Genetic study The Genetic Origin of the Indo-Europeans

That remains to be seen. This J2b-L283 sample dates from the late Yamna period (2900-2500 BCE) in Moldova (former Cucuteni-Trypillian area) and the authors say that it could also belong to the Chalcolithic Cernavodă culture. Unless I missed something from a previous study I don't think that any J2b-L283 has been found in the Pontic-Caspian Steppe itself or the North Caucasus during the Yamna and Maykop periods. Maykop had J2a, but not J2b.
Given that we find quite a lot of J2a branches from the CHG ancestral components to the CLV, and that we find L283 in the descendant components of CLV (Phase 3: Core Yamnaya). While we find both fully Progress-like J2a and J2b in the Caucasian HG in Kotias Kilde 10kbp, I think its a good bet to assume the L283 was probably among the CLV.

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Also we do have L283 with fully Caucasus autosomals and no connection to European branches in the North Caucasus. Coincidentally very close geographically to R1b-Z2103 samples:

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L283 in light green.

Now, while it is not impossible for Progress like L283 to have arrived in Europe and then been incorporated into core Yamnaya, it is unlikely. Not least because Progress itself was a core component of CLV. If we had already L283 with CHG+EEF autosomals in the Balkans your claim would make sense, but we see quite the opposite. We see 92% Core Yamnaya L283 that precedes the early high steppe L283 from Mokrin, Shkrel and Cetina, and a diminishing Core Yamnaya in all later samples.
 
That remains to be seen. This J2b-L283 sample dates from the late Yamna period (2900-2500 BCE) in Moldova (former Cucuteni-Trypillian area) and the authors say that it could also belong to the Chalcolithic Cernavodă culture. Unless I missed something from a previous study I don't think that any J2b-L283 has been found in the Pontic-Caspian Steppe itself or the North Caucasus during the Yamna and Maykop periods. Maykop had J2a, but not J2b.
@Archetype0ne has linked the samples in question. J2b-L283 has been found in the North Caucasus Bronze Age. BTW the oldest J2b2 (non L283) is a 10000 year old Caucasus Hunter Gatherer sample from Kotias Klde, Georgia. "Iran Neolithic" or "Cucuteni-Trypillia" are irrelevant for J2b-L283 early origins, since that is, I guess, what you're proposing (or have proposed). The Southern route i.e. CHG into ANF through Anatolia is not supported by the phylogeny of J2b-L283, nor auDNA of the many early IE related J2b-L283 samples in the Western Balkans and now from the southern Steppe itself.

I think the samples speak for themselves and there is all sorts of interesting stuff to come. I remember some interesting work and abstracts from last year and thus far for '24.
 
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Do we have Y-DNA from the Shulaveri-Shomu culture?

I always thought that R1b-M269 came from the northern Middle East via the Caucasus but I thought it more likely through the eastern side of the Caucasus how long do Caspian Sea (mainly because it's easier to cross there). This remains to be confirmed by ancient DNA
Yes, consistently J2B :) -- My hopes of finding L23 vanished long ago.
But Mtdna is seen in Northern Caucasus, H2+152, H15a, I1.
 
Yes, consistently J2B :) -- My hopes of finding L23 vanished long ago.
But Mtdna is seen in Northern Caucasus, H2+152, H15a, I1.
Could you please provide a source, this is quite interesting.
 
Just adding a point:
the paper uses Aknashen which is very early Shulaveri. Has they use Mentesh tepe (prob the french didn't allow them) and it would be even closer because there would be more CHG, since they were admixing with remaining CHG pop near the mountains.
When we get later Shulaveri, closer to vanishing abruptly in 4900 BC, it would willl be better.
Let us just wait.
 
@Archetype0ne has linked the samples in question. J2b-L283 has been found in the North Caucasus Bronze Age. BTW the oldest J2b2 (non L283) is a 10000 year old Caucasus Hunter Gatherer sample from Kotias Klde, Georgia. "Iran Neolithic" or "Cucuteni-Trypillia" are irrelevant for J2b-L283 early origins, since that that is, I guess, what you're proposing (or have proposed). The Southern route i.e. CHG into ANF through Anatolia is not supported by the phylogeny of J2b-L283, nor auDNA of the many early IE related J2b-L283 samples in the Western Balkans and now from the southern Steppe itself.

I think the samples speak for themselves and there is all sorts of interesting stuff to come. I remember some interesting work and abstracts from last year and thus far for '24.
This foreshadowing from Patterson, a co-author two years ago:

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Quite a big paper indeed.
 
@Olympus Mons

Do we know where the painting of the deceased in red ochre stems from?

preview.jpg


From Progress 2: https://plos.figshare.com/articles/...ial_Progress_2_Grave_37_i_in_situ_i_/13091228


From the L283 Yamnaya sample:

Grave 12:10 (individual ID I10206)

The grave was situated in the northeastern sector of the mound, 2.80 m away from the central landmark (R.C.), at a depth of 1.65 m. The burial chamber, rectangular with rounded corners and slightly arched sides, was oriented east-west, measuring 1.25 × 1.80 m. Based on the characteristics of the filling of the pit, the arrangement of skeletal remains, and the location of the grave goods, it can be inferred that grave no.10/M.10 was disturbed in ancient times. The skeletal remains were found scattered in different areas of the pit and at different depths.

