburg Culture; CHAHu – Chalcolithic Hungary; CWW – Corded Ware Culture west; CHABA – Chalcolitic Balkans; EBAG – Early Bronze Age Germany; FBC – Funnel Beaker Culture; IAG – Iron Age Germany; MNG – Middle Neolithic Germany; LBK – Linear Pottery Culture; LDN – Late Danubian Neolithic; MIC – Minoans; NEBA – Neolithic Balkans; PPNE – Pre-Pottery Near East; SCG – Schöningen group; SMC – Salzmünde Culture; AND – Andronovo Culture; BASI – Bronze Age Siberia; PWC – Pitted Ware Culture; HGE – eastern hunter-gatherers; NEUk- Neolithic Ukraine; HGS – southern hunter-gatherers; HGBal – Baltic hunter-gatheres; HGC – central huther-gatherers.

    Pairwise mtDNA-based FST values, visualized on MDS using the raw non-linearized FST (stress value = 0.099) (Fig. 4), also supported the PCA results and indicated that western and eastern Yamnaya horizon groups (YAW and YAE) were closer to people associated with the eastern Corded Ware culture (CWPlM) (FST = 0.00; FST = 0.01, respectively; both p > 0.05) and Baltic Corded Ware culture (CWBal) (FST = 0.00; FST = 0.00, respectively; both p > 0.05), than to populations associated with the western Corded Ware culture (CWW) (FST = 0.047 and FST = 0.059, respectively; both statistically significant p < 0.05). Western and eastern Yamnaya horizon groups also showed close genetic affinity to the Iron Age western Scythians (SCU) (FST = 0.0022 and FST = 0.006, respectively, both p > 0.05). The most distant populations to the Yamnaya horizon groups were western hunter-gatherers (HGW) (FST = 0.23 and FST = 0.15, p < 0.001). The FST-based MDS reflected the general European population history in the post-LGM period as the three highest FST scores were detected between western hunter-gatherers (HGW) and people associated with Linear Pottery culture (LBK) (FST = 0.33, p < 0.001), between eastern hunter-gatherers (HGE) and Baltic hunter-gatherers (HGBal) (FST = 0.35, p < 0.05), and between western (HGW) and eastern hunter-gatherers (HGE) (FST = 0.36, p < 0.05). The Yamnaya horizon groups (YAE and YAW) were placed centrally between northern hunter-gatherers (HGN) and Neolithic farmers (LDN), in direct proximity to the Bronze and Iron Age populations from Eastern Europe (SCU, BARu, SRU) and close to individuals associated with eastern and Baltic Corded Ware culture.

    yamna-corded-ware-pca
    Modified image, from the paper. In circles, relevant European groups for the question of ‘steppe ancestry’. MDS plot based on FST values calculated from mitochondrial genomes. Population abbreviations: BBC – Bell Beaker Culture; BAHu – Bronze Age Hungary; BARu – Bronze Age Russia; CWPlM – Corded Ware Culture from Poland and Moravia; CWW – western Corded Ware Culture; CWBal – Baltic Corded Ware Culture; EBAG – Early Bronze Age Germany; GAC – Globular Amphora Culture; HGE – eastern hunter-gatherers; HGN – northern hunter-gatherers; HGW – western hunter-gatherers; HGBal – Baltic hunter-gatherers; LBK – Linear Pottery Culture; LDN – Late Danubian Neolithic; MNE – Middle Neolithic; NENE – Near Eastern Neolithic; SCU – Scythians from Moldova and Ukraine; SRU – Rostov-Scythians, Samara.

    Among the analyzed samples, we identified two Catacomb culture-associated individuals (poz220 and poz221) belonging to hg X4. They are the first ancient individuals assigned to this particular lineage. Haplogroup X4 is rare among present day populations and has been found only in one individual each from Central Europe, Balkans, Anatolia and Armenia.

