“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

Dzudzuana, Sidelkino, and the Caucasus contribution to the Pontic-Caspian steppe

hunter-gatherer-pottery

It has been known for a long time that the Caucasus must have hosted many (at least partially) isolated populations, probably helped by geographical boundaries, setting it apart from open Eurasian areas.

David Reich writes in his book the following about India:

The genetic data told a clear story. Around a third of Indian groups experienced population bottlenecks as strong or stronger than the ones that occurred among Finns or Ashkenazi Jews. We later confirmed this finding in an even larger dataset that we collected working with Thangaraj: genetic data from more than 250 jati groups spread throughout India (…)

Rather than an invention of colonialism as Dirks suggested, long-term endogamy as embodied in India today in the institution of caste has been overwhelmingly important for millennia. (…)

The Han Chinese are truly a large population. They have been mixing freely for thousands of years. In contrast, there are few if any Indian groups that are demographically very large, and the degree of genetic differentiation among Indian jati groups living side by side in the same village is typically two to three times higher than the genetic differentiation between northern and southern Europeans. The truth is that India is composed of a large number of small populations.

There is little doubt now, based on findings spanning thousands of years, that the Mesolithic and Neolithic Caucasus hosted various very small populations, even if the ancestral components may be reduced to the few known to date (such as ANE, EHG, AME*, ENA, CHG, and other “deep” ancestral components).

NOTE. I will call the ancestral component of Dzudzuana/Anatolian hunter-gatherers Ancient Middle Easterner (AME), to give a clear idea of its likely extension during the Late Upper Palaeolithic, and to avoid using the more simplistic Dzudzuana, unless it is useful to mention these specific local samples.

dzudzuana-pca
Image modified from Lazaridis et al. (2018), including Caucasus, Don-Volga-Ural, and North Pontic Mesolithic-Neolithic populations. “Ancient West Eurasian population structure. (a) Geographical distribution of key ancient West Eurasian populations. (b) Temporal distribution of key ancient West Eurasian populations (approximate date in ky BP). (c) PCA of key ancient West Eurasians, including additional populations (shown with grey shells), in the space of outgroup f4-statistics (Methods).”

Genetic labs have a strong fixation with ancestry. I guess the use of complex statistical methods gives professionals and laymen alike the feeling of dealing with “Science”, as opposed to academic fields where you have to interpret data. I think language reveals a lot about the way people think, and the fact that ancestral components are called ‘lineages’ – while not wrong per se – is a clear symptom of the lack of interest in the true lineages: Y-DNA haplogroups.

Y-DNA bottlenecks

It has become quite clear that male-biased migrations are often the ones which can be confidently followed for actual population movements and ethnolinguistic identification, at least until the Iron Age. The frequently used Palaeolithic clusters offer a clear example of why ancestry does not represent what some people believe: They merely give a basic idea of sizeable population replacements by distant peoples.

Both concepts are important: sizeable and distant peoples. For example, during the Upper Palaeolithic in Europe there was a sizeable population replacement of the Aurignacian Goyet cluster by the Gravettian Vestonice cluster (probably from populations of far eastern Russia) coupled with the arrival of haplogroup I, although during the thousands of years that this material culture lasted, the previously expanded C1a2 lineages did not disappear, and there were probably different resurgence and admixture events.

Haplogroup I certainly expanded with the Gravettian culture to Iberia, where the Goyet ancestry did not change much – probably because of male-driven migrations -, to the extent that during the Magdalenian expansions haplogroup I expanded with an ancestry closer to Goyet, in what is called a ‘resurge’ of the Goyet cluster – even though there is a clear replacement of male lines.

The Villabruna (WHG) cluster is another good example. It probably spread with haplogroup R1b-L754, which – based on the extra ‘East Asian’ affinity of some samples and on modern samples from the Middle East – came probably from the east through a southern route, and not too long before the expansion of WHG likely from around the Black Sea, although this is still unclear. The finding of haplogroup I in samples of mostly WHG ancestry could confuse people that do not care about timing, sub-structured populations, and gene flow.

palaeolithic-expansions-reich
Image from David Reich’s Who We Are and How We Got Here. Having migrated out of Africa and the Near East, modern human pioneer populations spread throughout Eurasia (1). By at least thirty-nine thousand years ago, one group founded a lineage of European hunter-gatherers that persisted largely uninterrupted for more than twenty thousand years (2). Eventually, groups derived from an eastern branch of this founding population of European huntergatherers spread west (3), displaced previous groups, and were eventually themselves pushed out of northern Europe by the spread of glacial ice, shown at its maximum extent (top right). As the glaciers receded, western Europe was repeopled from the southwest (4) by a population that had managed to persist for tens of thousands of years and was related to an approximately thirty-five-thousand-year old individual from far western Europe. A later human migration, following the first strong warming period, had an even larger impact, with a spread from the southeast (5) that not only transformed the population of western Europe but also homogenized the populations of Europe and the Near East. At a single site—Goyet Caves in Belgium—ancient DNA from individuals spread over twenty thousand years reflects these transformations, with representatives from the Aurignacian, Gravettian, and Magdalenian periods.

