The spread of 'Steppe' DNA and autosomal best-fit analysis

The L51+ bottleneck, if you will, could be due to the Yersinia pestis outbreak that caused the downfall of the Neolithic farming communities across Europe (including non agriculturalists like R1b). I don't see why it has anything to do with France, in fact, the data has shown L51+ is well distributed across Europe and isn't linked to any specific region. L11+/P312+/U106+ would all be post-plague and in north-central Europe.

Are we sure Y-R1b-L51 knew a bottelneck phenomenon? What we a re sure is that it knew a founder effect, what doesn't everytime imply a previous bottelneck: only mutation a some stage (of L23 lineage here)... And some new discoveries in the northern southbaltic plain of Europe could provide us surprises.
I agree with you at a global level (the "bottleneck" theory isn't yours seemingly; I have to date no opinion about its reality); Concerning Balkans for L51 I'm not sure, maybe rather NW Ukraine not too far from N-E Carpathians? only a bet for the game.
 
The phylogeny points broadly to North of the West Med. (as I'll explain), but I associate it with Los Millares as that matches the profile we know from the Central Beakers perfectly (warlike, metallurgical (first in W. Europe), caste-like elite, that also was part of the Bell Beaker phenomenon later on).

So (I'm posting it as I've written it up twice on anthrogenica, just because so many get confused by it it seems):The first subclade to break away from L51 not related to L52, Z2118, dates back to only 400 years after the formation of L51 (5700 years ago, so well before the migrational period of L51 Beaker folk across Western and Central Europe). The men with this subclade, in modern times, are distributed mostly around Southern France and the Rhône region. Why is that the case, if not for that general area being L51's homeland? Why, during the great Beaker migrational period, would already differentiated Z2118 men "choose" to migrate to Southern France and not throughout the rest of Western and Central Europe? It would be like time-travelling to just before the great migrational period of the Beaker folk, marking those carriers of the subclade Z2118, and seeing that the vast majority migrated to that region North of the West Med. - that is ridiculously unlikely!

An Eastern European origin of L51 would require those with branches that split at an earlier date before the great migrational period of the Beaker folk (i.e. Z2118) to have preferentially, for some reason, migrated to the vicinity of the South of France, and not elsewhere, DESPITE having been present at the earliest stages in L51's Urheimat. It would be like travelling back in time to just before the supposed great migration of L51 Westwards from E. Europe, marking those who carried this haplogroup, and seeing that the vast majority of them ended up in Southern France and the areas nearby and not so much elsewhere. There IS no reason for that, there can't be!

I have little doubt you are right, and that the phylogenic evidence clearly indicates the common ancestors of extant L51 most recently lived in Western Europe, most likely France. I am intrigued by the less clear issues of how, when and why they got there.

There is some circumstantial yDNA, mtDNA and aDNA evidence pointing to possible associations with Southern Poland, Chalcolithic Bulgaria, Cucuteni Triploye, Globular Amphora and RRBP, so I feel the most likely route is mid/late millennium BC up Moldova and across Southern Poland and South/Central Germany; but I have not seen anything that looks conclusive.
 
Other than the French hotspot of L51, the track can be deduced by brother and uncle clades spanning with interesting frequencies in Albania or Italy, it's like a track of a migrating clan that I suppose were Neolithic herders coming from Anatolia.
 
That's interesting for me, because I can see my way of reasoning differs quite a lot from yours and that of other people. In my opinion, a calculated result of 96% of similarity between a R1a-M417 CWC individual dated to ~2500 B.C. in Germany and a R1a-M47 ~4000 B.C. sample in North Ukraine should raise a red flag because it's just possible, but unlikely and not that plausible especially considering the archaeological evidence that that was not an established culture with a fantastic population continuity, but rather a "new" and very expansionist culture that spread through previously settled territories and had its origins (as far as German CWC is concerned) nearly 1000km away - and all of that is assuming that they'd have mixed with no population along the way since the beginning of CWC (~2900 B.C.) and that they would've remained totally untainted by the Yamnaya expansion (~3.300 B.C. onwards) and by any earlier migration and admixture event from ~4000 B.C. to ~2900 B.C. I will accept those results if they end up being correct, but it'll be a big surprise for me from a linguistic (CWC IE branches supposedly much more divergent from other IE branches than previously thought, with the split dating to before 4000 B.C.) and archaeological point of view, and even from my own common sense. But for now I think a~65% match of Yamnaya looks much more reasonable at first sight, especially if you consider the possibility that the root of CWC was itself just a Yamnaya-ized and thus probably Yamnaya-admixed Sredny Stog population (and, as Dereivka shows, Sredny Stog already had at least some groups with a higher percentage of EEF ancestry).