    Moreover, we have reported mtDNA haplotypes that might be associated with the migration from the steppe and point to genetic continuity in the north Pontic region from Bronze Age until the Iron Age. These haplotypes were assigned to hgs U5, U4, U2 and W3. MtDNA hgs U5a and U4, identified in this study among Yamnaya, Late Eneolithic and Corded Ware culture-associated individuals, have previously been found in high frequencies among northern and eastern hunter-gatherers. Moreover, they appeared in the north Pontic region in populations associated with Mesolithic (hg U5a), Eneolithic (Post-Stog) (hg U4), Yamnaya (hgs U5, U5a), Catacomb (hgs U5 and U5a) and Iron Age Scythians (hg U5a), suggesting genetic continuity of these particular mtDNA lineages in the Pontic region from, at least, the Bronze Age. Hgs U5a and U4-carrying populations were also present in the eastern steppe, along with individuals from the Yamnaya culture from Samara region, the Srubnaya and the Andronovo from Russia. Interestingly, hg U4c1 found in the Yamnaya individual (poz224) has so-far been found only in two Bell Beaker- associated individuals and one Late Bronze Age individual from Armenia, which might suggest a steppe origin for hg U4c1. A steppe origin can possibly also be assigned to hg U4a2f, found in one individual (poz282) but not reported in any other ancient populations to date, and to U5a1- the ancestral lineage of U5a1b, reported for individual poz232, which was identified not only in Corded Ware culture-associated population from central and eastern Europe, but also in representatives of Catacomb culture from the north Pontic region, Yamnaya from Bulgaria and Russia, Srubnaya and Andronovo-associated groups. Hg U2e, reported for Late Eneolithic individual (poz090), was also identified in western Corded Ware culture-associated individual and in succeeding Sintashta, Potapovka and Andronovo groups, suggesting possible genetic continuity of U2e1 in the western part of the north Pontic region.

    Hgs W3a1 and W3a1a, found in two Yamnaya individuals from this study (poz208 and poz222), were also identified in Yamnaya-associated individuals from the Russia Samara region and later in Únětice and Bell Beaker groups from Germany, supporting the idea of an eastern European steppe origin of these haplotypes and their contribution to the Yamnaya migration toward the central Europe. The W3a1 lineage was not identified in Neolithic times and, thus, we assume that it appeared in the steppe region for the first time during the Bronze Age. Notably, hgs W1 and W5, which predate the Bronze Age in Europe, were found only in individuals associated with the early Neolithic farmers from Starčevo in Hungary (hg W5), early Neolithic farmers from Anatolia (hg W1-T119C), and from the Schöningen group (hg W1c) and Globular Amphora culture from Poland (hg W5).

    west-yamna-west-corded-ware

    Some comments

    The most recent radiocarbon dates show that Early Yamna expanded to the west with Repin settlers of the Lower Don ca. 3350/3300 BC. At the end of the 4th millennium, then, these settlers dominated over groups whose population had in turn also elevated ‘steppe ancestry’ (at least from ca. 4000 BC, as shown by Ukraine Eneolithic samples from the forest-zone), and probably replaced the male population completely, as evidenced by other Yamna and Poltavka, and later Bell Beaker, Catacomb, and Sintashta samples.

    The ‘second wave’ of expansion of Yamna settlers to the west, into east-central European steppes, began probably ca. 3100/3000 BC, and – based on material culture – stemmed mainly from the North Pontic area. The Yampil Barrow Complex on the Dnieper (which I recently wrote about) seems to be part of one of the groups of western Yamna migrants: those who migrated westward from the left bank of the Dniester to the west into the Prut-Siret region, and north along the Prut.

    This region is the key for population movements that gave rise to the Corded Ware culture (see another recent post on Corded Ware origins). It is quite likely that we will see a dance of late Trypillia / Usatovo, GAC, (Proto-)Corded Ware, and Yamna samples in this area. Judging by the clear-cut Y-DNA bottlenecks we are seeing in Neolithic populations, especially among steppe pastoralists, the difference between groups in recovered ancient samples will not only be clear from their culture, but also from their male lineages.