NOTE. If you don’t understand why ‘clusters’ that span thousands of years don’t really matter for the many Palaeolithic population expansions that certainly happened among hunter-gatherers in Europe, just take a look at what happened with Bell Beakers expanding from Yamna into western Europe within 500 years.

If we don’t thread carefully when talking about population migrations, these terms are bound to confuse people. Just as the fixation on “steppe ancestry” – which marks the arrival in Chalcolithic Europe of peoples from the Pontic-Caspian region – has confused a lot of researchers to this day.

When I began to write about the Indo-European demic diffusion model, my concern was to find a single spot where a North-West Indo-European proto-language could have expanded from ca. 2000 BC (our most common guesstimate). Based on the 2015 papers, and in spite of their conclusions, I thought it had become clear that Corded Ware was not it, and it was rather Bell Beakers. I assumed that Uralic was spoken to the north (as was the traditional belief), and thus Corded Ware expanded from the forest zone, hence steppe ancestry would also be found there with other R1a lineages.

With the publication of Mathieson et al. (2017) and Olalde et al. (2017), I changed my mind, seeing how “steppe ancestry” did in fact appear quite late, hence it was likely to be the result of very specific population movements, probably directly from the Caucasus. Later, Mathieson published in a revision the sample from Alexandria of hg R1a-M417 (probably R1a-Z645, possibly Z93+), which further supported the idea that the migration of Corded Ware peoples started near the North Pontic forest-steppe (as I included in a the next revision).

The question remains the same I repeated recently, though: where do the extra Caucasus components (i.e. beyond EHG) of Eneolithic Ukraine/Corded Ware and Khvalynsk/Yamna come from?

Steppe ancestry: “EHG” + “CHG”?

About EHG ancestry

From Lazaridis et al. (2018):

Considering 2-way mixtures, we can model Karelia_HG as deriving 34 ± 2.8% of its ancestry from a Villabruna-related source, with the remainder mainly from ANE represented by the AfontovaGora3 (AG3) sample from Lake Baikal ~17kya.

AG3 was likely of haplogroup Q1a (as reported by YFull, see Genetiker), and probably the ANE ancestry found in Eastern Europe accompanied a Palaeolithic migration of Q1a2-M25 (formed ca. 22600 BC, TMRCA ca. 14300 BC).

NOTE. You can read more about the expansion of Q lineages during the Palaeolithic.

Combined with what we know about the Eneolithic Steppe and Caucasus populations – it is likely that ANE ancestry remained the most important component of some of the small ghost populations of the Caucasus until their emergence with the Lola culture.

pca-caucasus-dzudzuana
Image modified from Wang et al. (2018). Samples projected in PCA of 84 modern-day West Eurasian populations (open symbols). Previously known clusters have been marked and referenced. Marked and labelled are the Balkan samples referenced in this text An EHG and a Caucasus ‘clouds’ have been drawn, leaving Pontic-Caspian steppe and derived groups between them. See the original file here. To understand the drawn potential Caucasus Mesolithic cluster, see above the PCA from Lazaridis et al. (2018).

The first sample we have now attributed to the EHG cluster is Sidelkino, from the Samara region (ca. 9300 BC), mtDNA U5a2. In Damgaard et al. (Science 2018), Yamnaya could be modelled as a CHG population related to Kotias Klde (54%) and the remaining from ANE population related to Sidelkino (>46%), with the following split events:

  1. A split event, where the CHG component of Yamnaya splits from KK1. The model inferred this time at 27 kya (though we note the larger models in Sections S2.12.4 and S2.12.5 inferred a more recent split time).
  2. A split event, where the ANE component of Yamnaya splits from Sidelkino. This was inferred at about about 11 kya.
  3. A split event, where the ANE component of Yamnaya splits from Botai. We inferred this to occur 17 kya. Note that this is above the Sidelkino split time, so our model infers Yamnaya to be more closely related to the EHG Sidelkino, as expected.
  4. An ancestral split event between the CHG and ANE ancestral populations. This was inferred to occur around 40 kya.

Other samples classified as of the EHG cluster:

  • Popovo2 (ca. 6250 BC) of hg J1, mtDNA U4d – Po2 and Po4 from the same site (ca. 6550 BC) show continuity of mtDNA.
  • Karelia_HG, from Juzhnii Oleni Ostrov (ca. 6300 BC): I0211/UzOO40 (ca. 6300 BC) of hg J1(xJ1a), mtDNA U4a; and I0061/UzOO74 of hg R1a1(xR1a1a), mtDNA C1
  • UzOO77 and UzOO76 from Juzhnii Oleni Ostrov (ca. 5250 BC) of mtDNA R1b.
  • Samara_HG from Lebyanzhinka (ca. 5600 BC) of hg R1b1a, mtDNA U5a1d.