I don't see any significant sign of Yamnayan admixture in core Corded Ware populations. Neither do I see much genetic input from core Sredy Stog, which was heavily WHG before it was largely replaced by Yamnaya, which itself withered a millennium or so later.

Combining 65% Yamnaya and 35% Dereivka Sredny Stog gives a projected autosomal reading for German CW that has 20 times greater variance from actual CW than that projected by the best-fit combination. To me, this does not look like a very reasonable match.

However, CW does not appear to have been totally untainted by Yamnaya at all. Its North Eastern corner shows strong signs of CW-Yamnaya admixture; and as this was the only CW population that appears to have thrived in Europe following Corded Ware's demise, I would say that the Yamnayan genetic footprint is probably greater in modern R1a-M417 populations than in most R1a-M417 CW populations.
 
But Suvorovo-Novodanilovka settlements have always been understood by archaeologists as intrusive (and accompanied by contemporaneous signs of widespread destruction and sociocultural rupture, with abandonment of tell settlements and other evidences of displacement/depopulation) in the Balkans, wasn't it? The traditional understanding of the culture's original movement would also fit with the gradual growth of steppe ancestry in the Balkans even during the Late Neolithic and Early Chalcolithic (the Varna outliers amidst the average Varna EEF individuals, and seemingly steppe-derived imports even before Suvorovo indicate that east-west movements already happened), as well as explain the particular "archaeological circumstances" in which Suvorovo sites appear in the Balkans, which arguably smell of invasion or a sudden and disruptive migration. As for your last point, I'll only say that genetics is awesome, but without archaeology, linguistics and other sources of knowledge the interpretation of the myriad of genetic data can become quite troublesome and even misleading.

Yes, Suvorovo might well have been predominantly intrusive and invasive, but there are also some signs of collaboration and admixture (perhaps a bit like Vikings turning into Normans?). I would say Varna looks to have witnessed some of the first signs of Suvorovo, rather than a predecessor of it. After the collapses of the Balkan Neolithic, Suvorovo populations look to have taken away with them both some DNA and some of what they had learned from places they had 'invaded' - we know they migrated to Central Ukraine (where they would have look genetically rather like M417 Corded Ware) and probably through Moldova to the Upper Dniester (where, upon admixture with some Globular Amphora, they would have looked genetically rather like L51 Bell Beaker).

Merging this analysis with what we think we know from archaeology, linguistics and other disciplines can sometimes assist, but can also sometimes muddy the clearer genetic waters.
 
Are we sure Y-R1b-L51 knew a bottelneck phenomenon? What we a re sure is that it knew a founder effect, what doesn't everytime imply a previous bottelneck: only mutation a some stage (of L23 lineage here)... And some new discoveries in the northern southbaltic plain of Europe could provide us surprises.
I agree with you at a global level (the "bottleneck" theory isn't yours seemingly; I have to date no opinion about its reality); Concerning Balkans for L51 I'm not sure, maybe rather NW Ukraine not too far from N-E Carpathians? only a bet for the game.
Extant L51 shows bottleneck signs now. It might not have been bottlenecked at the time; it might be that all the other strands of it died out some time later, but its absence from early archaeological samples and in matching archaeological aDNA would suggest a likely bottleneck at the time as well, I think.

Extant L51's origin might well have been NE Carpathian, but if so - it looks like the East Balkan samples with Steppe DNA would likely have been close relations of it.
 
L51 from France ??? Until 2600 BC, R1b is nowhere to be seen in western Europe, save a few outliers who had probably come along with the "farmers". After 2500 BC, they suddenly show up all over central and western Europe. Either L51 had been in Hungary since the old Suvorovo pocket for centuries, or they came along with Yamna Danube.