    Based on the number of burials studied from the different settlement regions for West Yamna migrants, the Prut-Siret group was one of the smallest new Yamna ‘provinces’ in south-eastern Europe, and was probably overrun early, although – since kurgan findings continued into the Catacomb culture in the Yampil complex – the Dnieper region was well-enough connected to the core North Pontic area to be kept into its retreating territory by 2500 BC, as was the Danube delta, in contrast with other east-central European areas.

    steppe-chalcolithic-migrations
    Steppe-related migrations ca. 3100-2600 BC with tentative linguistic identification.

    Taking into account that the earliest Corded Ware burials are from ca. 2900 BC (in the Single Grave culture), and that the earliest A-horizon pottery expanded from Lesser Poland (a syncretic pottery based on the previous GAC-type) a century later, it is likely that what this paper shows for Corded Ware in eastern Europe and the Baltic is what I have suggested many times (see here, or here) as the most likely reason for elevated steppe ancestry (and close PCA cluster) of the Baltic LN ‘outliers’: the exogamy of Corded Ware groups with females from Yamna or a North Pontic steppe culture with similar ancestry.

    If Proto-Corded Ware populations of the North Pontic region did not have an identical “steppe ancestry” to these eastern CW groups already during the Eneolithic (which is the other possibility), I might be right in their more recent exogamy, and this could be seen in this study by the close cluster of east Corded Ware (especially Baltic) mtDNA to GAC and Yamna West groups, and distant from previous hunter-gatherer populations of the area, which suggests that expanding males from the Volhynia/Podolia region practiced exogamy mainly with southern groups.

    I think this is probably related to demographic pressure imposed on other populations by the explosive expansion of pastoralists with their new subsistence economy (part of the “Secondary Products Revolution”), which the hunter-gatherer and farmer population of Europe could not keep up with (as seen later in the admixture of expanding East Bell Beakers), although studies on European prehistoric demography are scarce and too general to tell us anything relevant for this precise period and region.

    Related

    Sredni Stog, Proto-Corded Ware, and their “steppe admixture”

    steppe-eneolithic-migrations

    Once the haplogroups of the announced West Yamna and Yamna settlers in Hungary and Khvalynsk from Ekaterinovka appear, it is to be expected that there won’t be much discussion on the Y-DNA bottlenecks that affected Khvalynsk – Yamna migrations.

    So let’s cut to the chase and see where Corded Ware peoples (mainly of R1a-Z645 subclades) got their so-called “steppe admixture” different from that of Yamna. Because, as you might have realized by now, Sredni Stog – and consequently Corded Ware – remains nowadays an undefined (archaeological) mess.

    Rassamakin explains it quite well, in the chapter Eneolithic of the Black Sea Steppe; In Levine M., Rassamakin Yu., Kislenko A. and Tatarintseva N., 1999. Late Prehistoric Exploitation of the Eurasian Steppe. McDonald Institute Monographs, University of Cambridge.

    NOTE. These are only certain relevant excerpts. The whole chapter is worth a thorough read, whatever position you hold regarding steppe expansions. In fact, he supports the Skelya cultural (macro-)group instead of Khvalynsk-Novodanilovka vs. Sredni Stog, he does not believe in significant expansions from the east (but in local movements and a ‘general evolution’ of Pontic-Caspian steppe cultures to Early Yamna), and offers e.g. the presence of copper and trade from the west (and its poor presence in the east) as an example of the importance of the North Pontic area vis-à-vis Khvalynsk/Repin. Not an interested party in supporting Gimbutas or Anthony, then, if you fear that.

    Cultural groups in the North Pontic area

    Telegin divided the Sredny Stog culture into three local variants – the Dnepr Culture variant, the Oskol-Donets (Aleksandriya) Culture variant, and the Lower Don (Konstantinovka) Culture variant. He elaborated a periodization based on the evolution of decorative motifs in the pottery assemblages.

