Tripolje, Dereivka, and the Sredni Stog phenomenon (guest post)

This is another guest post by an anonymous contributor. Again, I don't necessarily agree with the author, but is he wrong? Feel free to let me know in the comments below.

The definition of Sredni Stog phenomenon (SSP) varies and is often a loosely applied term to refer to pretty much any individual in the Dnieper and Don regions between the ‘Neolithic’ and Yamnaya periods.

In order to elucidate the SSP, some brief remarks on the preceding Mariupol phenomenon are warranted. Understanding the Mariupol horizon is fairly straightforward – its development was catalysed by an intrusion of groups from somewhere west of the Dnieper, ~ 5500BC. The ‘proto-Mariupol’ group were genomic and economic ‘hunter-gatherers’, lacking any discernible EEF admixture, and with Y-hg I-L702 uniparental ‘trace-dye’. The Mariupol phenomenon predominantly impacted the lower reaches of the Dnieper and Azov steppe (Lower Dnieper- Azov group, ‘LDA’), but extended toward the Don, Volga and even the Kuban steppe in an attenuated form. The elevated levels of “Ukr N’ in Golubaya Krinitsa and the Y-hg I2a-L702 individual at Berezhnovka attest to this movement. The Mariupol phenomenon is associated with the development of formal cemeteries, linking them with Late Neolithic mixed farmer/HG groups in the northeast Balkans. Individuals were buried in a ‘supine straight-legged’ inhumation, with grave goods such as boar-tusk pendants for select males and adorning shell beads for females. This might signal the emergence of gender-differentiation in burials and the rise of local leaders or ‘chiefs’.

Data sets treat the ‘Neolithic’ in Ukraine as a monolithic phenomenon, however it is important to note that Dereivka stands apart – it is 200km north of other “Ukraine_N” sites such as Volnienski, Igren and Vovnigi, beyond the ‘Dnipro bend’. Moreover, male individuals from Dereivka are overwhelmingly assigned to Y-hg R1b-V88 and the burial pose (N-S) at Dereivka deviates from the more common E-W orientation seen elsewhere. Quite a few of the published ‘Sredni Stog’ individuals are from Dereivka, and often earlier than 4500BC, and N. Kotova assigns them to the Dnieper-Donets culture. Moreover, the recently published middle Don individuals, such as those from Golubaya Krinitsa and Vasilyevsky Cordon -17, are also not Sredni Stog, but can be thought of as ‘partially Mariupolised hunter-gatherers’. In another example, the (undated) ‘Sredni Stog’ individual I27930 from Igren was assigned to Y-hg Q and he can be modelled as a 2-way mix of EHG & WHG. This individual is actually from the Mesolithic.

So what occurred during the Sredni Stog period? In contrast to the Mariupol phase, the population dynamics associated with SSP are complex: at least three external flows can be highlighted (i) the advance of Tripolje communities from the Carpathians to the Dnieper (ii) arrival of South-Caucasian/CHG agro-pastoralists in the north Caucasus, and (iii) arrival of ‘central Asian’ populations in Volga-Caspian region (represented by “TTK individual’); in addition to intra-steppe shifts and flows. Notwithstanding, the ‘ideological background’ of SSP is rooted in the Mariupol horizon. The stereotypical SSP burials features individuals buried on their back, but increasingly with legs up-flexed. And we see the beginnings of kurgan constructions, which vary from stone cairns to soil-thrown barrows. Most are buried in simple pits, however some have more complex ‘catacomb’ pits.

What happened in the Dniester-Dnieper-Don region during the SSP? We can begin by orientating ourselves with a PCA to observe two main clines developing. One cline develops between ‘Ukr_N and EEF and a second cline pulls toward Lower-Volga Caucasus groups. The first cline mostly comprises of ‘Farmers’ from Tipolje and ‘hunter-gatherers’ from Dereivka. The second cline consists of individuals from Dereivka and the lower Dnieper-Azov group pulling toward Lower Volga-Caucasus groups.