Sample ID / genetic sex (M/F): I7041 / M
Find location: Szigetszentmiklós-Üdülősor
Country: Hungary
Associated label in publication: Hungary_BA
Date: 2500–2200 BCE
MtDNA haplogroup (mother): H1b1
Y-DNA haplogroup (father): R1b1a1a2 (L151)
Reference: Olalde et al. 2018
Colour group: Steppe (R1b)
Comments: null
Other references: null

Sample ID / genetic sex (M/F): I5666 / M
Find location: Lochenice
Country: Czech Republic
Associated label in publication: Beaker Central Europe
Date: 2500–1900 BCE
MtDNA haplogroup (mother): U4a2c
Y-DNA haplogroup (father): R1b1a1a2a1a2b1 (L2)
Reference: Olalde et al. 2018
Colour group: Steppe (R1b)

Sample ID / genetic sex (M/F): I5025, RISE567 / F
Find location: Kněževes
Country: Czech Republic
Associated label in publication: Beaker Central Europe
Date: 2500–1900 BCE
MtDNA haplogroup (mother): U5b2c
Y-DNA haplogroup (father): null
Reference: 1240k of shotgun data in AllentoftNature2015
Colour group: Steppe (autosomal)

Sample ID / genetic sex (M/F): I6480 / M
Find location: Velké Přílepy
Country: Czech Republic
Associated label in publication: Beaker Central Europe
Date: 2500–1900 BCE
MtDNA haplogroup (mother): U4a2
Y-DNA haplogroup (father): R1b1a1a2a1a2b1 (L2)
Reference: Olalde et al. 2018
Colour group: Steppe (R1b)

Sample ID / genetic sex (M/F): I7271 / M
Find location: Brandýsek
Country: Czech Republic
Associated label in publication: Beaker Central Europe
Date: 2500–2200 BCE
MtDNA haplogroup (mother): U4a2
Y-DNA haplogroup (father): R1b1a1a2a1a2b1 (L2)
Reference: Olalde et al. 2018
Colour group: Steppe (R1b)

Sample ID / genetic sex (M/F): I7212 / M
Find location: Radovesice
Country: Czech Republic
Associated label in publication: Beaker Central Europe
Date: 2500–2200 BCE
MtDNA haplogroup (mother): K1b1a1+199
Y-DNA haplogroup (father): R1b1a1a2a1a2b1 (L2)
Reference: Olalde et al. 2018
Colour group: Steppe (R1b)

Sample ID: RISE564.SG
Location: Osterhofen-Altenmarkt, Germany
Haplogroup name
R1b1a1a2a1 (L51)

Sample ID: I5529
Location: Osterhofen-Altenmarkt, Bavaria
Haplogroup name
R1b1a1a2a1a2b1 (L2)

Sample ID / genetic sex (M/F): I4132, RISE560 / M
Find location: Augsburg Sportgelände, Augsburg, Bavaria
Country: Germany
Associated label in publication: Beaker Central Europe
Date: 2500–2000 BCE
MtDNA haplogroup (mother): U5a1a1
Y-DNA haplogroup (father): R1b1a1a2a1a2 (P312)
Reference: 1240k of shotgun data in Allentoft Nature 2015
Colour group: Steppe (R1b)

Sample ID / genetic sex (M/F): I5748 / M
Find location: De Tuithoorn, Oostwoud, Noord-Holland
Country: The Netherlands
Associated label in publication: Beaker The Netherlands
Date: 2579–2233 calBCE (3945±55 BP, GrN-6650C)
MtDNA haplogroup (mother): X2b4
Y-DNA haplogroup (father): R1b1a1a2a1a2 (P312)
Reference: Olalde et al. 2018
Colour group: Steppe (R1b)

Sample ID / genetic sex (M/F): I5757 / M
Find location: Sion-Petit-Chasseur, Dolmen XI
Country: Switzerland
Associated label in publication: Beaker Central Europe
Date: 2469–1984 BCE
MtDNA haplogroup (mother): H3af
Y-DNA haplogroup (father): R1b1a1a2a1a (L151)
Reference: Olalde et al. 2018
Colour group: Steppe (R1b)

Sample ID / genetic sex (M/F): I2575 / M
Find location: La Fare, Forcalquier
Country: France
Associated label in publication: Beaker Southern France
Date: 2475–2210 calBCE (3895±40 BP, GrA-22988)
MtDNA haplogroup (mother): K1c1
Y-DNA haplogroup (father): no_data
Reference: Olalde et al. 2018
Colour group: Steppe (autosomal)

Sample ID / genetic sex (M/F): I6472 / M
Find location: La Magdalena, Madrid
Country: Spain
Associated label in publication: Beaker Iberia
Date: 2500–2000 BCE
MtDNA haplogroup (mother): HV0b
Y-DNA haplogroup (father): R1b1a1a2 (M269)
Reference: Olalde et al. 2018
Colour group: Steppe (R1b)

Sample ID / genetic sex (M/F): I6588 / M
Find location: Humanejos, Madrid
Country: Spain
Associated label in publication: Beaker Iberia
Date: 2500–2000 BCE
MtDNA haplogroup (mother): U5b2b3
Y-DNA haplogroup (father): R1b1a1a2a1a (L151)
Reference: Olalde et al. 2018
Colour group: Steppe (R1b)

