Updates to ASoSaH: new maps, updated PCA, and added newest research papers

steppe-ancestry-cut

The title says it all. I have used some free time to update the series A Song of Sheep and Horses:

I basically added information from the latest papers published, which (luckily enough for me) haven’t been too many, and I have added images to illustrate certain sections.

I have updated the PCAs by including North Caucasus samples from Wang et al. (2018), whose position I could only infer for older versions from previously published PCA graphs.

pca-steppe-eneolithic-early
PCA of ancient and modern Eurasian samples. Early Eneolithic admixture events in the steppe drawn.

I have also added to the supplementary materials the “Tip of the Iceberg” R1b tree by Mike Walsh from the FTDNA R1b group, with permission, because some relevant genetic sections are centered on the evolution of R1b lineages, and the reader can get easily lost with so many subclades.

I have also updated maps, including some of the Y-DNA ones, and managed to finish two new maps I was working on, and I added them to the supplementary materials and to the menu above:

One on Yamna kurgans in Hungary, coupled with contemporaneous sites of Baden-Boleráz or Kostolac cultures:

burials-yamnaya-hungary
Map of attested Yamnaya pit-grave burials in the Hungarian plains; superimposed in shades of blue are common areas covered by floods before the extensive controls imposed in the 19th century; in orange, cumulative thickness of sand, unfavourable loamy sand layer. Marked are settlements/findings of Boleráz (ca. 3500 BC on), Baden (until ca. 2800 BC), Kostolac (precise dates unknown), and Yamna kurgans (from ca. 3100/3000 BC on).

Another one on Steppe ancestry expansion, with a tentative distribution of “steppe ancestry” divided into that of Sredni Stog/Corded Ware origin vs. that of Repin/Yamna origin, a difference that has been known for quite some time already.

It is tentative because there hasn’t been any professional study or amateur attempt to date to differentiate both “steppe ancestries” in Yamna, and especially in Bell Beakers. So much for the call of professional geneticists since 2018 (see here and here) and archaeologists since 2017 (see e.g. here and here) to distinguish fine-scale population structure to be able to follow neighbouring populations which expanded with different archaeological (and thus ethnolinguistic) groups.

steppe-ancestry-corded-ware
Tentative map of fine-scale population structure during steppe-related expansions (ca. 3500–2000 BC), including Repin–Yamna–Bell Beaker/Balkans and Sredni Stog–Corded Ware groups. Data based on published samples and pairwise comparisons tested to date. Notice that the potential admixture of expanding Repin/Early Yamna settlers in the North Pontic area with the late Sredni Stog population (and thus Sredni Stog-related ancestry in Yamna) has been omitted for simplicity purposes, assuming thus a homogeneous Yamna vs. Corded Ware ancestry.

I think both maps are especially important today, given the current Nordicist reactionary trends arguing (yet again) for an origin of Indo-Europeans in The North™, now based on the Fearsome Tisza River hypothesis, on cephalic index values, and a few pairwise comparisons – i.e. an absolutely no-nonsense approach to the Indo-European question (LOL). At least I get to relax and sit this year out just observing how other people bury themselves and their beloved “steppe ancestry=IE” under so many new pet theories…

NOTE. Not that there is anything wrong with a northern origin of North-West Indo-European from a linguistic point of view, as I commented recently – after all, a Corded Ware origin would roughly fit the linguistic guesstimates, unlike the proposed ancestral origins in Anatolia or India. The problem is that, like many other fringe theories, it is today just based on tradition, or (even worse) ethnic, political, or personal desires, and it doesn’t make sense when all findings from disciplines involved in the Indo-European and Uralic questions are combined.

steppe-ancestry-modern-populations
Simple ancestry percentages in modern populations. Recent image by Iain Mathieson 2019 (min. 5.57). A simplistic “Steppe ancestry” defining Indo-European speakers…? Sure.

Within 20 or 30 years, when genetic genealogists (or amateur geneticists, or however you want to call them) ask why we had the opportunity since 2015 to sample as many Hungarian Yamnaya individuals as possible and we didn’t, when it is clear that the number of unscathed kurgans is diminishing every year (from an estimated 4,000 in the 20th century, of the original tens of thousands, to less than 1,500 today) the answer will not be “because this or that archaeologist or linguist was a dilettante or a charlatan‘, as they usually describe academics they dislike.

It will be precisely because the very same genetic genealogists – supposedly interested today in the origin of R1b-L151 and/or genetic marker associated with North-West Indo-Europeans – are obsessed with finding them anywhere else but for Hungary, and prefer to use their money and time to play with a few statistical tools within a biased framework of flawed assumptions and study designs, obtaining absurd results and accepting far-fetched interpretations of them, to be told exactly what they want to hear: be it the Franco-Cantabrian homeland, the Dutch or Moravian Beaker from CWC homeland, the Maykop homeland, or the Moon homeland.

Poetic justice this heritage destruction, whose indirect causes will remain written in Internet archives for everyone to see, as a good lesson for future generations.

