The Eneolithic cemetery at Khvalynsk on the Volga River

Anfänger

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New archeological paper about Khvalynsk. Shared by Lazaridis on Twitter.

Abstract
The genetically attested migrations of the third millennium BC have made the origins and nature of the Yamnaya culture a question of broad relevance across northern Eurasia. But none of the key archaeological sites most important for understanding the evolution of Yamnaya culture is published in western languages. These key sites include the fifth-millennium BC Khvalynsk cemetery in the middle Volga steppes. When the first part of the Eneolithic cemetery (Khvalynsk I) was discovered in 1977–1979, the graves displayed many material and ritual traits that were quickly recognized as similar and probably ancestral to Yamnaya customs, but without the Yamnaya kurgans. With the discovery of a second burial plot (Khvalynsk II) 120 m to the south in 1987–1988, Khvalynsk became the largest excavated Eneolithic cemetery in the Don-Volga-Ural steppes (201 recorded graves), dated about 4500–4300 BCE. It has the largest copper assemblage of the fifth millennium BC in the steppes (373 objects) and the largest assemblage of sacrificed domesticated animals (at least 106 sheep-goat, 29 cattle, and 16 horses); and it produced four polished stone maces from well-documented grave contexts. The human skeletons have been sampled extensively for ancient DNA, the basis for an analysis of family relationships. This report compiles information from the relevant Russian-language publications and from the archaeologists who excavated the site, two of whom are co-authors, about the history of excavations, radiocarbon dates, copper finds, domesticated animal sacrifices, polished stone maces, genetic and skeletal studies, and relationships with other steppe cultures as well as agricultural cultures of the North Caucasus (Svobodnoe-Meshoko) and southeastern Europe (Varna and Cucuteni-Tripol’ye B1). Khvalynsk is described as a coalescent culture, integrating and combining northern and southern elements, a hybrid that can be recognized genetically, in cranio-facial types, in exchanged artifacts, and in social segments within the cemetery. Stone maces symbolized the unification and integration of socially defined segments at Khvalynsk.

Link: https://doi.org/10.1515/pz-2022-2034



Edit: There are also genetic and skeletal studies in this paper but unfortunately it is not open access.
 
I'm temporarily out of juice, but thank you for posting that.

Too bad it isn't open access.
 
You’re welcome.

Seems like a big paper about the 5th and 4th millennum BC steppe is on it's way to being published. I'll keep you informed.
 
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Someone on AG has access to the paper. I'll post the Y-DNA and genetic info here:


khvalynsk.jpg


The Khvalynsk population was genetically admixed between northern and southern ancestry types, in general agreement with Khokhlov’s interpretation based on cra-nio-facial data. The northern type, Eastern Hunter-Gather-ers (EHG), evolved in northern Eurasia; and the southern type, designated Caucasus Hunter-Gatherers (CHG), was defined initially by Mesolithic and Early Neolithic inhab-itants of Georgia and western Iran. Both the EHG and CHG labels were first applied to hunter-gatherers, but afterwards were extended to genetically similar individuals regardless of economy. Wang et al. recognized that EHG & CHG ancestry like Khvalynsk was shared by Eneolithic individuals at Progress-2 (Figure 16) and Vonyuchka-1 (also known as Konstantinovskii-1) in the North Caucasus steppes. They are dated 4336–4173 calBCE (5397±28BP/MAMS-110563); and 4233–4047 calBCE (5304±25BP/MAMS-11210)

We do not know the proximate source of the CHG population that mixed with EHG to create the typical Khvalynsk/Progress-2 pattern of genetic ancestry. But it must have separated from other CHG populations in the Caucasus and western Iran before about 6500–6000 BCE, because after this date145 the CHG populations in the Caucasus and western Iran became admixed with Anatolian Farmer (AF) ancestry. By 4700 BCE, when the first farmers migrated from Georgia across the western North Caucasus Mountains and occupied sites on the north side of the North Caucasus ridge such as Meshoko and Svobodnoe, they had up to 50 % AF ancestry. The Progress-2/Khvalynsk steppe people had no AF ancestry, so they did not exchange mates with Meshoko farmers, even if archaeology shows that they did exchange material valuables (see Copper section above).