Admixture analysis with qpAdm reveals 3 groups within the 2 broad clines. The first group can be thought of as ‘core Sredni Stog’. These individuals are 2-way mixes of ‘Ukr_N’ and ‘Steppe Eneolithic’ (sometimes Progress works, sometimes Remontoye or Berezhnovka). They are both males and females. In our examples, the females are from Kopachiv Yar (4000 BC) and Dereivka (3500BC). The males come from Dereivka (4300BC), Moluykhiv Bugor (4000BC), Vynohrado (4000BC); they are all derived for Y-hg I2a-L703+. These results represent a blending of social networks between the LDA and various lower Volga-Caucasus groups, and the subsequent expansion by LDA further West. The terminus ante quem of 4300 BC matches the corrected dating of the Kuban steppe sites such as Progress & Vojnuchka.

Another subset comprises of individuals from Dereivka and Verteba cave who situated on an ‘Ukr-N’ < - - > EEF cline. Many of the earlier Dereivka individuals are almost 100% Ukr_N. Verteba cave Tripolje can be modelled as 80% EEF + 20% Ukr_N. One individual from Dereikva (I3719) falls outside the Dereivka <-> Tripolje clin, as he plots further ‘south’ with Balkan-LBK farmers. Consistently, he comes out as ~100% EEF with qpAdm. Dating to ~4700BC, he precedes the arrival of Tipolje groups to the region by hundreds of years. FtDNA have assigned him to I2-Z161- FTH81, which is distinctive to the LDA haplotypes and is phylogenetically linked to a Czech LBK individual.

A third group consists of individuals with more complex 3-way ancestries, consisting of EEF, Ukr-N and Steppe_En and/or Maikop. These come from late Dereivka and late Tripolje groups, in archaeological literature often termed as ‘Soldanesti’, ‘Zhivotilovksa-Volchansk’, Cernavoda (Kartal). Once again, the males from Soldanesti and Cernavoda derive from LDA-related Y-hg I2a-L703 in some shape or form.

Conclusions:

1) Firstly, we note that the Dereivka group was subject to early EEF influence, as soon as eastern LBK groups reached Ukraine after 5000BC. However, their main interaction occurred with the younger, Tripolje group, which expanded toward the Dnieper after ~ 4300 BC.

2) In the LDA group we observe patrilineal continuity. These clans created expansive social-networks. They initially mixed with groups from the lower Volga-Caucasus area. Some then moved west, and ‘took over’ the Tripolje region and acquired high levels of EEF.

3) As a third conclusion, we can reject the commonly held notion that Tripolje was ‘conquered by Yamnaya pastoralists’. Our analysis instead highlights that their core structure fragmented as they became intertwined with powerful networks to their west (Trpolje) and east (Sredni Stog). The ‘take-over’ was due to the expansion of LDA/ SS groups. Mixed groups emerged such as Cernavoda and Soldanesti, which retained Tripolje ancestry and some cultural traditions. By the time Yamnaya groups reached the Dniester forest steppe, Tripolje had been long gone.

See also...

‘Proto-Yamnaya’ Eneolithic individuals from Kuban steppe c. 3700 BC ? (guest post)

Dear Iosif...Yamnaya

Even though the Yamnaya culture probably originated in what is now Ukraine, the earliest Yamnaya samples currently available are from the modern-day Samara region of Russia. They mostly date to around 3,000 BCE. I can analyze their ancestry using Principal Component Analysis (PCA) data.

Target: RUS_Yamnaya_Samara
Distance: 3.2816% / 0.03281581
81.0 RUS_Progress_En
14.4 UKR_N
4.6 HUN_Vinca_MN
0.0 ARM_Aknashen_N
0.0 ARM_Masis_Blur_N
0.0 AZE_Caucasus_lowlands_LN
0.0 BGR_C
0.0 BGR_Dzhulyunitsa_N
0.0 IRN_Ganj_Dareh_N
0.0 IRN_Hajji_Firuz_C
0.0 IRN_Seh_Gabi_C
0.0 IRN_Tepe_Abdul_Hosein_N
0.0 IRN_Wezmeh_N
0.0 RUS_Darkveti-Meshoko_En
0.0 RUS_Maykop
0.0 RUS_Maykop_Late
0.0 RUS_Maykop_Novosvobodnaya

The above results show exactly zero ancestry from West Asia. Admittedly, both RUS_Progress_En and HUN_Vinca_MN are European ancients with significant West Asian-related ancestry. However, this ancestry is very distantly West Asian-related, and, for instance, it almost certainly has no relevance to the Indo-Anatolian homeland debate.