It was deduced that the deceased was originally placed at the bottom of the pit, with the head oriented to the east. Traces from the plant layer covering the pit bottom were found near and beneath the skeleton. Grave goods: a flint arrowhead of elongated triangular shape with slightly convex sides and a concave base. The artefacts’ length is 2 cm, width 1.4 cm, and thickness 0.5cm.

The skeleton, lacking anatomical connection, is incomplete and poorly preserved. Traces of bright red ochre were identified on all its elements, particularly intense on the skull. Other taphonomic changes observed include cracking, exfoliation (aerial weathering), and marks left by rodent teeth.

Anthropological sex: male.
Molecular sex: male, with an unusual Y-haplogroup J2b2a1 (J-L283).
Biological age at death: about 30 years (young adult).
Pathologies: supragingival calculus; active porotic hyperostosis (cribra cranii).

The skeleton is very robust, with extremely pronounced muscle insertions, with extensive enthesopathic changes on the humerus and femurs. The individual exhibits a very large skeletal stature, at least 181 cm.

Traumas: a perimortem fracture in the middle third of the right clavicle.

The burial must be attributed to Cernavodă-I or Yamnaya cultures.

Edit: It seems to have been mainstream in Yamnaya burials.

Regarding Yamnaya burials:
Characteristic for the culture are the burials in pit graves under kurgans (tumuli), often accompanied by animal offerings. Some graves contain large anthropomorphic stelae, with carved human heads, arms, hands, belts, and weapons. The dead bodies were placed in a supine position with bent knees and covered in ochre.

Regarding Progress 2:
This ancestry profile is known as 'Eneolithic Steppe' ancestry, or 'pre-Yamnaya ancestry', and is represented by ancient individuals from the Khvalynsk II and Progress 2 archaeological sites. These individuals are chronologically intermediate between EHGs and the later Yamnaya population, and harbour very variable proportions of CHG ancestry.[22][23][10]

This early, 'pre-Yamnaya' ancestry was first detected in Eneolithic individuals at the Khvalynsk II cemetery and directly north of the Caucasus mountains at the Progress 2 archaeological site; this ancestry is also detected in individuals of the Steppe Maykop culture, but with additional Siberian and Native American-related admixture.[25][5]

Three individuals from the Progress 2 site in the foothills north of the Caucasus also harbour EHG and CHG related ancestry, and are genetically similar to Eneolithic individuals from Khvalynsk II but with higher levels of CHG-related ancestry that are comparable to the later Yamnaya population.[5][26]

Archaeologist David Anthony speculates that the Khvalynsk/Progress-2 mating network, located between the middle Volga and the North Caucasus foothills, makes a "plausible genetic ancestor for Yamnaya".[26]

Admixture_proportions_of_Yamnaya_populations.png
 
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The oldest R1b-Z2103 (actually Z2103 > M12149 > Y467276 > Z2106) in this study comes from the Inland Caspian region (EBA Yamnaya) and dates from 5577 years before present (c. 3600 BCE). It is sample I33307. Interestingly it is even older than the R1b-M269 and R1b-L23 in this study.

There is an even older R1b-BY95094 (R-M269 > PF7562 > FGC31929 >...> BY95094) dating from 6100 BP (c. 4100 BCE). But according to Y-Full this deep clade only formed 1100 years ago. So it's either a contamination or a mistake.

We finally have R1b-L51 (the ancestor of Italic, Celtic and Germanic R1b) in the EBA Yamna. Not many samples compared to R1b-Z2103, but it confirms that R1b-L51 also originated in the Yamna culture. They were found in very different places: Kalmykia (4900 ybp), the Don region (4600 ybp), the Volga region (4600 ybp), but also in Yamna extension as far as Romania (4850 ybp) and Serbia (4710 ybp). The Serbian sample is the only one that is also positive for L52, so that may be the first clade born outside the Pontic Steppe.

Sample I3525 is reported as R1b-Y132510 (downstream of L21 and DF13!). It comes from Hungary and dates from 4600 ybp, but it's also extremely unlikely, as that clade is estimated to be only 2900 years old. EDIT: after double checking that sample is clearly indicated as contaminated.

The oldest R1a samples are two R1a* 5M459) from Golubaya Krinitsa in the Middle Don dating from 7400 years ago (Neolithic). There are only two other R1a samples, both Z93 from the MBA Fatyanovo culture further north (an offshoot of the Corded Ware culture).

There were actually far more I2a1b-L699 (descended from Neolithic Ukrainians) in the Don region during the Yamna period than R1b-L51 or R1b-L23.

Over 300 Steppe samples and there was not a single E1b1b and only one G2a (Neolithic G2a-L91 > Z42565) from Hungary. Likewise there was only one J2b2a (L283) from Moldova. This confirms that these three haplogroups were assimilated by the Steppe people after invading Old Europe.