    At first, the internal contradictions of the Sredny Stog culture were not accorded particular prominence, despite clear intimations of problems when, for example, sites like the settlement of Konstantinovka on the Lower Don, was identified as actually belonging to another, independent culture (Kiyashko 1974). The first real blow to the integrity of the Sredny Stog culture was dealt by Telegin himself, when he removed five Novodanilovka-type sites, and accorded them the status of an independent cultural phenomenon (Telegin 1985a). Until that point, sites of this type had were customarily considered to be within the framework of the late group of Sredny Stog cemeteries, despite having been for long regarded by some as representative of an earlier, independent cultural group (Movsha & Chebotarenko 1969; Zbenovich 1973, 74-5; Gimbutas, Merpert and Danilenko passim). Indeed, it was unclear why these cemeteries had initially been assigned to the Late, rather than the Early, Sredny Stog culture. Now these sites were mechanically stripped out of the one system, but their place in the new system was not clearly defined.

    rassamakin-skelya-culture
    Rassamakin (1999) concept of cultural development of burial rituals in the North Pontic steppe.

    Thus a paradox arose. Sites that had served to a considerable extent as the initial basis for the Sredny Stog culture, for the elaboration of its periodization and chronology, were now accepted as forming the core of an essentially different culture. Because of this, by the mid 1980s, both the Sredny Stog culture and the Eneolithic systematization as a whole were becoming rather amorphous. Essentially, the Sredny Stog culture was associated in the minds of many researchers solely with the settlement site at Dereivka, while they had only a confused and indistinct idea of the nature of early Sredny Stog sites. (…) Ultimately a situation developed where all attempts to view new evidence, new local groups, or even cultures through the prism of the Sredny Stog culture were futile, since researchers were unclear about of the essence of the culture itself, and often qualified themselves in footnotes with references to ‘preCorded Ware’ or ‘Corded Ware’ stages, or by relating their observations back directly to the specific sites of Sredny Stog II or Dereivka.

    Thus, in the Middle Eneolithic, a number of independent cultures (Kvityana, Repin, Konstantinovka, and, to some degree, Dereivka, Cernavoda and Lower Mikhailovka) emerged in the region that had in the Early Eneolithic been occupied by the Skelya culture, either as lineal successors to this culture or under its influence. But the principal stimuli in this period were the Tripolye tribes, direct imports from whom reach the southern zone of the Dnepr left bank.

    It is apparent that, for all their conceptual differences, if we remove Danilenko’s subdivisions, Telegin’s and Danilenko’s models are identical in terms of site periodization and sequence: first Kvityana, then Sredny Stog II, and finally Dereivka.

    The North Pontic area in the Eneolithic (4000-3500 BC)

    kvityana-dereivka-repin-trypolie-maykop
    The Kvityana and Dereivka cultures in relation to other sites: 1) Molyukhov Bugor (settlement); 2) Dereivka (settlement); 3) Aleksandriya (settlement); 4) Minevsky Yar (settlement); 5) Khutor Repin (settlement).

    Tripolye influence is seen most clearly in the development of the Lower Mikhailovka culture and a new burial rite which spread as far as the Molochnaya. Changes are apparent in kurgan architecture; that is, in the construction of stone chambers, sanctuaries comprising upright elements, and ring-shaped ditches (Rassamakin 1990; 1993; 1994; Pleshivenko & Rassamakin 1994 ). Lower Mikhailovka sites in the northwestern Black Sea coast region are known by a whole series of different designations, one of which, as I have already noted above, is ‘the Bessarabian variant of the Cernavoda I culture’ (Manzura 1993).

    The formation of the Kvityana culture should be considered both in the context of the development of the Lower Mikhailovka culture, and in terms of the influence of the Sredny Stog II pottery assemblage. The first is manifested in the development of the kurgan ritual itself, with such structural elements as cromlechs, orthostats and stone cists. These are most apparent to the south, in the zone of contact with the Lower Mikhailovka culture. The second is apparent in the similarity between Kvityana pottery and Sredny Stog II pottery, notably in a number of shared compositional and technical elements, despite the fact that the shapes, techniques and style are all quite different.

    As a whole, the Kvityana culture is notably conservative and archaic in appearance; this is manifest both in the preservation of a burial rite involving a supine position, and in the appearance of the pottery which, on the basis of the absence of corded or caterpillar track decoration, was until recently considered the earliest Sredny Stog ware.