Sample ID / genetic sex (M/F): I5665 / M
Find location: Virgazal, Tablada de Rudrón, Burgos
Country: Spain
Associated label in publication: Beaker Iberia
Date: 2280–1984 calBCE (3730±40 BP, Poz-49174)
MtDNA haplogroup (mother): K1a24a
Y-DNA haplogroup (father): R1b1a1a2a1a2 (P312)
Reference: Olalde et al. 2018
Colour group: Steppe (R1b)
Comments: null
Other references: null
 
Other than the French hotspot of L51, the track can be deduced by brother and uncle clades spanning with interesting frequencies in Albania or Italy, it's like a track of a migrating clan that I suppose were Neolithic herders coming from Anatolia.
Possibly, although it seems to have too much EHG to have been a part of anything pre-Suvorovo, in my view.

I tend not to look at frequencies, which can be misleadingly influenced by sudden population bursts, but more at phylogeny and intra-phylogenic diversity.
 
I have little doubt you are right, and that the phylogenic evidence clearly indicates the common ancestors of extant L51 most recently lived in Western Europe, most likely France. I am intrigued by the less clear issues of how, when and why they got there.

There is some circumstantial yDNA, mtDNA and aDNA evidence pointing to possible associations with Southern Poland, Chalcolithic Bulgaria, Cucuteni Triploye, Globular Amphora and RRBP, so I feel the most likely route is mid/late millennium BC up Moldova and across Southern Poland and South/Central Germany; but I have not seen anything that looks conclusive.

Why is it incorrect to reason as I did though? I'm always open to criticism, but nobody has told me why exactly I'm wrong. I don't claim to know the truth, like everyone I'm just speculating based on how I personally interpret the facts. As for haplogroups, I only really focus on Y DNA - in most cases, it isn't useful to look too much into mtDNA (try and make sense of many of the mtDNA maps on eupedia for example)

As for the circumstances, I have explained that pretty well I think - it follows the spread of copper metallurgy and warlike elites. The first copper metallurgy in Western Europe is from Los Millares, which was organised based on caste and often at war with surrounding tribes. Of note too, is that copper metallurgy definitely did not enter SE Spain (the area of Los Millares) from a continental route, as Southern Spain received copper significantly earlier than e.g. Northern Spain. Thus, copper metallurgy necessarily must have made its way to Los Millares across the Mediterranean (the only other alternative being an origin from North Africa, however that seems unlikely as copper metallurgy only existed in the Maghreb much later). And as far as I'm aware without exception, the spread of metallurgy always involves the spread of people, but in any case it is still usually the case, so we have a new people arriving from across the Mediterranean (and as quick evidence of this, Los Millares imported goods from the Middle East and the elites buried themselves in tholoi (West Asian in origin)). The further connections with anthropomorphic stelae, the phylogeny point mentioned, the lack of L51 in Steppe cultures, the fact that L21 and DF27 cannot be clearly associated to any IE language, and the fact that the cultural profile of Los Millares matches what we know of the Beaker folk (warlike, metallurgical elites that readily appropriate pre-existing communities they come across) all add to the theory.
 
L51 from France ??? Until 2600 BC, R1b is nowhere to be seen in western Europe, save a few outliers who had probably come along with the "farmers". After 2500 BC, they suddenly show up all over central and western Europe. Either L51 had been in Hungary since the old Suvorovo pocket for centuries, or they came along with Yamna Danube.
Before 2500 BC, they don't show up much in Central and Western Europe, nor in Eastern Europe, nor in the Steppe. Genetically, they probably looked more like those in the old Suvorovo pocket in Hungary, so perhaps they had been there for centuries. However, given that both the Suvorovo and Bell Beaker moved around a lot, I think it likely that they too moved around a lot. Indeed, the El Portalon sample shows a similar genetic footprint quite some distance away from Hungary.

The point about L51 and France is that the phylogeny demonstrates a clear predicted coalescence point there. It says nothing about where L51's ancestors were before that point.
 
Before 2500 BC, they don't show up much in Central and Western Europe, nor in Eastern Europe, nor in the Steppe. Genetically, they probably looked more like those in the old Suvorovo pocket in Hungary, so perhaps they had been there for centuries. However, given that both the Suvorovo and Bell Beaker moved around a lot, I think it likely that they too moved around a lot. Indeed, the El Portalon sample shows similar genetic footprint quite some distance away from Hungary.

