The Genetic Origin of the Indo-Europeans

[Archetype0ne]

snip

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.

[Capsian20]

To me it looks clear: all the living IE languages, including Armenian, Greek, and Albanian, stem from 3, and only Anatolian stems from 1. It means that after the phase 1, Pre-Proto-Anatolian branched off, and only after the phase 3, all the other branches (possibly excl. Tocharian, which is not living any longer) branched off.

so basically their take that area number 1 is the urheimat is based esclusively on the anatolian language  split.
Well anatolian language arriving from the balkans ( and later on) would have been brought by a population that was basically nearly 100% EEF so quite difficult to spot them.

Lordy these R1bV136 folks are the greatest language teachers in history. Despite being a very tiny minority among every branch of IE living languages they are considered by the authors as the sole creators of PIE. Look a lot like BS to me frankly

I know i can google, but where is R1bV136 more common today? Do we have any ancient historical population aDNA where this clade was common?

R-V1636 (arent R-V136) is rare subclade i see is spread in Europe and Near East and Central Asia
https://discover.familytreedna.com/y-dna/R-V1636/tree 

[Mitchell-Atkins]

Another quote from the paper about Corded Ware

...place of origin of the Corded Ware complex ...The Dnipro-Don area of the Serednii Stih culture fits the genetic data...

Serednii Stih=Sredny Stog

Phase II (according to Telegin, middle 3rd millennium BC) is represented by the Sredny Stog complexes of the Deriivka-Moliukhovyi Buhor type that used corded ware pottery which may have originated there.  Wiki

These sites are listed on this map from the paper (circled in red)

This is basically the area I mentioned on anthrogenica back in 2019...albeit a little north, but same river basin...and barely in the oval.
https://genoplot.com/discussions/topic/2...ll-beakers



In hindsight, based on this paper, it may be better labeled as "P310?" rather than "L51?"


While the actual SNP label locations may be a little off (imo P310 in place of L51), I think FTDNA's on the "right path" (pun intended) from Dnipro to Poland in their globetrekker tool.

[Riverman]

People just need to follow the breadcrumbs, because Yamnaya didn't use corded decorated pottery, but other Western steppe groups did, like Usatovo, Cernavoda and Cotofeni. That points to the Western complex being interrelated and in exchange, while the actual Yamnaya were intrusive and from a different subset of Sredny Stog descendants in the East.

[ANIEXCAVATOR]

There's a lot of speculation in this thread that is completely unfounded based on what the papers and the supplements say.

The authors are not saying that PIE is a "Aknashen_N language" or a "CHG language"--this is not provable using the current sampling and not even provable if there was good sampling. The authors are trying to discover the origins of the Yamnaya people, who have a little bit more Lower-Volga/N_Caucausus ancestry than even the most Lower-Volga/N_Caucausus-shifted people in the Stredny Stiih population. The transformation from Sredny Stiih to Yamnaya involved complete Y-chromosomal replacement and almost complete autosomal replacement, except for the Don Yamnaya, who admixed a little bit more with local Ukrainian-HG-shifted populations (i.e. currently sampled Sredny Stiih) and retained their I2 y-chromosomes. So the Sredny Stiih populations we have right now are not directly ancestral to Yamnaya, but Yamnaya emerged from somewhere just adjacent to them, possibly a Sredny Stiih population that is even more Lower-Volga/N_Caucausus-shifted than the most shifted ones we already have.

The populations of the Lower-Volga/N_Caucausus cline (PVgroup, BPgroup, together known as the CLV cline) all contain a fraction of Aknashen ancestry and Central Asian HG (Kelteminar-like) ancestry, but the majority of their ancestry is the traditional CHG and EHG we are familiar with. They contributed ~80% to the Yamnaya population. The earliest Corded Ware people from the Baltics who had no GAC ancestry are a little bit EHG- and WHG-shifted compared to the Yamnaya, and the later GAC-admixed CW from Germany and elsewhere got their Steppe ancestry from this slightly EHG- and WHG-shifted source, so the CW population probably came from a slightly less Lower Volga/N Caucasus-shifted version of a Yamnaya-like population. The CLV cline populations also contributed to the Central Anatolian Hittite-era population at low percentages. Because the CLV is the only think shared between Yamnaya/CW and BA Central Anatolian populations, the authors think the CLV population was PIE-speaking. You can get a Balkan population to model the BA Central Anatolian populations, but these models involve a direct admixture between the Balkan IE population + a Mesopotamian population, which is incredibly unrealistic, while the CLV model involves all the intermediate populations (a Mesopotamian population of Eastern origin + a Caucasus/Anatolian population, alongside CLV admixture).