    (…) we still lack sufficient evidence to trace in detail the path by which the Kvityana culture spread from the Dnepr into the Dnestr-Danube region. The southern steppe route is excluded, but Kvityana sites are recorded on the Southern Bug, in the Dnestr region, and even along the forest-steppe boundary on the Prut Gudging by the numerous excavations of kurgans in this belt. This route in some respects repeats that along which the Skelya elite groups moved. Southward movement along the Southern Bug and its tributaries into the steppe zone is indicated only by isolated sites, the number of which is far smaller than in the Dnestr-Danube region, despite the intensive excavation of kurgans in this region. Evidence for Kvityana penetration into the northwestern Black Sea coast is provided by the appearance in Usatovo assemblages of typical Kvityana figural tubular bone beads, with diagnostic lateral notches on the sides (Malyukevich & Petrenko 1993).

    kvityana-cucuteni-tripolye-expansions
    Expansions of Kvityana and Trypillia cultures. Rassamakin (1999)

    [The Dereivka] culture is currently only known only from settlement material, notably from sites in the Dnepr region (Dereivka and Molyukhov Bugor), but also from typologically distinctive pottery in the Eneolithic layer of the settlement of Aleksandriya on the Oskol. Dereivka culture pottery has also been recorded at a number of locations in the forest-steppe Dnepr region and the Seversky Donets, at Tetyanchino, Kamennye Pataki, and Minevsky Yar. The ceramic assemblage is well-defined and easily recognizable: vessels consistently display a weak profile and slightly elongated proportions, with high, straight mouths, evenly cut off at the rim, and conical bases (Fig. 3.23). The Dereivka culture occupies the southern part of the forest-steppe region and is bounded to the south by the Kvityana culture.

    Telegin rightly noted that Dereivka and Kvityana pottery bore some resemblance to one another. Several fragments of the latter were found in the Dereivka assemblage, and provide evidence for the contemporaneous existence of the two cultures. The Molyukhov Bugor pottery assemblage stands out in terms of a prevalence of pottery with corded decoration, which only occurs in insignificant amounts at Dereivka and Aleksandriya. However, artefactual analysis has not produced any clear guidance for a chronological organization of these sites, as was postulated by Telegin.

    sredni-stog-yamnaya
    Rassamakin’s (1999) periodization of the North Pontic cultures

    Late Phase and Final Eneolithic (3500-3000 BC)

    In the Dnestr region, southward pressure from Tripolye led to the formation of, firstly, Vykhvatinsk-type sites, and then, in the steppe zone, Usatovo-type sites, which had undoubtedly absorbed some features of the Lower Mikhailovka culture. In the Prut and Middle Dnestr regions, sites of the Gordineşti (Kasperovkao) types are formed (these correspond, in the Romanian Prut region and on the Siret, to sites of the Horodiştea-Erbiceni type, and on the Lower Danube to the Cernavoda III culture: Morintz & Roman 1968; Dinu 1980; 1987). Movsha considers that sites of this type (Kasperovkao in her terminology) also occur in the Southern Bug region.

    Sofievka-type sites emerge in the forest-steppes of the Middle Dnepr. A number of researchers (Zbenovich, Dergachev, and Sorokin), taking account of the change in the Tripolye culture at stage C2, propose a special division, considering sites of this period alone as ‘Late Tripolye’. In their view (with which I agree), stage Cl in culture-historical terms still corresponds to ‘Middle Tripolye’.

    (…) existing evidence allows us to put forward the following scheme (Fig. 3.49:2). To the east of the Usatovo sites, from the lower reaches of the Southern Bug to the Azov region, encroaching on the Crimean steppes, the Lower Mikhailovka culture remains intact. To the north, upstream along the Dnepr and its tributaries, the Kvityana culture survives in its initial core zone. Between the Southern Bug and the Dnepr, in the contact zone between the three cultures (Tripolye, Lower Mikhailovka and Kvityana) the Dnepr-Bug group of sites emerges, displaying mixed features (Nikolova & Rassamakin 1985; Rassamakin 1988) Tripolye influence on the Dereivka culture appears to increase, as manifested in the appearance of late cultural elements (corded decoration, plastic art, bowls). The fate of the Pivikha culture is unclear. On the Lower Don, the late phase of the Konstantinovka culture (corresponding to the settlements of Konstantinovka and Razdorskoe I: Level 7) continued.