From the analysis of Lazaridis et al. (2018), we have some details about their admixture:

dzudzuana-admixture-sidelkino
Image modified from Lazaridis et al. (2018). Modeling present-day and ancient West-Eurasians. Mixture proportions computed with qpAdm (Supplementary Information section 4). The proportion of ‘Mbuti’ ancestry represents the total of ‘Deep’ ancestry from lineages that split prior to the split of Ust’Ishim, Tianyuan, and West Eurasians and can include both ‘Basal Eurasian’ and other (e.g., Sub-Saharan African) ancestry. (Left) ‘Conservative’ estimates. Each population 367 cannot be modeled with fewer admixture events than shown. (Right) ‘Speculative’ estimates. The highest number of sources (≤5) with admixture estimates within [0,1] are shown for each population. Some of the admixture proportions are not significantly different from 0 (Supplementary Information section 4).

About Anatolia_Neolithic ancestry

About the enigmatic Anatolia_Neolithic-related ancestry found in Pontic-Caspian steppe samples, this is what Wang et al. (2018) had to say:

We focused on model of mixture of proximal sources such as CHG and Anatolian Chalcolithic for all six groups of the Caucasus cluster (Eneolithic Caucasus, Maykop and Late Makyop, Maykop-Novosvobodnaya, Kura-Araxes, and Dolmen LBA), with admixture proportions on a genetic cline of 40-72% Anatolian Chalcolithic related and 28-60% CHG related (Supplementary Table 7). When we explored Romania_EN and Greece_Neolithic individuals as alternative southeast European sources (30-46% and 36-49%), the CHG proportions increased to 54-70% and 51-64%, respectively. We hypothesize that alternative models, replacing the Anatolian Chalcolithic individual with yet unsampled populations from eastern Anatolia, South Caucasus or northern Mesopotamia, would probably also provide a fit to the data from some of the tested Caucasus groups.

Also:

The first appearance of ‘Near Eastern farmer related ancestry’ in the steppe zone is evident in Steppe Maykop outliers. However, PCA results also suggest that Yamnaya and later groups of the West Eurasian steppe carry some farmer related ancestry as they are slightly shifted towards ‘European Neolithic groups’ in PC2 (Fig. 2D) compared to Eneolithic steppe. This is not the case for the preceding Eneolithic steppe individuals. The tilting cline is also confirmed by admixture f3-statistics, which provide statistically negative values for AG3 as one source and any Anatolian Neolithic related group as a second source

yamnaya-caucasus-dzudzuana
Modified image from Wang et al. (2018). In blue, Yamna-related populations. In red, Corded Ware-related populations, and two elevated Anatolia_Neolithic values in Yamna. Notice how only GAC-related admixture increases the Anatolian_N-related ancestry in the Yamna outlier from Ozero, and the late Yamna sample from Hungary, related to the homogeneous Yamna population. “Supplementary Table 14. P values of rank=3 and admixture proportions in modelling Steppe ancestry populations as a four-way admixture of distal sources EHG, CHG, Anatolian_Neolithic and WHG using 14 outgroups.Left populations: Steppe cluster, EHG, CHG, WHG, Anatolian_Neolithic. Right populations: Mbuti.DG, Ust_Ishim.DG, Kostenki14, MA1, Han.DG, Papuan.DG, Onge.DG, Villabruna, Vestonice16, ElMiron, Ethiopia_4500BP.SG, Karitiana.DG, Natufian, Iran_Ganj_Dareh_Neolithic.”

Detailed exploration via D-statistics in the form of D(EHG, steppe group; X, Mbuti) and D(Samara_Eneolithic, steppe group; X, Mbuti) show significantly negative D values for most of the steppe groups when X is a member of the Caucasus cluster or one of the Levant/Anatolia farmer-related groups (Supplementary Figs. 5 and 6). In addition, we used f- and D-statistics to explore the shared ancestry with Anatolian Neolithic as well as the reciprocal relationship between Anatolian- and Iranian farmer-related ancestry for all groups of our two main clusters and relevant adjacent regions (Supplementary Fig. 4). Here, we observe an increase in farmer-related ancestry (both Anatolian and Iranian) in our Steppe cluster, ranging from Eneolithic steppe to later groups. In Middle/Late Bronze Age groups especially to the north and east we observe a further increase of Anatolian farmer related ancestry consistent with previous studies of the Poltavka, Andronovo, Srubnaya and Sintashta groups and reflecting a different process not especially related to events in the Caucasus.

(…) Surprisingly, we found that a minimum of four streams of ancestry is needed to explain all eleven steppe ancestry groups tested, including previously published ones (Fig. 2; Supplementary Table 12). Importantly, our results show a subtle contribution of both Anatolian farmer-related ancestry and WHG-related ancestry (Fig.4; Supplementary Tables 13 and 14), which was likely contributed through Middle and Late Neolithic farming groups from adjacent regions in the West. The discovery of a quite old AME ancestry has rendered this probably unnecessary, because this admixture from an Anatolian-like ghost population could be driven even by small populations from the Caucasus.

yamna-caucasus-cwc-anatolia-neolithic
Image modified from Wang et al. (2018). Marked are: 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 1128 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 (see also Supplementary Tables 10, 14 and 20).”