Nowhere in this paper have they said that pre-proto-IE is a "CHG language" or a "Aknashen language" because there is no way to make a deduction that goes so far back in time--they only localize proto-IE to CLV. It is true that CLV is a very CHG-rich population with less EHG than a lot of people would have expected the PIE population to be, and that the authors' conclusions also imply that the PIE population contained ancestry from South of the Caucasus (i.e. Aknashen), but even Davidski has basically reached the same conclusions as the authors of this paper quite some time ago--the fact that something Progress-like (very CHG-rich) contributed to most of the ancestry of the Yamnaya was already widely aknowledged on his blog a number of years ago, and that small amounts of ancestry from something Nalchik-like played a role as well in Yamnaya formation (i.e. ancestry from south of the Caucasus) via its input into Progress was acknowledged several weeks ago (check his comments on the Nalchik blog post).

The authors will not disagree with us on the fact that almost all the Y-chromosomes of the Yamnaya and their ancestors must have come from the EHG and WHG-related side of their ancestry, but they're not gonna make a claim that "pre-proto-PIE is an EHG language! IE must ultimately have gone all the way back to ANE ancestors!" because the evidential base for something like that is just not strong enough to satisfy anyone, geneticists, linguists or whoever. Though us amateurs can speculate and favor that theory of course, which is in all likelihood correct, but unprovable unless we had a time machine.

The conclusions of this paper clearly contradicts the South of the Caucausus theory and the "no EHG in BA Anatolia" finding in the Southern Arc paper, so some people have clearly changed their mind. I mean, the CLV population does overlap with the circle drawn in the map on the Southern Arc paper, so technically theres less contradiction, but we all know that the most important conclusions of that paper have been dropped. However, there's been a couple of important facts that have all been harmonized by these new aDNA results. We've known for some time that qpAdm models for Yamnaya and even Progress repeatedly fail with only CHG and EHG because of excess relatedness to something Central Asian-like, and the Southern Arc paper pointed to excess relatedness to Levant_N (implying in a distal model 1% ancestry from Levant_N), and these facts stubbornly refused to go away even if the people who stated them had an agenda (e.g. Vasishtha). This new paper, by presenting the CLV population which has affinities in both directions through Aknashen and Central Asian HG ancestry, resolves all these issues nicely.

[strawberry]

So GAC ancestry is absent in Yamnaya and its this Aknashen type,

They didn't model Yamnaya using GAC or any EEF population in place of Aknashen_N or alongside Aknashen_N. Check this model from Davidski:

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

https://eurogenes.blogspot.com/2022/09/d...mnaya.html

[ANIEXCAVATOR]

The issue is that Aknashen ancestry is present *within* Progress-related ancestry (now know as CLV ancestry), not outside and in addition to that.

Davidski said...
@Vladimir

Yes, I know the model is anachronistic. But the point was just to see how much more Anatolian ancestry NL122 had compared to Progress/Vonyuchka, rather than to calculate his actual Anatolian ancestry.

Progress/Vonyuchka actually do have some Anatolian ancestry, but I can't show that yet until new Steppe Eneolithic samples are published.

Once they're available it might turn out that Progress/Vonyuchka have ancestry from a population similar to NL122.

Let's wait and see.

February 13, 2024 at 4:41 AM

NL22 is Nalchik.

Davidski said...
No, I'm missing the hunter-gatherers of the Neolithic steppe.

Progress is Steppe hunter-gatherer plus something Nalchik-like.

North Caucasus hunter-gatherers were probably just CHG.