    The final stage

    The final stage of this period is characterized by two waves of migration, which properly speaking conclude the development of the Eneolithic.(…)

    The first migration is connected with the breakdown of that system of Late Tripolye forest-steppe sites of the Prut-Dnestr and Southern Bug regions, dealt with by Movsha within the framework of the Kasperovo local group (and termed Gordineşti by others such as Dergachev, Manzura, and Petrenko). Almost all researchers into the Tripolye culture note the widespread occurrence of diagnostic elements of this group in the south, in the zone of the Usatovo sites and, in the east and southeast, towards the Dnepr and its left bank (Movsha 1984; 1990; 1993; Subbotin & Petrenko 1986; Manzura 1990a). (…)

    The migrational wave that left Zhivotilovo-Volchanskoe-type burials in the steppe also linked up the forest-steppe Bug, Dnestr, and Prut regions with the Lower Don region, and, possibly with the North Caucasus, where the late stage of the Maikop culture (the Novosvobodnaya sites) continued. The identical rites of the Maikop culture and Zhivotilovo-Volchanskoe sites makes it difficult to establish the direction of migration, or which was the active side in the process. A number of researchers have given precedence to the Maikop culture. But the spread of the Tripolye assemblage unambiguously indicates the active involvement of the Tripolye tribes.

    north-pontic-kvityana-dereivka
    The system of the latest Eneolithic Pointic cultures and the sites of the Zhivotilovo-Volchanskoe type: 1) Volchanskoe; 2) Zhivotilovka; 3) Vishnevatoe; 4) Koisug

    The second migration, at the very end of the Eneolithic, is connected with the spread of the Repin culture (in its second phase) from the Middle Don. Sinyuk defined three main directions: north, to the Upper Don; southwest, into the Dnepr region; and south, to the Lower Don and the Lower Volga. Trifonov considers this broad expansion of the Repin culture to be colonization (Trifonov 1996). The Repin culture level at Razdorskoe I (Razdorskoe I: Level 8) overlies the Konstantinovka levels (Levels 6 and 7), signalling that the Konstantinovka culture had apparently ceased to exist (Kiyashko 1994, 80). It seems that the expansion of the Repin culture is also associated with a reduction in the territorial extent of the Kvityana and Dereivka cultures. Repin burial assemblages, settlements and temporary camps appear in the Seversky Donets basin and in the Eastern Azov region (at Trekhizbenka, Kapitanovo, Aleksandriya, and Razdolnoe). The same complexes are also widely distributed towards the Dnepr (Marina 1992). The most striking western manifestation of Repin elements is seen in the upper horizon of the middle level of the Mikhailovka settlement (Lagodovska et al. 1962, 39-46).

    Khvalynsk-Yamna and Sredni Stog-Corded Ware

    We already know that Ukraine Eneolithic samples showed steppe ancestry and had apparently began a process of convergence coinciding with (or after) the first Khvalynsk-related migrations. It is unclear what had happened before (i.e. how much “CHG ancestry” was absorbed by Ukraine Neolithic groups in their transition to the Eneolithic before ca. 4500 BC), although in principle we can assume that all Caucasus-related admixture received by North Pontic cultures ca. 4500-4000 BC was mediated by westward movements from Khvalynsk-related peoples.

    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.

    Contacts with (and later absorption of) Khvalynsk-Novodanilovka-related migrants, as well as heir cultures, like those in the steppe adjacent to the Black Sea coast, and also direct contacts with Caucasus-related populations through Zhivotilovo-Volchanskoe can justify a greater contribution of CHG ancestry ca. 4000-3500 BC. Close contacts with Cucuteni-Trypillia (through Mikhailovka and maybe Kvityana, possibly with WHG and CHG admixture related to Khvalynsk-Novodanilovka) and GAC peoples to the north are the obvious source of further similarities with Yamna. Distinct similarities, that is, if we take into account the different sources and timing of such ancestral components, and Y-chromosome bottlenecks…

    Therefore, after a process of convergence ca. 4500-4000 BC, and potentially more contacts with late Eneolithic North Pontic steppe cultures ca. 4000-3500 BC, Proto-Corded Ware peoples must have finally spread from the northernmost (forest-steppe) areas previously occupied by Dereivka, Pivikha, or Sofievka groups from ca. 3300 BC onwards – a date roughly coincident with the expansion of late Khvalynsk/Repin to the west developing the Early Yamna culture, with which it likely entered in contact (hence potentially a source for further admixture convergence ca. 3500-3000 BC).

