The Genetic Origin of the Indo-Europeans

[Archetype0ne]

Table S 44 A unified model of Indo-Anatolian and Indo-European origins from the supplement

[Jaska]

Ebizur:

I wonder what the climate in the Northeast Caucasus/Lower Volga region was like during the PIE era. - - Is that really the sort of region from which one should expect a major language family to have expanded?

I do not see how the environment is relevant. It is not the nature of the original homeland but the social factors, military technology etc. which determine which language prevails over others. Harsh or arid conditions might push masses moving, but how their language fares when they meet and admix with other folks is a totally different question.

[Archetype0ne]

Some other interesting bits:

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 .

[Moeca]

Ipatovo 3 ( 5400 yBP - Early Bronze Age ) Early Steppe Maykop Culture

IV3002 ( 5383 ± 64 yBP )
Kurgan: 2
Phase: 1st
Grave: 187 ( Founding grave of the entire mound )
Other dates: 5058 ± 223 yBP / 5328 ± 251 yBP / 4630 ± 50 yBP (radiocarbon)
Y-DNA: T1-CTS6004 (xT1a1a-CTS484, T1a1b-Y6031, T1a2b-FGC37316) Probably: T1a2a-Y8614
mtDNA: X1'2'3
Age at Death: 35-45
Burial: The first phase (burial 187) may have been an Early Maikop Culture burial, destroyed by later graves. An oval pit covered by stone slabs. No inventory was discovered.
Other IDs: IV3002.A0101 / "BZNK-293/1" / (ГИН-10297), (ГИН-10953) / GIN-10297
Autosomal notes: Steppe group 2, not closely related to Maykop nor Yamnaya.

[TanTin]

Why do you think that language would rather follow paternal lineages than autosomal ancestry components? There is no law of nature supporting that.

We do not know from which of the ancestry components the Indo-European language lineage was originally inherited. Surely we cannot ignore the fact that Europe was full of Paleo-European language families before the Indo-European and the Uralic expansions. These lost languages would be more likely associated with EHG and WHG ancestries, don't you think?

There are some laws for the pathernal or patriarchal type of society.
1. They do not marry between themselves,  they have to   find their women from outside to avoid  incest..
2. The older men and women have higher status in such communities. So the new arrived woman will need to accept the language of her husband,  the mother and the brothers.
3. The daughters in IE community will not stay in the same house with their parrents - daughters will have to go outside and live in other houses - in the same vilage or elsewhere.
4. The Mother of the man has still lot of power, especially to control the wifes of her son,  to delegate tasks, to control the home and to educate the kids.
5. One man may have several women (wifes).  But the power of old woman is still there, doesn't matter how many wifes .

The mens in IE society are also some kind of solders, so they may need to go and fight.. But the home of IE family is generally controlled by the mother of such man. So this is the reason why the language is the same - because the mother of the man is controlling all.

Next rule:  all the sons stay in the same home with their parrents util the death of the Oldest man: who is the patriarch. 
Once the oldest man is dead - all the sons will have to go out and to establish their own homes.
Only the older son is allowed to stay in their mother's house and the oldest son is taking care for his old mother..

So these are the basic rules for the IE family,  I may have some mistakes in discribing this.  And there could be some differences from one IE group to another, but many of these rules were same or similar for the IE people..  And we know, that most of other European before IE were Matriarchal families.

In general these patriarchal system was very stable and IE communities grew faster compared to the others.

[Moeca]

Why do you think that language would rather follow paternal lineages than autosomal ancestry components? There is no law of nature supporting that.

We do not know from which of the ancestry components the Indo-European language lineage was originally inherited. Surely we cannot ignore the fact that Europe was full of Paleo-European language families before the Indo-European and the Uralic expansions. These lost languages would be more likely associated with EHG and WHG ancestries, don't you think?

There are some laws for the pathernal or patriarchal type of society.
1. They do not marry between themselves,  they have to   find their women from outside to avoid  incest..
2. The older men and women have higher status in such communities. So the new arrived woman will need to accept the language of her husband,  the mother and the brothers.
3. The daughters in IE community will not stay in the same house with their parrents - daughters will have to go outside and live in other houses - in the same vilage or elsewhere.
4. The Mother of the man has still lot of power, especially to control the wifes of her son,  to delegate tasks, to control the home and to educate the kids.
5. One man may have several women (wifes).  But the power of old woman is still there, doesn't matter how many wifes .