The Afanasievo culture of Central Asia is regarded to have been an early offshoot of the Yamnaya culture. A good number of Afanasievo samples are available, so let's have a look if their results match those of the Yamnaya folks. And indeed they do, since BGR_C is very similar to HUN_Vinca_MN.

Target: RUS_Afanasievo
Distance: 3.4055% / 0.03405499
84.0 RUS_Progress_En
11.4 UKR_N
4.6 BGR_C
0.0 ARM_Aknashen_N
0.0 ARM_Masis_Blur_N
0.0 AZE_Caucasus_lowlands_LN
0.0 BGR_Dzhulyunitsa_N
0.0 HUN_Vinca_MN
0.0 IRN_Ganj_Dareh_N
0.0 IRN_Hajji_Firuz_C
0.0 IRN_Seh_Gabi_C
0.0 IRN_Tepe_Abdul_Hosein_N
0.0 IRN_Wezmeh_N
0.0 RUS_Darkveti-Meshoko_En
0.0 RUS_Maykop
0.0 RUS_Maykop_Late
0.0 RUS_Maykop_Novosvobodnaya

To try this at home, stick the PCA data in the text file here into the relevant fields here and cranck up the "Cycles" to 4X. You should see exactly zero ancestry from West Asia every time.

I can, more or less, reproduce these results with tools that are routinely used in peer reviewed papers. Below is a table of mixture models produced with the qpAdm software. I set the pass threshold to P ≥0.05, which is an arbitrary value, but the pattern is clear. The full output from each qpAdm run is available here.

Importantly, qpAdm needs to be fed the relevant "right pop" outgroups to be able to discriminate accurately between reference populations.

right pops:
CMR_Shum_Laka_8000BP
MAR_Taforalt
Levant_Natufian
IRN_Ganj_Dareh_N
Levant_PPNB
TUR_Marmara_Barcin_N
HUN_Starcevo_N
HUN_Koros_N
SRB_Iron_Gates_HG
Iberia_Southeast_Meso
RUS_Karelia_HG
RUS_West_Siberia_HG
RUS_Boisman_MN
MNG_North_N
TWN_Hanben
BRA_LapaDoSanto_9600BP

So, for instance, if one were to use in this role the modern-day Mbuti people, as opposed to, say, the ancient hunter-gatherers of Shum Laka, one might find that many models look statistically better than they should. And then one might also find that the Yamnaya samples carry significant West Asian ancestry.

Actually, I'm not opposed to the idea of some West Asian ancestry in Yamnaya. Indeed, considering the extraordinary mobility of the Yamnaya people and their Eneolithic predecessors on the Pontic-Caspian steppe, it would be unusual if they didn't come into close contact and mix, to some degree, with their neighbors from West Asia.

However, based on everything I've seen, from uniparental markers to different types of autosomal genetic tests, it's clear to me that there's no substantial West Asian ancestry in any Yamnaya samples, except for an outlier female from modern-day Ozera, Ukraine (see here).

Admittedly, ancient DNA does have a habit of throwing curveballs, so I'm eagerly awaiting new Eneolithic samples from the Pontic-Caspian steppe, particularly those associated with the Yamnaya-like Sredni Stog culture, to help finally settle this issue.

Believe it or not, a contact recently sent me a supposedly unpublished female sample from a ~4,200 BCE Sredni Stog burial in modern-day Igren, east central Ukraine. So what the hell, let's assume for the time being that this sample is genuine. This is how Miss Sredni Stog behaves in my PCA mixture test.

