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

[Pip]

I see, but, I mean, how likely is it that a R1b BB and a R1a CWC from, say, around 2500 BC would still have a very close match with a 5th millennium Bulgarian individual even after so intensive migrations, cultural changes and certainly lots of mixing with the local peoples (who, especially in the case of BB, they didn't seem to replace overwhelmingly)? Would very little mixing and autosomal change have happened in more than 1500 years even as the Bulgarian Suvorovo spread to lands very far away and already densely inhabited? I'd be very surprised if that did happen. I think the fact that the BB and CWC do not match as well as with the Sredny Stog and the Yamnaya samples may just result from the very likely and plausible fact that those were still "steppe proper", pre-expansion societies with much less admixture with the ANF+WHG mix dominant especially to the west of the Dniester.

We do not know how much people admixed until we examine the data. If we look at the R1a-M417 sample from circa 4,000 BC, for example, it differs little autosomally from the R1a-M417 samples in Corded Ware 1,500 years later. I would suggest this indicates that extant M417 admixed very little during this period, just as its females seemed to admix almost exclusively with M417 men. This is surely a more likely explanation than that it changed autosomally through admixture with Sredny Stog and Yamnaya before changing back to pretty much the same autosomal mix that it had before.

Sredny Stog and Yamnaya look to me like red herrings. They had different autosomal mixes from each other, and neither of their Steppe DNA mixes match the mixes in the Steppe components within BB or CW.

[epoch]

I think 'useless' is exaggerated, particularly as it is often verified with f3 or D-stats. In the absence of evidence to the contrary, it is at least better than nothing. It may be one admixture run, but it is a dataset in that the data has been obtained from many samples.

The way you use it is pretty much useless. The percentages you get aren't ancestry, they are distances in a forced fit. By the way, there are plenty of ADMIXTURE runs in different papers. Try some of these and see if the results are the same.

If I understand you correctly - if C's descendants A and B later merge, rather than admix with other populations, then they are likely to show a greater proportional descent from C. This is what I am measuring, rather than separation times.

A and B are full 100% unadmixted descendants of C. However, Measuring with Fst only one will consider a merged A+B closer to C than either A or B.

[Pip]

The way you use it is pretty much useless. The percentages you get aren't ancestry, they are distances in a forced fit. By the way, there are plenty of ADMIXTURE runs in different papers. Try some of these and see if the results are the same.

I could do, but usually the ranges in these papers are less extensive, there are fewer components, and there seems little point in repeating a laborious exercise if people consider it useless.

A and B are full 100% unadmixted descendants of C. However, Measuring with Fst only one will consider a merged A+B closer to C than either A or B.

If Bell Beaker and Corded Ware were A and B (both principally descended from Yamnayan C) that ended up coming back together in Germany, this would make German samples on Admixture appear closer to Yamnaya than they are to other BB or CW. However, exactly the opposite is the case - all such samples are significantly closer to each other than to Yamnaya in respect of all inter-component relationships. This would suggest that Yamnayans must be even less close ancestrally to BB and CW than my results estimate.

If there are other studies which show that Yamnaya has a better fit with BB and CW than Suvorovo or Khvalynsk does, then this would make me less confident that the spread of Steppe DNA occurred before Yamnaya arose. In the absence of such studies, it makes more sense to me to go with the clear results obtained from the best fit analysis - that BB and CW each look Suvorovo/East Balkan in origin. Not only does this tie up with my yDNA branching estimates (that the splits between Eastern and Western branches of R1b-L23 and R1a-M417 preceded the Yamnayan westward expansions), but it would also explain what happened to all the DNA from the Suvorovo/early Steppe people that we know had spread far and wide over an area encompassing Hungary, Romania, Bulgaria, Moldova and Eastern Ukraine by 4,000 BC (and even, it appears, North Central Spain by the mid fourth millennium BC).

[Pip]

For what it's worth, I ran best-fit calculations for some other early Steppe component populations:
1. The Steppe component in Northern Funnel Beaker comes out as early/indigenous and unrelated to Suvorovo or Yamnaya.
2. Balkan Yamnaya comes out as unrelated to Suvorovo/Corded Ware/Bell Beaker and best fits with an admixture between various Eastern peoples (Russian Yamnaya, Maykop and Dereivka) with Cucuteni Tripolye.

My reading of the autosomal data is as follows:
Early Scandinavian Steppe-like DNA most likely arrived during the Mesolithic with basal clades of R1a.
Khvalynsk-like people of mixed yDNA (Suvorovo) arrived in the Western Pontic and admixed with locals to form proto-BB/CW/Vucedol. Corded Ware admixed little, Bell Beaker was the product of some admixture with Globular Amphora, and Vucedol with various Southern European EEF.
About 1,500 years later, another Khvalynsk-descendant population (Yamnaya) moved westwards into the Balkans, together with some Maykop and Dereivka, and admixed with Cucuteni-Tripolye people. Yamnaya was probably the catalyst, as the resulting culture was theirs and they also seem to have moved into Corded Ware territory in the Eastern Baltic. These moves displaced Corded Ware westwards, which in turn displaced Bell Beaker further westwards.

I would be interested to hear of any evidence that might lead me to revise or refine these tentative estimates.

[gidai]

It is about autosomal or Y-ADN?
I think, it's a very complex process like one of hydrodynamics, with little data about the ancient DNA. Y-DNA is too little of the genetic code. Ancient cultures could spread from one population to another without massive population movements, the same as Y-dna, almost without to afect autosomal DNA.

[epoch]

I could do, but usually the ranges in these papers are less extensive, there are fewer components, and there seems little point in repeating a laborious exercise if people consider it useless.

Frankly, I respect the effort you put into this. You might want to take the steep learning curve and learn how to use the Reich labs tools or ADMIXTURE yourself. The datasets are mostly freely available. But this is simply not going to fly. I might be wrong in this but then prudence would dictate that you at least verify it with other ADMIXTURE runs to show that these produce similar results.

If Bell Beaker and Corded Ware were A and B (both principally descended from Yamnayan C) that ended up coming back together in Germany, this would make German samples on Admixture appear closer to Yamnaya than they are to other BB or CW. However, exactly the opposite is the case - all such samples are significantly closer to each other than to Yamnaya in respect of all inter-component relationships. This would suggest that Yamnayans must be even less close ancestrally to BB and CW than my results estimate.

They all have picked up farmer ancestry, which is highly related.

If there are other studies which show that Yamnaya has a better fit with BB and CW than Suvorovo or Khvalynsk does, then this would make me less confident that the spread of Steppe DNA occurred before Yamnaya arose. In the absence of such studies, it makes more sense to me to go with the clear results obtained from the best fit analysis - that BB and CW each look Suvorovo/East Balkan in origin.

No, they look more East Balkans in one of the ADMIXTURE runs that Gentiker made. The K=14 run.

Not only does this tie up with my yDNA branching estimates (that the splits between Eastern and Western branches of R1b-L23 and R1a-M417 preceded the Yamnayan westward expansions), but it would also explain what happened to all the DNA from the Suvorovo/early Steppe people that we know had spread far and wide over an area encompassing Hungary, Romania, Bulgaria, Moldova and Eastern Ukraine by 4,000 BC (and even, it appears, North Central Spain by the mid fourth millennium BC).

[epoch]

@Pip

Two notes: Models are just proving that a certain ancestry is genetically feasable, they aren't definitive proof. Also, the more methods you use the more evidence you muster.

Take as a lesson for instance how amateurs started to notice that admixture runs showed some European admixture in American Indians. It was replicated in Treemix. However, it turned out that this was shared ancient ancestry which we now call ANE. Only with a combination of genetics, archaeology and common sense you can rebuild an ancestry tree this deep.

[Pip]

Frankly, I respect the effort you put into this. You might want to take the steep learning curve and learn how to use the Reich labs tools or ADMIXTURE yourself. The datasets are mostly freely available. But this is simply not going to fly. I might be wrong in this but then prudence would dictate that you at least verify it with other ADMIXTURE runs to show that these produce similar results.

I don't think I really have the time, and am in any case suspicious of admixture, which is easier to manipulate than yDNA. I was only really looking at it to see whether it matches what yDNA seems to indicate, and to help fill in the gaps caused by bottlenecks in yDNA.

They all have picked up farmer ancestry, which is highly related.

The point is that it only seems to be Khvalynsk admixed with this EEF (and not Yamnaya admixed with EEF) that provides close fits with Bell Beaker and Corded Ware.

No, they look more East Balkans in one of the ADMIXTURE runs that Gentiker made. The K=14 run.

I don't know what they look like in the other admixture runs. I use the K=14, as that is where Genetiker publishes the most extensive database.

[Pip]

@Pip

Two notes: Models are just proving that a certain ancestry is genetically feasable, they aren't definitive proof. Also, the more methods you use the more evidence you muster.

Agreed. If people think there will ever be definitive proof about exactly who bred with whom and where they were located when they did, they will ultimately be disappointed. I'm just looking at what is feasible, given the data, and what is infeasible. From the data I have seen, I would say that the Yamnaya (as generally understood) are only a remote possibility as the major source of pan-European Steppe DNA, and not what I would call a credible one.

The gaps I was keen to fill were between extant Western and Eastern branches of R1b-M269 and R1a-M417, which I estimate to have split during the 5th millennium BC. Most of the major branches of these haplogroups seem to coalesce to either Western Europe or the Caucasus, so what was the substantial path between the two? My combination SNP/STR analysis indicates Poland or North Western Ukraine as the slightly most likely overall origin point, with an eastwards branching shortly afterwards; whereas Genetiker's K=14 autosomal data indicates something similar, but in reverse.

For R1b-M269 the most likely major route to me now looks something like this - Azov split mid-fifth millennium BC.
Azov>W Pontic>Moldova>Poland>S/C Germany>N France (R1b Bell Beaker expansion point)
Azov>Steppe Maykop>Caucasus>NE Turkey/Armenia

For R1a-M417, the most likely major route looks like - North Ukraine split circa 4,000 BC (Corded Ware expansion point some time later)
Azov>W Pontic>North Ukraine>S Baltic>Scandinavia
Azov>W Pontic>North Ukraine>E Baltic>Poland>E Europe
Azov>W Pontic>North Ukraine>E/C Europe>Caspian>Middle East/N India

Other branches look largely to have shrivelled and died out.

I am interested in any data that might cause these estimates to be revised or refined.

[Pip]

I might be wrong in this but then prudence would dictate that you at least verify it with other ADMIXTURE runs to show that these produce similar results.

I cannot find other published admixture databases that have the same extensive coverage as Genetiker's K=14 (including over 3,000 samples). He has also compiled K=16 etc., but with insufficient samples and breakdowns to replicate my K=14 best-fit tests, so I will have to leave them as they stand.

[Pip]

Corded Ware samples are nearly all R1a-M417. According to yfull, everyone with M417 descends from a single man who lived 3,500 BC.
But we have a North Ukrainian M417 sample dated to 4,000 BC. This means that either all of his descendants died out, or that he personally was paternal ancestor to all M417 people today. The culture in which he lived exhibit the first-known corded patterns on pottery. His autosomal DNA shows an almost perfect (96%) match with German Corded Ware - in fact, it is no different to German Corded Ware than German Corded Ware samples are to each other.

1. Is there any reason to think that the 4,000 BC ancestor to R1a-M417 Corded Ware was anyone other than the man in this sample or someone very much like him?

2. As there is no discernible change in autosomal DNA between 4,000 BC and the German Corded Ware period, and as nearly all Corded Ware yDNA is from the same M417 subclade of R1a, is there any reason to think there was any significant admixture with other people during this period of development leading up to full Corded Ware?

3. In particular, given that (i) almost all Yamnayan samples are from a different yDNA group (R1b-Z2103) and bear only a 65% match to the North Ukrainian and German Corded Ware samples, and (ii) Yamnaya only arose circa 3,300 BC (700 years after the closely-matching Ukrainian M417 man lived), is there any reason to think that German Corded Ware was derived from Yamnayan admixture to any significant degree, rather than by direct descendance from the community in North Ukraine in which the 4.000 BC M417 man lived?

[markod]

Corded Ware samples are nearly all R1a-M417. According to yfull, everyone with M417 descends from a single man who lived 3,500 BC.
But we have a North Ukrainian M417 sample dated to 4,000 BC. This means that either all of his descendants died out, or that he personally was paternal ancestor to all M417 people today. The culture in which he lived exhibit the first-known corded patterns on pottery. His autosomal DNA shows an almost perfect (96%) match with German Corded Ware - in fact, it is no different to German Corded Ware than German Corded Ware samples are to each other.

1. Is there any reason to think that the 4,000 BC ancestor to R1a-M417 Corded Ware was anyone other than the man in this sample or someone very much like him?

2. As there is no discernible change in autosomal DNA between 4,000 BC and the German Corded Ware period, and as nearly all Corded Ware yDNA is from the same M417 subclade of R1a, is there any reason to think there was any significant admixture with other people during this period of development leading up to full Corded Ware?

3. In particular, given that (i) almost all Yamnayan samples are from a different yDNA group (R1b-Z2103) and bear only a 65% match to the North Ukrainian and German Corded Ware samples, and (ii) Yamnaya only arose circa 3,300 BC (700 years after the closely-matching Ukrainian M417 man lived), is there any reason to think that German Corded Ware was derived from Yamnayan admixture to any significant degree, rather than by direct descendance from the community in North Ukraine in which the 4.000 BC M417 man lived?

I think the people at Harvard & Max Planck have been asking themselves similar questions, hence the hints at a very early source population of PIE in the Near East. The normal Chalcolithic timeframe contradicts their hypothesis, because there's no way Yamnaya, BBC, CWC & the early Anatolian speakers derive from one population that lived in the copper age.

Other than that I don't see why you'd want to discuss these things now - trying to pin migration routes with so little data is a waste of time.

[Pip]

I think the people at Harvard & Max Planck have been asking themselves similar questions, hence the hints at a very early source population of PIE in the Near East. The normal Chalcolithic timeframe contradicts their hypothesis, because there's no way Yamnaya, BBC, CWC & the early Anatolian speakers derive from one population that lived in the copper age.

Other than that I don't see why you'd want to discuss these things now - trying to pin migration routes with so little data is a waste of time.

Yes, let's forget migration routes for a moment - the point is that the community of the North Ukrainian M417 sample (and not Yamnaya) looks like the only credible (and pretty much exclusive) source for later Corded Ware populations, given the data we have available. Unless I'm missing another candidate that is anywhere near as likely?

However, why was this the only sample that had its DNA published, out of 39 samples discovered at the site? What was the yDNA and autosomal mix of the other 38? This might help determine whether its non-Steppe components were likely to have been of Cucuteni-Tripolye, Suvorovo, Anatolian or other origin.

All we do know is that its mtDNA was H2a1a, a haplogroup principally from South of the Caucasus, and a downstream subclade of the mtDNA of the Khvalynsk R1b sample. So it looks pretty much like Suvorovo to me, especially as its autosomal DNA fits best with a South East Balkan origin, and we already know that there were several contemporaneous Suvorovo sites that had arisen in the same area east of the Dnieper.

We may have relatively little data, but the data that we do have looks instructive enough for us to make an informed guess.

[markod]

Yes, let's forget migration routes for a moment - the point is that the community of the North Ukrainian M417 sample (and not Yamnaya) looks like the only credible (and pretty much exclusive) source for later Corded Ware populations, given the data we have available. Unless I'm missing another candidate that is anywhere near as likely?

However, why was this the only sample that had its DNA published, out of 39 samples discovered at the site? What was the yDNA and autosomal mix of the other 38? This might help determine whether its non-Steppe components were likely to have been of Cucuteni-Tripolye, Suvorovo, Anatolian or other origin.

All we do know is that its mtDNA was H2a1a, a haplogroup principally from South of the Caucasus, and a downstream subclade of the mtDNA of the Khvalynsk R1b sample. So it looks pretty much like Suvorovo to me, especially as its autosomal DNA fits best with a South East Balkan origin, and we already know that there were several contemporaneous Suvorovo sites that had arisen in the same area east of the Dnieper.

We may have relatively little data, but the data that we do have looks instructive enough for us to make an informed guess.

The eastern Balkanic steppe region is definitely in need of investigation. Not only Cernavoda and Ezero, but the LN cultures that prececde those as well; in particular Hamangia and Boian culture. The former contributed to the later steppe culture the typical anthropomorphic stelae. They appear already fully developed in LN Hamangia, barely distinguishable from the later stelae in the steppe:



Another thing worthy of consideration are the autosomally divergent samples from Mikhailovska, some of which seem to show elevated CHG affinity which could or could not indicate that there existed a cline within Ukraine, perhaps running from south to north. In fact the earliest steppe culture in which all constituent elements (herding, stelae, kurgans) exist side by side is Kemi Oba concentrated in the Crimean flatlands. Culturally and artistically Kemi Oba appears to be of seminal importance for developments in the wider steppe region, so hopefully we'll get some samples from those sites.



At least as far as archaeology is concerned, your hypothesis of south-eastern European influence in the steppe seems to have some support. I'm not entirely sure whether that influence will show up in the DNA, but it will be interesting to see what the earliest stelae builders of the Black Sea look like genetically nonetheless. Their artistic traditions spread far and wide for sure, adopted by Scythians, Mongols, Turks and most of the other nomadic societies of the Eurasian plains.

[ToBeOrNotToBe]

The eastern Balkanic steppe region is definitely in need of investigation. Not only Cernavoda and Ezero, but the LN cultures that prececde those as well; in particular Hamangia and Boian culture. The former contributed to the later steppe culture the typical anthropomorphic stelae. They appear already fully developed in LN Hamangia, barely distinguishable from the later stelae in the steppe:



Another thing worthy of consideration are the autosomally divergent samples from Mikhailovska, some of which seem to show elevated CHG affinity which could or could not indicate that there existed a cline within Ukraine, perhaps running from south to north. In fact the earliest steppe culture in which all constituent elements (herding, stelae, kurgans) exist side by side is Kemi Oba concentrated in the Crimean flatlands. Culturally and artistically Kemi Oba appears to be of seminal importance for developments in the wider steppe region, so hopefully we'll get some samples from those sites.



At least as far as archaeology is concerned, your hypothesis of south-eastern European influence in the steppe seems to have some support. I'm not entirely sure whether that influence will show up in the DNA, but it will be interesting to see what the earliest stelae builders of the Black Sea look like genetically nonetheless. Their artistic traditions spread far and wide for sure, adopted by Scythians, Mongols, Turks and most of the other nomadic societies of the Eurasian plains.

Very similar stelae appear in Western Europe (mostly around the Western Med.) during the Bell Beaker period at least as early as 3000 BC

Too early to be from Yamnaya, for sure.

[markod]

Very similar stelae appear in Western Europe (mostly around the Western Med.) during the Bell Beaker period at least as early as 3000 BC

Too early to be from Yamnaya, for sure.

Yeah, Remedello culture in Italy has the earliest western stelae. That's why scholars used to think those people were steppe IE migrants.

[ToBeOrNotToBe]

Yeah, Remedello culture in Italy has the earliest western stelae. That's why scholars used to think those people were steppe IE migrants.

I think they're earlier in the South of France, but that intrigues me - do we know what Remedello DNA was? Similar to Iberia at the time?

[markod]

I think they're earlier in the South of France, but that intrigues me - do we know what Remedello DNA was?

Y-DNA I2 EEF types. Probably Megalithic Iberian colonist looking for tin.

Seems quite random that they would adopt the stelae tradition with such fidelity from the Black Sea I guess. Who knows what Chalcolithic people were thinking.

[ToBeOrNotToBe]

Y-DNA I2 EEF types. Probably Megalithic Iberian colonist looking for tin.

Seems quite random that they would adopt the stelae tradition with such fidelity from the Black Sea I guess. Who knows what Chalcolithic people were thinking.

Eh, I think that those stelae are signs of R1b L51. They're in full military attire when they depict men, and I know this is stereotyping but that isn't something you'd expect from Megalithic folk.

[markod]

Eh, I think that those stelae are signs of R1b L51. They're in full military attire when they depict men, and I know this is stereotyping but that isn't something you'd expect from Megalithic folk.

Afaik the R1b-rich Iron Gates people were typical Mesolithic 'goddess' worshippers. I don't think there's a link between warriorhood and R1b.

[Angela]

The eastern Balkanic steppe region is definitely in need of investigation. Not only Cernavoda and Ezero, but the LN cultures that prececde those as well; in particular Hamangia and Boian culture. The former contributed to the later steppe culture the typical anthropomorphic stelae. They appear already fully developed in LN Hamangia, barely distinguishable from the later stelae in the steppe:



Another thing worthy of consideration are the autosomally divergent samples from Mikhailovska, some of which seem to show elevated CHG affinity which could or could not indicate that there existed a cline within Ukraine, perhaps running from south to north. In fact the earliest steppe culture in which all constituent elements (herding, stelae, kurgans) exist side by side is Kemi Oba concentrated in the Crimean flatlands. Culturally and artistically Kemi Oba appears to be of seminal importance for developments in the wider steppe region, so hopefully we'll get some samples from those sites.



At least as far as archaeology is concerned, your hypothesis of south-eastern European influence in the steppe seems to have some support. I'm not entirely sure whether that influence will show up in the DNA, but it will be interesting to see what the earliest stelae builders of the Black Sea look like genetically nonetheless. Their artistic traditions spread far and wide for sure, adopted by Scythians, Mongols, Turks and most of the other nomadic societies of the Eurasian plains.

Statue Stelae of the Lunigiana:




People used to embed them in walls, use them as doorstops, whatever...



This is their distribution.



Jean Manco used to think that they were a sign of R1b Indo-Europeans coming across the Balkans, and then the Po and then onto the western Med, particularly at such an early time.

[IMG][/IMG]

I didn't and don't see why they would have just gone due west like that. In actuality, the area where they are found is on the route from north of the Alps into Italy. It follows the rivers, then the mountain passes over the Apennines and then the Magra south either into Italy itself or the Mediterranean. Or vice versa, of course. It's basically the same route as the later Via Francigena.

[ToBeOrNotToBe]

Jean Manco used to think that they were a sign of R1b Indo-Europeans coming across the Balkans, and then the Po and then onto the western Med, particularly at such an early time.



I didn't and don't see why they would have just gone due west like that. In actuality, the area where they are found is on the route from north of the Alps into Italy. It follows the rivers, then the mountain passes over the Apennines and then the Magra south either into Italy itself or the Mediterranean. Or vice versa, of course. It's basically the same route as the later Via Francigena.

There's actually one in Northern Saudi Arabia of all places (see below), of the same style dated to between 4000 and 3000 BCE(!!!), and supposedly (according to Russian Wikipedia, make of that what you will), the Kemi Oba stelae have links to the Caucasus.



I personally feel that matches well with this speculative map I made, with the origin of this anthropomorphic stelae folk at the point marked by L23:

[Pip]

Y-DNA I2 EEF types. Probably Megalithic Iberian colonist looking for tin.

Running my autosomal best-fit calculator on them, they come out as majority Bulgarian Neolithic, minority Iberian Neolithic (perhaps unsurprising, given Italy's position between the Balkans and Iberia). No noticeable Steppe ancestry. If their given yDNA readings are correct, they are of a variety of I2 that looks Iberian, with no ancient extant branches coalescing in Italy. My guess is that Remedello were principally Iberian men who mixed with local women, and ultimately died out.

I've no insights into how they would have appropriated stelae.

[markod]

Running my autosomal best-fit calculator on them, they come out as majority Bulgarian Neolithic, minority Iberian Neolithic (perhaps unsurprising, given Italy's position between the Balkans and Iberia). No noticeable Steppe ancestry. If their given yDNA readings are correct, they are of a variety of I2 that looks Iberian, with no ancient extant branches coalescing in Italy. My guess is that Remedello were principally Iberian men who mixed with local women, and ultimately died out.

I've no insights into how they would have appropriated stelae.

Well if they did have Balkan moms and Iberian Megalithic fathers that would be very interesting for sure, and it would provide at least some explanation as to how this very specific tradition spread to Western Europe. As Angela pointed out, many people were convinced that the Remedello guys were steppe immigrants because the stelae are nigh indistinguishable from those on the steppe.

One thing Anthony, Mallory and others didn't take into account is that they appear in Romania before they are adopted in Crimea whence the tradition spreads to the wider steppe region. I believe this to be an unfortunate result of the hyperfocus on the steppe; lots of nuance is lost due to this.

[Angela]

Well if they did have Balkan moms and Iberian Megalithic fathers that would be very interesting for sure, and it would provide at least some explanation as to how this very specific tradition spread to Western Europe. As Angela pointed out, many people were convinced that the Remedello guys were steppe immigrants because the stelae are nigh indistinguishable from those on the steppe.

One thing Anthony, Mallory and others didn't take into account is that they appear in Romania before they are adopted in Crimea whence the tradition spreads to the wider steppe region. I believe this to be an unfortunate result of the hyperfocus on the steppe; lots of nuance is lost due to this.

I completely agree.

Also, in terms of Remedello itself, like Baden, the assumption that they were steppe people came from the fact that their cultures exhibited many of the "hallmarks" of steppe culture. Whether it was a function of the fact that sometimes "pots are indeed just pots", and culture can travel without genes (westward), or a lot of it, like the stelae, might have developed first in the Balkans and then moved east, I don't know.