The point about L51 and France is that the phylogeny demonstrates a clear predicted coalescence point there. It says nothing about where L51's ancestors were before that point.

It doesn't predict coalescence there, it predicts an origin there. The subclade that split from the rest of L51 at the earliest point (400 years only after L51 was born) is majorly distributed around Southern France and surrounding regions about the West Med. And what do we know, those stelae I mentioned earlier have a huge hotspot in S. France, and these stelae date to the late neo/copper age transitional period...

https://i.imgur.com/scFbZpS.png

Below is Coon's (not fully accurate) interpretation (ignore dating, this was before the discovery of radiocarbon dating):

455c2-troemap2.jpg


troe-V-map.jpg


Not because I saw this beforehand, look where the arrow across the Mediterranean (indicating Beaker origins) traces back to - exactly the region I labelled in my map as L23's Urheimat, and exactly the same region where Leyla-Tepe in the Caucasus was thought to be founded from (i.e. from Eastern Anatolia to Leyla Tepe bringing copper metallurgy - another non-coincidence, Leyla-Tepe had the first metallurgy in the Caucasus). If I haven't made it clear, I'm talking about the arrow that enters Cyprus.
 
Combining 65% Yamnaya and 35% Dereivka Sredny Stog gives a projected autosomal reading for German CW that has 20 times greater variance from actual CW than that projected by the best-fit combination. To me, this does not look like a very reasonable match..

Well, but that wouldn't be a realistic model, and it's too simplistic, anyway. The vast majority of the Yamnaya samples are of Northeastern (Don-Volga) Yamnaya, not its western or southernmost part, and that's still a hindrance to a better understanding of these migrations in my opinion. We shouldn't expect to find excellent fits for all populations using just a few samples and very simple two-way admixtures or the like. As for Dereivka, its much higher than average EEF input may not be representative of the northern Sredny Stog people that may have given birth to the Yamnaya-Sredny Stog pre-CWC mix, maybe near Pripyat. In my opinion, admixture (especially due to female exogamy and male-biased conquests) was much more common than some are still willing to think. Therefore, of course I'm just speculating here, but if we're talking of a demographic path between Eneolithic (~4000 B.C.) and Early Bronze Age CWC in Germany (~2500 B.C.), I'd expect something much more complex. Importantly, Germany CWC wouldn't have been just a transplant of the North Ukraine Yamnaya-ized population, but the final result of an expansion of that population first northward, then westward, possibly absorbing different EEF and also EHG-enriched populations. I think simple models, even if they have a good statistical fit, are a bit implausible especially if you're talking about two populations very distant in time (by ~1500 years) and geography (by ~1500 km).

There is also the problem that, according to your hypothesis, Suvorovo-Novodanilovka is "the key", but your main fit for CWC is basically a miraculously unmixed (for 1500 years and 1500km) Ukraine Eneolithic sample, which, if it's the one I think, is dated to 5000-3500 BCE (average ~4300 BCE), roughly when Suvorovo-Danilovka influence starts to appear in the archaeological record. But they don't seem to have been much influenced by the transformations brought by them and passed their genetic makeup on to Germany 1500 years almost totally unmixed, or so it seems according to your analysis. Would Suvorovo-Novodanilovka have just spread their culture via diffusion with little or no genetic impact, and also not changing their Y-DNA lineages at all?
 
Well, but that wouldn't be a realistic model, and it's too simplistic, anyway. The vast majority of the Yamnaya samples are of Northeastern (Don-Volga) Yamnaya, not its western or southernmost part, and that's still a hindrance to a better understanding of these migrations in my opinion. We shouldn't expect to find excellent fits for all populations using just a few samples and very simple two-way admixtures or the like. As for Dereivka, its much higher than average EEF input may not be representative of the northern Sredny Stog people that may have given birth to the Yamnaya-Sredny Stog pre-CWC mix, maybe near Pripyat. In my opinion, admixture (especially due to female exogamy and male-biased conquests) was much more common than some are still willing to think. Therefore, of course I'm just speculating here, but if we're talking of a demographic path between Eneolithic (~4000 B.C.) and Early Bronze Age CWC in Germany (~2500 B.C.), I'd expect something much more complex. Importantly, Germany CWC wouldn't have been just a transplant of the North Ukraine Yamnaya-ized population, but the final result of an expansion of that population first northward, then westward, possibly absorbing different EEF and also EHG-enriched populations. I think simple models, even if they have a good statistical fit, are a bit implausible especially if you're talking about two populations very distant in time (by ~1500 years) and geography (by ~1500 km).