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 (I): Y-DNA haplogroups among Indo-Europeans (apart from R1b-L23)

eneolithic-early-admixture-steppe-ancestry

Given my reduced free time in these months, I have decided to keep updating the text on Indo-European and Uralic migrations and/or this blog, simultaneously or alternatively, to make the most out of the time I can dedicate to this. I will add the different ‘A Song of Sheep and Horses (ASoSaH) reread’ posts to the original post announcing the books. I would be especially interested in comments and corrections to the book chapters rather than the posts, but any comments are welcome (including in the forum, where comments are more likely to stick).

This is mainly a reread of iv.2. Indo-Anatolians and vi.1. Disintegrating Indo-Europeans.

Indo-Anatolians and Late Indo-Europeans

I have often written about R1b-L23 as the majority haplogroup among Late Proto-Indo-Europeans (see my predictions for 2018 and my summary of 2018), but always expected other haplogroups to pop up somewhere along the way, in Khvalynsk, in Repin, in Yamna, and in Bell Beakers (see e.g. the post on common fallacies of R1a/IE-fans).

Luckily enough – for those of us who want precise answers to our previous infinite models of Indo-European language expansions (viz. GAC-associated expansion, IE-speaking Old Europe, Anatolian homeland, Iran homeland, Maykop as Proto-Anatolian, Palaeolithic Continuity Theory, Celtic in the Atlantic façade, etc.) – the situation has been more clear-cut than expected: it turns out that, especially during population expansions, acute Y-chromosome bottlenecks were very common in the past, at least until the Iron Age.

Khvalynsk and Repin-Yamna expansions were no different, and that seems quite natural in hindsight, given the strong familial ties and aversion to foreigners proper of the Late Proto-Indo-European society and culture – probably not really that different from other contemporary societies, like the neighbouring Late Proto-Uralic or Trypillian ones.

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

Y-DNA haplogroups

During the expansion of early Khvalynsk, the most likely Indo-Anatolian culture, the society of the Don-Volga area was probably made up of different lineages including R1b-V1636, R1b-M269, R1a-YP1272, Q1a-M25, and I2a-L699 (and possibly some R1b-V88?), a variability possibly greater than that of the contemporary north Pontic area, probably a sign of this region being a sink of different east and west migrations from steppe and forest areas.

During its expansion, the Khvalynsk society saw its haplogroup variability reduced, as evidenced by the succeeding expansive Repin culture:

Afanasevo, representing Pre-Tocharian (the earliest Late PIE dialect to branch off), expanded with R1b-L23 – especially R1b-Z2103 – lineages, while early Yamna expanded with R1b-L23 and I2a-L699 lineages, which suggests that these are the main haplogroups that survived the Y-DNA bottleneck undergone during the Khvalynsk expansion, and especially later during the late Repin expansion. Nevertheless, other old haplogroups might still pop up during the Repin and early Yamna period, such as the R1b-V1636 sample from Yamna in the Northern Caucasus.

It is still unclear if R1b-L23 sister clade R1b-PF7562 (formed ca. 4400 BC, TMRCA ca. 3400 BC), prevalent among modern Albanians, expanded with Yamna migrants, or if it was part of an earlier expansion of R1b-M269 into the Balkans, and represent thus Indo-Anatolian speakers who later hitchhiked the expansion of the Late PIE language from the north or west Pontic area. The early TMRCA seems to suggest an association with Repin (and therefore Yamna), rather than later movements in the Balkans.

chalcolithic-early-y-dna
Y-DNA samples from Yamnaya and neighbouring cultures. See full version here.

‘Yamnaya’ or ‘steppe’ ancestry?

After the early years when population genetics relied mainly on modern Y-DNA haplogroups, geneticists and amateurs have been recently playing around with testing “ancestry percentages”, based on newly developed free statistical tools, which offer obviously just one among many types of data to achieve a proper interpretation of the past.

Today we have quite a lot Y-DNA haplogroups reported for ancient samples of more recent prehistoric periods, and they seem to offer (at least since the 2015 papers, but more evidently since the 2018 papers on Bell Beakers and Europeans, Corded Ware, or Fennoscandia among others) the most straightforward interpretation of all results published in population genomics research.

NOTE. The finding of a specific type of ancestry in one isolated 40,000-year-old sample from Tianyuan can offer very interesting information on potential population movements to the region. However, the identification of ethnolinguistic communities and their migrations among neighbouring groups in Neolithic or Bronze Age groups is evidently not that simple.

PCA-caucasus-steppe-all
Yamnaya (Indo-European peoples) and their evolution in the steppes, together with North Pontic (eventually Uralic) peoples.Notice how little Indo-European ancestry changes from Khvalynsk (Indo-Anatolian) to Yamna Hungary (North-West Indo-Europeans) Image modified from Wang et al. (2018). See more on the evolution of “steppe ancestry”.

It is becoming more and more clear with each paper that the true “Yamnaya ancestry” – not the originally described one – was in fact associated with Indo-Europeans (see more on the very Yamnaya-like Yamna Hungary and early East Bell Beaker R1b samples, all of quite similar ancestry and PCA cluster before their further admixture with EEF- and CWC-like groups).