February 13, 2024 at 5:45 PM

https://eurogenes.blogspot.com/2024/02/t...prise.html

The authors of this paper did not have access to the Nalchik genome, but that genome shows the spread of Aknashen-related ancestry started ~8-7000BP into the CLV area, explaining how the CLV population formed. Like Davidski says this is too early for a PIE homeland south of the Caucasus and does not support the Southern Arc theory, but it does capture how ancestry from south of the Caucasus did end up in Yamnaya and Yamnaya-related populations (e.g. proto-CW) via its Progress-related ancestors.

[Andar]

The 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

So GAC ancestry is absent in Yamnaya and its this Aknashen type, so basically BP Group ie Eneolithic Steppe types are the "PIE" since they are the vector of this ancestry for Indo Anatolians, is that what they are implying?  Thanks for clarifying! 

My second question is then , if Corded Ware stems from these Dnipro groups, where is the R1a?  I am astounded by the lack of R1a in the samples its overwhelmingly R1b.

Some authors of the paper still try hard to find a direct connection with Armenia when there really is none. Armenia_N is in large parts made of ancestry that didn't exist or was very low in Eneolithic Steppe (EEF/East Anatolian and North Mesopotamian) so it can't be on a cline with Progress. Rather Steppe Eneolithic and Armenia_N shared ancestry from an older Mesolithic source of a CHG-type that lived north of Caucasus.

About R1a if we look at modern R1a clades that formed in HG Era before pastoralism we see around 3-5 relatively big clades with M417 making basically 99% of modern R1a so R1a should be very small in numbers and only in 3500 BC really started to grow probably among one clan that transitioned to pastoralism around (Middle) Dnjepr region.

[old europe]

The issue is that Aknashen ancestry is present *within* Progress-related ancestry (now know as CLV ancestry), not outside and in addition to that.

Davidski said...
@Vladimir

Yes, I know the model is anachronistic. But the point was just to see how much more Anatolian ancestry NL122 had compared to Progress/Vonyuchka, rather than to calculate his actual Anatolian ancestry.

Progress/Vonyuchka actually do have some Anatolian ancestry, but I can't show that yet until new Steppe Eneolithic samples are published.

Once they're available it might turn out that Progress/Vonyuchka have ancestry from a population similar to NL122.

Let's wait and see.

February 13, 2024 at 4:41 AM

NL22 is Nalchik.

Davidski said...
No, I'm missing the hunter-gatherers of the Neolithic steppe.

Progress is Steppe hunter-gatherer plus something Nalchik-like.

North Caucasus hunter-gatherers were probably just CHG.

February 13, 2024 at 5:45 PM

https://eurogenes.blogspot.com/2024/02/t...prise.html

The authors of this paper did not have access to the Nalchik genome, but that genome shows the spread of Aknashen-related ancestry started ~8-7000BP into the CLV area, explaining how the CLV population formed. Like Davidski says this is too early for a PIE homeland south of the Caucasus and does not support the Southern Arc theory, but it does capture how ancestry from south of the Caucasus did end up in Yamnaya and Yamnaya-related populations (e.g. proto-CW) via its Progress-related ancestors.

but progress can be modeled as having ancestry from the Dneper Don foragers. So the don cluster could have expanded to the north caucasus before progress expanded into Ukraine

Target: RUS_Progress_LNG2004
Distance: 4.8991% / 0.04899101 | R5P
58.8 Russia_N_Golubaya_Krinitsa_Lower_Don
23.2 TJK_Tutkaul_Meso
9.6 IRN_Ganj_Dareh_Meso
8.4 ARM_Aknashen_N

arget: Ukraine_Eneolithic_CernavodăI_KartalA:KTL001.merge
Distance: 3.9577% / 0.03957715 | R5P
39.0 Russia_N_Golubaya_Krinitsa_Lower_Don
21.4 ARM_Aknashen_N
21.2 Baltic_LTU_meso
10.8 BGR_MP_N
7.6 TJK_Tutkaul_Meso