NOTE. For a detailed account of the possibilities regarding this differential admixture in the North Pontic area in contrast to the Don-Volga-Ural region, you can read the posts Sredni Stog, Proto-Corded Ware, and their “steppe admixture”, and Corded Ware culture origins: The Final Frontier.

While it is not yet fully clear, the increased Anatolian_Neolithic-like ancestry in Ukraine_Eneolithic samples (see below) makes it unlikely that all such ancestry in Corded Ware groups comes from a GAC-related contribution. It is likely that at least part of it represents contributions from populations of the Caucasus, based on the mostly westward population movements in the steppe from ca. 4600 BC on, including the Suvorovo-Novodanilovka expansion, and especially the Kuban-Maykop expansion during the final Eneolithic into the North Pontic area.

NOTE. Since CHG-like groups from the Caucasus may have combinations of AME and ANE ancestry similar to Yamna (which may thus appear as ‘steppe ancestry’ in the North Pontic area), it is impossible to interpret with precision the following ADMIXTURE graphic:

ukraine-whg-ehg-steppe
Modified image from Mathieson et al. (2018). Supervised ADMIXTURE analysis, modelling each ancient individual (one per row) as a mixture of population clusters constrained to contain northwestern-Anatolian Neolithic (grey), Yamnaya from Samara (yellow), EHG (pink) and WHG (green) populations. Dates in parentheses indicate approximate range of individuals in each population.

North-Eastern Technocomplex

The East Asian contribution to samples from the WHG samples (like Loschbour or La Braña), as specified in Fu et al. (2016), does not seem to be related to Baikal_EN, and appears possibly (in the ADMIXTURE analysis) integrated into he Villabruna component. I guess this implies that the shared alleles with East Asians are quite early, and potentially due to the expansion of R1b-L754 from the East.

It would be interesting to know the specific material culture Sidelkino belonged to – i.e. if it was related to the expansion of the North-Eastern Technocomplex – , and its Y-DNA. The Post-Swiderian expansion into eastern Europe, probably associated with the expansion of R1b-P297 lineages (including R1b-M73, found later in Botai and in Baltic HG) is supposed to have begun during the 11th millennium BC, but migrations to the Urals and beyond are probably concentrated in the 9th millennium, so this sample is possibly slightly early for R1b.

NOTE. User Rozenfeld at Anthrogenica posted this, which I think is interesting (in case anyone wants to try a Y-SNP call):

there is something strange with Sidelkino EHG: first, its archaeological context is not described in the supplementary. Second, its sex is not listed in the supplementary tables. Third, after looking for info about this sample, I found that: “Сиделькино-3. Для снятия вопроса о половой принадлежности индивида была проведена генетическая экспертиза, выявившая принадлежность останков мужчине.”(translation: Sidelkino-3. To resolve the question about sex of the remains, the genetic analysis was conducted, which showed that remains belonged to male), source: http://static.iea.ras.ru/books/7487_Traditsii.pdf

So either they haven’t mentioned his Y-DNA in the paper for some reason, or there are more than one Sidelkino sample and the male one has not yet been published. The coverage of the Sidelkino sample from the paper is 2.9, more than enough to tell Y-DNA haplogroup.

zaliznyak-post-swiderian
The map of spreading of Post-Swiderian and Post-Krasnosillian sites in Mesolithic of Eastern Europe in the 8th millennia BC. From Zaliznyak (see here).

My speculative guess right now about specific population movements in far eastern Europe, based on the few data we have:

  • The expansion of the North-Eastern Technocomplex first around the 9th millennium BC, most likely expanded R1b-P279 ca. 11300 BC, judging by its TMRCA, with both R1b-M73 (TMRCA 5300) and R1b-M269 (TMRCA 4400 BC) info (with extra El Mirón ancestry) back, and thus Eurasiatic.
  • The expansion of haplogroup J1 to the north may have happened before or after the R1b-P279 expansion. Judging by the increase in AG3-related ancestry near Karelia compared to Baltic_HG, it is possible that it expanded just after R1b-P279 (hence possibly J1-Y6304? TMRCA 9700 BC). Its long-lasting presence in the Caucasus is supported by the Satsurblia (ca. 11300 BC) and the Dolmen BA (ca. 1300 BC) samples.
  • The expansion of R1a-M17 ca. 6600 BC is still likely to have happened from the east, based on the R1a-M17 samples found in Baikalic cultures slightly later (ca. 5300 BC). The presence of elevated Baikal_EN ancestry in Karelia HG and in Samara HG, and the finding of R1a-M417 samples in the Forest Zone after the Mesolithic suggests a connection with the expansion of Hunter-Gatherer pottery, from the Elshanka culture in the Samara region northward into the Forset Zone and westward into the North Pontic area.
  • The expansion of R1b-M73 ca. 5300 BC is likely to be associated with the emergence of a group east of the Urals (related to the later Botai culture, and potentially Pre-Yukaghir). Its presence in a Narva sample from Donkalnis (ca. 5200 BC) suggest either an early split and spread of both R1b-P297 lineages (M73 and M269) through Eastern Europe, or maybe a back-migration with hunter-gatherer pottery.
  • R1b-M269 spread successfully ca. 4400 BC (and R1b-L23 ca. 4100 BC, both based on TMRCA), and this successful expansion is probably to be associated with the Khvalynsk-Novodanilovka expansion. We already know that Samara_HG ca. 5600 was R1b1a, so it is likely that R1b-M269 appeared (or ‘resurged’) in the Volga-Ural region shortly after the expansion of R1a-M17, whose expansion through the region may be inferred by the additional AG3 and Baikal_EN ancestry. Interesting from Samara_HG compared to the previous Sidelkino sample is the introduction of more El Mirón-related ancestry, typical of WHG populations (and thus proper of Baltic groups).