The so-called “steppe ancestry”, on the other hand, reflects the contribution of a Northern Caucasus-related ancestry to expanding Khvalynsk settlers, who spread through the steppes more than a thousand years before the expansion of Late Proto-Indo-Europeans with late Repin, and can thus be found among different groups related to the Pontic-Caspian steppes (see more on the emergence and evolution of “steppe ancestry”).

In fact, after the Yamna/Indo-European and Corded Ware/Uralic expansions, it is more likely to find “steppe ancestry” to the north and east in territories traditionally associated with Uralic languages, whereas to the south and west – i.e. in territories traditionally associated with Indo-European languages – it is more likely to find “EEF ancestry” with diminished “steppe ancestry”, among peoples patrilineally descended from Yamna settlers.

Y-DNA haplogroups, the only uniparental markers (see exceptions in mtDNA inheritance) – unlike ancestry percentages based on the comparison of a few samples and flawed study designs – do not admix, do not change, and therefore they do not lend themselves to infinite pet theories (see e.g. what David Reich has to say about R1b-P312 in Iberia directly derived from Yamna migrants in spite of their predominant EEF ancestry): their cultural continuity can only be challenged with carefully threaded linguistic, archaeological, and genetic data.

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

Iron Age bottleneck of the Proto-Fennic population in Estonia

tarand-graves-estonia-early-late

Demographic data and figures derived from Estonian Iron Age graves, by Raili Allmäe, Papers on Anthropology (2018) 27(2).

Interesting excerpts (emphasis mine):

Introduction

Inhumation and cremation burials were both common in Iron Age Estonia; however, the pattern which burials were prevalent has regional as well temporal peculiarities. In Estonia, cremation burials appear in the Late Bronze Age (1100–500 BC), for example, in stone-cist graves and ship graves, although inhumation is still characteristic of the period [28, 18]. Cremation burials have occasionally been found beneath the Late Bronze Age cists and the Early Iron Age (500 BC–450 AD) tarand graves [30, 28]. In south-eastern Estonia, including Setumaa, the tradition to bury cremated human remains in pit graves also appeared in the Bronze Age and lasted during the Pre-Roman period (500 BC–50 AD) and the Roman Iron Age (50–450 AD), and even up to the medieval times [30, 23, 33, 9]. During the Early Iron Age, cremations appear in cairn graves and have occasionally been found in many Pre-Roman early tarand graves where they appear with inhumations [27, 28, 19, 20, 21, 22, 24]. In Roman Iron Age tarand graves, cremation as well inhumation were practiced [28, 36, 37]; however, cremation was the prevailing burial practice during the Roman Iron Age, for example, in tarand graves in south-eastern Estonia [30, 28]. Roman Iron Age (50 AD–450 AD) burial sites have not been found in continental west Estonia [28, 38]). At the beginning of the Middle Iron Age (450–800 AD), burial sites, for example stone graves without a formal structure, like Maidla I, Lihula and Ehmja ‘Varetemägi’, appear in Läänemaa, west Estonia; in these graves cremations as well inhumations have been found [39, 48]. Like underground cremation burial, the stone grave without a formal structure was the most common grave type during the Late Iron Age (800– 1200 AD) in west Estonia [39, 35, 48]. In south-eastern and eastern Estonia, sand barrows with cremation burials appeared at the beginning of the Middle Iron Age. Cremation barrows are attributed to the Culture of Long Barrows and are most numerous in the villages Laossina and Rõsna in northern Setomaa, on the western shore of Lake Peipsi [8, 48]. Apparently during the Iron Age, the practiced burial customs varied in Estonia.

cist-grave-tarand
Typical prehistoric Estonian graves. Top: Cist-graves common during the Bronze Age, by Terker (GNU FDL 1.2). Bottom: Tarand graves of the Iron Age, by Marika Mägi (2017)

Abstract:

Three Iron Age cremation graves from south-eastern Estonia and four graves including cremations as well inhumations from western Estonia were analysed by osteological and palaeodemographic methods in order to estimate the age and sex composition of burial sites, and to propose some possible demographic figures and models for living communities.

The crude birth/death rate estimated on the basis of juvenility indices varied between 55.1‰ and 60.0‰ (58.5‰ on average) at Rõsna village in south-eastern Estonia in the Middle Iron Age. The birth/death rates based on juvenility indices for south eastern graves varied to a greater degree. The estimated crude birth/death rate was somewhat lower (38.9‰) at Maidla in the Late Iron Age and extremely high (92.1‰) at Maidla in the Middle Iron Age, which indicates an unsustainable community. High crude birth/death rates are also characteristic of Poanse tarand graves from the Pre-Roman Iron Age – 92.3‰ for the 1st grave and 69.6‰for the 2nd grave. Expectedly, newborn life expectancies are extremely low in both communities – 10.8 years at Poanse I and 14.4 years at Poanse II. Most likely, both Maidla I and Poanse I were unsustainable communities.

tarand-graves-estonia
Locations of the investigated Estonian Iron Age graves. Map by R. Allmäe

According to the main model where the given period of grave usage is 150 years, the burial grounds were most likely exploited by communities of 3–14 people. In most cases, this corresponds to one family or household. In comparison with other graves, Maidla II stone grave in western Estonia and Rõsna-Saare I barrow cemetery in south-eastern Estonia could have been used by a somewhat larger community, which may mean an extended family, a larger household or usage by two nuclear families.