Target: UKR_Deriivka_En:ukr104
Distance: 1.7624% / 0.01762434 | R5P
63.6 Russia_N_Golubaya_Krinitsa_Lower_Don
14.2 UKR_Deriivka_N
8.0 IRN_Tepe_Abdul_Hosein_Meso
7.8 Hungary_N
6.4 RUS_Karelia_HG_Meso

Target: UKR_LN_Sredni_Stog_En_Igren_Dnepr-4200BC
Distance: 3.6107% / 0.03610693 | R5P
69.0 Russia_N_Golubaya_Krinitsa_Lower_Don
13.4 TJK_Tutkaul_Meso
9.6 ARM_Masis_Blur_N
4.6 AZE_Caucasus_lowlands_LN
3.4 ARM_Aknashen_N

Target: Ukraine_LN_Usatovo_Majaky:MAJ023.merge
Distance: 1.9711% / 0.01971095 | R5P
39.0 ARM_Aknashen_N
25.2 BGR_MP_N
15.6 Russia_N_Golubaya_Krinitsa_Lower_Don
14.6 RUS_Vologda_Veretye_Meso
5.6 TJK_Tutkaul_Meso

Target: RUS_Vonyuchka_LN:VJ1001
Distance: 5.2306% / 0.05230616 | R5P
57.6 Russia_N_Golubaya_Krinitsa_Lower_Don
24.6 IRN_Ganj_Dareh_Meso
10.4 TJK_Tutkaul_Meso
7.4 ARM_Aknashen_N

Target: RUS_Khvalynsk_LN:I11837
Distance: 3.3479% / 0.03347887 | R5P
49.8 Russia_N_Golubaya_Krinitsa_Lower_Don
15.4 TJK_Tutkaul_Meso
15.0 RUS_Vologda_Veretye_Meso
13.2 RUS_Karelia_HG_Meso
6.6 IRN_Ganj_Dareh_Meso

[Sephesakueu]

So GAC ancestry is absent in Yamnaya and its this Aknashen type,

They didn't model Yamnaya using GAC or any EEF population in place of Aknashen_N or alongside Aknashen_N. Check this model from Davidski:

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

https://eurogenes.blogspot.com/2022/09/d...mnaya.html

Davidsky really? Do you think that northeast euro ethno nationalist will embrace his   Aknashen    mesopotamian ancestry ?

[old europe]

Every ancient post neolithic european population can be modeled with this. And this a figure from a peer reviewed paper

[ANIEXCAVATOR]

The issue is that Aknashen ancestry is present *within* Progress-related ancestry (now know as CLV ancestry), not outside and in addition to that.

Davidski said...
@Vladimir

Yes, I know the model is anachronistic. But the point was just to see how much more Anatolian ancestry NL122 had compared to Progress/Vonyuchka, rather than to calculate his actual Anatolian ancestry.

Progress/Vonyuchka actually do have some Anatolian ancestry, but I can't show that yet until new Steppe Eneolithic samples are published.

Once they're available it might turn out that Progress/Vonyuchka have ancestry from a population similar to NL122.

Let's wait and see.

February 13, 2024 at 4:41 AM

NL22 is Nalchik.

Davidski said...
No, I'm missing the hunter-gatherers of the Neolithic steppe.

Progress is Steppe hunter-gatherer plus something Nalchik-like.

North Caucasus hunter-gatherers were probably just CHG.

February 13, 2024 at 5:45 PM

https://eurogenes.blogspot.com/2024/02/t...prise.html

The authors of this paper did not have access to the Nalchik genome, but that genome shows the spread of Aknashen-related ancestry started ~8-7000BP into the CLV area, explaining how the CLV population formed. Like Davidski says this is too early for a PIE homeland south of the Caucasus and does not support the Southern Arc theory, but it does capture how ancestry from south of the Caucasus did end up in Yamnaya and Yamnaya-related populations (e.g. proto-CW) via its Progress-related ancestors.

but progress can be modeled as having ancestry from the Dneper Don foragers. So the don cluster could have expanded to the north caucasus before progress expanded into Ukraine