There is also a single R-PF7562 sample(I10500). I believe this branch represents the forefathers of the Brygians and becomes Brygian by forming Bubanj-Hum culture and Armenchori culture. Armenchori branch might play a big role in indo-europeanizing the J2a locals who become the Greek speakers. There will be sequel paper as was hinted by Kristiansen, we are going to find eventually.
 
@Olympus Mons

Do we know where the painting of the deceased in red ochre stems from?

preview.jpg


From Progress 2: https://plos.figshare.com/articles/...ial_Progress_2_Grave_37_i_in_situ_i_/13091228


From the L283 Yamnaya sample:



Edit: It seems to have been mainstream in Yamnaya burials.

Regarding Yamnaya burials:


Regarding Progress 2:






Admixture_proportions_of_Yamnaya_populations.png
Nice catch! From The History of the Caucasus by Christoph Baumer, Chapter 2.1 Chalcolithic settlements and early low burial mounds. Credits go to @Polska, I remember you stating the RISE408 J2b-L283 sample being likely a descendant of Trialeti. The Ochre spraying ritual links Nalchik with Bedeni and Trialeti. Recalling the BA Nalchik J2b-L283 sample here.

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From The History of the Caucasus by Christoph Baumer, Chapter 2.1 Chalcolithic settlements and early low burial mounds. Credits go to @Polska, I remember you stating the RISE408 J2b-L283 sample being likely a descendant of Trialeti. The Ochre spraying ritual links Nalchik with Bedeni and Trialeti. Recalling the BA Nalchik J2b-L283 sample here.

View attachment 16018
Wait, Nalchik? This same Nalchik?

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The green dot is the L283 fully Caucasian sample BTW. Although same location it seems the older Nalchik graves have not been excavated for aDNA.

From the supplements of The Genetic Origin of Indo-Europeans:
2.10 Progress-2 and VonyuchkaSummary by D. Anthony

Progress-2 was excavated 2009-2010 under the direction of S.Y. Berzina. The multi-period kurgan cemetery was located on the left bank of the Malka River, itself a left tributary of the Terek River in the central Caucasus steppes east of Piatigorsk. The central Caucasus steppes consist of high grass-covered ridges that form a watershed, cut by streams flowing northeast into the Caspian (through the Terek River), north into the Manych Depression, or northwest into the Sea of Azov/Black Sea (through the Kuban), with the glaciated peaks of the Caucasus visible 100kmto the south. Progress-2 is one of many Eneolithic grave sites including Vonyuchka concentrated in the central Caucasus steppes, the upper Terek tributaries, and the Manych Depression. Very few Eneolithic graves are found in the NW Caucasus Kuban drainage.

The oldest and largest of these Eneolithic cemeteries was excavated in 1929-30 in the city of Nalchik and yielded one date of 4840–4820 BCE (GrA-24442, 5910 ± 45 BP). Nalchik has not been sampled for aDNA. Nalchik differed from other Eneolithic sites in its size (121 burials, while the later Eneolithic cemeteries such as Progress-2 usually have 2-4 individuals) and in the factthat 75% of the Nalchik burials were posed contracted on the left (mainly females) or right (mainly males) sides, while Progress-2 and almost all later Eneolithic graves were posed supine with raised knees, like most Khvalynsk and Serednii Stih graves (although contracted-on-the-side graves continued at Khvalynsk as a small minority). Burials contracted on one side were later also38typical of the Maikop culture.

About 25% of the graves at Nalchik in which a specific pose was clear were posed supine with raised knees. They can be interpreted as later graves in a multi-component cemetery, showing a shift in funeral ritual, a hypothesis supported by the discovery in one of the supine-with-raised knees graves (#83) of the only copper artifact found at Nalchik, a ring, and a serpentine stone bracelet like one found at Khvalynsk, dated 4500-4300 BCE there. Or the raised-knee graves at Nalchik could be older than Khvalynsk, contemporary with the raised-knee graves at LebyazhinkaV:12 and Ekaterinovka Mys in the Samara region, dated 4700-4500 BCE. More radiocarbon dates are needed to identify the oldest raised-knee graves, but Nalchik shows that the raised-knee posture was an Eneolithic innovation that replaced an older ritual (contracted on the side) in the North Caucasus.

Now from this other source: The Eneolithic cemetery at Khvalynsk on the Volga River by Anthony
Now from this other source:

Other Eneolithic cemeteries were smaller than Khvalynsk, usually less than 30 graves. Khlopkov Bugor,130 km south of Khvalynsk on the west Volga bank, had24 excavated Eneolithic graves, including one person who was a 2nd-degree relative of a person buried at Khvalynsk(see below). A cemetery on the Volga 160 km north of Khvalynsk, at Ekaterinovka Mys, dated 150–200 years earlier, had more than 100 excavated graves25. Nalchik in the North Caucasus, approximately contemporary with Khvalynsk, had 121 graves 26. A few other large Eneolithic cemeteries are known in the Volga-Caucasus steppes, but Khvalynsk was the largest.
Context to compare to the largest cemetery at Khvalynsk:
"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."
Rassamakin74 proposed that the Dnieper Rapidsregion emerged in this era as a secondary center of ‘Skelya-culture’ metalworking between Varna and the North Caucasus steppes. Most of the Khvalynsk copper is consistent with this kind of secondary source, among local steppe artisans. This could also be the source of a copper bead found at Svobodnoe, made of Balkan copper75. Svobodnoewas one of a series of agricultural settlements established in the Kuban River drainage after 4700 BCE by immigrant farmers who crossed the North Caucasus peaks from Georgia76. They participated in the trading network that brought Balkan copper into the steppes. Svobodnoe also produced many polished greenstone axes with faceted butts, like the axe found at Khvalynsk in grave I:105, probably made in the North Caucasus. A polished serpentine bracelet at Khvalynsk found in grave I:8 probably was made in the North Caucasus (Figure 9: middle panel);it was like bracelets at Nalchik. The Khvalynsk population was active in inter-regional exchange systems (Danube-Dnieper-Caucasus-Volga) that were stimulated by the heightened production of Balkan copper after 4500 BCE
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In the North Caucasus steppes at older Eneolithic cemeteries such as Nalchik (4840–4820 BCE, GrA-24442,5910 ± 45 BP), the flexed pose, contracted on the side, was used for most individuals, but even here a few individuals were buried on the back with raised knees. When the first small earthen mounds, or kurgans, began to appear in the North Caucasus steppes during the Eneolithic, after4500 BCE, they were erected over graves in which the deceased was positioned supine with raised knees, usually oriented to the east127. The Eneolithic individuals at Progress-2 and Vonyuchka in the North Caucasus steppes who had genetic ancestry similar to Khvalynsk were buried in the Khvalynsk position, in graves intensely colored with red ochre, beneath small (less than 1m high, ca. 15 m diameter) earthen mounds (Figure 16). These mounds were among the oldest kurgans in the Pontic-Caspian steppes 128; the other region where small kurgans appeared this early was in the steppes north of the Danube delta, as at Suvorovo129, again at a cultural, economic, and genetic border (Figure 2). Although they were small compared to later Yamnaya kurgans, the Eneolithic kurgans in the upper Tersek steppes east of the Svobodnoe-Meshoko agriculturalists perhaps were a boundary-marking practice that emerged during the late fifth millennium BCE. This was a millennium before the Yamnaya culture made the kurgan type of funeral monument universal across the Pontic-Caspian steppes.The Skelya and Sredni Stog cultures in Ukraine, contemporary with Khvalynsk, also used the supine-with raised-knee posture, unlike the supine-extended burial pose in the Dnieper Neolithic cemeteries130. Sredni Stog individuals also had genetic ancestry more like Khvalynsk and Progress-2 than the Dnieper Neolithic ancestry type(see below). Sredni Stog lithics also were similar to Khvalynsk, particularly the use of large lanceolate projectile points and long unifacial lamellar flint blades. Sredni Stog pottery was tempered with crushed shell, like Khvalynsk pottery, and unlike the Neolithic pottery of the Dnieper valley, where sand or mineral temper had been used. The high percentage of horse bones, averaging more than 50 % of all animal bones in Dnieper-valley Sredni Stog sites, was consistent with the high importance of horses in the diets of many fifth- and fourth-millennium BCE Ene127 Korenevskii 2012; 2016.128 Korenevskii 2012.129 Anthony 2007, 253.130 Telegin/Potekhina 1987.olithic settlements in the Volga and Don valleys131. Many traits indicate ‘eastern’ influences on Sredni Stog material culture, economy, and genetic ancestry, and the supine with-raised-knee burial pose is one of these.
Note: the Core Yamnaya L283 was indeed buried with a projectile point.
Now one of the preprints from upcoming papers:
Human Genetics Perspective of the Carpathian Basin at the Dawn of the 3rd Millennium BCE
Anna Szécsényi-Nagy, János Dani, Gabriella Kulcsár, Dániel Gerber, Tamás Hajdu, Alena Šefčáková, Ron Pinhasi, Balázs Gusztáv Mende, Mária Bondár, David Reich

This paper explores the archaeogenomic landscape of the Carpathian Basin around the transition from the 3rd millennium BCE, leveraging both published and new ancient DNA data. During the final phase of the Late Copper Age, archaeological cultures such as the Baden Complex and the Coţofeni culture witnessed the emergence of kurgan burials from the westernmost communities related to the Yamnaya culture across the Pannonian Plain and Transylvania. The arrival of these communities is traced back to the northeast Balkans around 3100/3000 BCE. Our analyses indicate that Yamnaya-associated individuals from Hungary and certain burials of the so-called ‘East Slovakian Mound cultures’ are genetically indistinguishable, reflecting the genetic makeup of the Yamnaya group from the Samara region in Russia. Interestingly, the introgression of additional farmer ancestry within the Yamnaya sample set appears sporadic, while dating of these introgressions reveals local contributions in only a few instances. Our paper presents evidence of potential sex-biased diffusion of Yamnaya-related ancestry into the Carpathian Basin and the dynamics of the early 3rd millennium BCE spread of steppe components in
the area.
Concurrent with the cultural and genetic impacts of the Yamnaya and Corded Ware, the Vučedol culture of Slavonia/north Balkans emerged, significantly influencing the Early Bronze Age community formation in southern Transdanubia. The Vučedol culture was noted for introducing steppe-related genetic ancestry into their region in the early 3rd millennium BCE, thereby diversifying ancestry
proportions within the population. These new findings provide significant insight into the complex genetic and cultural transformations within East-Central Europe during this pivotal era, directly contributing to our understanding of Early Bronze Age population formations.

