Shift from G2 to I2 dominance and WHG resurgence between the Early_N and the Late_N

Times of Neolithic Transition along the Western Mediterranean

Editors: García Puchol, Oreto, Salazar García, Domingo Carlos (Eds.)

​Discusses the expansion of farming at the beginning of the Neolithic in the Western and Central Mediterranean
Highlights the most recent advances in archaeological and scientific research aimed at better understanding the neolithisation process in the Mediterranean
Focuses on the Neolithic transition in the Western and Central Mediterranean regions
Brings together different novel methods and techniques applied to research assessing the neolithisation of the Mediterranean

https://www.springer.com/gp/book/9783319529370


This 2017 book looked into the demographic transition from G2a to I2a. G2a is present in studied Anatolian populations and also in Western, Central and Southeastern Neolithic cultures. However, basal levels of hg I and its subclade I2a are also detected, which are present in Mesolithic Scandinavia (Motala, Loschbour). I and I2a are both present in the Anatolian Neolithic, making it difficult to ascertain if they were introduced through demographic diffusion or if they signify acculturation of local hunter-gatherer groups. One of the samples from the Körös culture (KO2) is more closely related to the Anatolian and Greek Early Neolithic, while the other skelton from the same culture displays a strong Metholithic signature with hg I2a, clustering with Scandinavian hunter-gatherers (Gamba et al. 2014).
 
@Riverman
This supposed movement West would be kind of a little "G2a resurgence" then. Lol
If true, these people may have helped to brought this type of farmer component which is supposedly stronger in N. Italy and Sicily nowadays, more shifted to East (Balkan) than to West, I suppose.

Ygorcs started an interesting thread about it, by the way. His findings seem to fit well with the map I posted above.
He also mentions Remedello as a "Western" type, which also makes sense imo. Here:
https://www.eupedia.com/forum/threa...tempt-to-distinguish-Italics-from-Tyrrhenians

Isn't it more likely that the explanation is they moved around older G2a strongholds and allied up with others? Exactly like the Corded Ware did with GAC!
Could you elaborate this?
 
@Riverman
This supposed movement West would be kind of a little "G2a resurgence" then. Lol

Yes, that is true actually and its not the only case. Many haplotypes survived in a refuge, just to come back big time afterwards, if their position was not that bad for a big comeback after all. In Europe some of the best examples are I1 and E-V13. Almost dead, but coming back again.

If true, these people may have helped to brought this type of farmer component which is supposedly stronger in N. Italy and Sicily nowadays, more shifted to East (Balkan) than to West, I suppose.
Ygorcs started an interesting thread about it, by the way. His findings seem to fit well with the map I posted above.
He also mentions Remedello as a "Western" type, which also makes sense imo. Here:
https://www.eupedia.com/forum/threa...tempt-to-distinguish-Italics-from-Tyrrhenians

Ygorcs might be onto something and knows for sure more than me about this. My idea is too for some time that Etruscans might have come from a group within the Baden sphere with Yamnaya influences. But that's just speculation really. Yet I think a great deal of the linguistic diversity, even of the one which made it to the far west, was born in the area between Pannonia-Carpathians and Balkans, were the mixtures and alliances of the post-steppe world were particularly complicated and interwoven. Yet when and how exactly the modern Eastern Mediterranean/Levantine influences spread is also open to debate (?).

Could you elaborate this?

If a new player enters a region, he has three choices how to deal with locals:
- attack and subjugate or destroy them
- ally up and try to make a deal, especially if you need allies, probably against other foes
- trying to avoid them altogether, exploiting niches the others don't occupy or moving on, to new pastures so to say.

Yet what you do will always depend on how strong you are and how strong the others seem to be. If you reign supreme, the first option will be your first choice most of the time. But if you are not as sure, you might try the second and if that doesn't work out or you have a better destination in mind try to fall back to number three.

This is what the Corded Ware people did as well, and the reason why there was, initially, such a patch-work of cultures after the big first steppe expansion. Actually even more of a patchwork before. Because the newcomers did cut into the soft parts and most valuable parts for them, while bypassing all areas not interesting or with a strong and hostile population. The Corded Ware people seem to have done all three things in relation to GAC, depending on the circumstances. They did annihilate some tribes, they did ally up and assimilated some others, while bypassing other groups where it seemed to have been the better option. These were sometimes much later integrated into the broader steppe sphere.
An example might be from the perpetrators of the Eulau massacre, most likely coming from the Sch?nfelder Culture, which was part of this TRB-GAC related horizon and persisted for quite long beside their Corded Ware neighbours, even if violence seems to have errupted occasionally, as is evidenced by finds in Eulau.

What I wanted to propose is that whereever this I2a/WHG enriched Middle Neolithic movement started (I'd assume in the North but with new South Eastern influences coming up as well), the logical assumption is they acted the same, using all three options where it seemed suitable. E.g. in parts of Bohemia, Switzerland, Sardinia and so on, the local G2a Neolithic communities were strong enough for option 2 and 3, resulting in more mixed communities with a lower share or even a relative persistence of the locals. In some other places and groups, especially for the GAC/KAK groups, it seems to have been a more exclusive club.
 
Times of Neolithic Transition along the Western Mediterranean

Editors: García Puchol, Oreto, Salazar García, Domingo Carlos (Eds.)

​Discusses the expansion of farming at the beginning of the Neolithic in the Western and Central Mediterranean
Highlights the most recent advances in archaeological and scientific research aimed at better understanding the neolithisation process in the Mediterranean
Focuses on the Neolithic transition in the Western and Central Mediterranean regions
Brings together different novel methods and techniques applied to research assessing the neolithisation of the Mediterranean

https://www.springer.com/gp/book/9783319529370


This 2017 book looked into the demographic transition from G2a to I2a. G2a is present in studied Anatolian populations and also in Western, Central and Southeastern Neolithic cultures. However, basal levels of hg I and its subclade I2a are also detected, which are present in Mesolithic Scandinavia (Motala, Loschbour). I and I2a are both present in the Anatolian Neolithic, making it difficult to ascertain if they were introduced through demographic diffusion or if they signify acculturation of local hunter-gatherer groups. One of the samples from the Körös culture (KO2) is more closely related to the Anatolian and Greek Early Neolithic, while the other skelton from the same culture displays a strong Metholithic signature with hg I2a, clustering with Scandinavian hunter-gatherers (Gamba et al. 2014).