The mens in IE society are also some kind of solders, so they may need to go and fight.. But the home of IE family is generally controlled by the mother of such man. So this is the reason why the language is the same - because the mother of the man is controlling all.

Next rule:  all the sons stay in the same home with their parrents util the death of the Oldest man: who is the patriarch. 
Once the oldest man is dead - all the sons will have to go out and to establish their own homes.
Only the older son is allowed to stay in their mother's house and the oldest son is taking care for his old mother..

So these are the basic rules for the IE family,  I may have some mistakes in discribing this.  And there could be some differences from one IE group to another, but many of these rules were same or similar for the IE people..  And we know, that most of other European before IE were Matriarchal families.

In general these patriarchal system was very stable and IE communities grew faster compared to the others.

The language of a new child is always the language of the mother ................only if the father makes an appearance and stays there is a chance that the child will learn the father language
But history shows , most men did not stay

[Riverman]

The language of a new child is always the language of the mother ................only if the father makes an appearance and stays there is a chance that the child will learn the father language
But history shows , most men did not stay

That's not true for the past, the men did not just stay, but they formed the society. It wasn't the individual choice of a mother or child which language was spoken. The most you can get out of it, from the female substrate side of things, is that for a short period of time there was a bilingual environment and some influences from the substrate made it to the language of the ruling elites and males.
Anything else is theoretically possible, and we know of cases in which the bilingual state lasted longer or the females had more influence, but whenever that happened, it was truly exceptional and is therefore very rare.

The PIE society was even agnatic, which means that the female ancestry didn't count in their general system. The male lineage determined ancestry and the status of relatives primariily.

The best argument for PIE being an EHG language is that out of the main three male lineages, 3 out of 3 are derived from them and not the Caucasian side or the assimilated people of the West.

When in later periods some Westenr lineages gained prominence, like I-M253, E-V13 etc., the Indoeuropeanisation of the populations was already, by and large, finished and it was a rise to prominence from within an IE speaking society.

One EHG lineage means nothing, might be chance, but 3 out of 3 from the PIE stage, that's significant.

[old europe]

Table S 44 A unified model of Indo-Anatolian and Indo-European origins from the supplement

now I see their hypothesis. Pay attention everybody. Basically what they are saying is that WHG/EHG on the Dneper Don and even full fledged EHG on the Volga were not PIE. For them PIE starts when the CHG/TTK kicks in to form the northern caucasus-lower Volga cline. So the cluster that makes PIE is the CHG/TKK
so what the  paper is all about is a revisited "southern arc hypothesis" For them PIE is born north of the caucasus but thanks to a southern caucasian influence.
The news about the haplo discoverd are finer and they did a real good job. The conclusion of the paper is a total garbage

[Archetype0ne]

They leave all scenarios open, but claim based on the current evidence that hypothesis B is a bit more likely. I highly suggest you read the last 20-40 pages of the supplement, everything will become clearer.
The way I understood some of the rationale, is that BPgroup is mix of Khvalyns + Caucasus. And BPgroup + SShi (Sredni Stih) gave raise to Core Yamnaya. "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." And BPgroup inflow is necessary to model Anatolian BA. 

Between the North Caucasus piedmont and Lower Volga population, mixtures were taking place during the 5 th and 4th millennia BCE between populations with ancestry characteristic of the Neolithic and Eneolithic Caucasus (represented by Aknashen and Maikop) and Lower Volga-North Caucasus Eneolithic populations (represented by BPgroup), both of which had substantial proportions of CHG ancestry from the earlier hunter-gatherer periods. Steppe ancestry was present in the later Maikop (Table S 9) and Remontnoye (Table S 6, Table S 7) individuals from this area. It was also present further south in Chalcolithic Armenia at Areni-1 cave, but there it was added to a different, Masis Blur Neolithic population (Table S 11). 5 Thus, the Caucasus area was seeing admixture in both south-north and north-south directions: Remontnoye had Aknashen Neolithic/Maikop ancestry and Maikop and Armenian Chalcolithic had steppe ancestry.

People out of this Caucasus-Lower Volga admixture zone people flowed outwards: along the Volga where all the “southern” ancestry can be well explained as of BPgroup origin alone (Table S 14) and into the Don-Dnipro area where it interacted with the descendants of hunter-gatherers of the Dnipro-Don forming the Serednii-Stih cline. 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.