Target: UKR_Sredni_Stog
Distance: 4.0769% / 0.04076877
75.6 RUS_Progress_En
17.8 UKR_N
6.6 HUN_Vinca_MN
0.0 ARM_Aknashen_N
0.0 ARM_Masis_Blur_N
0.0 AZE_Caucasus_lowlands_LN
0.0 BGR_C
0.0 BGR_Dzhulyunitsa_N
0.0 HUN_Vinca_MN
0.0 IRN_Ganj_Dareh_N
0.0 IRN_Hajji_Firuz_C
0.0 IRN_Seh_Gabi_C
0.0 IRN_Tepe_Abdul_Hosein_N
0.0 IRN_Wezmeh_N
0.0 RUS_Darkveti-Meshoko_En
0.0 RUS_Maykop
0.0 RUS_Maykop_Late
0.0 RUS_Maykop_Novosvobodnaya

Wow, just wow. Have we actually found Miss Proto-Yamnaya? What does qpAdm have to say in the matter?

UKR_Sredni_Stog
HUN_Vinca_MN 0.034±0.028
RUS_Progress_En 0.796±0.045
UKR_N 0.170±0.034
P-value 0.41088

Again, this is an excellent match with the results from my PCA test, especially if we take into account the standard errors. However, with qpAdm it's also possible to model this individual's ancestry as part West Asian.

UKR_Sredni_Stog
AZE_Caucasus_lowlands_LN 0.056±0.039
RUS_Progress_En 0.761±0.061
UKR_N 0.183±0.036
P-value 0.465667

As I pointed out above, it's plausible for such people to harbor some West Asian ancestry, but I'm very sceptical that this is really the case here, despite the rather solid qpAdm statistical fit. That's because UKR_Sredni_Stog is not a high quality sample, and, from my experience, qpAdm often has problems analyzing fine scale ancestry in singletons or even small groups that show excess DNA damage and/or offer much less than a million markers.

See also...

Dear Iosif, about that ~2%

But Iosif, what about the Phrygians?

The PIE homeland controversy: February 2022 status report

I think we'll see the emergence of two main competing proto-Indo-European (PIE) homeland theories over the next few years:

- a homeland in the Eneolithic North Caucasus, and the spread of Anatolian languages into West Asia with Maykop-related ancestry

- a homeland in the North Pontic region, possibly within the Eneolithic Sredny Stog archeological culture, and the spread of Anatolian languages into West Asia via the Balkans.

Both theories have support from ancient DNA. Some of it has already been published (for instance, see here).

At this point, I can see myself firmly in the North Pontic camp, even if it turns out that North Pontic-related ancestry only made a fleeting impact on Bronze Age Anatolia.

After all, there's no direct relationship between genes and languages, so to prove that Anatolian languages came from the North Pontic, there's no need for North Pontic-related ancestry to persist in Anatolia, as long as we have solid evidence that people with this type of ancestry moved there at the right time.

In my mind, for now, the Maykop culture provides an excellent explanation for non-Indo-European influences in PIE, and there's no need to make it Indo-European speaking, let alone PIE speaking.

See also...

The PIE homeland controversy: June 2021 status report

Looking forward to a post-Covid world

I was hoping that the Covid-19 pandemic wouldn't have an immediate impact on the publication of ancient DNA papers and new data, but considering how much things have slowed down in this respect, it seems that I was fooling myself.

So let's take a break until early next year, and then see what happens.

Trust me, we've got a lot to look forward to in the post-Covid-19 world. Based on what I've heard from various sources, here are some predictions about what we might see:

- the search for the Proto-Indo-European homeland will shift west to the North Pontic steppe

- on the other hand, the search for the Proto-Uralic homeland will move deep into Siberia

- the key role of the Single Grave (westernmost Corded Ware) culture in the population history of Western Europe will finally get some attention

- following on from the above, Y-haplogroup R1b-L51 will be revealed as a Single Grave marker

- the idea that the Pontic-Caspian steppe was colonized by migrants from Mesopotamia during the Bronze Age will be forgotten, and, ironically, we'll instead learn that there was a significant influx of steppe ancestry into ancient Mesopotamia

- Old Kingdom Egyptians will come out less Sub-Saharan African than present-day Egyptians.

I probably shouldn't blab everything out, so that's all you're getting from me for now. You'll just have to wait for the rest until next year, or perhaps even the year after that.

See also...

The PIE homeland controversy: August 2019 status report

Archeologist David Anthony has a new paper on the Indo-European homeland debate titled Archaeology, Genetics, and Language in the Steppes: A Comment on Bomhard. It's part of a series of articles dealing with Allan R. Bomhard's "Caucasian substrate hypothesis" in the latest edition of The Journal of Indo-European Studies. It's also available, without any restrictions, here.