Great info, Third Term. Thanks. :)
 
My real point was that the early Neolithic got more violent and in the transitional period the even more aggressive and warlike hunter-derived clans took over.

The Neolithic people became more violent during the times of scarcity after climate change and crop failures. That happens in every culture. There is absolutely no indication anywhere that it was because of battles between I2a "clans" and G2a "clans".

Once again, please point me to the papers which show that hunter-derived "clans" murdered the G2a clans to take over.

I keep asking, and you keep failing to do it, and the reason is that no such evidence exists. You are just hypothesizing that it "must have" happened that way.


It proves that most changes were caused by violent, aggressive behaviour, by wars. Why should the transition from I2a/WHG at the end of the early Neolithic, in one of the most brutal times of European history, be any different? That's what my argument was about. The LBK fought with their teeth and claws against each other, mutilated, utterly annihilated their enemies, even the young and women more than once, but suddenly they say friendly hello to the new forager males and tell them to take their land and wives?

Who says they didn't just absorb the people whom they encountered? They took forager women as wives, why not include some I2a forager men too? They needed strong backs. What else was that hg male doing in the Hungarian Neolithic village in Gambas et al.


I can't stop saying what I never said. I said they were not nomadic and they did crop farming too, but at a lower rate in comparison to animal husbandry than LBK, that's what I said. They were more mobile and pastoralist than early farmers, but they were still agro-pastoralists with houses and no pastoralists living only in tents and caves.

True nomadism is exceptional, semi-nomadism and transhumance is more common. The GAC practised, most likely, transhumance, its written in the paper I quoted. But they were not nomadic, I NEVER EVER said so, this is what you quoted from me yourself, I will bold what's decisive:

So probably my wording was bad, I wanted to say that they did have (smaller, poorer) houses and were no nomads and they did practise crop farming. If the sentence was wrong, I'm sorry for the misunderstanding. :ashamed2:


Apologies accepted. For the record this is just cut and paste.

"Whether the GAC were full blown pastoralists is up to debate, but by most accounts they were not. They were just agro-pastoralists which shifted in the direction of pastoralism in comparison to the preceding early Neolithic cultures in particular. They were no nomadic herders with no crop farming and without houses, even if the buildings found so far were rather poor in comparison to LBK houses.


I wouldn't say they killed all G2a men everywhere, but the shift was happening violently and was drastic enough. There was an increase of violence and competition from the start to the end of the early Neolithic. When the dust settled, better organised, more mobile and warlike agro-pastoralist societies emerged, dominated by I2a. The earlier Neolithic people might have been as violent, but they were not as good at it, at least that's what the end result is suggesting. Of course, part of the reason for the increase in violence is their failure to pacify the border ones and to fully exploit the ecological zone of Northern Europe. The situation must have been pretty dire in some places, especially if the climate changed just a little bit, with, like I said, malnourished and inbred people surrounded by foragers in the woods and fighting with each other for what's left.
That's when the second stage of the Neolithisation with increased livestock breeding kicks in.

You apparently haven't read the literature, and you didn't read my comments carefully. Livestock breeding was extremely important in the Balkans from the earliest days of the Neolithic. Some areas were almost completely dependent on livestock herding of cattle. NO, the I2A men did not introduce it. They were virtually all G2a at that point. So, you're wrong.

As for the rest of the paragraph it's sheer speculation. What are you basing it on? What papers? What archaeology shows this?

The GAC samples which I quoted were all I2a (forager) yDNA and the complete mtDNA variation was Neolithic. It is also crucial when and where the mtDNA was acquired. I know of no place where the drastic increase of I2a/WHG correlated with a synchronous rise of HG mtDNA. I'm sorry, I don't have that information and couldn't find it. Whether some Iberian Neolithics got local mtDNA in a preceding period is not relevant, its about the timing. Of course, the GAC had mtDNA of foragers too, like U5b, and I wouldn't say that it was only males coming in, but the its the ratio. The ratio is completely on the paternal side, even more so if considering that the preceding and not as I2a dominated groups had U lineages as well. So the presence doesn't explain the change.

All you had to do was look on our own site.

This looks like a genocidal wipe out to you? The mtdna is 14% HG.

'No Y-chromosomal DNA from the Funnelbeaker itself has been tested to date. The following samples are from the related Baalberge and Salzmünde cultures in central-east Germany, but may not necessarily reflect the paternal lineages found in Scandinavia and northern Germany at the time. Indeed the mtDNA of these various groups displays considerable differences.
  • Baalberge group (5,800 to 5,350 ybp ; central Germany): I, R1
  • Salzmünde group (5,400 to 5,000 ybp : East Germany): G2a2a (x2), I2a1b1a (x2)
mtDNA


  • Funnelbeaker Culture (samples from Sweden) : H (x3), H1, H24, J1d5, J2b1a, K1a5, T2b
    • Baalberge group (c. 5,800 to 5,350 ybp ; central-east Germany): H (x3), H1e1a, H7d5, HV, J, K1a (x2), N1a1a, T1a1, T2b, T2c (x2), T2e1, U5b2a2, U8a1a, X, X2c
    • Walternienburg-Bernburg group (c. 5,100 to 4,700 ybp ; central-east Germany): H, H1e1a3, H5, K1, K1a (x2), T2b, U5a, U5b, U5b1c1, U5b2a1a, V, W, X
    • Salzmünde group (5,400 to 5,000 ybp : central-east Germany): H (x2), H3 (x2), H5, HV, HV0, J, J1c (x2), J2b1a, K1, K1a, K1a4a1a2, N1a1a1a3 (x2), T2b (x2), U3a, U3a1, U5b, V, X2b1'2'3'4'5'6
    • Outliers from Gotland, Sweden (5,300 to 4,700 ybp): H7d, HV0a, J1c5 (2x), J1c8a, K1a2b (2x), K2b1a, T2b8"

Globular Amphora: Let's remember where they were located please.

"In a 2017 genetic study published in Proceedings of the Royal Society B, the fifteen samples of mtDNA was extracted. The majority of the samples belonged to subclades of U and Haplogroup H (mtDNA), while J, W and K was also detected. The remains were found to closely related to Neolithic European farmers and Western Hunter-Gatherers, with little genetic relations to the Yamnaya culture in the east. The authors of the study suggested that the Globulara Amphora culture was non-Indo-European-speaking, but with cultural influences from Yamnaya.[4]

A February 2018 study published in Nature included an analysis of eight males of the Globular Amphora culture. Three of them carried haplogroup I2a2a1b and a subclade of it; two carried I2a2; one carried I2; one carried BT and one carried CT.[5]

In a 2019 genetic study published in the Proceedings of the National Academy of Sciences of the United States of America, 15 skeletons from the Koszyce mass grave in southern Poland, which is ascribed t
to the Globular Amphora culture. The individuals were all shown to be members of an extended family...all were found to belong to I2a-L801."

Please look at the autosomal dna of Globula Amphora...now look back at LBK. You see evidence of genocide and take over there between the two periods?? You have to be kidding.
Genetic-affinities-of-the-Koszyce-individuals-and-other-GAC-groups-here-including-Zlota.png


From Shroeder et al 2019...

As per data set 2, no cattle remains, only pig.

Look at data set 5. These people were not fair haired and blue eyed. Have no idea about the ones from the prior papers.

The reason that they were mobile is unknown. Did they learn it from the steppe people with whom they were in contact? Did the influence go the other way? There's no way of telling at the present time. There's certainly no proof that the male ancestors of these people killed all the G2a men with whom they came into contact. Maybe they just liked EEF women better, and kept on mating with them, and that's how they wound up 70% EEF.

Be aware too that GAC is not Iberian Neolithic or English Neolithic.

You are right if you look at the WHG alone, but I'm pretty sure its much more. However, my assumption is that a people which was probably more around 50:50 expanded on. Similar to a comparison of Yamnaya -> Corded Ware -> CE Bell Beaker -> Iberian Bell Beaker. At the end of the chain of expansions the steppe was already fairly low. But my assumption would be more like the original WHG led group being probably 30-40 percent additional WHG. This means its no big deal in terms of the big ancestral components, but it could mean a quite drastic regional replacement nevertheless.

There's no need to respond. You're right. It's all assumptions.


Ok, I just think there is no better explanation out there. If there is, I never heard about it and the question was raised, once more, not even by myself. The alternatives are heard are all far fetched. Definitely not more probable than my proposition - at least to me and for the moment. I'm however open to all arguments and facts which might contradict the hypothesis I deem the most probable and hoping for new inputs to the discussion. Peace :)

Well, that's certainly a more measured approach.
 
Who says they didn't just absorb the people whom they encountered? They took forager women as wives, why not include some I2a forager men too?

Because foreign men being always more a problem and less of a win, unless they bring in a real benefit for the group. Despite this, that happened, in different regions at a different ratio. But G2a was always keeping its dominant position in the pre-transitional phase. That's the hallmark of small scale assimilation: The yDNA changes only slowly and the original founders don't disappear, keep staying on top. We also have the studies which prove the structure of the early Neolithics being centered around male clans. Those won't disappear without being broken, from which direction it happened in every region, that's what we need to clear up.
But I have to agree with you that for being sure, that for saying it definitely, more data is needed, especially for the evolution of TRB.

What else was that hg male doing in the Hungarian Neolithic village in Gambas et al.

Single individuals can be everything, even prisoners of wars and human sacrifices. But yes, I know it from different places too, individuals were assimilated. But there is a difference between individuals and the kind of change and overtake we can observe at the transition from the early to the middle Neolithic period.

You apparently haven't read the literature, and you didn't read my comments carefully. Livestock breeding was extremely important in the Balkans from the earliest days of the Neolithic. Some areas were almost completely dependent on livestock herding of cattle.

I was speaking especially about LBK and the West. TRB-GAC being always brought up as an example with increased livestock breeding and use of cattle in comparison to the preceding cultures. Its entirely possible that parts of that transition were actually introduced from the South East. But we deal with big changes in the North, at the time the proposed transition is taking place, and the most likely scenario is still that it was Northern hunter lineages being the avantgarde THERE.

Please look at the autosomal dna of Globula Amphora...now look back at LBK. You see evidence of genocide and take over there between the two periods?? You have to be kidding.

Well, no genocide in the sense of the preceding culture disappearing, but a take over nevertheless. Just look at this like if a group with more like 50:50 have taken over, the original carriers were more WHG, but mixed with farmer wives and they began to dominate other groups, some were broken, others allied up. Between LBK and TRB its doubled, to GAC tripled. Autosomal data interpretation would be much easier in many parts of Europe if the ethnic population differences would be as big as between LBK and GAC. The PCA is quite striking and they seem to represent completely different ethnicities.

As per data set 2, no cattle remains, only pig.

Remains and burials differed, but GAC was, you can see this in the various newer articles, online entries and the literature, centered around cattle breeding. They had pigs and others animals too of course.

Look at data set 5. These people were not fair haired and blue eyed. Have no idea about the ones from the prior papers.

Its not relevant to the current debate, but some other samples other than the victims of the massacre were. I never said that all GAC were light, I know they weren't.

The reason that they were mobile is unknown.

One of the main reasons for mobility is the ability of a people to be mobile. Because of their subsistence and strength. As well as a need for it, like finding better and new pastures. Like explained above I don't mean they were nomadic pastoralists, but they used transhumance and colonised new regions.

Did they learn it from the steppe people with whom they were in contact? Did the influence go the other way? There's no way of telling at the present time.

Its even possible that both the Northern agro-pastoralists of TRB-GAC and the steppe people were influenced by the same source. Because for the steppe people its clear that they adopted cattle herding from their Neolithic neighbours in the Western Ukrainian region.

There's certainly no proof that the male ancestors of these people killed all the G2a men with whom they came into contact.

Never said so. They just have to break their rule in some regions, ally up in others. In the course of this a lot of G2a men were killed, but certainly not all and everywhere. Never said that. What I said that without a violent expansion, the spread of a new group with increased I2a/WHG, this pattern is not explainable.

Maybe they just liked EEF women better, and kept on mating with them, and that's how they wound up 70% EEF.

Yes, but how should they have got them? They took the land, the livestock and the women. That's not for free and I doubt the LBK-related G2a clans just agreed on such scenario...

And this is also what I see for the whole Megalithic sphere and Iberia. It might be quite complicated, but we deal with new incomers.

I agree with the uncertainties. It is a pity that a lot of data both from the archaeological as well as the ancient DNA side is still missing to complete the picture and explain what really happened in detail. Let's just hope new data makes better explanations possible.
 
Great info, Third Term. Thanks. :)
I confess I missed the detail posted by ThirdTerm, and you brought me into it.
Does it mean that some Y-DNA I2 and perhaps pre-I1 may have been absorved by early farmers already in Balkan and then just expanded together, "founding" some areas such UK, for example? Not sure about GAC, since you suggested the samples belonged to a same family, if I understand it right.

Below is an interesting graph posted by Bicicleur in another thread, regarding the drop of Y diversity all over the world after 10k*, culminating in ~5k ybp, and then super-fast growing again. The R-L151 TMRCA, for example, is 4800 ybp (it roughly coincides to Cucuteni-Trypillia's "death", so to speak), not even far in time from those samples in Switzerland, suggestive of a rapid expansion, after thousands of years of relatively normal contacts with "Old Europe" (late Trypillia even had some Steppe ancestry, while farmer mtDNA was already present in Steppe; if I'm not missing something).

img.jpg


*While curiously mtDNA kept somewhat stable, just a bit "flattered" (not sure it's the right word). I didn't get the differences of pop size in the two graphs though; I'll try to read the paper later in order to understand it better.

So, perhaps the question is not only how they dropped, and more than one (in)direct factor is naturally possible (the worldwide phenomenon is intriguing), but how they re-expanded after the supposed crisis? Some tribes may have taken advantage of it, which doesn't mean "technology" explains it all, neither physical advantage (such few cm in height, for example, related perhaps to a diet richer in protein). Perhaps an example of that is Central/East Asian in Europe from Iron Age onwards, or even Mongol expansions, with an important difference related to population size of invaded areas. Afaik, they were not technologically or physically advantaged, neither the Romans themselves were "taller" than the conquered people. It's possibly a non-issue.

Anyway, replacements are well characterized by two scenarios from different times and same place in which the features are changed, as in LN Switzerland, for instance. Here's my question: is it noticeable when it comes to this theoretical WHG resurgence? Like a previous pop rich in G2a and poor in WHG component being replaced in certain place by another pop poor in G2a, rich in I2 and substantially richer in WHG, before the Bronze Age? Perhaps in Scandinavia? Iberia?
Another question would be: when and where exactly did the biggest changes from mtDNA U and types of K to H-T-J etc. start to happen, and how? It may be relevant knowing it. Yamnaya, for instance, already had mtDNA T and others, whereas the Y-DNA pool was dominated by R-Z2103 (again, by memory).

By the way, I remember that some researchers in the past tried to check these sex biases through biases in X chromosome (don't remember if farmers vs. WHG or farmers vs. Steppe). Not sure, but I guess it was Reich Lab. IIRC, they found no bias, but another lab would have found it later. Then, again, Reich's said no, there was not. Finally, this another lab disagreed and reinforced its first findings. je je je
Well, so far back in time, so many variables, so many "ifs"/assumptions... If it's complicated for professionals, it's especially complicated for newbies, as myself and probably big part of members here. It's still fascinanting though. :)
 
I confess I missed the detail posted by ThirdTerm, and you brought me into it.
Does it mean that some Y-DNA I2 and perhaps pre-I1 may have been absorved by early farmers already in Balkan and then just expanded together, "founding" some areas such UK, for example? Not sure about GAC, since you suggested the samples belonged to a same family, if I understand it right.
Below is an interesting graph posted by Bicicleur in another thread, regarding the drop of Y diversity all over the world after 10k*, culminating in ~5k ybp, and then super-fast growing again. The R-L151 TMRCA, for example, is 4800 ybp (it roughly coincides to Cucuteni-Trypillia's "death", so to speak), not even far in time from those samples in Switzerland, suggestive of a rapid expansion, after thousands of years of relatively normal contacts with "Old Europe" (late Trypillia even had some Steppe ancestry, while farmer mtDNA was already present in Steppe; if I'm not missing something).
img.jpg

*While curiously mtDNA kept somewhat stable, just a bit "flattered" (not sure it's the right word). I didn't get the differences of pop size in the two graphs though; I'll try to read the paper later in order to understand it better.
So, perhaps the question is not only how they dropped, and more than one (in)direct factor is naturally possible (the worldwide phenomenon is intriguing), but how they re-expanded after the supposed crisis? Some tribes may have taken advantage of it, which doesn't mean "technology" explains it all, neither physical advantage (such few cm in height, for example, related perhaps to a diet richer in protein). Perhaps an example of that is Central/East Asian in Europe from Iron Age onwards, or even Mongol expansions, with an important difference related to population size of invaded areas. Afaik, they were not technologically or physically advantaged, neither the Romans themselves were "taller" than the conquered people. It's possibly a non-issue.
Anyway, replacements are well characterized by two scenarios from different times and same place in which the features are changed, as in LN Switzerland, for instance. Here's my question: is it noticeable when it comes to this theoretical WHG resurgence? Like a previous pop rich in G2a and poor in WHG component being replaced in certain place by another pop poor in G2a, rich in I2 and substantially richer in WHG, before the Bronze Age? Perhaps in Scandinavia? Iberia?
Another question would be: when and where exactly did the biggest changes from mtDNA U and types of K to H-T-J etc. start to happen, and how? It may be relevant knowing it. Yamnaya, for instance, already had mtDNA T and others, whereas the Y-DNA pool was dominated by R-Z2103 (again, by memory).
By the way, I remember that some researchers in the past tried to check these sex biases through biases in X chromosome (don't remember if farmers vs. WHG or farmers vs. Steppe). Not sure, but I guess it was Reich Lab. IIRC, they found no bias, but another lab would have found it later. Then, again, Reich's said no, there was not. Finally, this another lab disagreed and reinforced its first findings.
je je je
Well, so far back in time, so many variables, so many "ifs"/assumptions... If it's complicated for professionals, it's especially complicated for newbies, as myself and probably big part of members here. It's still fascinanting though. :)

I remember the back and forth. :) I don't know if I'm remembering the Reich Lab position precisely, however. Did the Reich people say the steppe men didn't absorb farmer women, or just that they also brought women with them ?

This relates to the point raised by Third Term, which is that it's sometimes difficult to know where a particular dna line originated. Some yDna I2a was already in Anatolia, never mind the Balkans, as was pre-I. Are the samples resolved enough, or could they even ever be resolved enough, given that they are ancient samples, that it would be possible to determine if certain I2a yDna lines or even pre I lines were local to Europe or originated in Anatolia, or could it even be both?

Then there's Cardial to consider. I think that looking at the trail it seemed to me at one point that they picked up I2a along the way, so it could have been a founder effect in Spain, in the Megalithic etc. Of course, the same could be true of cultures like GAC. As LBK dissolved, some of the men might have carried I2a, and being patriarchal and patrilocal you get the high I2a in family settlements. We have the I2a man of a reasonable age from the Hungarian Neolithic (Gambas et al), so they weren't all killed.

I said Europe deliberately, because there weren't very many hunter gatherers in the Balkans to begin with, other than pretty far north around the Iron Gates, right? As in many parts of Europe, they were clustered around the good fishing areas there. Plus, didn't Mathiesen find that the Balkan Neolithic people didn't expand much outside the Balkans? Not that I think that Mathiesen is infallible.

If he was correct, LBK was from a slightly different group and a lot of the expansion would be from there?

I think people also forget the cave burial where we find both hunter-gatherers and farmers of the same period cozily buried side by side. Related through a bride exchange, perhaps?

As to the mtDna we have the same issue. Did some of the "European farmer" mtDna migrate out to the steppe in small amounts and then migrate back in?

That's why I think the researchers were looking at the X. If the Reich Lab is correct that "softens" the kill all the men, steal all the women scenario, although bride kidnapping during raids by young men is certainly a documented part of Indo-European culture. It would also mean that women might indeed have died at the same rates as their men from the plague.

Your questions about the turnover are important ones. I'll respond in the next post.

As you say, very complex. I don't see how one can be dogmatic about how this particular scenario played out.

Another complexity can be found in the fact that while there is certainly plenty in the record to show invading men trying to wipe out local men, and indeed we saw it in the Balkans barely 60 years ago, and in the conflict between the Hulu and the Tutsis, there are some indications from ancient genetics that this wasn't invariably the case.

As I pointed out in the mta thread, the wealthy tomb of an Ostrogoth "Chief", contained his bedecked skeleton, and he turns out to be closest to modern Pontic Greeks. The Ostrogoths seem to have been pretty inclusive. The same is true of the Scythians. You have "eastern" type Scythians, "northern" type Scythians, and "Italian" like Scythians. I'm pretty close to some of the latter myself. Sarmatians, on the other hand, were not inclusive at all. Nor were Langobards for the most part. Moving forward a little in history, the Huns seem to have absorbed a lot of men of different tribes, and the Alans too as well, yes?

So, different scenarios at different times and places.
 
I remember the back and forth. :) I don't know if I'm remembering the Reich Lab position precisely, however. Did the Reich people say the steppe men didn't absorb farmer women, or just that they also brought women with them ?

This relates to the point raised by Third Term, which is that it's sometimes difficult to know where a particular dna line originated. Some yDna I2a was already in Anatolia, never mind the Balkans, as was pre-I. Are the samples resolved enough, or could they even ever be resolved enough, given that they are ancient samples, that it would be possible to determine if certain I2a yDna lines or even pre I lines were local to Europe or originated in Anatolia, or could it even be both?

Then there's Cardial to consider. I think that looking at the trail it seemed to me at one point that they picked up I2a along the way, so it could have been a founder effect in Spain, in the Megalithic etc. Of course, the same could be true of cultures like GAC. As LBK dissolved, some of the men might have carried I2a, and being patriarchal and patrilocal you get the high I2a in family settlements. We have the I2a man of a reasonable age from the Hungarian Neolithic (Gambas et al), so they weren't all killed.

I said Europe deliberately, because there weren't very many hunter gatherers in the Balkans to begin with, other than pretty far north around the Iron Gates, right? As in many parts of Europe, they were clustered around the good fishing areas there. Plus, didn't Mathiesen find that the Balkan Neolithic people didn't expand much outside the Balkans? Not that I think that Mathiesen is infallible.

If he was correct, LBK was from a slightly different group and a lot of the expansion would be from there?

I think people also forget the cave burial where we find both hunter-gatherers and farmers of the same period cozily buried side by side. Related through a bride exchange, perhaps?

As to the mtDna we have the same issue. Did some of the "European farmer" mtDna migrate out to the steppe in small amounts and then migrate back in?

That's why I think the researchers were looking at the X. If the Reich Lab is correct that "softens" the kill all the men, steal all the women scenario, although bride kidnapping during raids by young men is certainly a documented part of Indo-European culture. It would also mean that women might indeed have died at the same rates as their men from the plague.

Your questions about the turnover are important ones. I'll respond in the next post.

As you say, very complex. I don't see how one can be dogmatic about how this particular scenario played out.

Another complexity can be found in the fact that while there is certainly plenty in the record to show invading men trying to wipe out local men, and indeed we saw it in the Balkans barely 60 years ago, and in the conflict between the Hulu and the Tutsis, there are some indications from ancient genetics that this wasn't invariably the case.

As I pointed out in the mta thread, the wealthy tomb of an Ostrogoth "Chief", contained his bedecked skeleton, and he turns out to be closest to modern Pontic Greeks. The Ostrogoths seem to have been pretty inclusive. The same is true of the Scythians. You have "eastern" type Scythians, "northern" type Scythians, and "Italian" like Scythians. I'm pretty close to some of the latter myself. Sarmatians, on the other hand, were not inclusive at all. Nor were Langobards for the most part. Moving forward a little in history, the Huns seem to have absorbed a lot of men of different tribes, and the Alans too as well, yes?

So, different scenarios at different times and places.
Thanks for the insights!

As for the X vs. Autosomal bias, I just checked. It was about Farmers vs. Steppe (not WHG). And you're right. It seems I missed the fact that Reich was not the first to check this possible bias. Actually Reich and Lazaridis just "failed" to replicate the findings of a previous paper (Goldberg et al.). Here are more infos:

https://www.eupedia.com/forum/threads/33749-Failure-to-replicate-sex-bias-in-steppe-migrations
 
@Angela: The steppe mtDNA issue being complicated by the fact that some of the later, post-Mesolithic and Near Eastern derived haplogroups were coming from the East (CHG), as well as from the West (EEF). You know that of course, but there are still debates going on about whether this kind of T was taken up from farmers further West or was present among the early foragers in the North Pontic which were ancestry to the later steppe people.

As I pointed out in the mta thread, the wealthy tomb of an Ostrogoth "Chief", contained his bedecked skeleton, and he turns out to be closest to modern Pontic Greeks. The Ostrogoths seem to have been pretty inclusive. The same is true of the Scythians. You have "eastern" type Scythians, "northern" type Scythians, and "Italian" like Scythians. I'm pretty close to some of the latter myself. Sarmatians, on the other hand, were not inclusive at all. Nor were Langobards for the most part. Moving forward a little in history, the Huns seem to have absorbed a lot of men of different tribes, and the Alans too as well, yes?

Another aspect of this is how they were living with each other. Like they could form different castes, like the Lombards in Pannonia, or they could live side by side for quite long, but were still people apart. Like the Avars actually used Slavic tribesmen as cannon fodder, the Suebians and other Germanics had Sarmation groups among them. On the long run, most of these people mixed, but they could also live together for a time, as long as they were on campaign, as long as the alliance was working, and splitting afterwards. So I would be very cautious if its about some burials. They could have had special roles or they might have been a different tribus within the alliance. The best proof for a real panmixture and full scale integration on equal terms is always reciprocal marriage. This means we find men of group X and women of group Y as couples and vice versa. Its the vice versa which makes the difference. Because if we find only males taking the women from group Y, they were not on equal terms, they were dependent, probably even slave-like.
Similarly, if e.g. one elite Sarmation knight was accepted as a Germanic or Slavic upper class member, this doesn't have to mean ALL of his kinsmen would have been equally accepted. It might just mean that he, as an exceptional and exceptionally valuable member of the group, was fully accepted.

The Avars had all kind of people around them, but their elite was fairly Eastern steppe for quite some time and most of their tributaries, especially the Slavs, which rose up against their rule later, were treated quite bad. So having foreign people around you doesn't mean you treat them well or accept them as group members. Like African slaves in 17th century America. You could find burials of masters and slaves side by side in some areas probably, but they were not really related or had the same status.

The study on the Lombards was really great, in this respect, not the way it was communicated, as it showed that first some mixed individuals, later even predominantely Southern Europeans, could make it to the higher ranks, form their own family dynasty in the Lombard reign. This happened in Italy, over time, but much less so in Pannonia - at least according to the preliminary results. Without such careful and extensive analyses, definitive conclusions about the social and ethnic relations are quite often hard to make.
 
@Riverman
Regarding the "counterpart" you mentioned, the way I see it, "Old Europe" collapsed at certain point anyway, and so several Y lineages apparently, of all kinds. If diseases played a role against Old Europe, they don't seem to have been the only factor - as per that graph I posted above (already assuming it's accurate). How the "re-expansion" happened would be another story, and may also help to explain the apparent bias. It possibly doesn't "prove" there was no counterpart before Steppe expansions though. Maybe there was (some sort of) counterpart, maybe there wasn't. I don't know, hence the references and questions in my previous email, and hence the importance of this research on X chromosome.

ED: Agree that may be difficult to distinguish between CHG and EEF mtDNA sometimes. More ancient DNAs may provide a better idea on the lineages already present before the rapid expansion.

By the way, this is a recent paper on contacts between Farmer and Steppe folks in the East:
Gene-flow from steppe individuals into Cucuteni-Trypillia associated populations indicates long-standing contacts and gradual admixture
https://www.nature.com/articles/s41598-020-61190-0

Thanks for the insights!
As for the X vs. Autosomal bias, I just checked. It was about Farmers vs. Steppe (not WHG). And you're right. It seems I missed the fact that Reich was not the first to check this possible bias. Actually Reich and Lazaridis just "failed" to replicate the findings of a previous paper (Goldberg et al.). Here are more infos:
https://www.eupedia.com/forum/threads/33749-Failure-to-replicate-sex-bias-in-steppe-migrations
Forgot to mention that not even Goldberg et al. found sex bias in Neolithic itself. It says:
"we find no evidence of sex-biased admixture during the migration that spread farming across Europe during the early Neolithic"

I didn't read the paper, so I don't know if they checked for farmer male vs. whg female (possibly, since it mentions "early Neolithic"), for the other way around or for both ways. "If" it's accurate, and we don't know it (given the opposition by big figures such Reich and Lazaridis regarding the supposed Steppe male vs. Farmer female bias), it could be relevant for the debate in this thread, either regarding the way farmers expanded and/or the WHG resurgence.
 
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@Angela: The steppe mtDNA issue being complicated by the fact that some of the later, post-Mesolithic and Near Eastern derived haplogroups were coming from the East (CHG), as well as from the West (EEF). You know that of course, but there are still debates going on about whether this kind of T was taken up from farmers further West or was present among the early foragers in the North Pontic which were ancestry to the later steppe people.



Another aspect of this is how they were living with each other. Like they could form different castes, like the Lombards in Pannonia, or they could live side by side for quite long, but were still people apart. Like the Avars actually used Slavic tribesmen as cannon fodder, the Suebians and other Germanics had Sarmation groups among them. On the long run, most of these people mixed, but they could also live together for a time, as long as they were on campaign, as long as the alliance was working, and splitting afterwards. So I would be very cautious if its about some burials. They could have had special roles or they might have been a different tribus within the alliance. The best proof for a real panmixture and full scale integration on equal terms is always reciprocal marriage. This means we find men of group X and women of group Y as couples and vice versa. Its the vice versa which makes the difference. Because if we find only males taking the women from group Y, they were not on equal terms, they were dependent, probably even slave-like.
Similarly, if e.g. one elite Sarmation knight was accepted as a Germanic or Slavic upper class member, this doesn't have to mean ALL of his kinsmen would have been equally accepted. It might just mean that he, as an exceptional and exceptionally valuable member of the group, was fully accepted.

The Avars had all kind of people around them, but their elite was fairly Eastern steppe for quite some time and most of their tributaries, especially the Slavs, which rose up against their rule later, were treated quite bad. So having foreign people around you doesn't mean you treat them well or accept them as group members. Like African slaves in 17th century America. You could find burials of masters and slaves side by side in some areas probably, but they were not really related or had the same status.

The study on the Lombards was really great, in this respect, not the way it was communicated, as it showed that first some mixed individuals, later even predominantely Southern Europeans, could make it to the higher ranks, form their own family dynasty in the Lombard reign. This happened in Italy, over time, but much less so in Pannonia - at least according to the preliminary results. Without such careful and extensive analyses, definitive conclusions about the social and ethnic relations are quite often hard to make.


the goths where split into 2 groups in early 150BC ............the Ostrogoth's mixed with sarmatians, settled on the black sea then went to Italy ( the only goths that went to Italy ).

the Visigoths stayed in modern Poland and went directly to Iberia ( first mixing with the Germanic Suebi tribes of germany ), later north-africa and these goths destroyed Rome .............the visigoths did not mix with sarmatians

these 2 goth groups would have different types of DNA
 
Regio: Another question would be: when and where exactly did the biggest changes from mtDNA U and types of K to H-T-J etc. start to happen, and how? It may be relevant knowing it. Yamnaya, for instance, already had mtDNA T and others, whereas the Y-DNA pool was dominated by R-Z2103 (again, by memory).

In a relatively early 2012 paper by Fu, Q, they looked at the whole set of Neolithic versus hunter-gatherer dna, so not quite what we want, but...
https://europepmc.org/article/med/22427842

So, we have an average of 12% mtDna U.

xEALz9z.png


Some sites, though, have no mtDna U at all, including the Cucuteni Tripolye site of Verteba in the Ukraine, which is right before the arrival of the steppe people.
https://www.biorxiv.org/content/10.1101/217109v1.full.pdf

This paper does have a chart with the results of some Neolithic sites. It's on page 33. However, believe it or not, they don't break out mtDna U.

In 2013 we get another pie chart of averages, but at least it shows the differences before and after 4500 BC. In the "pioneer" days it seems the farmer communities took on some HG women, but no farmer women went to HG communities. After that, the percentage of HG mtDna went up in the farmer communities, but the increase of farmer mtDna in hunter communities increased more, so at the end this chart would make it appear that the exchange went both ways, and was at fairly equal levels, I think, but averaging maybe 25%?
https://www.karger.com/Article/PDF/356933

DZxBz4m.png



I think the mtDna I might be steppe, Maciamo thinks so, and it's unclear to me whether some of the other HG lineages might also have come from the steppe. I would think perhaps a few of them.

The early Iberian Neolithic had more HG than the LBK and descendant groups, but I don't remember them having a lot more HG mtDna.

Anyway, the Tassi et al al paper's Supplement has a list of mtDna sequences of HG and Neolithic sites.(the GAC paper). It's Table S6.
https://figshare.com/collections/Su...he_spread_of_Indo-European_languages_/3928333

It doesn't look to me like a lot of mtDna U anywhere, but on the other hand the total number of samples is small, so it may add up to about 12% in the early period. I don't understand how the prior paper could get 25% from the samples listed in this paper. Do you see any LN actual farmer culture obviously missing from the list?

Unless someone knows of a compendium which has been kept up to date of all mtDna and was done by someone who is precise and careful, that's all I have.
 
the goths where split into 2 groups in early 150BC ............the Ostrogoth's mixed with sarmatians, settled on the black sea then went to Italy ( the only goths that went to Italy ).

the Visigoths stayed in modern Poland and went directly to Iberia ( first mixing with the Germanic Suebi tribes of germany ), later north-africa and these goths destroyed Rome .............the visigoths did not mix with sarmatians

these 2 goth groups would have different types of DNA

That's right, and there might have been even more groups of Goths around, even though we primarily refer to Western and Eastern Goths, which fought with each other on the Catalanaunian plains.

the visigoths did not mix with sarmatians

The Ostrogoths mixed with all kinds of people. The Visigoths might have mixed with Sarmatians as well, because the Germanic tribes had often allied Sarmatian riders with them. At the battle of Adrianople, both the Goths and the Romans had Sarmatians in their forces, especially among their cavalry. This was not just true for the Ostrogoths, but for many Germani tribes, like the Suebi (which I actually referred to) and the Vandals:
Upon the Hunnic defeat of the Goths on the Pontic Steppe around 375 AD, many of the Alans migrated westwards along with various Germanic tribes. They crossed the Rhine in 406 AD along with the Vandals and Suebi, settling in Orl?ans and Valence. Around 409 AD, they joined the Vandals and Suebi in the crossing of the Pyrenees into the Iberian Peninsula, settling in Lusitania and Carthaginensis.

As the Roman Empire continued to decline, the Alans split into various groups; some fought for the Romans while other joined the Huns, Visigoths or Ostrogoths.

https://en.wikipedia.org/wiki/Alans

In the long term, some Alans should have made it into the West Gothis/Spanish elite. Funnily they might have brought additional lines of G back to Iberia :LOL:
 
That's right, and there might have been even more groups of Goths around, even though we primarily refer to Western and Eastern Goths, which fought with each other on the Catalanaunian plains.



The Ostrogoths mixed with all kinds of people. The Visigoths might have mixed with Sarmatians as well, because the Germanic tribes had often allied Sarmatian riders with them. At the battle of Adrianople, both the Goths and the Romans had Sarmatians in their forces, especially among their cavalry. This was not just true for the Ostrogoths, but for many Germani tribes, like the Suebi (which I actually referred to) and the Vandals:




https://en.wikipedia.org/wiki/Alans

In the long term, some Alans should have made it into the West Gothis/Spanish elite. Funnily they might have brought additional lines of G back to Iberia :LOL:

doubt the Visgoths mixed at that time with sarmatians .............visigoth means pure goth ...............ostrogoths mixed with everyone ( bastard goths )

a good read is...........The Goths in ancient Poland by Jan Czarnecki
or try
The well spring of the Goths by I. Nordgren
 
That's right, and there might have been even more groups of Goths around, even though we primarily refer to Western and Eastern Goths, which fought with each other on the Catalanaunian plains.



The Ostrogoths mixed with all kinds of people. The Visigoths might have mixed with Sarmatians as well, because the Germanic tribes had often allied Sarmatian riders with them. At the battle of Adrianople, both the Goths and the Romans had Sarmatians in their forces, especially among their cavalry. This was not just true for the Ostrogoths, but for many Germani tribes, like the Suebi (which I actually referred to) and the Vandals:




https://en.wikipedia.org/wiki/Alans

In the long term, some Alans should have made it into the West Gothis/Spanish elite. Funnily they might have brought additional lines of G back to Iberia :LOL:


the Vandals are not one tribe, they are a confederation on many tribes


sometimes known as the Vindili confederation
 
the Vandals are not one tribe, they are a confederation on many tribes
sometimes known as the Vindili confederation

Well, most Germanic "tribes" of the early Medieval times were confederations of older, different ethnic/tribal units. One of the oldest were the Suebi and one of the latest the Baiuvarii, the ancestral group to modern Bavarians and (most of the) Austrians. The Allemanni refers to something similar, as it means just "all men" (= all men fighting with us = a confederation/alliance for a campaign originally most likely).

doubt the Visgoths mixed at that time with sarmatians .............visigoth means pure goth ...............ostrogoths mixed with everyone ( bastard goths )

The Ostrogoths were more mixed, even with Asians, yes, but that doesn't mean the Visigoths didn't mix at all, besides I didn't specify a time, but said it happened "on the long run" (in Iberia). The first potentially (Visi-) Gothic samples from Iberia don't look unmixed Germanic...
 
In a relatively early 2012 paper by Fu, Q, they looked at the whole set of Neolithic versus hunter-gatherer dna, so not quite what we want, but...
https://europepmc.org/article/med/22427842

So, we have an average of 12% mtDna U.

xEALz9z.png


Some sites, though, have no mtDna U at all, including the Cucuteni Tripolye site of Verteba in the Ukraine, which is right before the arrival of the steppe people.
https://www.biorxiv.org/content/10.1101/217109v1.full.pdf

This paper does have a chart with the results of some Neolithic sites. It's on page 33. However, believe it or not, they don't break out mtDna U.

In 2013 we get another pie chart of averages, but at least it shows the differences before and after 4500 BC. In the "pioneer" days it seems the farmer communities took on some HG women, but no farmer women went to HG communities. After that, the percentage of HG mtDna went up in the farmer communities, but the increase of farmer mtDna in hunter communities increased more, so at the end this chart would make it appear that the exchange went both ways, and was at fairly equal levels, I think, but averaging maybe 25%?
https://www.karger.com/Article/PDF/356933

DZxBz4m.png



I think the mtDna I might be steppe, Maciamo thinks so, and it's unclear to me whether some of the other HG lineages might also have come from the steppe. I would think perhaps a few of them.

The early Iberian Neolithic had more HG than the LBK and descendant groups, but I don't remember them having a lot more HG mtDna.

Anyway, the Tassi et al al paper's Supplement has a list of mtDna sequences of HG and Neolithic sites.(the GAC paper). It's Table S6.
https://figshare.com/collections/Su...he_spread_of_Indo-European_languages_/3928333

It doesn't look to me like a lot of mtDna U anywhere, but on the other hand the total number of samples is small, so it may add up to about 12% in the early period. I don't understand how the prior paper could get 25% from the samples listed in this paper. Do you see any LN actual farmer culture obviously missing from the list?

Unless someone knows of a compendium which has been kept up to date of all mtDna and was done by someone who is precise and careful, that's all I have.
Awesome data, Angela. Thank you again.
These are a lot of info to "digest".

As for Cucuteni-Trypillia, I think they found U, yes. U8, supposedly of HG origin. I know there're more than one paper with Trypillian mtDNA.
 
the first continnetal Bell Beakers to arrive in Iberia had women with them
some early continental Bell Beaker females in Iberia had 80 % continental Bell Beaker ancestry

as for Cardial ware, the HG haplogroups that joined them were the haplogroups observed in the Danube Gorge : R1b-V88 and I2a-Z161
R1b-V88 has also been found amongst LBK

none of the typical meagalithic farmer haplogroups I2a1b, I2a1a1-M26 nor I2a-Y6098 were found along Cardial Ware
they were found in EN Els Trocs and in MN Iberia, and also in 6,6 ka megalithic Provence, SE France, in 6,1 ka EN Britain, in the megalithic TRB and in Nuragic Sardegna

it looks like these haplogroups mixed with Cardial Ware women (they had 70-80 % EEF), but Y-DNA remained seperate, at least among megalithic elite (there was some G2a2 in megalithic Provence though)
how and why this exchange of women happened, I have no clue
those megalithic farmers were quite dominant, early 6,3 ka TRB was not megalithic, but after 5,9 ka certain areas became megalithic, it was quite intrusive,
but pottery style remained allbeit more decorated, it looks like the original farmers still existed, but they were dominated by a new elite
and we also know from DNA that the megalithic elite was based on male kinship

earliest megalithic structure in Europe is probably the Almendres Cromlech in the Portuguese Alentejo, 8 ka
the nearby estuaries were populated by fishers/hunters since 8,5 ka, they introduced geometric microlith tools, a technology which spread in Europa along with the expansion of the Villabruna clade
their autosomal was 80 % Magdalenian (El Miron clade) and 20 % Villabruna, so they were probably I2a males admixed with El Miron women (this El Miron admixture is what makes Iberian famer DNA different from European farmer DNA)

7,5 ka Cardial Ware arrived in the area, and by 7 ka most of the fishers/hunters were gone
 
Has there been any published or ongoing effort to understand where the WHG resurgence and boom of I2 to the detriment of Anatolia_N lineages like G2, T and H2 originated and spread from, and how profound its overall genetic (autosomal) and sociocultural impact must've been between the Middle Neolithic and the Early Neolithic? Was the change toward more agro-pastoral economies in several parts of Europe even well before the westward expansion of steppe people related to that genetic shift? Sometimes I feel that the fact that what happened involved basically the same autosomal admixtures, but in different proportions (less ANF and more WHG), is a complicating factor and causes most people (including scholars) to completely ignore what happened, but it must've been nearly as important as the arrival of steppe admixture in parts of Europe.

The Cassidy thesis (available as of yesterday) suggested the resurgence was due to patrilocality, basically confirming what Riverman is suggesting above. Atlantic Europe, from Iberia to Britain/Ireland seems to have this effect, as well as nordic Europe. However, the LBK did not. (based on data so far) I'm curious why that would be. I also suspect that the bulk of the neolithic lineages arrived to Britain/Ireland from Iberia->France, which explains the dominance. However, I suspect the steppe lineages R1b in particular, brought LBK related lineages afterwards (G-P303 on the male side) during the Bronze or even later periods.

An interesting thing is tha the Atlantic-Neolithic lineages certainly took a major hit to near extinction. Many of the living I2 lines are from areas which were not under the Atlantic-Neolithic umbrella, such as among the PC steppe (I2-L701, L801), or even I1 who seems to have expanded during the Germanic migration period. What could have cause the I2a (Neolithic branches) near extinction?
 

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