In the aDNA literature, “steppe ancestry” is a phrase used since Allentoft et al. and Haak et al. to refer to the typical Yamnaya pattern of genetic ancestry. The principal components of steppe ancestry were EHG & CHG, each in robust proportions, like Khvalynsk, although often with more CHG than in the Khvalynsk/Progress-2 population, with an added component of Anatolian Farmer (AF) ancestry (5–15 %) that was absent from the Khvalynsk/Progress-2 populations. Also, the Khvalynsk/Progress-2 mating network has not yet yielded the Y-haplogroup mutations that were directly ancestral to the typical Yamnaya form of R1b (R-Z2103). The R-V1636 form of R1b, found in males at Khvalynsk, Ekaterinovka Mys, Berezhnovka II, and Progress-2, identifies a branch that split from the Yamnaya branch defined by R-P297 > R-M269 > R-L23 > R-Z2103 (yfull.com). This entire branch is absent from the sampled Eneolithic males from the steppes, appearing for the first time in Yamnaya males. The evolution of Yamnaya Y-haplogroup ancestry occurred in a still-unsampled Eneolithic population.
 
Y-DNA distribution according to yfull:

R1b-L754-L389-V1636:
https://www.yfull.com/tree/R-V1636/

Q1-L472-M25-YP1669:
https://www.yfull.com/tree/Q-YP1669/

J1-CTS1026:
https://www.yfull.com/tree/J-CTS1026/

R1a-M459:
https://www.yfull.com/tree/R-M459/

I2a-L699:
https://www.yfull.com/tree/I-L699/

All of them look like EHG derived haplogroups, except maybe J1-CTS1026. R1a-M459 is interesting because it was found in an Ukraine_Mesolithic sample from Zaporizhzhia Oblast west of the Don River. Let's take a look at his Autosomal DNA:


Target: UKR_Meso:I1819
Distance: 0.0389% / 0.03886458
69.3RUS_Karelia_HG
25.6WHG
5.1Progress_CHG

Interestingly, he picks up some Progress_CHG but it has too much WHG to be the source for the steppe groups.

Target: UKR_Meso:I1819
Distance: 0.0404% / 0.04037524
72.8RUS_Samara_HG
24.6WHG
2.6TUR_Barcin_N

With EHG from Samara the Progress_CHG disappears instead there is 2,5% Anatolian Neolithic. Maybe this is some old Dzudzuana influence that we detect in EHGs?

Edit: I tried to remodel the Ukraine_Mesolithic sample with the same proxies, but I get different results. Barcin_N and Progress_CHG completely disappeared. What has happened ? I start to think that G25 has many flaws.:unsure:

Target: UKR_Meso:I1819
Distance: 0.0779% / 0.07793249
83.8RUS_Karelia_HG
16.2WHG
 
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Very interesting NEW genetic information:

The Eneolithic populations around the Dnieper Rapids were even more different from Yamnaya. All those sampled were admixtures of EHG (primarily) and Western European Hunter-Gatherers (WHG) similar to the Iron Gates Mesolithic populations. Among 30 published individuals from three Neolithic and Eneolithic cemeteries (Dereivka-1, Vil’nyaka, and Vovnigi) in the Dnieper River valley dated 5200–4400 BC, assigned to the Dnieper-Donets culture, there were a few individuals with minor (<10 %) CHG ancestry, but most had none. The Khvalynsk/Progress-2 populations had substantial CHG ancestry but no WHG ancestry, ubiquitous in Dnieper-valley populations. This indicates that the Dnieper-Donets mating network did not extend eastward to the Volga, nor westward to the Criş and early Tripol’ye farmers, whose ancestry was typical of European farmers (AF or EEF)153. The Dnieper-Donets people seem to have been an endogamous population focused on the rich resources of the Dnieper Rapids. Their substantial WHG ancestry, nearly absent in Yamnaya individuals, rules them out from being a major source for the Yamnaya. The Sredni Stog culture succeeded and replaced the Dnieper-Donets culture in the strategic Dnieper Rapids and throughout the steppes of Ukraine beginning around 4500–4300 BCE and ending in the late fourth millennium BCE with the appearance of Yamnaya. Unpublished Sredni Stog male genomes exhibit admixture ‘cocktails’ with the same basic elements as Yamnaya (EHG & CHG & AF). The CHG & EHG component was like Khvalynsk/ Progress-2, suggesting an eastern origin for at least part of the Sredni Stog population, and the AF component could have come from either the early Maikop or Tripol’ye populations. Sredni Stog introduced into the Ukrainian steppes new funeral customs (the Khvalynsk or ‘Yamnaya’ position), ceramic types (shell-tempered like Khvalynsk), and economies (large numbers of horse bones) that had appeared earlier on the Volga. Sredni Stog has for decades been recognized as an Eneolithic ancestor of Yamnaya influenced by late Khvalynsk, early Maikop, and the Tripol’ye and Varna cultures. But neither R1b Z-2108 nor its immediate ancestral forms are found among sampled Sredni Stog males, most of whom belonged to the R1a or I haplogroups, unlike Volga males. The sampled Sredni Stog populations included individuals who autosomally resembled Yamnaya a millennium before the Yamnaya culture appeared. But within that population the Yamnaya Y-haplogroup patriline evolved in a region that has not been sampled.