It seems to me that the need for metal tools and ore was a core driver.

^ 5:45 to 7:00

First tech wise, North Caucasus to Steppe. But then ore wise, Steppe to Balkans.
 
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The origins of Remontnoye: half Maikop/Aknashen Neolithic and half Lower Volga Eneolithic Thus, we have arrived at the Remontnoye+SShi model as the only plausible solution, given the available sampled individuals, for the origins of the core Yamnaya population. It is possible that other such models would work but we lack the sources for them in our dataset; for example, models with SSmed+X as sources could be possible but we have not sampled an unknown “X” population. This is of course, a general problem in admixture modeling not unique to our study. However, with these limitations, we know for a fact that core Yamnaya are not a clade with any of the Serednii Stih subsets and Remontnoye is one source of extraSerednii Stih ancestry that results in a successful model. So, regardless of what other possible combinations may have occurred historically, investigating the origins of Remontnoye is important as its composition will contain forms of ancestry not represented in the sampled Serednii Stih. Thus, we turn to the question of the origins of Remontnoye itself.
We show in Table S 6 the feasible models and the results of the tournament comparing them. The models are all quite similar; they involve one population from the south (either Maikop, from the North Caucasus, or Aknashen Neolithic from Armenia or Azerbaijan Neolithic/Chalcolithic, in the South Caucasus) and one population from the north (BPgroup or PVgroup or Steppe Maikop or WSHG).

The best fitting models both in terms of the tournament score and in terms of p-value involve BPgroup and either Aknashen Neolithic or Maikop.


The results of the full tournament are presented in Table S 7. The best fitting models with greatest difference between wins and losses the BPgroup+Aknashen or BPgroup+Azerbaijan ones, which the BPgroup+Maikop model is almost as good (one fewer win and one more draw) and is also geographically and temporally more plausible. Strictly speaking, however, the non-BPgroup-related source for the Remontnoye must ultimately have had genetic roots further south, which is consistent with the observation that the models with Neolithic southern Caucasus sources have the most wins in the tournament. The models with either Steppe Maikop or WSHG as the northern source consistently fail in the competition. When did the Aknashen-related ancestry reach the north Caucasus and via it steppe populations like Remontnoye? All sampled individuals from the north Caucasus are from the 5th millennium BCE or later, while Aknashen is from the beginning of the 6th millennium BCE. We know that the native population of the South Caucasus consisted of Caucasus hunter-gatherers as late as the Kotias individual19 of the mid-8 th millennium BCE. Thus, the genetic Neolithization of the South Caucasus must have occurred sometime between the 8th and 5 th millennium BCE, i.e., during the flourishing of the Shulaveri–Shomu culture Neolithic culture. This ancestry could then have reached the north Caucasus by the time of the Unakozovskaya individuals (ref.8 and this study) of the mid-5 th millennium BCE. Thus, the time window of ~6000-4500BCE appears to be a plausible time frame for the spread of Aknashen-related ancestry into the North Caucasus. We note that Aknashen itself differs from the Masis Blur Neolithic of a few centuries later (mid-6 th millennium BCE),4 in which the CHG-related ancestry is greatly reduced. Thus, it seems probable that the Aknashen-related ancestry reached the North Caucasus before the 5633-5532 calBCE date of the Masis Blur individual when the (undiluted) Aknashen-related ancestry still existed in the South Caucasus.

Regardless of the temporality of this process, all fitting models agree on is that Remontnoye was formed as a mixture of people of the Caucasus-Volga north (BPgroup or PVgroup) and Neolithic-Chalcolithic south (from Maikop or Armenia-Azerbaijan).
p 160 supplement
The origins of Maikop: Aknashen Neolithic with minor BPgroup influence The Maikop population is very similar to that of Aknashen and the only feasible models for Maikop involve admixture between Aknashen and populations of eastern Europe.