More papers on the same subject by the author – who participated in the recent Mittnik et al. (2018) paper – include Observations On Estonian Iron Age Cremations (2013), and The demography of Iron Age graves in Estonia (2014).

Fast life history in Iron Age Estonia

While the demographic situation in the Gulf of Finland during the Iron Age is not well known – and demography is always difficult to estimate based on burials, especially when cremation is prevalent – , there is a clear genetic bottleneck in Finns, which has been estimated precisely to this period, coincident with the expansion of Proto-Fennic.

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

The infiltration of N1c lineages in Estonia – the homeland of Proto-Fennic – happened during the Iron Age – as of yet found in 3 out of 5 sampled Tarand graves – , while the previous period was dominated by 100% R1a and Corded Ware + Baltic HG ancestry. With the Iron Age, a slight shift towards Corded Ware ancestry can be seen, which probably signals the arrival of warrior-traders from the Alanino culture, close to the steppe. They became integrated through alliances and intermarriages in a culture of chiefdoms dominated by hill forts. Their origin in the Mid-Volga area is witnessed by their material culture, such as Tarand-like graves (read here a full account of events).

This new political structure, reminiscent of the chiefdom system in Sintashta (with a similar fast life history causing a bottleneck of R1a-Z645 lineages), coupled with the expansion of Fennic (and displaced Saamic) peoples to the north, probably caused the spread of N1c-L392 among Finnic peoples. The linguistic influence of these early Iron Age trading movements from the Middle Volga region can be seen in similarities between Fennic and Mordvinic, which proves that the Fenno-Saamic community was by then not only separated linguistically, but also physically (unlike the period of long-term Palaeo-Germanic influence, where loanwords could diffuse from one to the other).

NOTE. Either this, or the alternative version: an increase in Corded Ware ancestry in Estonia during the Iron Age marks the arrival of the first Fennic speakers ca. 800 BC or later, splitting from Mordvinic? A ‘Mordvin-Fennic’ group in the Volga, of mainly Corded Ware ancestry…?? Which comes in turn from a ‘Volga-Saamic’ population of Siberian ancestry in the Artic region??? And, of course, Palaeo-Germanic widely distributed in North-Eastern Europe with R1a during the Bronze Age! Whichever model you find more logical.

Related

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

indo-european-uralic-migrations-corded-ware

This is the first of four posts on the Corded Ware—Uralic identification:

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

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

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

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

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

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

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

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

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

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

Corded Ware – Yamna similarities?

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

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

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

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

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

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

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

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

Archaeology

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Genetic similarities

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

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

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

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

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

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

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

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

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

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

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

Corded Ware and ‘steppe ancestry’

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

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

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

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

uralic-steppe-ancestry

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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Trypillia and Greece Neolithic outliers: the smoking gun of Proto-Anatolian migrations?

neolithic-migrations-khvalynsk-novodanilovka-anatolian

(Continued from the post Corded Ware culture origins: The Final Frontier).

Looking at the PCA of Wang et al. (2018), I realized that Sredni Stog / Corded Ware peoples seem to lie somewhere between:

  • the eastern steppe (i.e. Khvalynsk-Yamna); and
  • Lower Danube and Balkan cultures affected by Anatolian- and steppe-related (i.e. Khvalynsk-Novodanilovka) migrations.

This multiethnic interaction of the western steppe fits therefore the complex archaeological description of events in the North Pontic, Lower Danube, and Dnieper-Dniester regions. Here are some interesting samples related to those long-lasting contacts:

1. I3719 (mtDNA H1, Y-DNA I2a2a) Ukraine Neolithic sample from Dereivka ca. 4949–4799 BC, described in Mathieson et al. (2018) as of “entirely northwestern-Anatolian-Neolithic-related ancestry”.

2. ANI163 from Varna I ca. 4711–4550 BC (mtDNA H7a1), and I2181 from Balkans Chalcolithic (Smyadovo, in Bulgaria) ca. 4500 BC (mtDNA HV15, Y-DNA R) show the first steppe ancestry in regions known to be affected by the expansion of Suvorovo chiefs.

3. The Yamna Bulgaria outlier (Y-DNA I2a2a1b1b), 3012-2900 calBCE, shows apparently an admixture with cultures of that region (but 1,500 years later).

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

Trypillia and Corded Ware

4. There is one ‘Trypillia outlier’ among five samples from the Verteba cave in Wang et al. (2018): I1927 (Y-DNA G2a2b2a1a1b1a1a1, mtDNA H1b), ca. 3619-2936 BC, a sample published previously in Nikitin et al. (2017) and Mathieson et al. (2017). We were very quick to dismiss Trypillia (three samples of haplogroup G2a, one sample E) and GAC as a source of Corded Ware admixture, but archaeology clearly shows important population movements at the end of the fourth millennium between late Trypillia groups, GAC, and post-Sredni Stog populations, and genetics is showing that in both cultures, too.

I am not a fan of the ‘lack of samples’ argument, but (similar to Old Hittite samples related to all Anatolian speakers) one site is not enough to describe a culture that spanned millennia and many different early and late groups. One among five Trypillian samples (from a single site), showing a late date (ca. 3228 BC) compared to the other samples (ca. 3700 BC), and quite close to the only three Ukraine Eneolithic samples we have may mean much more than what we may a priori think, i.e. some simplistic unidirectional punctual ‘intrusion’ of steppe ancestry, and instead hint at the known close contacts of late Trypillian groups and North Pontic cultures, including also the Caucasus.

NOTE. The big difference in PCA among GAC-like Hungary LCA – EBA samples (see above two star symbols close to Ukraine Neolithic outlier in the PCA, in contrast with the other three at the bottom) may also be significant, although we don’t have any data about their culture, sites, or the relationship between them.

trypillia-verteba-cave
Location of Verteba Cave in relation to different stages and neighbouring groups of the Cucuteni-Trypillia culture. Image from the paper A Subterranean Sanctuary of the Cucuteni-Trypillia Culture in Western Ukraine, by Kadrow and Pokutta (2016).

Greece Neolithic outlier: Proto-Anatolians?

5. Especially interesting is I6423, one of the Greece Neolithic samples referred to in Wang et al. (2018), which is obviously an outlier among the three used in the paper. It does not seem to correspond to any of the ancient DNA samples published to date; it is not in Hofmanova et al. (2016), in Lazaridis et al. (2017), or in Mathieson et al. (2018).

Since the Neolithic in Greece could mean any period from ca. 6500 BC to ca. 3200 BC, I guess we are talking here about some migration related to the expansion of Khvalynsk-Novodanilovka chieftains after ca. 4500 BC, because it appears on the PCA precisely on the same spot as Varna and Smyadovo outliers, and its ADMIXTURE shows similar components

admixture-ukraine-eneolithic-greece-neolithic
Image modified from Wang et al. (2018). “ADMIXTURE results of relevant prehistoric individuals mentioned in the text (filled symbols)”. ‘Outlier’ samples referred to in this post have been marked in red. See the original file here.

So, this may be the smoking gun of Proto-Anatolian (or maybe early Common Anatolian) expansion with steppe migrants up to the border of Western Anatolia, and we may be able to get rid of those unfounded doubts about Anatolian origins once and for all…

NOTE. Also interesting seems another Greece Neolithic sample, I6420, in ADMIXTURE, although its position in the PCA (near Minoans and Mycenaeans) does not necessarily point to potential steppe influence, but rather to the extra ‘eastern (Caucasus/Iran-related) ancestry’ contribution found in Minoans and in Mycenaeans (and Anatolia Chalcolithic) compared to previous samples of the region. The third Greece Nelithic sample, I5427 (mtDNA K1a24), from Diros, Alepotrypa Cave, is dated 6005-5879 BC (mean 5892 BC), and appeared first in Mathieson et al. (2018).

greece-neolithic
Modified from Wang et al. (2018) (Greece_Neolithic in red). Supplementary Table 6. P values of rank=1 in modelling the two-way admixture in the Caucasus cluster. 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. Source 2 populations in bold print are used as examples in modelling the Caucasus cluster groups (See Supplementary Table 7).
greece-neolithic-caucasus
Modified from Wang et al. (2018) (Greece_Neolithic in red). Supplementary Table 10. P values of rank=2 and ancestry proportions in modelling a three-way admixture in the Caucasus cluster testing additional contribution from Iran_ChL. Here, we used an extended set of outgroup populations populations to constrain standard errors: 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, EHG, WHG, Levant_N.

If this Greece Neolithic sample is not related to Yamna migrations – and its use for statistical analysis of Caucasus samples from Wang et al. (2018) suggests that it is not – , it may have important consequences:

If it is located near the Western Anatolian coast – especially near Troy – there won’t be much to add about the potential site of entry of Common Anatolian languages into Anatolia… I have read some comments about how ‘impossible’ it was for steppe migrants and their language to ‘invade’ the more advanced cultures of Anatolia from the west, but it seems as ‘impossible’ as it was for Barbarians to invade the Roman Empire and impose their language as elites in certain regions. (And yes, we have at least one important weak political period among Middle Eastern cultures in the early 3rd millennium BC, similar to the period of the Fall of the Western Roman Empire).

indo-european-anatolian-uralic-migrations
Most likely Proto-Anatolian expansion in the North Pontic and Balkan area with Khvalynsk-Novodanilovka chieftains, including ADMIXTURE data from Wang et al. (2018).

If it is located somewhere more ‘central’ in the Greek Peninsula, then it could also be used to support the Anatolian nature of the controversial Pre-Greek (‘Pelasgian’) substrate. While we know that Greek (at least since Mycenaean) shows a strong Pre-Greek cultural and linguistic heritage (also reflected in its genetic continuity), the nature of that language is usually believed to be non-Indo-European, and Anatolian contacts are rather few and coincident with the Mycenaean period. I don’t think this sample can tell much about the Pre-Greek language, though, because – if it is really Neolithic, and comparing it with later Minoan and Mycenaean samples – it seems a clear outlier.

suvorovo-novodanilovka-expansion-europe
Heyd (2016): The Southeast European distribution of graves of the Suvorovo-Novodanilovka group and such unequipped ones mentioned in the text which can be attributed by burial custom and stratigraphic position in the barrow, plus zoomorphic and abstract animal head sceptres as well as specific maceheads with knobs as from Decea Maresului (mid-5th millennium until around 4000 BC). The site in the south-west Balkans is Suvodol-Šuplevec, Northern Macedonia (FYROM).

If it is, however, related to later Yamna migrations after ca. 3000 BC (and, like the ‘Ukraine Eneolithic’ sample that is likely from Catacomb, it is classified as Neolithic just because it cannot be attributed to precise Helladic periods), then we may be in front of the first obvious Yamna migrants in Greece. If that is the case (which I doubt), the sample wouldn’t be so informative for PIE dialectal expansions, because by now it is evident that we will find steppe ancestry and R1b-Z2103 subclades accompanying Yamna migrants in the southern Balkans, and probably well into Mycenaean Greece.

NOTE. Whatever the case, I am sure that for those fond of absurd autochthonous continuity theories, as well as for anti-steppe conspirationists, this sample will be just another good way of arguing for anything, ranging from a rejection of the Middle PIE – Late PIE division, to a support for some mythic ancient autochtonous Proto-Graeco-Anatolian group, or maybe some ancient Graeco–Indo-Slavonic split, or whatever new dialectal stage one can invent to support the own genealogical fantasies…

So, if it actually is a Neolithic sample, let’s hope that it shows a clear R1b-M269 (xL23 or early L23) subclade distinct from those (likely Z2103) expanded later with Late PIE-speaking Yamna (and probably to be found among Mycenaeans), so that there can be no more place for ethnic fantasies.

EDIT (28 JUL 2018): Added information on Greece Neolithic and Trypillia samples

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Corded Ware culture origins: The Final Frontier

corded-ware-yamna-bell-beaker

As you can imagine from my latest posts (on kurgan origins and on Sredni Stog), I am right now in the middle of a revision of the Corded Ware culture for my Indo-European demic diffusion model, to see if I can add something new to the draft. And, as you can see, even with ancient DNA on the table, the precise origin of the Corded Ware migrants – in spite of the imaginative efforts of the Copenhagen group to control the narrative – are still unknown.

Corded Ware origins

The main objects of study in Corded Ware origins are necessarily the region where the oldest Corded Ware vessels appeared, Lesser Poland, as well as the adjacent (traditionally considered Proto-Corded Ware regions) Volhynia, Podolia, and upper Dniester river basin. These are some relevant points, continuing where I left the Eneolithic steppe developments (following Szmyt 1999, Rassamakin 1999, Kadrow 2008, Furholt 2014):