    Only later happened the great migration ca. 3000-2800 BC of Classical Corded Ware culture migrants, at the same time as Early Yamna migrants expanded to the west, and some groups also to the north along the Prut (possibly directly connected to the admixture found in the two Baltic LN/CWC ‘outliers’).

    steppe-chalcolithic-migrations
    Steppe-related migrations ca. 3100-2600 BC with tentative linguistic identification.

    We didn’t know much about Sredni Stog or Corded Ware, and we still don’t. I can’t see the future, and I don’t have access to information from Reich-Jena or Copenhagen groups, and never have. But I just don’t see the need to explain Corded Ware as derived from (coeval) Early Yamna, and I haven’t since the 2015 papers. It was not the best explanation for the data that was published, and the more information we receive, the less sense this theory makes.

    However, I guess we will see some groups still resorting to the good old Yamnaya ancestral component™ = Indo-European no matter what, consciously ignoring that a proportion of ancestral components (some combination of EHG:CHG:WHG in this case) means nothing without a proper explanation of their precise temporal and regional origin, and how they connect with Yamna; just like the CHG ancestry = Indo-European trend we are living right now does not make any sense.

    Publishing only selected results after trying every possible combination of samples with bioinformatic tools does not help strengthen this connection, either.

    Related

    Cereal cultivation and processing in Trypillian mega-sites

    eneolithic-trypillia-cucuteni-steppe

    New paper (behind paywall) Where are the cereals? Contribution of phytolith analysis to the study of subsistence economy at the Trypillia site Maidanetske (ca. 3900-3650 BCE), central Ukraine, by Dal Corso et al. Journal of Arid Environments (2018).

    Interesting excerpts (only introduction and conclusions, emphasis mine):

    Archaeological setting at the site of Maidanetske, Ukraine

    From ca. 4800 to 3350 BCE, Trypillia settlements were widespread over parts of eastern Romania, Moldova and Ukraine (Menotti and Korvin-Piotrovskiy, 2012; Müller et al., 2016; Videiko, 2004). Maidanetske (Fig. 1B) is one of the so-called “mega-sites” which developed during ca. 3900–3400 BCE in central Ukraine, in the Uman region (Cherkasy district) (Müller and Videiko, 2016; Müller et al., 2017). In this region, nine of these “mega-sites” have been found. Mega-sites are characterized by a regular plan with concentric rings of houses around a large empty central space, additional quartiers, with radial and peripheral track ways (Fig. 1B). The three mega-sites Maidanetske, Taljanky and Dobrovody, lay ca. 15 km apart from each other (Fig. 1A); other mega-sites are located within a 50 km radius around Maidanetske. Archaeologically, these mega-sites consist of the remains of buildings most of them burnt, although a minority of unburnt buildings is known of as well (Burdo and Videiko, 2016; Müller and Videiko, 2016; Ohlrau, 2015). Most of these buildings have a standardized regular size (average 6×12 m) and architecture including domestic installations and a standardized assemblage of artifacts. At Maidanetske beside normal sized houses there are few larger rectangular buildings that are located regularly along the main pathways. Further archaeological contexts include pits, pottery kilns, and peripheral ditches. A huge variety of mostly painted pottery (including many with figurative animal and plant motives), some flint artifacts, rare copper objects, querns, adzes and a broad range of anthropomorphic and zoomorphic figurines are attested within houses and mega-structures. In terms of organic remains, animal bones are fairly common, while botanical macro-remains appear to be scarce and poorly preserved (Kirleis and Dal Corso, 2016; Pashkevich and Videjko, 2006).

    trypillia-maidanetske-north-pontic-steppe-forest
    The location of the Chalcolithic site of Maidanetske and of other sites mentioned in the text within the map of the natural vegetation (modified after Kirleis
    and Dreibrodt, 2016, graphic K. Winter, Kiel University).