Any thoughts? Feel free to share them in the comments below. Admittedly, I found this part somewhat puzzling (emphasis is mine):

It was the faint trace of WHG, perhaps 3% of whole Yamnaya genomes, that identified this admixture as coming from Europe, not the Caucasus, according to Wang et al. (2018). Colleagues in David Reich’s lab commented that this small fraction of WHG ancestry could have come from many different geographic places and populations.

I think that's highly optimistic. It really should be obvious by now thanks to archeological and ancient genomic data, including both uniparental and genome-wide variants, that the Yamnaya people were practically entirely derived from Eneolithic populations native to the Pontic-Caspian (PC) steppe. So, in all likelihood, this was also the source of their minor WHG ancestry.

Indeed, they clearly weren't some mishmash of geographically, culturally and genetically disparate groups that had just arrived in Eastern Europe, but the direct descendants of closely related and already significantly Yamnaya-like peoples associated with long-standing PC steppe archeological cultures such as Khvalynsk and Sredny Stog. I discussed this earlier this year, soon after the Wang et al. paper was published:

On Maykop ancestry in Yamnaya

I hope I'm wrong, but I get the feeling that the scientists at the Reich Lab are finding this difficult to accept, because it doesn't gel with their theory that archaic Proto-Indo-European (PIE) wasn't spoken on the PC steppe, but rather south of the Caucasus, and that late or rather nuclear PIE was introduced into the PC steppe by migrants from the Maykop culture who were somehow involved in the formation of the Yamnaya horizon.

Inexplicably, after citing Wang et al. on multiple occasions and arguing against any significant gene flow between Maykop and Yamnaya groups, Anthony fails to mention Steppe Maykop. But the Steppe Maykop people are an awesome argument against the idea that there was anything more than occasional mating between the Maykop and Yamnaya populations, because they were wedged between them, and yet clearly distinct from both, with a surprisingly high ratio of West Siberian forager-related ancestry (see here and here).

Despite all the talk lately about the potential cultural, linguistic and genetic ties between Maykop and Yamnaya, including claims that the latter possibly acquired its wagons from the former, my view is that the Steppe Maykop and Yamnaya wagon drivers may have competed with each other and eventually clashed in a big way. Indeed, take a look at what happens after Yamnaya burials rather suddenly replace those of Steppe Maykop just north of the Caucasus around 3,000 BCE.

Yamnaya_RUS_Caucasus
RUS_Progress_En_PG2001 0.808±0.058
RUS_Steppe_Maykop 0.000
UKR_Sredny_Stog_II_En_I6561 0.192±0.058
chisq 13.859
tail prob 0.383882
Full output

Yep, total population replacement with no significant gene flow between the two groups. Apparently, as far as I can tell, there's not even a hint that a few Steppe Maykop stragglers were incorporated into the ranks of the newcomers. Where did they go? Hard to say for now. Maybe they ran for the hills nearby?

Intriguingly, Anthony reveals a few details about new samples from three different Eneolithic steppe burial sites associated with the Khvalynsk culture:

The Reich lab now has whole-genome aDNA data from more than 30 individuals from three Eneolithic cemeteries in the Volga steppes between the cities of Saratov and Samara (Khlopkov Bugor, Khvalynsk, and Ekaterinovka), all dated around the middle of the fifth millennium BC.

...

Most of the males belonged to Y-chromosome haplogroup R1b1a, like almost all Yamnaya males, but Khvalynsk also had some minority Y-chromosome haplogroups (R1a, Q1a, J, I2a2) that do not appear or appear only rarely (I2a2) in Yamnaya graves.

As far as I can tell, he suggests that they'll be published in the forthcoming Narasimhan et al. paper. If so, it sounds like the paper will have many more ancient samples than its early preprint that was posted at bioRxiv last year.

For me the really fascinating thing in regards to these new samples is how scarce Y-haplogroup R1a appears to have been everywhere before the expansion by the putative Indo-European-speaking steppe ancestors of the Corded Ware culture (CWC) people. It's basically always outnumbered by other haplogroups wherever it's found prior to about 3,000 BCE, even on the PC steppe. But then, suddenly, its R1a-M417 subclade goes BOOM! And that's why I call it...