gac-trypillia-yamna-usatovo
Kadrow (2008). Cultural interactions around Carpathians at the beginnings of the 3rd millennium BC: 1 – Globular Amphora culture; 2 – Sofievka group of Trypillia culture; 3 – Funnel Beaker culture; 4 – Baden culture; 5 – Kostolac culture; 6 – Coţofeni culture; 7 – Cernavoda II culture; 8 – Yamnaya culture and Usatovo group of Trypillia culture (apud Kadrow, 2001).
  • More frequent contacts were seen ca. 3500-3000 BC, with an interaction showing multidirectional migrations of larger human groups in the centuries around 3000 BC, involving a significant part of the population of central-east Europe.
  • The easternmost area of the Funnel Beaker culture had become more Baden-like with the expansion of the Baden culture in its western area ca. 3300-2900 BC (with findings up to 2600 BC), and these younger groups with Baden features moved increasingly into the western part of Volhynia.
  • The influence of the neighbouring Trypillian culture is seen in the eastern parts of Volhynia, from ca. 3000 BC, either from a younger phase CII (cf. Troyaniv, Koshilivtsy, Brînzeni, Zhvaniets, or Vychvatintsy) or later groups (cf. Gorodsk, Kasperivtsy, Sofievka, Horodiştea-Folteşti, Usatovo).
  • In the forest-steppe zone, herding and hunting activities intensified, while agricultural traditions were preserved, as shown by the Sofievka, Kasperivtsy, and Gorodsk groups. From the end of the 4th millennium BC mobile parts of the late Trypillian populations moved to the steppe zone, absorbing more and more steppe elements; among others, cord ornamentation (in Vykhvatintsy, Troyaniv, and Gorodsk groups), pottery forms (Vykhvatintsy, which served as prototype for the Thuringian Apmphorae, dispersed along the Dniester river, too), flat burials with bodies in contracted position on the left or right side (Vykhvatintsy, reminding of Polgár culture different male-female position, and later Corded Ware burials, and also Lower Mikhailovka, under a mound without stone constructions). At the end of the Trypillia culture, its agricultural system collapsed completely.
gac-trypillia-usatovo-corded-ware
Globular Amphorae culture „exodus” to the Danube Delta: a – Globular Amphorae culture; b – GAC (1), Gorodsk (2), Vykhvatintsy (3) and Usatovo (4) groups of Trypillia culture; c – Coţofeni culture; d – northern border of the late phase of Baden culture;red arrows – direction of Globular Amphora culture expansion; blue arrow – direction of „reflux” of Globular Amphora culture (apud Włodarczak, 2008, with changes).
  • Slash and burn techniques of agriculture – especially those practiced by Trypillian and Funnel Beaker populations – must have intensified effects of natural growth of humidity (ca. 3400-2400), increasing fluvial activities in west Ukrainian river valleys, and increasing deforestation processes, which favoured pastoralism and nomadisation of the settlement system, and a consequent change of the social structure
  • At the same time, Yamna communities expanded along the lower and central Danube to the west, while the populations of the late phase of the Baden culture took the opposite direction and reached as far as Kiev in the north-east, contributing to the culture of the Sofievka group.
  • Globular Amphora communities migrated from the north-west, from eastern Poland, towards the Danube Delta and as far as the Dnieper in the east, destroying the primary structures of the communities in the supposed cradle territories of the Corded Ware culture. These communities found refuge and conditions for further development in south-eastern margin zone of the Funnel Beaker culture territories, penetrating at first the upper parts of the loess uplands like typical Funnel Beaker sites, but on the margins of their range, and also on areas avoided by Funnel Beaker settlement agglomerations. They brought with them the so-called Thuringian amphora up to Lesser Poland, borrowed from the late Trypillian Usatovo group. This resulted in the Złota culture, which eventually gave rise to the A-Amphorae.
funnelbeaker-trypillia-corded-ware
Map of territorial ranges of Funnel Beaker Culture (and its settlement concentrations in Lesser Poland), local Tripolyan groups and Corded Ware Culture settlements (■) at the turn of the 4th/3rd millennia BC.

In the end, we are left with this information about the oldest CWC (Furholt 2014):

  • The earliest radiocarbon-dated groups associated with the Corded Ware culture come from new single graves from Jutland in Denmark and Northern Germany, ca. 2900 BC. This Early Single Grave culture is associated with the appearance of individual graves (some time after the decline of the megalithic constructions), composed of a small round barrow and a new gender-differentiated burial practice emphasising male individuals orientated west-east (with regional exceptions), combined with the internment with new local battle-axe types (A-Axe). However, there is no single type of burial or burial custom in Corded Ware:
    • In southern Sweden the prevailing orientation is north-east – south-west, and south-north, contrary to the supposed rule male individuals are regularly deposited on their left and females on their right side.
    • In the Danish Isles and north-eastern Germany, the Final Neolithic / Single Grave Period is characterized by a majority of megalithic graves, with only some single graves from typical barrows. In south Germany, west-east and collective burials prevail, while in Switzerland no graves are found.
    • In Kujawia (south-eastern Poland), Hesse (Germany), or the Baltic, west-east orientation and gender differentiation cannot be proven statistically.
corded-ware-regions-main
Furholt (2014). Map of the Corded Ware regions of central Europe. The dark shading indicates those regions where Corded Ware burial rituals are present regularly
  • The oldest Corded Ware vessels (the A-Amphorae, which define the A-Horizon of the CWC) come probably from the Złota (or a related) group in Lesser Poland, where a mixed archaeological culture connecting Funnel Beaker, Baden, Globular Amphorae and Corded Ware appears ca. 2900-2600 BC. No cultural (typological) break is seen between earlier Globular Amphorae and the first Corded Ware Amphorae, but rather a continuum of traits and characteristics among the recovered vessels. This strengthens the connection of Corded Ware with Globular Amphorae peoples. The A-horizon expanded thus probably from Lesser Poland ca. 2800-2600, as seen in local contexts.
  • And of course we have a third way of defining Corded Ware individuals, which is the presence of herding, and thus a transition from hunter-gatherers to agropastoralists. This is how some Baltic Late Neolithic individuals with no archaeological data have been classified as members of the Corded Ware culture: Even though no cultural remains were extracted with the two ‘outlier’ individuals, their haplogroup and ancestry point to a direct origin in or around the steppe and forest-steppe region (yes, that risks circular reasoning).
globular-amphorae-corded-ware-zlota-amphorae
Correspondence analysis of amphorae from the Złota-graveyards reveals that there is no typological break between Globular Amphorae and Corded Ware Amphorae, including ‘Strichbündelamphorae’ (after Furholt 2008)

Corded Ware peoples in genetics

So, no clear origin of Corded Ware migrants, a lot of data pointing to intense migrations and interaction among GAC, Trypillia and the western steppe population (remember Kristiansen’s ‘long-lasting GAC-CWC connection’, now ignored to favour their Yamnaya admixture™ concept), and also three ways of defining Corded Ware culture…

Maybe genetics can help:

Ukraine Neolithic cultures – mainly from Dereivka – show haplogroups R1b-V88, R1a1, and R1b-L754 (xP297, xM269), which is similar to the haplogroup distribution found in Ukraine Mesolithic, but apparently with an expanding group marked by haplogroup I2a2a1b1 (possibly I2a2a1b1b).