Target: RUS_Progress_LNG2004
Distance: 4.8991% / 0.04899101 | R5P
58.8 Russia_N_Golubaya_Krinitsa_Lower_Don
23.2 TJK_Tutkaul_Meso
9.6 IRN_Ganj_Dareh_Meso
8.4 ARM_Aknashen_N

arget: Ukraine_Eneolithic_CernavodăI_KartalA:KTL001.merge
Distance: 3.9577% / 0.03957715 | R5P
39.0 Russia_N_Golubaya_Krinitsa_Lower_Don
21.4 ARM_Aknashen_N
21.2 Baltic_LTU_meso
10.8 BGR_MP_N
7.6 TJK_Tutkaul_Meso

Target: UKR_Deriivka_En:ukr104
Distance: 1.7624% / 0.01762434 | R5P
63.6 Russia_N_Golubaya_Krinitsa_Lower_Don
14.2 UKR_Deriivka_N
8.0 IRN_Tepe_Abdul_Hosein_Meso
7.8 Hungary_N
6.4 RUS_Karelia_HG_Meso

Target: UKR_LN_Sredni_Stog_En_Igren_Dnepr-4200BC
Distance: 3.6107% / 0.03610693 | R5P
69.0 Russia_N_Golubaya_Krinitsa_Lower_Don
13.4 TJK_Tutkaul_Meso
9.6 ARM_Masis_Blur_N
4.6 AZE_Caucasus_lowlands_LN
3.4 ARM_Aknashen_N

Target: Ukraine_LN_Usatovo_Majaky:MAJ023.merge
Distance: 1.9711% / 0.01971095 | R5P
39.0 ARM_Aknashen_N
25.2 BGR_MP_N
15.6 Russia_N_Golubaya_Krinitsa_Lower_Don
14.6 RUS_Vologda_Veretye_Meso
5.6 TJK_Tutkaul_Meso

Target: RUS_Vonyuchka_LN:VJ1001
Distance: 5.2306% / 0.05230616 | R5P
57.6 Russia_N_Golubaya_Krinitsa_Lower_Don
24.6 IRN_Ganj_Dareh_Meso
10.4 TJK_Tutkaul_Meso
7.4 ARM_Aknashen_N

There is nothing against the general point that EHG-related populations, specifically Don and Ukraine-related, could have contributed to the CLV population--the authors of course also aknowledge this. Check page 164 of the supplementary material.

pegasus


Some authors of the paper still try hard to fund a connection with Armenia when there really is none. Armenia_N is in large parts made of ancestry that didn't exist or was very low in Eneolithic Steppe (EEF/East Anatolian) so it can't be on a cline with Progress. Rather Steppe Eneolithic and Armenia_N shared ancestry from an oldef Mesolithic source of a CHG-type that lived north of Caucasus.
About R1a if we look at modern R1a clades that formed in HG Era before pastoralism we see around 3-5 relatively big clades with M417 making basically 99% of modern R1a so R1a should be very small in numbers and only in 3500 BC really started to grow probably among one clan that transitioned to pastoralism around (Middle) Dnjepr region.

The new samples in this paper and Nalchik disprove this directly. The CLV cline runs between South of Caucasus populations / Aknashen/Maikop and Lower Volga/Caspian populations like Remontnoye and Berezhnovka, so Aknashen-related ancestry was quite extensively present and Progress was only a part of it.

[ANIEXCAVATOR]

There is a lot of confusion in this thread: the issue is that Progress is now the population that carries Aknashen-related ancestry. It is not that Yamnaya contains Aknashen-related ancestry *recently*, it is that some Eneolithic populations contemporary with Sredny Stog and just before the Yamnaya genesis, like Progress, were part of a cline of populations, known as CLV populations, that run in a line between the Lower Volga area and populations from South of the Caucasus and so carry ancestry from South of the Caucasus. Progress-related populations (in this paper Remontnoye and Berezhnovka) that produce the best fits for Yamnaya when the other source is Sredny Stog already contain this South of Caucasus ancestry.