The beast among Y-haplogroups

At this stage, I'm not sure how to interpret the presence of Y-haplogroup J in the Khvalynsk population. It may or may not be important to the PIE homeland debate. Keep in mind that J is present in two foragers from Karelia and Popovo, northern Russia, dated to the Mesolithic period and with no obvious foreign ancestry. So it need not have arrived north of the Caspian as late as the Eneolithic with migrants rich in southern ancestry from the Caucasus or what is now Iran. In other words, for the time being, the steppe PIE homeland theory appears safe.

Update 20/12/2019: A note on Steppe Maykop

See also...

Is Yamnaya overrated?

The PIE homeland controversy: January 2019 status report

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

On Maykop ancestry in Yamnaya

What Maykop ancestry in Yamnaya? There is none, or at least not enough worth discussing, except in one highly unusual female outlier from a burial in what is now eastern Ukraine. But apparently this is still up for debate? Well it shouldn't be.

To anyone with even a passing interest in the Yamnaya culture, it should be rather obvious that it formed during the tail end of the Eneolithic on the Pontic-Caspian steppe, as basically a direct offshoot of the earlier Repin culture, but perhaps also with significant influences from the earlier still Khvalynsk and Sredny Stog cultures. So why should its population history be much different from this?

It isn't, and this is fairly easy to demonstrate now despite the still rather poor sampling of Eneolithic remains from the Pontic-Caspian steppe.

Below is a series of qpAdm analyses in which I modeled several Yamnaya groups, as well as the closely related Afanasievo and Poltavka populations, exclusively and successfully as two- and three-way mixtures of a few Eneolithic singletons from various parts of the Pontic-Caspian steppe (obviously, I'd love to use homogeneous population sets instead, but, as per my point above, that's not possible yet). The models are sorted by their statistical fits, best to worst. Also note the large number and wide range of right pops or outgroups. I wanted to make sure that I wasn't missing anything.

Yamnaya_Samara
Dereivka_I_I4110 0.324±0.035
Progress_Eneolithic_PG2004 0.676±0.035
chisq 6.797
tail prob 0.976979
Full output

Afanasievo
Progress_Eneolithic_PG2004 0.638±0.038
Sredny_Stog_II_I6561 0.362±0.038
chisq 10.855
tail prob 0.818366
Full output

Yamnaya_Ukraine
Progress_Eneolithic_PG2001 0.655±0.073
Sredny_Stog_II_I6561 0.345±0.073
chisq 12.676
tail prob 0.696277
Full output

Poltavka
Dereivka_I_I4110 0.324±0.038
Progress_Eneolithic_PG2004 0.676±0.038
chisq 12.895
tail prob 0.680437
Full output

Yamnaya_Caucasus
Khvalynsk_Eneolithic_I0122 0.086±0.054
Sredny_Stog_II_I6561 0.221±0.070
Vonyuchka_Eneolithic_VJ1001 0.693±0.101
chisq 13.113
tail prob 0.593562
Full output

So, you might ask, is there any way to add Maykop to these models? Nope, it's pointless, because it doesn't improve the stats (for instance, see here, here and here). In other words, the situation is this: I already have awesome models, and I can't readily fit Maykop into my framework, so why do it? But if anyone out there wants to try, then by all means, and feel free to share the results with us in the comments.

Of course, the fact that most of these Yamnaya and Yamnaya-related populations are best modeled with somewhat different Eneolithic steppe singletons doesn't mean that they have radically different origins. In fact, they're all very closely related and they're basically like one Bronze Age steppe family. They just harbor somewhat different ratios of the same ancient ancestral components.

For the sake of being thorough, as per scientific literature, I pooled all of the above Afanasievo, Poltavka and Yamnaya samples into a Steppe_EMBA set and analyzed it with several genetically and geographically matching pairs of the Eneolithic singletons. This was one of the best fitting models, which I think is interesting, because the region roughly between the burial sites of these pairs of Eneolithic individuals was the home of the Repin culture.