The first thing that stands out about Ukraine Eneolithic samples is that only two of them can be said to be really Ukraine Eneolithic (i.e. from “Sredni Stog”-related groups):

  • I5876 (Y-DNA R1a-Z93(Y3+), mtDNA U5a2a), from Alexandria, 4045-3974 calBCE (5215±20BP, PSUAMS-2832)
  • I4110 (mtDN AJ2b1), from Dereivka, 3634-3377 calBCE (4725±25 BP, UCIAMS-186349), J2b1

The other two samples are quite late, and in fact one of them is clearly too late (maybe from the Catacomb culture):

  • I5882 (mtDNA U5a2a), from Dereivka, 3264-2929 calBCE (4420±20BP, PSUAMS-2826)
  • I3499 (Y-DNA R1b-Z2103, mtDNA T2e), from Dereivka, 2890-2696 calBCE (4195±20BP, PSUAMS-2828)

Corded Ware samples from Mittnik et al. (2018) offer very wide radiocarbon dates, so it is unclear which of them may be the oldest one. Most of them cluster closely to the older Ukraine Eneolithic sample I5876, but also to later steppe_MLBA samples i.e. Sintashta, Potapovka, and especially Srubna and Andronovo). This points to a genetic continuity from Pre-Corded Ware to Classic and late Corded Ware peoples. Therefore, much like Khvalynsk-Yamna and apparently many other Neolithic cultures, these peoples did not really admix; at least not with the male population.

pca-mittnik-late-neolithic
File modified by me from Mittnik et al. (2018) to include the approximate position of the most common ancestral components, and an identification of potential outliers. Zoomed-in version of the European Late Neolithic and Bronze Age samples. “Principal components analysis of 1012 present-day West Eurasians (grey points, modern Baltic populations in dark grey) with 294 projected published ancient and 38 ancient North European samples introduced in this study (marked with a red outline).

Lucky for us, even though the culture remains undefined, haplogroup R1a-Z645 seems like a unifying trait, as I said long ago, so we only have to wait for more samples to trace their origin. Nevertheless, it is clear that Corded Ware may not have been as genetically homogeneous as Khvalynsk, Yamna and Yamna-related cultures, further supporting its archaeological complexity:

  • Jagodno1 and Jagodno2 (Silesia), dated ca. 2800 BC, show haplogroup G? and I/J? – compatible with an origin of CWC in common with Trypillia (which shows 3 samples of haplogroup G2a2b2a, and one E) and Ukraine Neolithic (showing the expansion of I2a2a1b1 subclades).
  • I7272, from Brandýsek (Czech Republic), dated ca. 2900-2200 BC shows haplogroup I2a2a2 (compatible with an origin in Ukraine Neolithic peoples – this haplogroup is also found in Yamna Kalmykia and in the Yamna Bulgaria outlier, i.e. late western samples from the Early Yamna culture).

NOTE. This precise subclade is only present to date in Chalcolithic samples from Iberia, which points (possibly like the Esperstedt family) to local Central European haplogroups integrated in a mixed Proto-Corded Ware population. The upper subclade I2a2a is found in Neolithic samples from Iberia, the British Isles, Hungary (Koros EN, ALPc), and also south-east European Mesolithic and Neolithic samples.

  • RISE1, from Oblaczkowo (Greater Poland), ca. 2865-2578 BC, shows haplogroup R1b1.
  • The Esperstedt family samples have been analysed as R1a-M417 (xZ645), although the supposed ‘xZ645’ has not been confirmed – not even in the risky new Y-calls from Wang et al. (2018) supplementary materials.
corded-ware-regions-network
Network analysis based on the quantitative occurrence of Corded Ware pottery forms, pottery ornamentation styles, tools,
weapons and ornaments as stated in Table 1, based on the catalogues given in Table 2, line thickness representing similarity

Maybe this heterogeneity is a problem of better defining the culture, but from what we can see the oldest CWC regions and the unifying ‘Corded Ware province’ – formed after ca. 2700 BC by Jutland and Northern Germany, the Netherlands, Saale, Bohemia, Austria and the Upper Danube regions – are for the moment not the most genetically homogeneous groups.

Homogeneity comes later – which we may tentatively identify with the expansion of the A-horizon from the northern Dnieper-Dniester and Lesser Poland area – , as seen around the Baltic (like the Battle Axe culture) with R1a-Z283 subclades, and around Sintashta (i.e. probably Abashevo – Balanovo) with R1a-Z93 subclades, which is compatible with the late spread of different Z645 groups (and potentially a unifying language) .

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