We have thus argued that Yamnaya was formed when people of “eastern” Aknashen-Maikop/BPgroup origins (of proximate Remontnoye-related origins) moved westward and admixed with people of the Serednii Stih culture. Motivated by this, we tested whether we could use this framework to jointly model all these populations involved in our reconstruction:

Observe also the previously mentioned contrast between Eneolithic populations of the Don-Volga who did not require Aknashen/Maikop ancestry but can be modeled with BPgroup/PVgroup ancestry alone (Fig. S 2, Fig. S 3) with those of the Serednii Stih culture of the Don-Dnipro area that also had Aknashen-related ancestry (Fig. S 9).

In Table S 27 that only models involving Aknashen_B/Azerbaijan_C and BPgroup/PVgroup and Igren_o ancestries have no losses in the tournament, the full results of which are shown in Table S 28. While no unique solution emerges out of this tournament, it is useful to weigh our confidence in the different models. With the knowledge that we have only partially sampled the genetic variation of the Caucasus, Lower Volga, and Dnipro-Don areas, it is nonetheless interesting that the 3-way models directly recapitulate the conclusions we reached by exploring 2-way models: that all three of these areas contributed to the formation of the Yamnaya.

Finally, we plot in Fig. S 11 the proportions of Table S 29 which visually demonstrates all our inferences. The contrast between the Don-Dnipro and Volga in terms of Ukraine_N vs. EHG ancestry; the presence of Aknashen ancestry in Maikop, Remontnoye, the Serednii Stih, and the Yamnaya, but not in Golubaya Krinitsa or the Volga; and the contribution of a population like BPgroup of ancestry in populations across the entire region except the far north of eastern Europe (Karelia) and starting from at least the mid-6 th millennium date of the Golubaya Krinitsa individuals, one of which has it (GK1) while the other one does not (GK2).

Hypothesis B harmonizes with all known facts and the results of our reconstruction of Yamnaya origins strengthen it, as the Yamnaya do indeed have ancestry from the south: both early ones via their BPgroup ancestors which experienced gene flow from the Caucasus and contributing to the Serednii Stih and Volga clines; but, also later ones via the migration of Remontnoye-related people (who also had Maikop/Aknashen ancestry). What was only indistinct before (the CHG ancestry in the Eneolithic steppe and the extra Anatolian-Levantine ancestry in the Yamnaya4 ) has now come into better focus.

Hypothesis B: a Caucasus-West Asian origin of Indo-Anatolian origins is strengthened by the finding of early migrations from the Caucasus into the Volga/Don-Dnipro Eneolithic populations followed by later Maikop/Armenian Neolithic ancestry into the ancestors of the Yamnaya. This 214 hypothesis also maps to the transformation of Chalcolithic and Bronze Age central/western Anatolia which saw half to all its Neolithic population replaced.

It can all get very confusing. Had to read the analysis part multiple times, and still not sure I understand it right.

[AimSmall]

Ok guys.... let's not repeat and repost every post every time. Edit your replies to thin them out. Don't need the last five posts included in every reply.

[Ebizur]

Ebizur:

I wonder what the climate in the Northeast Caucasus/Lower Volga region was like during the PIE era. - - Is that really the sort of region from which one should expect a major language family to have expanded?

I do not see how the environment is relevant. It is not the nature of the original homeland but the social factors, military technology etc. which determine which language prevails over others. Harsh or arid conditions might push masses moving, but how their language fares when they meet and admix with other folks is a totally different question.

The environment determines the carrying capacity of a territory.

The carrying capacity of a territory limits the manpower that can be mustered by the inhabitants to defend themselves from invasion or to invade foreign territories.

I do not know what the environment in the area between the Northeast Caucasus and the Volga River has been like during the time when the Proto-Indo-European language has been spoken. However, if you asked me to consider the likelihood that any population of present-day Europe might expand their territory and spread their language over the entire (sub)continent in the future, the Kalmyks would not be the first people to come to mind.