So Sredni-Stog also wasn't an ancestor to Yamnaya ? Where the hell the Yamnaya then come from ?
 
thanks for sharing anfanger(y)
those tables from the paper j1 is interesting
j1 was found in eastern hunter gatherers before and now this sample I6735

p.s
as i always said j came to the levant from the north
all those early samples from northern latitude
not even mantion the caucasus hunter gatherers who also belonged to y haplogroup j
 
thanks for sharing anfanger(y)
those tables from the paper j1 is interesting
j1 was found in eastern hunter gatherers before and now this sample I6735

p.s
as i always said j came to the levant from the north
all those early samples from northern latitude
not even mantion the caucasus hunter gatherers who also belonged to y haplogroup j

I think that J1 and J2 are CHG/Iran and/or Dzudzuana derived haplogroups, EHGs have minor affinity to the Caucasus. They could have went North very early. Oldest J1 is Satsurbila IIRC.
 
Very interesting NEW genetic information:



So Sredni-Stog also wasn't an ancestor to Yamnaya ? Where the hell the Yamnaya then come from ?

Same autosomal signature, though, just a different uniparental. The question indeed is what was the source of that uniparental signature. It certainly didn't change the autosomes very much. As to the J found, I'm sure you remember the assurances that no J, not J1 or J2, would be found on the steppe. :)

"But neither R1b Z-2108 nor its immediate ancestral forms are found among sampled Sredni Stog males, most of whom belonged to the R1a or I haplogroups, unlike Volga males. The sampled Sredni Stog populations included individuals who autosomally resembled Yamnaya a millennium before the Yamnaya culture appeared. But within that population the Yamnaya Y-haplogroup patriline evolved in a region that has not been sampled."




 
Same autosomal signature, though, just a different uniparental. The question indeed is what was the source of that uniparental signature. It certainly didn't change the autosomes very much. As to the J found, I'm sure you remember the assurances that no J, not J1 or J2, would be found on the steppe. :)

"But neither R1b Z-2108 nor its immediate ancestral forms are found among sampled Sredni Stog males, most of whom belonged to the R1a or I haplogroups, unlike Volga males. The sampled Sredni Stog populations included individuals who autosomally resembled Yamnaya a millennium before the Yamnaya culture appeared. But within that population the Yamnaya Y-haplogroup patriline evolved in a region that has not been sampled."





Yes, I remember. :) I am pretty sure upcoming papers about the *Volga Delta will refute further ideas of him, too.

The R1a they mentioned in the paper obviously also isn't ancestral to Corded ware or Balto-Slavic and Indo-Iranian R1a. Where could the PIE markers R1b and R1a have been ? Repin ?

Edit: *Volga Delta and/or Lower Don
 
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Very interesting NEW genetic information:



So Sredni-Stog also wasn't an ancestor to Yamnaya ? Where the hell the Yamnaya then come from ?

Not so fast, don't believe everything you read. A sample from the region

https://homeland.ku.dk/

Sample ID / genetic sex (M/F): I5884 / M
Find location: Dereivka
Country: Ukraine
Associated label in publication: Ukraine_Eneolithic
Date: 2890-2696 calBCE (4195±20BP, PSUAMS-2828)
MtDNA haplogroup (mother): U5a2b
Y-DNA haplogroup (father): R1b1a1a2a2 (Z2103)
Reference: Mathieson et al. 2018
Colour group: Steppe (R1b)
Comments: A mixture of western hunter-gatherer, Steppe, and Anatolian farmer ancestry
Other references: null
 
Very interesting NEW genetic information:



So Sredni-Stog also wasn't an ancestor to Yamnaya ? Where the hell the Yamnaya then come from ?
I0443 is also L23+ and negative for- Z2103



R1b-tree-Sergey-Malyshev.jpg
 
Not so fast, don't believe everything you read. A sample from the region

https://homeland.ku.dk/

He is dated to about 3000BC that's just the time of Yamnaya, he can´t be a source to any steppe group because of his WHG and DATE. He is probaly just an Yamnaya+Dereivka admixed sample.

IMO after this paper there are two options:

- Not yet sampled burials INSIDE Sredni Stog will turn out to be R1b-M269 and R1a-M417

OR

- There is another population with basically the same Autosomal DNA with different uniparentals. Maybe Repin?
 
Yes, I remember. :) I am pretty sure upcoming papers about the Volga Delta will refute further ideas of him, too.

The R1a they mentioned in the paper obviously also isn't ancestral to Corded ware or Balto-Slavic and Indo-Iranian R1a. Where could the PIE markers R1b and R1a have been ? Repin ?

Forget about R1a and J and whatever else they try throwing in the mix to try create confusion. I find it easier to stick to R1b-L23-Z2103. They admit they have not found the source population of Yamnaya, they also don't connect any dna evidence of the domesticated animals with the cultures they are trying to include/exclude.

Here is another interesting paper, on the domestication of the horse, Dom2
https://www.nature.com/articles/s41586-021-04018-9#Sec2
The C-PONT group not only possessed moderate NEO-ANA ancestry, but also was the first region where the typical DOM2 ancestry component (coloured orange in Fig. 1e, f) became dominant during the sixth millennium BC. Multi-dimensional scaling further identified three horses from the western lower Volga-Don region as genetically closest to DOM2, associated with Steppe Maykop (Aygurskii), Yamnaya (Repin) and Poltavka (Sosnovka) contexts, dated to about 3500 to 2600 BC (Figs. 2a, b, 3a). Additionally, genetic continuity with DOM2 was rejected for all horses predating about 2200 BC, especially those from the NEO-ANA group (Supplementary Table 2), except for two late Yamnaya specimens from approximately 2900 to 2600 BC (Turganik (TURG)), located further east than the western lower Volga-Don region (Figs. 2a, b, 3a). These may therefore have provided some of the direct ancestors of DOM2 horses.
 
He is dated to about 3000BC that's just the time of Yamnaya, he can´t be a source to any steppe group because of his WHG and DATE. He is probaly just an Yamnaya+Dereivka admixed sample.

IMO after this paper there are two options:

- Not yet sampled burials INSIDE Sredni Stog will turn out to be R1b-M269 and R1a-M417

OR

- There is another population with basically the same Autosomal DNA with different uniparentals. Maybe Repin?

There is no evidence of horse domestication in the earliest R1a samples.

Early ruminant dairying on the western steppe has also been inadequately demonstrated, as human stable isotope data from the region suggests—but cannot confirm—dairy consumption21,22. Palaeoproteomics, which is the only method that is able to evince individual dairy consumption (rather than milk production) and provide taxonomic resolution, has so far been minimally applied to steppe populations. Across Yamnaya and Afanasievo populations, dairying evidence is available only for a few individuals from the eastern steppe who have ancestry from western steppe groups; the earliest individual provides only a taxonomically ambiguous ruminant (Ovis/Bos) peptide result19.
To address the heavily debated question of what drove Yamnaya expansions across the steppe6,23,24,25, we conducted proteomic analysis of dental calculus sampled from 56 steppe individuals who span the Eneolithic to Late Bronze Age, and who date from between 4600 and 1700 BC. Our samples from the Eneolithic (about 4600 to 3300 BC) are from 19 individuals from 5 sites: Murzikha 2 (6 individuals), Khvalynsk 1 and Khvalynsk 2 (9 individuals), Ekaterinovka Mys (1 individual), Lebyazhinka 5 (1 individual) and Khlopkov Bugor (2 individuals) (Fig. 1, Supplementary Fig. 1a). Ancient DNA results from Khvalynsk and other Eneolithic sites in the Volga and northern Caucasus2,7,26 support the existence of an Eneolithic population across this region that was genetically similar to the Yamnaya population, but who lacked the additional farmer (Anatolian) ancestry that would arrive later on the steppe7. Published stable isotope and archaeological studies applied to Eneolithic populations from the Pontic region point to an economy based on fishing, the gathering of local plants and the keeping of domesticated animals6,21,27,28.Given the importance of the horse in reconstructions of early pastoralist expansions, we also examined dental calculus from two individuals from the well-known site of Botai. With faunal assemblages dominated by horse remains11,12,13 and early lipid studies of ceramics indicating horse milking at the site by 3500 BC13, the site is central to discussions of early horse milking and dairying in the Eurasian steppe.

https://www.nature.com/articles/s41586-021-03798-4
 
Y-DNA Update:

I6735 from Khvalynsk has just been added. This individual is J-CTS1026.
The Afanasievo sample from Mongolia (SHT002) is also CTS1026 and more precisely pre-Y136727, it's likely this individual is Y136727 as well. This branch must have been involved in IE dispersals since the earliest stage and is undoubtedly tied to the CHG component in Khvalynsk and other Eneolithic populations from the Pontic-Caspian steppe.
 
No R1b-M269, no R1a.
They appeared later on the Indo-European scene.
Ripin, IMO.
 

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