Model competition does not help us distinguish between these models, but they all agree in that the Maikop population is very similar to Aknashen but with some additional northern influence. In particular, the model that fits the Remontnoye (BPgroup+Aknashen) also fits the Maikop, but with substantially more Aknashenrelated ancestry (86.2±2.9% in Maikop vs. 44.6±2.7% in Remontnoye). The Maikop group here includes individuals: OSS001, OSS002.B0101, I1720, I6266, I6267, I6268, I6272. We separately analyzed I4429 (a PCA ancestry outlier) which we left out of the main group in our fitting analyses. We show the fits of the Aknashen+BPgroup model for all Maikop individuals (Table S 9), demonstrating the relative homogeneity of the Maikop label and the poor fit and higher BPgroup-related ancestry of the outlier individuals that we did not include in the Maikop label used for the model fitting.
The admixture in Remontnoye is geographically plausible: Maikop is modeled as having formed from more ancient populations of the Caucasus (similar to the earliest Neolithic of the southern Caucasus sampled in Aknashen in Armenia), but is in contact and experienced admixture with steppe Eneolithic populations like BPgroup which geographically spanned at least the area from the Lower Volga (where the four Berezhnovka individuals were sampled) and the North Caucasus piedmont (the site of Progress-2). Remontnoye is the result of one such admixture, with roughly half of its ancestry modeled as being of Maikop/Aknashen origin, and the other half derived from the BPgroup.
The origins of the Lower Volga-North Caucasus Eneolithic: more CHG-admixed than the rest of the Volga cline We have so far shown that core Yamnaya was formed by admixture between SShi- and Remontnoye-related sources, and that Remontnoye was consistent with being part of a cline (together with Maikop) of admixture between people of the Caucasus (most robustly represented by Aknashen Neolithic) and people of the north (most robustly represented by BPgroup).

What of the origins of BPgroup? Several models fit the ancestry of BPgroup (Table S 12). The only model that does not lose any matches is the one that involves Khlopkov Bugor and PVgroup ancestries.
All but one models involve ancestry from the north (Khi, Kmed, or Khlopkov Bugor) and southern ancestry (CHG or PVgroup). The one exception is the Krivyansky+TTK model which captures a west (Don/Krivyansky) to east (Central Asia/TTK) distinction. This model loses many matches in the tournament, but this is because TTK is from a Central Asian ancestry source that likely diverged from the TTK-related ancestors of BP-group in Neolithic times at the most recent, and thus underestimates later shared genetic drift between BPgroup and other Eneolithic populations of the Volga.
Quite a lot to digest and warp ones head around.
Edit: That was from the supplement where they go more in deatil. This below is from the paper itself.
Yamnaya could have been formed from diverse (but similar) distal sources which include populations of (i) Neolithic or Chalcolithic age from Armenia6,9 472 and Azerbaijan43,44 473 representing the “Caucasus Neolithic”, (ii) GK2, UNHG, or Serednii Stih 474 representing the Dnipro-Don area, and (iii) BPgroup or PVgroup representing the Lower Volga475 north Caucasus Eneolithic. What is invariant among the class of 2- and 3-way models for the 476 Core Yamnaya is that they posit their descent from people of the CLV Cline (the remaining four 477 fifths of their ancestry) who admixed with Dnipro-Don people of substantial UNHG ancestry. 478 479 Our results show that movement of people and culture we document as having occurred along the 480 CLV Cline was the vector by which Caucasus-derived ancestry like that present in the Aknashen Neolithic population flowed into the steppe and into the ancestors of the Yamnaya45 481 .

David Anthony vindicated or what?


TLDR:
The origins of the Lower Volga-North Caucasus Eneolithic: more CHG-admixed than the rest of the Volga cline We have so far shown that core Yamnaya was formed by admixture between SShi- and Remontnoye-related sources, and that Remontnoye was consistent with being part of a cline (together with Maikop) of admixture between people of the Caucasus (most robustly represented by Aknashen Neolithic) and people of the north (most robustly represented by BPgroup).

What of the origins of BPgroup? Several models fit the ancestry of BPgroup (Table S 12). The only model that does not lose any matches is the one that involves Khlopkov Bugor and PVgroup ancestries.
All but one models involve ancestry from the north (Khi, Kmed, or Khlopkov Bugor) and southern ancestry (CHG or PVgroup).

What is invariant among the class of 2- and 3-way models for the 476 Core Yamnaya is that they posit their descent from people of the CLV Cline (the remaining four 477 fifths of their ancestry) who admixed with Dnipro-Don people of substantial UNHG ancestry. 478 479 Our results show that movement of people and culture we document as having occurred along the 480 CLV Cline was the vector by which Caucasus-derived ancestry like that present in the Aknashen Neolithic population flowed into the steppe and into the ancestors of the Yamnaya45 481 .

Also gotta love that Kotias Klde is directly referenced as to how long unadmixed Caucasian HG were present in the Caucasus.
 
After reading the paper there is something that doesn’t seem to be clear to me. The Berezhnovka BP group is modeled as CHG+EHG+Tutkaul. How do they explain the CHG-related ancestry that far north in the Volga? They say there is cline but the supposed cline explanation is too complicated and weird to me and it doesn’t make much sense.
 
After reading the paper there is something that doesn’t seem to be clear to me. The Berezhnovka BP group is modeled as CHG+EHG+Tutkaul. How do they explain the CHG-related ancestry that far north in the Volga? They say there is cline but the supposed cline explanation is too complicated and weird to me.
Archetype0ne
Archetype0ne
It took me quite a few hours of reading and re-reading... and it might look ugly... but I hope this helps others (and that I did not make any mistakes).

dhh5knS.png

Cline SShi <-> DonVolga
SShi = Don-DniproHG + BPgroup
BPgroup = KhlopkovBugor + CHG/PVgroup
DonVolga = BPgroup + PVgroup
DonDniproEN = BPgroup + PVgroup +ASH
Remotnoye = BPgroup + ASH (44.6%)
A: CoreYamnaya = SShi + Remotnoye
B: CoreYamnaya = Aknashen + BPgroup + Igren_o (EHG?)


Now...

Deconstructing A:
CoreYamnaya = ((Don-DniproHG + (KhlopkovBugor + CHG/PVgroup)) + (( KhlopkovBugor + CHG/PVgroup) + ASH)

Deconstructing B:
CoreYamnaya = Aknashen + 2x*(KhlopkovBugor + CHG/PVgroup) + Igren_o
*my emphasis
8sotBm0.png


And the tournament results:

W1YtF8z.png


Assuming ASH as the standing for CHG, BPgroup as 50/50 CHG/EHG, and Igren_o as EHG, the numbers for CoreYamnaya make a lot of sense

21Aknashen + 56BPgroup + 23Igren_o

Edit: Although I recall reading that BPgroup was a three way of Lower Volga + CHG + Central Asia, so it might not be exactly 50-50.


Archetype0ne
pegasusThe labels are confusing, what is the Dnipro cline exactly ? Do they have BPgroup ancestry? Are they suggesting this BPgroup is what gave rise to Indo Anatolian languages by interacting with Aknashen type groups? Hope the coordinates come out today.
There is this Sredni Stih cline between Don-Volga that is BP + PV groups on the one end, towards the Don-Dnipro where it intermixed with additional Don-Dnipro Hg. BPgroup is present in both as is PV. However the Don-Dnipro in the Eneolithic saw additional Aknashen admixture. They call this the Srednih Stih cline.

They present this scenario in page 182 (183 of the PDF supplement).

Edit: It seems that they tried to model Core Yamnaya with this Don-Dnipro mix of BP + PV + Don-Dnipro HG, and they could not without adding an additional source of Aknashen (possibly through Maykop) + BP, for which they argue Remotnoye is the stand-in.

This helps visualize it
8sotBm0.png
Archetype0ne
Jerome
From a fine-print,they aren't claiming BP groups to be PIE,but rather the general 'CLV' people.
BP is just a proxy to them from what it seems.

What they are considering PIE is what they call as 'proto-yamnaya' which also gave ancestry to sredny stog (bringing aknashen type Ancestry into sredny stog)
They date this Ancestry influx and mixing to around 4200-4000 BC,same time at which proto-yamnaya/Yamnaya recived aknashen Ancestry according to the DATES output in their supplementery data
More specifically they think Remotnoye like ancestry (The dot in the middle of the CLV) is what transformed "Pre"-Yamnaya into Core-Yamnaya, as these two occupy different ends of the Srednih Stih cline.

Remotnoye as that stand-in contains the additional BP and Akshanen needed, which defines the CLV, without which you can model the former BUT not the later.


• No pre-Yamnaya populations seem to match exactly the core Yamnaya
A “Pre-Yamnaya” population quite like the Core Yamnaya was at the other end of 183 the cline (Table S 20) and Serednii Stih people had therefore not only BPgroup-related ancestry from the south (as people on the Volga did), but also some Aknashen/Maikop-related ancestry.
I have not seen a "Proto-Yamnaya" reference in the supplement, but the "Pre-Yamnaya" is different from the CLV they seem to identify as the IA-IE vector. Edit: Although they do leave all hypotheses open, this seems to be the horse they are backing.
 
The BPgroup is KhlopkovBugor(EHG)+ CHG/PVgroup that’s what I am also getting from the paper but that’s a lot of CHG-related ancestry that far up the Volga. They are not really answering this, it might seem there could be an
1) older cline with CHG on side and EHG on the other side, with additional Central Asian/Tutkaul or
2) Berezhnovka is an intrusive population from the south.
They should have tested some older samples from the steppe

I am not thinking about the formation of Yamnaya because it’s quite clear now that it is from the region of the CLV cline in much later times.
 
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The thing is they are differentiating the CHG related ancestry. The CHG that far north was quite ancient source wise from HG themselves, so the Progress profile was quite common from the Volga group all the way to the Dnieper group, spanning the whole cline already by this point. The CHG that came right before IA-IE spread however was Armenian Neolithic like very distinguishable from the earlier HG one.

Archetype0ne

It is interesting that the earlier CHG already present in the region saw a decline in favor of Aknashen right before these Indo-Anatolian / Indo-European initial movements... I wonder what is up with these Aknashen, or if anybody has any sources on them?

The origins of Krivyansky on the Lower Don: a Seredniih Stih population with excess Caucasus-related ancestry

The Krivyansky individual does not fit the Remontnoye+SShi model of Table S 5 well (p=0.015) and it also does not fit the model of Table S 21 well (p=0.025). This individual is also unusual as it belonged to Y-haplogroup J2 suggesting Caucasus/West Asian connections. The results of the model tournament (Table S 22) show that relative to populations of the Dnipro-Don Serednii Stih and Golubaya Krinitsa, Krivyansky has an excess of CHG-related ancestry. The presence of this ancestry in the lower Don at a higher level than in the middle Don (at the GK1 subset of Golubaya Krinitsa parallels the situation of the Volga where CHG-related ancestry was higher in the lower Volga (at Berezhnovka) than in the middle Volga (Fig. S 2) and of the Caucasus where CHG-related ancestry in Maikop and (in slight excess Unakozovskaya) was higher in the North Caucasus than in the South Caucasus where the Aknashen earliest known Neolithic had seen its CHG-related ancestry diluted twice in Masis Blur and Areni-1 Neolithic and Chalcolithic populations. While CHG-related ancestry refers to the preNeolithic inhabitants of Georgia33 in the South Caucasus, it seems that it is in the triangle formed by the lower Don (at Krivyansky), the lower Volga (at Berezhnovka), and north Caucasus (at Unakozovskaya) that it remained a strong source of ancestry in the Eneolithic while CHG-related ancestry diminished upriver along both Don and Volga to the north, and across the Caucasus to the south.
We also explored 3-way models for Krivyansky after fixing CHG as one source (to account for this population’s excess CHG ancestry) and Ukraine_N as another (as a stand-in for the source that generates the Serednii Stih cline). All fitting models (Table S 23) involve some “eastern” ancestry (in addition to the CHG and Ukraine_N fixed sources; Table S 23). Three models have no losses in the tournament (Table S 23) and these involve Lower Volga-North Caucasus Eneolithic ancestry (BPgroup or PVgroup) or the SShi subset of Serednii Stih. Thus, Krivyansky appears like a population of “western” affinity (due to its Ukraine_N-related ancestry) but also “eastern” affinity (due to its Lower Volga-North Caucasus Eneolithicrelated ancestry) and also “southern” affinity (due to its CHG-related ancestry). As we will see further on, the people of the Serednii Stih and the Yamnaya share two of these ancestries but their southern source is related to Neolithic and post-Neolithic people of the Caucasus, not the much earlier CHG as is the case for Krivyansky.

These pre-Yamnaya (blue) already had CHG from the older source. The real question is why did the Eneolithic CHG rich Remotnoye (46%) impact mobility so much to what would become Core Yamnaya?

46nl6MT.png

IBD map. Notice no >20cm IBD for the blue, implying very ancient ties, and very heavy IBD links on a far more geographically expansive network for the red.


I have spent more time than I would like to admit reading and re-reading the supplement.
 
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Thanks, interesting read. So they are going for a cline of CHG from the Lower Don (Krivyansky) to Lower Volga and North Caucasus for this older CHG ancestry. The second Aknashen_N ancestry they call CHG-related is a bit weird because it is very little CHG-like but already very West Asian with a lot of the Anatolian_N and Iran_N:

Bildschirmfoto 2024-04-20 um 17.36.05.png


Yes, I had to use shitty G25 because I have no idea how to run admixtools on my laptop.
 
Thanks, interesting read. So they are going for a cline of CHG from the Lower Don (Krivyansky) to Lower Volga and North Caucasus for this older CHG ancestry. The second Aknashen_N ancestry they call CHG-related is a bit weird because it is very little CHG-like but already very West Asian with a lot of the Anatolian_N and Iran_N:

View attachment 16040

Yes, I had to use shitty G25 because I have no idea how to run admixtools on my laptop.
Luckily they did model it in a note:
We can model Masis Blur in the framework used here as a mixture of Aknashen Neolithic and 30.6±4.8%TUR_Marmara_Barcın_N (p=0.36) or 37.2±4.7% TUR_C_Çatalhöyük_N (p=0.42). However, we can alsomodel Aknashen Neolithic as 69.1±4.5% Masis Blur Neolithic and 30.9±4.5% CHG ancestry. The onlyfeasible for both Aknashen and Masis Blur Neolithic populations that does not involve them mutuallymodeling each other includes TUR_SE_Çayönü_PPN and CHG ancestry. Masis Blur has 13.7±3.8% CHG(p=0.13) and Aknashen has 42.0±3.7% CHG (p=0.09) ancestry according to this model. It is unclear4if thisrepresents population change during the 6th millennium BCE in Armenia or pre-existing populationstructure. That Masis Blur and Aknashen share the majority of their ancestry could be consistent with being201part of a genetically variable population that shared a language, although their genetic differentiation couldalso be consistent with language shift in the area.

But yeah this Aknashen population seems fascinating.
 
Looks like j2b l283 may have been CHG that mixed with south steppe people to become Yamnaya. They then moved to south and central europe bringing j2b l283 there and must have had a founder effect in west Balkans because it was a minor lineage among yamnaya

Indo Europeans that expanded to north and west Europe had less CHG ancestry and didn't carry j2b l283, they had also had less z2103 more L51
 

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