Steppe_EMBA
North_Pontic_Eneolithic_I4110-I656 0.313±0.027
Progress_Eneolithic_PG2001-PG2004 0.687±0.027
chisq 15.378
tail prob 0.497157
Full output

Again, adding Maykop to this model makes no sense (see here, here and here). Clearly, I'd have to come up with a very different framework to successfully model Steppe_EMBA with a Maykop population. However, it's unlikely that such a model would make much sense in the context of various other types of genetic analyses and archeological data.

See also...

Yamnaya: home-grown

Big deal of 2018: Yamnaya not related to Maykop

Yamnaya isn't from Iran just like R1a isn't from India

Yamnaya: home-grown

I have some interesting news. It looks like Khvalynsk_Eneolithic I0434 can be used as essentially a perfect proxy for the Eneolithic steppe trio from Wang et al. 2018 when modeling the ancestry of the Yamnaya people of what is now the Samara region of Russia. Consider the qpAdm mixture models below, sorted by taildiff.

One of the best fitting models that also fairly closely matches archeological data, which suggest that Yamnaya was an amalgamation of the Khvalynsk, Repin and Sredny Stog cultures, is in bold. The worst fitting, and basically failed, models are listed below the dotted line. Note that almost all of these models feature reference populations from West and Central Asia.

Khvalynsk_I0434 + Iberia_ChL 0.681534184 > full output

Khvalynsk_I0434 + Globular_Amphora 0.525961242 > full output

Khvalynsk_I0434 + Iberia_Central_CA 0.515960444 > full output

Khvalynsk_I0434 + Sredny_Stog_I6561 0.485311962 > full output

Khvalynsk_I0434 + Varna 0.430411416 > full output

Khvalynsk_I0434 + Blatterhole_MN 0.328782809 > full output

Khvalynsk_I0434 + Baden_LCA 0.234307235 > full output

Khvalynsk_I0434 + Protoboleraz_LCA 0.231310724 > full output

Khvalynsk_I0434 + ALPc_MN 0.200002422 > full output

Khvalynsk_I0434 + Trypillia 0.193900977 > full output

Khvalynsk_I0434 + Balaton_Lasinja_CA 0.187031564 > full output

Khvalynsk_I0434 + Tiszapolgar_ECA 0.153940224 > full output

Khvalynsk_I0434 + Tisza_LN 0.145465993 > full output

Khvalynsk_I0434 + Balkans_ChL 0.111720163 > full output

...

Khvalynsk_I0434 + Armenia_EBA 0.0108890099 > full output

Khvalynsk_I0434 + Armenia_ChL 0.00882375703 > full output

Khvalynsk_I0434 + Levant_BA_North 0.0078751978 > full output

Khvalynsk_I0434 + Minoan_Lasithi 0.0675240088 > full output

Khvalynsk_I0434 + Peloponnese_N 0.046998906 > full output

Khvalynsk_I0434 + Hajji_Firuz_ChL 0.00269860335 > full output

Khvalynsk_I0434 + Shahr_I_Sokhta_BA1 0.00261908387 > full output

Khvalynsk_I0434 + Sarazm_Eneolithic 0.00120345503 > full output

Khvalynsk_I0434 + Seh_Gabi_ChL 0.00111898703 > full output

Khvalynsk_I0434 + Geoksiur_Eneolithic 0.000178295163 > full output

Khvalynsk_I0434 + Tepe_Hissar_ChL 0.000163698274 > full output

Khvalynsk_I0434 + Bustan_BA 0.000151088148 > full output

Why is this potentially important? Because unless Khvalynsk_Eneolithic I0434 was a recent migrant from the North Caucasus piedmont steppe, which is where the remains of the Eneolithic steppe trio were excavated, then Yamnaya's ethnogenesis might not have anything at all to do with Asia or even the Caucasus region. At least not within any reasonable time frame anyway. Here's a map showing the geographic locations of all of the populations relevant to the highlighted mixture model above.

I won't be fussed if it turns out that the majority of the ancestry of the Yamnaya, Corded Ware and other closely related ancient peoples was sourced from the Eneolithic populations of the North Caucasus piedmont steppe. But I think it's useful to make the point that there are still very few ancient samples available from the steppes between the Black and Caspian seas, so we don't yet have much of a clue how the groups living throughout this region during the Eneolithic and earlier fit into the grand scheme of things.