This is what Davidski agrees with in his replies. The other models showing that no South of Caucasus ancestry is needed in Yamnaya when Progress is included doesn't contradict the authors of this paper, because Progress-related ancestry *already* contains the Aknashen-related ancestry, which was present in a large number of Eneolithic pre-Yamnaya samples from the Lower Volga area down to the Northern Caucasus and which also admixed into the Sredny Stiih people already (who form a cline between Ukraine Neolithic and the CLV populations).

On this point none of the G25 models disagree either. This South-of-Caucasus ancestry is present already in Progress (alongside Central Asian Tutkaul-related ancestry) in all of the G25 models posted on this thread, even when the other half of their ancestry is from Don foragers, implying that it is present in Yamnaya as well.

[pegasus]

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

So GAC ancestry is absent in Yamnaya and its this Aknashen type, so basically BP Group ie Eneolithic Steppe types are the "PIE" since they are the vector of this ancestry for Indo Anatolians, is that what they are implying?  Thanks for clarifying! 

My second question is then , if Corded Ware stems from these Dnipro groups, where is the R1a?  I am astounded by the lack of R1a in the samples its overwhelmingly R1b.

Some authors of the paper still try hard to fund a connection with Armenia when there really is none. Armenia_N is in large parts made of ancestry that didn't exist or was very low in Eneolithic Steppe (EEF/East Anatolian) so it can't be on a cline with Progress. Rather Steppe Eneolithic and Armenia_N shared ancestry from an oldef Mesolithic source of a CHG-type that lived north of Caucasus.

About R1a if we look at modern R1a clades that formed in HG Era before pastoralism we see around 3-5 relatively big clades with M417 making basically 99% of modern R1a so R1a should be very small in numbers and only in 3500 BC really started to grow probably among one clan that transitioned to pastoralism around (Middle) Dnjepr region.

Yeah that make sense for R1a its amazing how rapidly it just exploded  but is very low prior to 3500 BCE. Still its so weird to see how few samples carry it before then. 

Wrt to Armenia_N , Lazardis is very firm , this ancestry was picked up by the BPgroup and this group admixed with UNHG , which already has 8-10% EEF it seems.  It helps explain away the Indo Anatolian connection though using the BP group.  I think the coordinates are needed , the labels are confusing to me.

[ANIEXCAVATOR]

People just need to follow the breadcrumbs, because Yamnaya didn't use corded decorated pottery, but other Western steppe groups did, like Usatovo, Cernavoda and Cotofeni. That points to the Western complex being interrelated and in exchange, while the actual Yamnaya were intrusive and from a different subset of Sredny Stog descendants in the East.

Agree, great observation.

Something people who insist on drawing the genesis of the Yamnaya-like/CW-like populations to the West (on the boundaries of the Ukraine HGs and European farmers), instead of more Eastwards like this paper does (where the PIE population emerges from the interaction of Volga-N Caucasus CLV populations with Dnepr-Don HGs) would have a really hard time explaining is the shape of the three clines on the Steppe at this time. 

There's a cline from Upper Volga forest cultures to lower Volga peoples on the Steppe (running through Khvalynsk etc), another cline from Caucasus to the lower Volga (the CLV cline running through Maykop, Progress etc), and a third cline linking Ukrainian foragers on the Dnepr and Don to these two clines on the east. These clines contain tens or hundreds of samples from many sites, indicating that they are the product of long-term interaction. The Yamnaya and Yamnaya-like populations like the earliest CW all fit neatly on one place in the clines (the far Eastern end of the third cline). Something that doesn't exist is clines linking European farmers from Trypillia or Euro Farmer-HG mixes to later populations like Yamnaya. There are sporadic pulses of ancestry from the East that we can see in the Nitikin paper, for example the Csongrad individual, or the Eastern ancestry found throughout Trypillia. But there's no cline that connects them to later Yamnaya and Yamnaya-like populations.

Ukraine HGs were definitely involved in Yamnaya formation through the Y-chromosomes and through their ancestry in Sredny Stog and potentially CLV populations. But this paper shows that the immediate ancestors of the Yamnaya, and the place where Yamnaya formed was quite far from where Ukraine HGs and Euro farmers had their stronghold.