NOTE. The TMRCA dates are obviously gross approximations, because a) the actual rate of mutation is unknown and b) TMRCA estimates are based on the convergence of lineages that survived. The potential finding of R1a-Z645 (possibly Z93+) in Ukraine Eneolithic (ca. 4000 BC), and the potential finding of R1b-L23 in Khvalynsk ca. 4250 BC complicates things further, in terms of dates and origins of any subclade.

The question thus remains as it was long ago: did R1b-M269 lineages expand (‘return’) from the east, near the Urals, or directly from the north? Were they already near Samara at the same time as the expansion of hunter-gatherer pottery, and were not much affected by it? Or did they ‘resurge’ from populations admixed with Caucasus-related ancestry after the expansion of R1a-M17 with this pottery (since there are different stepped expansions from the Samara region)? We could even ask, did R1a-M17 really expand from the east, i.e. are the dates on Baikalic subclades from Moussa et al. (2016) reliable? Or did R1a-M17 expand from some pockets in the Pontic-Caspian steppe, taking over the expansion of HG pottery at some point?

hunger-gatherer-pottery
Early Neolithic cultures in eastern and central Europe: 1–Yelshanian; 2–North Caspian; 3–Rakushechnyj Yar; 4–Surskian; 5–Dnieper-Donetsian; 6– Bug-Dniesterian; 7–Upper Volga; 8–Narvian; 9–Linear Pottery. White arrows: expansion of early farming; black arrows: spread of pottery-making traditions. From Dolukhanov et al. (2009).

Maglemose-related migrations

The most interesting aspect from the new paper (regarding Indo-Uralic migrations) is that Ancestral Middle Easterner ancestry will probably be a better proxy for the Anatolia_Neolithic component found in Ukraine Mesolithic to Eneolithic, and possibly also for some of the “more CHG-like” component found among Pontic-Caspian steppe populations, all likely derived from different admixture events with groups from the Caucasus.

NOTE. Even the supposed gene flow of Neolithic Iranian ancestry into the Caucasus can be put into question, since that means possibly a Dzudzuana-like population with greater “deep ancestry” proportion than the one found in CHG, which may still be found within the Caucasus.

If it was not clear already that following ‘steppe ancestry’ wherever it appears is a rather lame way of following Indo-European migrations, every single sample from the Caucasus and their admixture with Pontic-Caspian steppe populations will probably show that “steppe ancestry” is in fact formed by a variety of steppe-related ancestral components, impossible to follow coherently with a single population. Exactly what is happening already with the Siberian ancestry.

If the paper on the Dzudzuana samples has shown something, is that the expansion of an ANE-like population shook the entire Caucasus area up to the Zagros Mountains, creating this ANE – AME cline that are CHG and Iran_N, with further contributions of “deep ancestries” (probably from the south) complicating the picture further.

If this happens with few known samples, and we know of an ANE-like ghost population in the Caucasus (appearing later in the Lola culture), we can already guess that the often repeated “CHG component” found in Ukraine_Eneolithic and Khvalynsk will not be the same (except the part mediated by the Novodanilovka expansion).

This ANE-like expansion happened probably in the Late Upper Palaeolithic, and reached Northern Europe probably after the expansion of the Villabruna cluster (ca. 12000 BC), judging by the advance of AG3-like and ENA-like ancestry in later WHG samples.

The population movements during the Mesolithic and Early Neolithic in the North Pontic area are quite complicated: the extra AME ancestry is probably connected to the admixture with populations from the Caucasus, while the close similarity of Ukraine populations with Scandinavian ones (with an increase in Villabruna ancestry from Mesolithic to Neolithic samples), probably reveal population movements related to the expansion of Maglemose-related groups.

maglemose-mesolithic
Etno-cultural situation in Central and Eastern Europe in the Late Mesolithic — Early Neolithic (VI—V Mill. BC) (after Конча 2004: 201, карта 1; made after ideas by L. L. Zaliznyak). Legend: 1 — Maglemose circle in the VII Mill. BC (after Gr. Clark); 2—7 — Mesolithic cultures of the Post-Maglemose tradition, VI Mill. BC (after S. Kozłowsky, L. L. Zaliznyak): 2 — de Leyen-Wartena; 3 — Oldesloe — Godenaa; 4 — Chojnice — Peńki; 5 — Janisłavice; 6 — finds of Janisłavice artefacts outside of the main area; 7 — Donets culture; 8 — directions of the settling of Janisłavice people (after S. Kozłowsky and L. L. Zaliznyak); 9 — the south border of Mesolithic and Early Neolithic cultures of post-Swidrian and post-Arensburgian traditions; 10 — northern border of settlement of the Balkan-Danubian farmers; 11 — Bug- Dniester culture; 12 — Neolithic cultures emerged on the ethno-cultural basis of post-Maglemose: Э — Ertebölle-Ellerbeck, Н — Neman, Д — Dnieper-Donets, М — Mariupol (western variants). From Klein (2017).