Update 24/12/2018: I decided to repeat the analysis, but this time with Caucasus Hunter-Gatherers (CHG) as one of the outgroups (or right pops). The reason I initially didn't include CHG in the outgroups was because I didn't want to discriminate, perhaps unfairly, against West and Central Asians with high levels of CHG-related ancestry, and in favor of Europeans with no or minimal CHG-related input. But in my opinion, the new results clearly make more sense, with Sredny Stog and Varna at the top of the list.

Khvalynsk_I0434 + Sredny_Stog_I6561 0.410719649 > full output

Khvalynsk_I0434 + Varna 0.394089365 > full output

Khvalynsk_I0434 + Iberia_ChL 0.16554258 > full output

Khvalynsk_I0434 + Globular_Amphora 0.128348823 > full output

Khvalynsk_I0434 + Iberia_Central_CA 0.126100242 > full output

Khvalynsk_I0434 + Trypillia 0.135306664 > full output

Khvalynsk_I0434 + Baden_LCA 0.0853031796 > full output

Khvalynsk_I0434 + Protoboleraz_LCA 0.0766892008 > full output

Khvalynsk_I0434 + Tisza_LN 0.0661622403 > full output

Khvalynsk_I0434 + Tiszapolgar_ECA 0.0626469042 > full output

Khvalynsk_I0434 + Balaton_Lasinja_CA 0.0536293042 > full output

Khvalynsk_I0434 + ALPc_MN 0.0505788809 > full output

...

Khvalynsk_I0434 + Minoan_Lasithi 0.0439451605 > full output

Khvalynsk_I0434 + Balkans_ChL 0.0436885241 > full output

Khvalynsk_I0434 + Blatterhole_MN 0.0329758292 > full output

Khvalynsk_I0434 + Peloponnese_N 0.0181930605 > full output

Khvalynsk_I0434 + Armenia_EBA 0.014715999 > full output

Khvalynsk_I0434 + Armenia_ChL 0.0060437014 > full output

Khvalynsk_I0434 + Levant_BA_North 0.00514574731 > full output

Khvalynsk_I0434 + Shahr_I_Sokhta_BA1 0.00350059625 > full output

Khvalynsk_I0434 + Hajji_Firuz_ChL 0.00228771991 > full output

Khvalynsk_I0434 + Seh_Gabi_ChL 0.00117061206 > full output

Khvalynsk_I0434 + Sarazm_Eneolithic 0.001118931 > full output

Khvalynsk_I0434 + Bustan_BA 0.00021203609 > full output

Khvalynsk_I0434 + Tepe_Hissar_ChL 0.000200643323 > full output

Khvalynsk_I0434 + Geoksiur_Eneolithic 0.000175941977 > full output

Update 17/02/2019: I basically managed to confirm my analysis with samples from the Wang et al. Caucasus paper. See here.

See also...

Big deal of 2018: Yamnaya not related to Maykop

"The Homeland: In the footprints of the early Indo-Europeans" time map

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

The beast among Y-haplogroups

A lot has been written about Y-haplogroup R1a over the years. Sadly, most of it was wrong, such as its posited Pleistocene origin in the Indian subcontinent and subsequent migration to Europe.

In all likelihood, R1a was born somewhere in North Eurasia. More importantly, its R1a-M417 subclade, which encompasses almost 100% of modern-day R1a lineages, no doubt came into existence somewhere on the Pontic-Caspian (or Western) steppe in what is now Ukraine and southern Russia just 7,000-6,000 years ago.

And within a couple of thousand years it expanded in almost all directions, probably on the back of the early Indo-European dispersals (see here), to cover a massive range from Scandinavia to South Asia. It is the beast among Y-haplogroups.

The most common subclade of R1a-M417 in South Asia today is R1a-Z93, and, realistically, it couldn't have arrived there earlier than about 2,000BC. So much for the Pleistocene.

See also...

Y-haplogroup R1a and mental health

The Poltavka outlier

Yamnaya isn't from Iran just like R1a isn't from India