    Environmental setting at Maidanetske

    The Trypillia sites in central Ukraine, including Maidanetske, are located in a semi-arid forest-steppe ecozone, a mosaic-like ecosystem stretched between the dry steppe grasslands in the south and temperate woodland biomes in the north (Fig. 1A). In this transitional zone the natural vegetation is supposed to be patchy and sensitive to climate and topography (Feurdean et al., 2015; Molnàr et al., 2012; Walter, 1974). Since most of the accessible plateaus are converted to agricultural land and the scarce broadleaf woodlands are managed, the natural landscape heterogeneity is difficult to trace within the current landscape (Kuzemko et al., 2014). Besides agricultural fields and villages, narrow river valleys incised into the loess plateaus are present, with riparian vegetation and artificial lakes. This western Pontic area has a humid continental climate with wet winters and warm summers (Köppen and Geiger, 1939), which corresponds to a semi-arid 0.2–0.5 aridity index value according to UNEP (1997). Nevertheless, the reconstruction of past climatic as well as environmental conditions is not straightforward, since undisturbed archives for pollen analysis are lacking in the region and published climatic reconstructions combine evidences from peripheral areas (Gerasimenko, 1997; Harper, 2017; Kirleis and Dreibrodt, 2016). In the Transylvanian forest-steppe region, palynological investigations suggest that dry grasslands have expanded since the end of the 4th millennium BCE, fostered by Bronze Age forest clearance, while before this the area was largely forested (Feurdean et al., 2015). In the Hungarian forest-steppe, the mixed oak forest on Loess almost disappeared by the end of the 18th century AD, hampered by factors such as fragmentation, slow regeneration, spread of invasive species and lowering of the water table due to increased aridity (Molnàr et al., 2012). It is clear that forest-steppe environments are very sensitive to aridity and land use practices. To understand whether similar landscape change can have occurred in central Ukraine already at the time of Chalcolithic mega-sites, an understanding of the extent of crop growing and deforestation is crucial.

    The site of Maidanetske is situated on a plateau covered by Loess deposited during the Last Glaciation. This plateau is dissected by valleys of different sizes with perennial rivers present within the large valleys. One of these rivers passes the site in a distance of less than 500 m. The soils that are present nowadays are Chernozems. They show dark greyish-brown A-horizons of thicknesses between 30 and 50 cm and a texture dominated by silt. Numerous filled crotowinas indicate an intensive bioturbation during the formation of these soils. The Chernozems cover the archaeological record. The variations in thickness of the A-horizon are probably reflecting post-depositional soil erosion processes. Buried soils discovered at lower slope positions below colluvial layers show properties of Cambisols, thus pointing towards a forested past of the surrounding landscape (Kirleis and Dreibrodt, 2016).

    trypillia-maidanetske-mega-city-settlement-old-europe
    The reconstruction of Maidanetske based on geomagnetic survey (modern and from the 1970s by
    Dudkin), with the position of the trenches mentioned in this study.

    Conclusion

    At the site of Maidanetske, the phytolith record from different contexts including multiple houses, was studied, which confirmed cereal cultivation as part of the subsistence economy of the site. Furthermore, phytoliths gave information about wild grasses, whereas dicotyledonous material was scarce. For the house structures cereal byproducts, chaff and straw were identified as material selected for tempering daub for the wall construction. Ash layers in a pit filled with house remains show similar pattern. Daub fragments and pit filling are the most promising archives for further phytolith work on cereals at Trypillia sites. The sediment inside four burnt houses and the areas outside two houses, where also grinding stones were sampled, showed little presence of the remains of final cereal processing, suggesting that either the surfaces were cleaned and the chaff was collected after dehusking, or the cereal processing activity took place somewhere else. Specific archaeological contexts, such as vessels and grinding stones, did not differ much from the control samples from archaeological sediment nearby, suggesting disturbance of the record.(…)

    Related