So given you're basically suggesting L51 in Western Yamnaya and Z2103 in Eastern Yamnaya, how does that square with the fact that the Hungarian BBs clearly show a mixing of Z2103 from the East and L51 from the West? This was what made Davidski turn away from Yamnaya recently (to the Single Grave culture), and it was heavily alluded to in one of the OREA talks also recently uploaded to youtube. And the fact that all samples from the Balkans with a possible Yamnaya origin have belonged either to I2a2 or Z2103 (e.g. Vucedol being Z2103)? And the fact that Z2103 still has a relatively strong presence showing its approximate trail following the Danube, with L51 leaving none?

Haplogroup-R1b-Z2103.png
 
So given you're basically suggesting L51 in Western Yamnaya and Z2103 in Eastern Yamnaya, how does that square with the fact that the Hungarian BBs clearly show a mixing of Z2103 from the East and L51 from the West? This was what made Davidski turn away from Yamnaya recently (to the Single Grave culture), and it was heavily alluded to in one of the OREA talks also recently uploaded to youtube. And the fact that all samples from the Balkans with a possible Yamnaya origin have belonged either to I2a2 or Z2103 (e.g. Vucedol being Z2103)? And the fact that Z2103 still has a relatively strong presence showing its approximate trail following the Danube, with L51 leaving none?

Haplogroup-R1b-Z2103.png

Well, I'm not suggesting that at all, since I'm talking about the origins of CWC and more specifically German CWC. But I actually think L51 was probably a Yamnaya-ized .

Do you know why they're claiming that Z2103 came from the East and L51 from the West even in the absence of any pre-BB sample of L51 in the West? I saw some people in Eurogenes claiming that the fact that much of the EEF in Yamnaya_Hungary is pretty "northern" discards a relationship with Yamnaya because it should be closer to Yamnaya_Bulgaria. But I see no reason for that, honestly. Yamnaya_Hungary could well have been formed directly from Ukaine crossing the Carpathians. Throughout history there were peoples (the Magyars themselves, as well as Turks, Jasz etc.) that arguably invaded the Pannonian Basin from the north, via Western Ukraine/Slovakia, and not via Bulgaria and then northwards via the Danubian valleys.

If L51 was, as some assume, a western, more "Transcarpathian" brother of Z2103, maybe initially absorbed by Cucuteni-Tripolye before its Indo-Europeanization and subsequent Yamnaya-ization in the latest phase of Cucuteni-Tripolye, it would make sense that it would spread along the Carpathians and from there to Yamnaya_Hungary picking up exactly that GAC-like "northern EEF" along the way.

It still remains to me, for now, a bit hard to believe that L51 and Z2103 were very far from each other in Europe even if their split date was just a few centuries before Z2103 starts to be found in aDNA samples in Eastern Europe.

Carpathians_dem.jpg
 
Yes, Suvorovo might well have been predominantly intrusive and invasive, but there are also some signs of collaboration and admixture (perhaps a bit like Vikings turning into Normans?). I would say Varna looks to have witnessed some of the first signs of Suvorovo, rather than a predecessor of it. After the collapses of the Balkan Neolithic, Suvorovo populations look to have taken away with them both some DNA and some of what they had learned from places they had 'invaded' - we know they migrated to Central Ukraine (where they would have look genetically rather like M417 Corded Ware) and probably through Moldova to the Upper Dniester (where, upon admixture with some Globular Amphora, they would have looked genetically rather like L51 Bell Beaker).

Merging this analysis with what we think we know from archaeology, linguistics and other disciplines can sometimes assist, but can also sometimes muddy the clearer genetic waters.

Let me see if I understood your point: Early Suvorovo was intrusive in the Balkans and came from the steppes very early (Late Neolithic) - which would explain their high EHG -, mixed there with Balkan_Neolithic people and changed some of its cultural ways and later migrated back to Ukraine and replaced all the local steppe and EEF (Cucuteni-Tripolye included) virtually without any mixing at all, before they subsequently spread to other parts of Western & Central Europe?
 
Well, I'm not suggesting that at all, since I'm talking about the origins of CWC and more specifically German CWC. But I actually think L51 was probably a Yamnaya-ized .

Do you know why they're claiming that Z2103 came from the East and L51 from the West even in the absence of any pre-BB sample of L51 in the West? I saw some people in Eurogenes claiming that the fact that much of the EEF in Yamnaya_Hungary is pretty "northern" discards a relationship with Yamnaya because it should be closer to Yamnaya_Bulgaria. But I see no reason for that, honestly. Yamnaya_Hungary could well have been formed directly from Ukaine crossing the Carpathians. Throughout history there were peoples (the Magyars themselves, as well as Turks, Jasz etc.) that arguably invaded the Pannonian Basin from the north, via Western Ukraine/Slovakia, and not via Bulgaria and then northwards via the Danubian valleys.

If L51 was, as some assume, a western, more "Transcarpathian" brother of Z2103, maybe initially absorbed by Cucuteni-Tripolye before its Indo-Europeanization and subsequent Yamnaya-ization in the latest phase of Cucuteni-Tripolye, it would make sense that it would spread along the Carpathians and from there to Yamnaya_Hungary picking up exactly that GAC-like "northern EEF" along the way.

It still remains to me, for now, a bit hard to believe that L51 and Z2103 were very far from each other in Europe even if their split date was just a few centuries before Z2103 starts to be found in aDNA samples in Eastern Europe.

Carpathians_dem.jpg

I don't think they're claiming L51 came from Iberia/France, just that it arrived at Hungary (as BBs) from the West. And I have no idea, but I'm guessing that it's not baseless, so I'm going to go along with it - I'd logically guess they think L51 came from the East earlier, before U152 "turns around" and so comes back to Hungary from the West. And I'm not a big fan of this autosomal fitting that goes on, broadly it is of course important, but this is multivariable analysis so there's inevitably going to be many possible combinations when trying to tie down a specific type of admixture to specific groups (an example of this error is modelling Corded Ware as mostly Yamnaya-derived rather than Yamnaya-ised and simply having broadly similar overall ancestry). Looking at Y DNA never has this kind of error, it's 100% distinct, and for this reason I think should be looked at more than autosomal DNA when tracing migrations (Y DNA could thus be used as a marker).

And about the idea that it's unlikely that L51 and Z2103 could have their Urheimats so far apart despite being sibling subclades under L23, to me it's fine, as for example by the Beaker period of expansion their common ancestor with the Yamnaya men would have been 2000 years in the past. That is plenty of time for one branch of an originally Syro-Anatolian L23 to migrate to the Caucasian Leyla-Tepe (and then to the Steppe, maybe via Maykop), and for another branch of this L23 to migrate across the Mediterranean until hitting South-Eastern Spain. Even if we decide not to associate this with R1b-L23+, the spread of copper metallurgy certainly follows such paths as described, so it isn't unreasonable at all. I mean, Z2103 made it all the way to China during early Yamnaya lol, that distance is far greater than island-hopping across the Mediterranean

Copper_Age_Europe.png


(Also, does anybody know why Germany and the surrounding regions receive copper so early in this map? I know it isn't an error, but I can't think of what culture it would be associated with)
 
(an example of this error is modelling Corded Ware as mostly Yamnaya-derived rather than Yamnaya-ised and simply having broadly similar overall ancestry). Looking at Y DNA never has this kind of error, it's 100% distinct, and for this reason I think should be looked at more than autosomal DNA when tracing migrations (Y DNA could thus be used as a marker).

I think all genetic and non-genetic data are important and only contribute to a realistic and probable conclusion about what must've happened if they're used in combination. For instance, using the same example of yours, by autosomal analysis we can establish that the bulk of the CWC ancestry came from broadly the same genetic structure where Yamnaya also arose (though it was a bit too bold to claim they were the Yamnaya, which usually means "East Yamnaya" people like Samara and the like), and that information could never be found if you just looked at their Y-DNA phylogeny and, by the huge prevalence of R1a-M417, you could well believe they would have nothing to do with the overwhelmingly R1b-Z2103 Yamnaya and could not have come from a broadly similar autosomal makeup and nearby region. All the evidences fit together, and one of them may help dispel some misleading conclusions given by other evidences alone.
 
I think all genetic and non-genetic data are important and only contribute to a realistic and probable conclusion about what must've happened if they're used in combination. For instance, using the same example of yours, by autosomal analysis we can establish that the bulk of the CWC ancestry came from broadly the same genetic structure where Yamnaya also arose (though it was a bit too bold to claim they were the Yamnaya, which usually means "East Yamnaya" people like Samara and the like), and that information could never be found if you just looked at their Y-DNA phylogeny and, by the huge prevalence of R1a-M417, you could well believe they would have nothing to do with the overwhelmingly R1b-Z2103 Yamnaya and could not have come from a broadly similar autosomal makeup and nearby region. All the evidences fit together, and one of them may help dispel some misleading conclusions given by other evidences alone.