[Jerome]

Progress was predominantly EHG:

Target: RUS_Progress_EnG2004
Distance: 4.6770% / 0.04676998
48.8 RUS_Samara_HG
24.4 TJK_Sarazm_En
21.0 GEO_CHG
3.6 RUS_Tyumen_HG
2.2 UKR_Meso

Target: RUS_Progress_EnG2001
Distance: 4.2178% / 0.04217797
45.4 RUS_Samara_HG
26.0 GEO_CHG
23.8 TJK_Sarazm_En
3.6 ARM_Aknashen_N
1.2 RUS_Tyumen_HG

Sredny Stog and Yamnaya were mostly Progress with some local admixture:

Target: Ukraine_Eneolithic_Sredni_Stog
Distance: 3.7253% / 0.03725306
62.2 RUS_Progress_En
19.4 UKR_Meso
10.6 UKR_Globular_Amphora
4.8 TJK_Sarazm_En
3.0 RUS_Tyumen_HG

Target: Ukraine_Eneolithic_Post-Sredni_Stog
Distance: 3.3464% / 0.03346432
83.6 RUS_Progress_En
13.2 UKR_Trypillia
3.2 UKR_Meso

Target: Yamnaya_Samara
Distance: 2.9344% / 0.02934434
79.0 RUS_Progress_En
11.8 UKR_Meso
9.2 UKR_Trypillia

The paper, if I understand correctly, claims that it was the Progress/Vonyuchka type people (the CLV cline) who spoke Basal PIE (i.e. PIE before the Anatolian branch split and headed for central Anatolia), making the N Caucasus/Lower Volga region, NOT Sredny Stog/Dnieper-Don, the overall PIE urheimat. I'll have to read the paper first, though, heh. I wonder if they explained that Central Asian ancestry in IE speaking nomads.

Actually on Qpadm modelling there's a lot of WSHG that dosent appear on g25,or using sarazm cuts it out.
Here's a model with mesohoko/seh Gabi,the ratio of EHG/WSHG is like 65/35

[TanTin]

The language of a new child is always the language of the mother ................only if the father makes an appearance and stays there is a chance that the child will learn the father language
But history shows , most men did not stay

You can not prove this. We live in 21 century, now the reality is very different from what was the life 5000 years ago. The family of today is not what was a family 5000 years ago.
People in the past lived in small houses, living for example 5 families together under the same roof.
The women could be from different origin, speaking different languages, but the language in the house is the language of the oldest people: the man ( patriarch) and the old woman ( grand-mother).
The old man is the master for the work on the field, for the battle or for other tasks outside the home. The old woman was the master in the home..
This system of relations survived almost until 19 century. We may report such relations in many IE countries. Same system survived also in some countries which are not IE..
So the language was not really the leading principle.. There was such type of relationship, social structure, which helped to keep the language and all the rules for very long time.
Some of the IE people used collective system to take care for the kids. All the adults were busy with work and survival tasks. Old women were mainly responsible to take care for their grand-kids..
And this system also survived until 19 century.. We still may find some remains of such relations . But it almost dissapeared.
So this is to explain why the language of the mother was not necesserily transferred to the kids. Because if we have 10 mothers under the same roof, if all these people stay all the time together, if they speak together, eat and sleep together - there is almost no time when a single mother will stay isolated and speak to her son or daughter.. People in the past did not have their own room, people in the past did not have their own privacy.. They lived in the communities. And the community was rulled by the oldest people .. Until the dead of the old man.. When the old man ( patriarch ) is dead - the rule was : to separate the sons and to have them create their own new houses where the same social system was reproduced..
In general the rules were not too complicated, same rules can bee seen from India to all across Europe.
It is imposible to apply the rules of today and to make conclusions for the past time.. You need to know a lot more for the past to be able to make any valid conclusions.

[Jerome]

The only good thing is that they confirm the WSHG/central Asian Ancestry in Progress/Yamnaya/IE.

I think this Ancestry is from kelteminar people of urals/tobol river and not from TTK_Proper.

Kelteminar was likely TTK like( a mix of 25+35% iran_n/jeitun + ANE)

Also,It's interesting that the ANE ancestry we see in those maykop guys and kumsay guys,is likely from the same people that gave ANE to Progress.

[Jerome]

What they seem to be saying is that EHG isn't pie but Something like aknashen/Mesopotamian+Kelteminar/TTK+CHG+EHG is PIE.?

I think what they are trying to do is just mix up/fit the southern arc with david anthony's hypothesis of Volga -caucasus and Sredny

They are just trying to keep all parties happy,so they just mix it all up and force-fit theories.