These Maglemose-related groups were probably migrants from the north-west, originally from the Northern European Plains, who occupied the previous Swiderian territory, and then expanded into the North Pontic area. The overwhelming presence of I2a (likely all I2a2a1b1b) lineages in Ukraine Neolithic supports this migration.

The likely picture of Mesolithic-Neolithic migrations in the North Pontic area right now is then:

  1. Expansion of R1a-M459 from the east ca. 12000 BC – probably coupled with AG3 and also some Baikal_EN ancestry. First sample is I1819 from Vasilievka (ca. 8700 BC), another is from Dereivka ca. 6900 BC.
  2. Expansion of R1b-V88 from the Balkans in the west ca. 9700 BC, based on its TMRCA and also the Balkan hunter-gatherer population overwhemingly of this haplogroup from the 10th millennium until the Neolithic. First sample is I1734 from Vasilievka (ca. 7252 BC), which suggests that it replaced the male population there, based on their similar EHG-like adxmixture (and lack of sizeable WHG increase), and shared mtDNA U5b2, U5a2.
  3. Expansion of I2a-Y5606 probably ca. 6800 based on its TMRCA with Janislawice culture. Supporting this is the increase in WHG contribution to Neolithic samples, including the spread of U4 subclades compared to the previous period.
  4. Expansion of R1a-M17 starting probably ca. 6600 BC in the east (see above).

NOTE. The first sample of haplogroup I appears in the Mesolithic: I1763 (ca. 8100 BC) of haplogroup I2a1, probably related to an older Upper Palaeolithic expansion.

janislawice
Distribution of archeological cultures in the North Pontic Region during the Mesolithic (7th – 6th millennium BCE). Dotted, dashed and solid lines with corresponding arrows indicate alternative models of the spread of the Grebenyky culture groups. (After Bryuako IV., Samojlova TL., Eds, Drevnie kul’tury Severo-­‐Zapadnogo Prichernomor’ya, Odessa: SMIL, 2013.) Nikitin – Ivanova 2017.

Conclusion

It is becoming more and more clear with each new paper that – unless the number of very ancient samples increases – the use of Y-chromosome haplogroups remains one of the most important tools for academics; this is especially so in the steppes, in light of the diversity found in populations from the Caucasus. A clear example comes from the Yamna – Corded Ware similarities:

After the publication of the 2015 papers, it was likely that Yamna expanded with haplogroup R1b-L23, but it has only become crystal clear that Yamna expanded through the steppes into Bell Beakers, now that we have data about the strict genetic homogeneity of the whole Yamna population from west to east (including Afanasevo), in contrast with contemporary Corded Ware peoples which expanded from a different forest-steppe population.

The presence of haplogroups Q and R1a-M459 (xM17) in Khvalynsk along with a R1b1a sample, which some interpreted as being akin to modern ‘mixed’ populations in the past, is likely to point instead to a period of Khvalynsk-Novodanilovka expansion with R1b-M269, where different small populations from the steppe were being integrated into the common Khvalynsk stock, but where differences are seen in material culture surrounding their burials, as supported by the finding of R1b1 in the Kuban area already in the first half of the 5th millennium. The case would be similar to the early ‘mixed’ Icelandic population.

Only after the emergence of the Samara culture (in the second half of the 6th millennium BC), with a sample of haplogroup R1b1a, starts then the obvious connection with Early Proto-Indo-Europeans; and only after the appearance of late Sredni Stog and haplogroup R1a-M417 (ca. 4000 BC) is its connection with Uralic also clear. In previous population movements, I think more haplogroups were involved in migrations of small groups, and only some communities among them were eventually successful, expanding to be dominant, creating ever growing cultures during their expansions.