Yup, totally agreed - I don't think autosomal DNA is useless for obvious reasons (you can have any form of autosomal ancestry and have similar Y DNA to someone else with completely different ancestry, and the opposite too). But as you suggest, I think autosomal ancestry is only useful when looking at the broad picture - unless it can be made much more precise (like 23andme), however I'm guessing that's statistically impossible with the comparatively few samples we have.
 
And about the idea that it's unlikely that L51 and Z2103 could have their Urheimats so far apart despite being sibling subclades under L23, to me it's fine, as for example by the Beaker period of expansion their common ancestor with the Yamnaya men would have been 2000 years in the past. That is plenty of time for one branch of an originally Syro-Anatolian L23 to migrate to the Caucasian Leyla-Tepe (and then to the Steppe, maybe via Maykop), and for another branch of this L23 to migrate across the Mediterranean until hitting South-Eastern Spain. Even if we decide not to associate this with R1b-L23+, the spread of copper metallurgy certainly follows such paths as described, so it isn't unreasonable at all. I mean, Z2103 made it all the way to China during early Yamnaya lol, that distance is far greater than island-hopping across the Mediterranean

Copper_Age_Europe.png


(Also, does anybody know why Germany and the surrounding regions receive copper so early in this map? I know it isn't an error, but I can't think of what culture it would be associated with)

As for the Z2103 x L51 split, my main quibble with that big geographical distance is that they probably split around ~4500-4000 B.C., but IIRC there is already 1 Ukraine Eneolithic sample that is already and fully Z2103. That means that L23 would've split in Eastern Anatolia with Z2103 going to form Leyla-Tepe in the Caucasus and then migrated to the Northern Caucasus and from there westward to the Ukraine in a couple of centuries (and we already know from the Greater Caucasus paper that that wasn'tcommon stuff, the Caucasus seems to have acted as an effective barrier to any significant gene flow at least from the Eneolithic onward). In my opinion, where Z2103 was found in higher frequency earliest is probably not too far from where L51 also split.

I keep thinking you're associating it the Indo-Europeans with a sort of technological avant-garde and proto-civilization status that the earliest Indo-European cultures did not have until they clearly mixed heavily with and absorbed previous Mediterranean cultures (or do you think the more advanced L23 people absorbed the language of the less advanced steppe hunter-gatherers and therefore became Indo-European only there?).
 
As for the Z2103 x L51 split, my main quibble with that big geographical distance is that they probably split around ~4500-4000 B.C., but IIRC there is already 1 Ukraine Eneolithic sample that is already and fully Z2103. That means that L23 would've split in Eastern Anatolia with Z2103 going to form Leyla-Tepe in the Caucasus and then migrated to the Northern Caucasus and from there westward to the Ukraine in a couple of centuries (and we already know from the Greater Caucasus paper that that wasn'tcommon stuff, the Caucasus seems to have acted as an effective barrier to any significant gene flow at least from the Eneolithic onward). In my opinion, where Z2103 was found in higher frequency earliest is probably not too far from where L51 also split.

I keep thinking you're associating it the Indo-Europeans with a sort of technological avant-garde and proto-civilization status that the earliest Indo-European cultures did not have until they clearly mixed heavily with and absorbed previous Mediterranean cultures (or do you think the more advanced L23 people absorbed the language of the less advanced steppe hunter-gatherers and therefore became Indo-European only there?).

When does this Ukrainian sample date to?

And I don't think L23+ folk were some kind of supermen, but it's impossible to deny that they dominated most of the cultures they came across. I (and a few others, Tomenable was where I first saw this idea) have linked this to the spread of copper smelting and warlike elites (and during the Metal Ages, having that sort of knowledge automatically gave you huge social status, so it all fits in). I have repeatedly, however, attributed the sophistications of civilisation to Near Eastern pred. Y DNA J folk (almost everyone was illiterate back then, but the Indo-Europeans didn't even have a writing script and no concrete abstract "theory of knowledge" that was clearly present in the Middle East with, for example, the invention of mathematics), so it isn't as if anything great in prehistory and history has to be related to R1b. I have no personal bias towards R1b either, as I'm a Y DNA I Jew.

The ostensibly closest I get to "R1b supremacism" is with my belief in them even making up the elite in certain South American cultures lol, but even then virtually anybody from West Eurasia could have done the same if they crossed the Atlantic. We now know East Asians, albeit at a much later date, did the same across Polynesia, which I think is much more impressive than trans-Atlantic travel (which one man, Thor Heyerdahl, managed to do using very primitive rafting technology)
 

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