Indeed, if you think in terms of Uralic and Indo-European just as converging languages, and forget their potential genetic connection, then the genetic + linguistic picture becomes simplified, and the upper frontier of the 6th millennium BC with a division North Pontic (Mariupol) vs. Volga-Ural (Samara) is enough. However, tracing their movements backwards – with cultural expansions from west to east (with the expansion of farming), and earlier east to west (with hunter-gatherer pottery), and still earlier west to east (with the north-eastern technocomplex), offers an interesting way to prove their potential connection to macrofamilies, at least in terms of population movements.

corded-ware-uralic-qpgraph
Modified image from Tambets et al. (2018) Proportions of ancestral components in studied European and Siberian populations and the tested qpGraph model. a The qpGraph model fitting the data for the tested populations. Colour codes for the terminal nodes: pink—modern populations (‘Population X’ refers to test population) and yellow—ancient populations (aDNA samples and their pools). Nodes coloured other than pink or yellow are hypothetical intermediate populations. We putatively named nodes which we used as admixture sources using the main recipient among known populations. The colours of intermediate nodes on the qpGraph model match those on the admixture proportions panel. The NeolL (Neolithic Levant) ancestry selected in this qpGraph is likely to correspond (at least in part) to a specific Dzudzuana-like component present in the CHG-like population that admixed in the North Pontic area.

I am quite convinced right now that it would be possible to connect the expansion of R1b-L754 subclades with a speculative Nostratic (given the R1b-V88 connection with Afroasiatic, and the obvious connection of R1b-L297 with Eurasiatic). Paradoxically, the connection of an Indo-Uralic community in the steppes (after the separation of Yukaghir) with any lineage expansion (R1a-M17, R1b-M269, or even Q, I or J1) seems somehow blurrier than one year ago, possibly just because there are too many open possibilities.

David Reich says about the admixture with Neanderthals, which he helped discover:

At the conclusion of the Neanderthal genome project, I am still amazed by the surprises we encountered. Having found the first evidence of interbreeding between Neanderthals and modern humans, I continue to have nightmares that the finding is some kind of mistake. But the data are sternly consistent: the evidence for Neanderthal interbreeding turns out to be everywhere. As we continue to do genetic work, we keep encountering more and more patterns that reflect the extraordinary impact this interbreeding has had on the genomes of people living today.

I think this is a shared feeling among many of us who have made proposals about anything, to fear that we have made a gross, evident mistake, and constantly look for flaws. However, it seems to me that geneticists are more preoccupied with being wrong in their developed statistical methods, in the theoretical models they are creating, and not so much about errors in the true ancient ethnolinguistic picture human population genetics is (at least in theory) concerned about. Their publications are, after all, constantly associating genetic finds with cultures and (whenever possible) languages, so this aspect of their research should not be taken lightly.

Seeing how David Anthony or Razib Khan (among many others) have changed their previously preferred migration models as new data was published, and they continue to be respected in their own fields, I guess we can be confident that professionals with integrity are going to accept whatever new picture appears. While I don’t think that genetic finds can change what we can reconstruct with comparative grammar, I am also ready to revise guesstimates and routes of expansion of certain dialects if R1a-Z645 is shown to have accompanied Late Proto-Indo-Europeans during their expansion with Yamna, and later integrated somehow with Corded Ware.

However, taking into account the obsession of some with an ancestral, uninterrupted R1a—Indo-European association, and the lack of actual political repercussion of Neanderthal admixture, I think the most common nightmare that all genetic researchers should be worried about is to keep inflating this “Yamnaya ancestry”-based hornet’s nest, which has been constantly stirred up for the past two years, by rejecting it – or, rather, specifying it into its true complex nature.

This succession of corrections and redefinitions, coupled with the distinct Y-DNA bottleneck of each steppe population, will eventually lead to a completely different ethnolinguistic picture of the Pontic-Caspian region during the Eneolithic, which is likely to eventually piss off not only reasonable academics stubbornly attached to the CWC-IE idea, but also a part of those interested in daydreaming about their patrilineal ancestors.

Sometimes it’s better to just rip off the band-aid once and for all…

Featured image from The oldest pottery in hunter-gatherer communitiesand models of Neolithisation of Eastern Europe (2015), by Andrey Mazurkevich and Ekaterina Dolbunova.

Related

Interesting is today’s post in Ancient DNA Era: Is Male-driven Genetic Replacement always meaning Language-shift?

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

Recent archaeological finds near Indo-European and Uralic homelands

eneolithic-forest-zone

The latest publication of Documenta Praehistorica, vol. 44 (2017) is a delight for anyone interested in Indo-European and Uralic studies, whether from a linguistic, archaeological, anthropological, or genetic point of view. Articles are freely downloadable from the website.

The following is a selection of articles I deem more interesting, but almost all are.

On the Corded Ware culture

Do 14C dates always turn into an absolute chronology? The case of the Middle Neolithic in western Lesser Poland, by Marek Novak:

In the late 5th, 4th, and early 3rd millennia BC, different archaeological units are visible in western Lesser Poland. According to traditional views, local branches of the late Lengyel-Polgár complex, the Funnel Beaker culture, and the Baden phenomena overlap chronologically in great measure. The results of investigations done with new radiocarbon dating show that in some cases a discrete mode and linearity of cultural transformation is recommended. The study demonstrates that extreme approaches in which we either approve only those dates which fit with our concepts or accept with no reservation all dates as such are incorrect.

baden-corded-ware-culture
Territory of western Lesser Poland and the main archaeological units in the late 5th, 4th and early 3rd millennia BC: 1 borders of the area discussed in the paper; 2 sites of the Lublin-Volhynian culture; 3 the Wyciąże-Złotniki group; 4 the Funnel Beaker culture (a dense settlement typical of ‘loess’ upland; b more dispersed settlement typical of foothills, alluvial plains/basins and ‘jurassic zones; c highly dispersed settlement typical mainly of mountainous zone); 5 sites with the Wyciąże/Niedźwiedź materials; 6 the Baden culture, 7 the Beaker/Baden assemblages; 8 Corded Ware culture (a relatively dense settlement typical mainly of ‘loess’ upland; b highly dispersed settlement typical of other ecological zones).

This article brings new data against David Anthony’s new IECWT model, suggesting later dates for the Corded Ware Culture group of Lesser Poland, and thus an earlier origin of their nomadic herders in the steppe, forest-steppe or forest zone to the east and south-east.

On the Pontic-Caspian steppe and forest-steppe

First isotope analysis and new radiocarbon dating of Trypillia (Tripolye) farmers from Verteba Cave, Bilche Zolote, Ukraine, by Lillie et al.:

This paper presents an analysis of human and animal remains from Verteba cave, near Bilche Zolote, western Ukraine. This study was prompted by a paucity of direct dates on this material and the need to contextualise these remains in relation both to the transition from hunting and gathering to farming in Ukraine, and their specific place within the Cucuteni-Trypillia culture sequence. The new absolute dating places the remains studied here in Trypillia stages BII/CI at c. 3900–3500 cal BC, with one individual now redated to the Early Scythian period. As such, these finds are even more exceptional than previously assumed, being some of the earliest discovered for this culture. The isotope analyses indicate that these individuals are local to the region, with the dietary stable isotopes indicating a C3 terrestrial diet for the Trypillia-period humans analysed. The Scythian period individual has δ13C ratios indicative of either c. 50% marine, or alternatively C4 plant inputs into the diet, despite δ18O and 87Sr/86Sr ratios that are comparable to the other individuals studied.

trypillia-steppe-forest-steppe
Map showing the extent of the Trypillia culture of Ukraine and
neighbouring countries, key sites and the location of Verteba Cave ©WAERC
University of Hull.

New data on one of the cultures that was very likely a close neighbour of Corded Ware peoples.

Chronology of Neolithic sites in the forest-steppe area of the Don River, by Smolyaninov, Skorobogatov, and Surkov:

The first ceramic complexes appeared in the forest-steppe and forest zones of Eastern Europe at the end of the 7th–5th millennium BC. They existed until the first half of the 5th millennium BC in the Don River basin. All these first ceramic traditions had common features and also local particularities. Regional cultures, distinguished nowadays on the basis of these local particularities, include the Karamyshevskaya and Middle Don cultures, as well pottery of a new type found at sites on the Middle Don River (Cherkasskaya 3 and Cherkasskaya 5 sites).

Radiocarbon chronology of Neolithic in the Lower Don and North-eastern Azov Sea, by Tsybryi et al.:

So far, four different cultural-chronological groups of sites have been identified in the North-eastern Azov Sea and Lower Don River areas, including sites of the Rakushechny Yar culture, Matveev Kurgan culture, Donets culture, and sites of the Caspian-Ciscaucasian region. An analysis of all known dates, as well as the contexts and stratigraphies of the sites, allowed us to form a new perspective of the chronology of southern Russia, to revise the chronology of this region, and change the concept of unreliability of dates for this area.

On the Forest Zone

The past in the past in the mortuary practice of hunter-gatherers: an example from a settlement and cemetery site in northern Latvia, by Lars Olof Georg Larsson:

During excavations of burials at Zvejnieki in northern Latvia, it transpired that the grave fill included occupation material brought to the grave. It contained tools of a type that could not be contemporaneous with the grave. This is confirmed by the dating of bone tools and other bone finds in the fill. The fill was taken from an older settlement site a short distance away. The fill also included skeletal parts of humans whose graves had been destroyed with the digging of the grave for a double burial. This provides an interesting view of the mortuary practice of hunter-gatherers and an insight into the use of the past in the past.

zvejnieki-site
The Zvejnieki site with the location of the burial ground, the settlements,
the farmhouse on the site and the gravel pit.

I keep expecting that more information is given regarding the important sample labelled “Late Neolithic/Corded Ware Culture” from Zvejnieki ca. 2880 BC. It seems too early for the Corded Ware culture in the region, clusters too close to steppe samples, and the information on it from genetic papers is so scarce… My ad hoc explanation of these data – as a product of recent exogamy from Eastern Yamna -, while possibly enough to explain one sample, is not satisfying without further data, so we need to have more samples from the region to have a clearer picture of what happened there and when. Another possibility is a new classification of the sample, compatible with later migration events (a later date of the sample would explain a lot). Anyway, this article won’t reveal anything about this matter, but is interesting for other, earlier samples from the cemetery.

Other articles on the Forest Zone include:

Other articles include studies on Neolithic sites, potentially relevant for Indo-European migrations, such as Anatolia, Greece, southern or south-eastern sites in Europe. Check it out!

Related: