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

I'm not sure I'd call 15-20% WHG showing up eventually in LN farmers in Europe is nearly as important as percentages ranging from 55% down to 25-30% for steppe admixture. Plus, that 15-20% admixture would have come from HG women as well.

If I recall correctly, in Gambas et al there was one I1a who was hunter-gatherer autosomally, but he was lower status, perhaps a servant, and there was no admixture in any of the other inhabitants from that time period.

I mean, since this process took place after more than one millennium of fast spread of ANF admixture throughout Europe, it is quite unlikely IMHO that the peoples that were involved in this "WHG resurgence" were still typical WHG or even mostly WHG. They might have been like some of the more ANF-rich individuals in Blatterhöhle_MN, i.e. ~40% WHG + ~60% ANF. In that case, if you consider that some parts of Europe had ~0-10% WHG ancestry in the Early Neolithic and that increased to ~10-25% in the LN in many parts of Europe, the demographic change may have been pretty significant between the EN and the LN in some parts of Europe. Assuming the "WHG resurgence" involved a people that was ~40% WHG, and the pre-MN/LN populaton was ~10% WHG, changing to ~20% WHG in the LN, that would've meant a ~35% population replacement, which is as high or even higher than the steppe input in many parts of Europe.

For instance, I can see that Blatterhöhle_MN (~40% WHG), used as a proxy for a WHG-rich EEF population, would be needed to model many MN and LN European samples (population averages) not in small proportions, and interestingly it seems the change happened mostly along the Atlantic/North Sea Façade (where megalithism also seems to have spread from).

Results of the genetic ancestry model: https://imgur.com/a/QiD0PnC

Also, from what I can tell, we're seeing I2a showing up relatively early in western Europe where there's no indication of "pastoralism" before the arrival of steppe people to my knowledge; domesticated animals, yes, and maybe some transhumance as all farmers practice, but not pastoralism. Yes, there's I2a in the border areas of central Europe, among people who might have traded animals to the steppe and maybe even did some initial pastoralism, but I doubt it was a Europe wide phenomenon, or at least I've seen no papers indicating that it is.

Yes, that was just a speculation of mine, considering that the final phase of cultures like Cucuteni-Tripolye and Southeastern European/Balkanic Neolithic/Chalcolithic cultures seems to have been particularly more pastoral than the earlier stages of the same cultures or former cultures in the same place. There was also the GAC, whose way of life was pastoral, patriarchal and semi-nomadic enough for them to be mistaken for Indo-Europeans for a long time (and they also had been a clear example of the "WHG resurgence"). And that was even before steppe admixture became significant enough in any of them. That may have been caused by climate change, not necessarily a foreign influence and a change in the demographic makeup. But in any case the WHG resurgence seems to have happened contemporaneously with the spread of megalithism and perhaps, I think, a more maritime culture in parts of the continent.
 
@Ygorcs,

I don't know if you've been looking at the French Brunel et al paper. It seems, going by their graphic, WHG admixture after the Neolithic was sort of hit or miss. I'm still trying to figure out if the samples in the chart are arranged in chronological order. Their Tables leave out a lot of information even if you hunt through all of them. You'd think it would be easy to have a master excel sheet with sample number, culture, location, y and mtDna and calibrated date, but I guess not. :)

Michelsberg is interesting, with an influx of U5b. The north and east are pretty varied, but that isn't the case for Southern France.
 
I think the results of this paper (https://advances.sciencemag.org/content/6/22/eaaz5344) are of importance for the whole debate:
However, the situation is different in today?s France, where we observe not only the highest HG ancestry proportion overall when compared to other regions in Europe but it is also found in the oldest individuals from the southern sites of PEN and LBR. This observation is also supported by uniparental markers. Y chromosome lineages in western early farmers in the southern region are exclusively derived from HG (I2a; table S5 and text S5). In contrast, mitochondrial DNA results show a more universal Neolithic diversity profile, as previously reported [i.e., (4, 5)], with only two haplotypes (U5 and U8) that are potentially of HG origin (table S4 and text S4).

[...]

Two individuals from the Middle Neolithic OBN site located immediately west of the Rhine (Fig. 1A) also show a high proportion of the HG ancestry component (OBN B), in contrast to LBK sites east of the Rhine. To quantify the extra-HG proportion, we modeled LBK from Germany as a mixture of Anatolia_Neolithic and European_HG, and subsequently OBN subgroups as a more proximal mixture model of LBK and European_HG. We find this model well supported (table S11), and obtained up to 31.8% of excess HG ancestry for OBN B. Given the absence of a strong HG contribution in LBK groups east of the Rhine, we assume this to be a local contribution during the centuries following the arrival of first farmers. Moreover, male individuals from OBN carry exclusively the Y chromosome haplogroups I2a1a2 and C1a2b, attributed to HG groups (text S5), providing further evidence for a greater amount of the HG contribution in this region.

Archaeological research has argued for increased interaction between incoming farmers and indigenous HG in the western Mediterranean during a second stage of the Neolithization process and especially in areas with higher HG population densities, e.g., the Tosco-Emilian Apennine and Po plain (18). We are now able to confirm that these contacts left a traceable biological signal during the Neolithic expansion in southern France. From an archaeological perspective, this suggests that HG have contributed to the clear changes observed within the material culture postdating the pioneer phase.

Note that ICC individuals from the eastern Adriatic coast have only a very small amount of HG ancestry with a greater affinity to central European groups (see table S8). This fits with the hypothesis of a differentiation of technical traditions within material cultures observed from both sides of Apennine Mountains in Italy: an Adriatic tradition connected to the Balkans and a Tyrrhenian one whose origin is still unknown (41). It is tempting to associate such a strong HG component on the Tyrrhenian side with the characteristic/specific pottery traditions observed in this same region and to consider these original traditions the result of a HG reinterpretation (41). However, the scarcity of genomic data available from central and southern Italy currently does not allow this hypothesis to be tested directly.
Moreover, ICC individuals from the Iberian Peninsula also carry less HG ancestry. Together, this rejects the hypothesis that ICC-associated individuals represent a uniform genetic horizon per se and argues for more regionally nuanced scenarios of interaction.

In accordance with the established chronology of first Neolithic settlements in the French territory, the overlapping/synchronous date estimates obtained for southern ICC sites are consistent with the signal of a first HG contribution in the south of France, followed by a subsequent northward expansion of groups carrying this HG legacy

So here we have such a case, a forager clan taking Neolithic wives and adopting the Neolithic culture. From this or a similar group the shift from G2/H2 to I2a was initiated on a grand scale.

Figure S13 is interesting, showing which alliances were formed and that G2 and H2 marched in the West together:
https://advances.sciencemag.org/content/suppl/2020/05/21/6.22.eaaz5344.DC1/aaz5344_SM.pdf

Also:
We caution that because the amount of HG ancestry is very low in many Neolithic groups (<10%), it remains difficult to characterize the ultimate source reliably. Nevertheless, our admixture patterns from supported models show clear geographic signals. Neolithic groups associated with the LBK in central Europe (Hungary, Austria, and Germany) carry a small HG proportion, which was likely derived from admixture with HG individuals of the EHG-WHG cline and could have occurred in southeastern Europe during a preceding phase of the Neolithic expansion around 6000?5400 BCE. When using f-statistics of the form f4(Mbuti, test; BDB001, KO1) with our new Mesolithic genome (BDB001) from the Middle-Elbe-Saale region in central Germany as a geographically local HG proxy (instead of using Loschbour, which is located west of the Rhine), we do not find support for a local attraction for LBK groups, but the same pattern as for Loschbour (fig. S5C). This suggests that additional gene flow from neighboring Loschbour-like HG such as BDB001 in central Europe was negligible in the first Neolithic groups. However, the German Baalberge group (4000?3500 BCE) shows a marked increase of such HG ancestry, as well as individuals from the Bl?tterh?hle group, as has been suggested (5, 6), compared to a combination of both KO1-like and Loschbour-like ancestries for LBK groups (6). We can now show that this increase in WHG ancestry (up to 21.3 ? 1.5% in Baalberge; table S13) is driven by either local Loschbour-like ancestry or an expansion of farming groups from the west carrying this signal during the fifth millennium BCE, as suggested by archaeological data (36). For all studied Neolithic groups west of the Rhine, we observe a different pattern with a higher HG ancestry proportion, even for earliest groups that appears to be of a local (Loschbour-like) HG origin, consistent with archaeological data (2).

So the small amount of HG ancestry before the expansion of HG dominated groups from the West and possibly North is different and came from the early contacts of farmers with SEE foragers.

Particularly interesting is the continuity of a I2a Neolithic clans in Southern France through the ages. They established themselves and stood their ground. By 4000 BC the G2/H clans were reduced to a few places in Europe. We can now be pretty sure these were completely different ethnicities.

Also worthwhile to note is that they detect Pitted Ware ancestry further South, coming from a potential expansion from Northern Europe. But right now France seems to be the primary place where HG lineages took over and expanded from, replacing first farmer lineages, going after the results of this study which contributes a lot to the discussed topic.
 
My speculation is likely wrong. G-PF3346 is present in this study. It's not in line with the idea of a later arrival from the East. It opens the possibility they (or at least most part of them) were already in the West.
The locations of these samples (Gurgy "les Noisats" and Halberstadt) are close to one of the areas where G-L497 STR diversity peaks (in West Germany), for example. It looks possible this clade - or the subclade G-Z1816 especially - expanded from around there after Neolithic.
If I find a time I'll check BAM files to see if some of those G-L140 in the study could be categorized more deeply.
 
Riverman,

"Archaeological research has argued for increased interaction between incoming farmers and indigenous HG in the western Mediterranean during a second stage of the Neolithization process and especially in areas with higher HG population densities, e.g., the Tosco-Emilian Apennine and Po plain (18). We are now able to confirm that these contacts left a traceable biological signal during the Neolithic expansion in southern France. From an archaeological perspective, this suggests that HG have contributed to the clear changes observed within the material culture postdating the pioneer phase.
Note that ICC individuals from the eastern Adriatic coast have only a very small amount of HG ancestry with a greater affinity to central European groups (see table S8). This fits with the hypothesis of a differentiation of technical traditions within material cultures observed from both sides of Apennine Mountains in Italy: an Adriatic tradition connected to the Balkans and a Tyrrhenian one whose origin is still unknown (41). It is tempting to associate such a strong HG component on the Tyrrhenian side with the characteristic/specific pottery traditions observed in this same region and to consider these original traditions the result of a HG reinterpretation (41). However, the scarcity of genomic data available from central and southern Italy currently does not allow this hypothesis to be tested directly.
Moreover, ICC individuals from the Iberian Peninsula also carry less HG ancestry. Together, this rejects the hypothesis that ICC-associated individuals represent a uniform genetic horizon per se and argues for more regionally nuanced scenarios of interaction.




You're certainly consistent: always categorizing groups into ever smaller, separate, "ethnic" groups. I wouldn't personally consider someone who is 15-20% WHG and the remainder Anatolia Neolithic, a "different ethnic group" from someone who is 30% WHG and the remainder Anatolia Neolithic, which by the end of the process, i.e. by the Middle Neolithic, is what you're looking at all across Europe.

As for the excerpts above, half of my ancestry is from the Tosco-Emilian Apennines, so I try to keep abreast of the samples, and I'm not aware that any ancient samples from that area have ever been analyzed. Unless they've come across some, which would be great, there's no way they could know the proportion of WHG. It would make sense that they would adopt more foraging up there, it's true; it's terrible land for farming. Snows six months a year, and the soil is poor, as is the case in most mountainous areas, so about the only thing you can do is raise some cows. Nobody got rich off the land up there, ever. I will admit that I told Moesan repeatedly that it wasn't true that there was no "Mesolithic" survival in terms of phenotype in Italy and posted some pretty hair raising pictures of local men, and women, to prove it. :) Maybe there are some I2a men up there, although I've never seen it. Mostly R1b U-152.
 
My speculation is likely wrong. G-PF3346 is present in this study. It's not in line with the idea of a later arrival from the East. It opens the possibility they (or at least most part of them) were already in the West.
The locations of these samples (Gurgy "les Noisats" and Halberstadt) are close to one of the areas where G-L497 STR diversity peaks (in West Germany), for example. It looks possible this clade - or the subclade G-Z1816 especially - expanded from around there after Neolithic.
If I find a time I'll check BAM files to see if some of those G-L140 in the study could be categorized more deeply.

Just so I'm clear.

So, you're saying a clade which was absorbed in Central Europe by steppe admixed people and then moved forward with them?
 
My speculation is likely wrong. G-PF3346 is present in this study. It's not in line with the idea of a later arrival from the East. It opens the possibility they (or at least most part of them) were already in the West.
The locations of these samples (Gurgy "les Noisats" and Halberstadt) are close to one of the areas where G-L497 STR diversity peaks (in West Germany), for example. It looks possible this clade - or the subclade G-Z1816 especially - expanded from around there after Neolithic.
If I find a time I'll check BAM files to see if some of those G-L140 in the study could be categorized more deeply.
GRG003: negative for L497 equiv.
GRG008: I have no BAM
GRG016: I have no BAM
GRG021: negative for L497 equiv.
GRG023: negative for L497 equiv.
GRG027: negative for L497 equiv.
GRG035: negative for L497 equiv.
GRG043: negative for L497 equiv.
GRG047: negative for L497 equiv.
GRG052: negative for L497 equiv.
GRG057: I have no BAM
HBS002: negative for L497 equiv.
HBS004: negative for L497 equiv.
HBS005: I have no BAM
HBS006: negative for L497 equiv. Has a positive result below CTS342
HBS009: I have no BAM

It doens't seem to explain G-L497 then. Apparently the only relevant G-L497 ancient DNA keeps being that one from Baden Culture in Hungary, cal date from 5340 to 4945 years before present, so way more recent than these samples above.
 
Just so I'm clear.

So, you're saying a clade which was absorbed in Central Europe by steppe admixed people and then moved forward with them?
My new speculation was that G-L497, for example, developed locally in Neolithic and then expanded from there later, for whatever reason, but it seems most of these Neolithic samples are negative for L497 equivalent(s) after all, at the same time there is this ancient G-L497 individual way more recent, from Baden Culture, so I don't know anymore... :)

G-L497 is one of the best to check imo in BAM files because L497 SNP has lots of equivalents. It helps.
 
My new speculation was that G-L497, for example, developed locally in Neolithic and then expanded from there later, for whatever reason, but it seems most of these Neolithic samples negative for L497 equivalent(s) after all, at the same time there is this ancient G-L497 individual way more recent, from Baden Culture, so I don't know anymore... :)

G-L497 is one of the best to check imo in BAM files because L497 SNP has lots of equivalents. It helps.

Thanks for the clear answer.
 
Anyway, the study proves that at least part of G-L140 per se reached the West in Neolithic already. Given the absence of G-U1 thus far in Neolithic Europe, those were possibly G-CTS342.

Someone in another forum provided a list of deeper categorizations, but it doesn't seem reliable at all. Just an example: GRG003 was categorized in there as G-Y10621, however, that's not possible. Firstly, the structure was ignored. I mean, the individual is negative for upstream SNPs such L42 itself and L497 equiv., which evidenced the false positive (single read) for Y10621 (false positives are not unusual in ancient DNA). Additionally, GRG003 is something about 6500 years old, whereas the formation of G-Y10621 likely happened between 4300-3100 years ago. People should be more careful. :)
 
You're certainly consistent: always categorizing groups into ever smaller, separate, "ethnic" groups. I wouldn't personally consider someone who is 15-20% WHG and the remainder Anatolia Neolithic, a "different ethnic group" from someone who is 30% WHG and the remainder Anatolia Neolithic, which by the end of the process, i.e. by the Middle Neolithic, is what you're looking at all across Europe.

The highest WHG, at the start of the development, might have been even higher, but that's not decisive. What is decisive is that a significant portion of WHG was there and paternal lineages, the male clans were exclusively derived of foragers. These people were living apart, later mixture seems to have been very limited to non-existent, so they are in my book not just a subpopulation, after the Neolithic transition, but a people, a population very much apart. The material culture, even though within the wider sphere of ICC, was also different. But its still remarkable that they largely adopted the Cardial customs, they just altered and transformed them, to make them fit for their own needs. So even limited differences in the material culture could mean a lot, really a lot, in terms of ancestry and ethnicity. That's what many people claimed in the past, and its being vindicated. At the same time the cultural diffusion is not completely off too, as this locals accepted new ways on a bigger scale than aknowledged before.
The true ICC people however moved around them, over sea and land, reaching Iberia in the West. So we deal with a situation in which, most likely, the foragers were strong enough to make a stand. The farmers tried to come to terms with them, one way or another, and the locals had enough time to adapt and, again one way or another, taking in farmer women.

As for the excerpts above, half of my ancestry is from the Tosco-Emilian Apennines, so I try to keep abreast of the samples, and I'm not aware that any ancient samples from that area have ever been analyzed. Unless they've come across some, which would be great, there's no way they could know the proportion of WHG. It would make sense that they would adopt more foraging up there, it's true; it's terrible land for farming. Snows six months a year, and the soil is poor, as is the case in most mountainous areas, so about the only thing you can do is raise some cows. Nobody got rich off the land up there, ever. I will admit that I told Moesan repeatedly that it wasn't true that there was no "Mesolithic" survival in terms of phenotype in Italy and posted some pretty hair raising pictures of local men, and women, to prove it. :) Maybe there are some I2a men up there, although I've never seen it. Mostly R1b U-152.

Like the I2a foragers, the incoming steppe people took in local women. That's how this ancestry survived, for the most part.
 
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.
There could also be other theories why HG presence increase with time. One of the facts we know is that HG and Farmer or Pastoralist did not mix at all for a very long time. There are studies which show that HG and Farmers lived for thousands of years without mixing.

"It is commonly assumed that the European hunter-gatherers disappeared soon after the arrival of farmers", said Dr Ruth Bollongino, lead author of the study. "But our study shows that the descendants of the first European humans maintained their hunter-gatherer way of life, and lived in parallel with the immigrant farmers, for at least 2,000 years. The hunter-gathering way of life only died out in Central Europe around 5,000 years ago, much later than previously thought"
https://www.ucl.ac.uk/news/2013/oct/european-hunter-gatherers-and-immigrant-farmers-lived-side-side

The reason why HG reemerged could be that due to some climate conditions HG had advantage over farmers or whatever other reasons that we are not aware of now, not necessarily taking over by force.

I for instance imagine that HG had very different life style - they could not hold property and lived in small families, women did not have many children compared to farmers and they had to be very well adapted to nature, silent, unnoticeable, whereas farmers were smelly, laud, moved in large numbers, burned forest, kept animals.

HG and Pastoralist (who for sure must have been warlike) did not had "a war" when pastoralist moved in the Eastern Baltics. In the beginning they too lived in different places (HG near rivers and lakes) pastoralist where the fertile lands were.

CWC groups who lived at the coastline ate pigs, cattle (either goat or sheep), but also seals, and wild roe deer. Besides, they were fishing too (bones of lagoon fish were found: pike, pikeperch, bream) So the CWC economy could be different, and in some locations domestic animals were raised alongside seal hunting and fishing. No remains of cultivated plants were found only some burnt hazelnut shells and fruit of an unidentified species of sorrel (the settlement dates to 2600 BC)
It is just an example that under some conditions HG way of live could be better adapted to local conditions, and other groups like CWC started fishing/hunting practices too that they could not do on the steppe.
 
There could also be other theories why HG presence increase with time. One of the facts we know is that HG and Farmer or Pastoralist did not mix at all for a very long time. There are studies which show that HG and Farmers lived for thousands of years without mixing.

"It is commonly assumed that the European hunter-gatherers disappeared soon after the arrival of farmers", said Dr Ruth Bollongino, lead author of the study. "But our study shows that the descendants of the first European humans maintained their hunter-gatherer way of life, and lived in parallel with the immigrant farmers, for at least 2,000 years. The hunter-gathering way of life only died out in Central Europe around 5,000 years ago, much later than previously thought"
https://www.ucl.ac.uk/news/2013/oct/european-hunter-gatherers-and-immigrant-farmers-lived-side-side

The reason why HG reemerged could be that due to some climate conditions HG had advantage over farmers or whatever other reasons that we are not aware of now, not necessarily taking over by force.

I for instance imagine that HG had very different life style - they could not hold property and lived in small families, women did not have many children compared to farmers and they had to be very well adapted to nature, silent, unnoticeable, whereas farmers were smelly, laud, moved in large numbers, burned forest, kept animals.

HG and Pastoralist (who for sure must have been warlike) did not had "a war" when pastoralist moved in the Eastern Baltics. In the beginning they too lived in different places (HG near rivers and lakes) pastoralist where the fertile lands were.

CWC groups who lived at the coastline ate pigs, cattle (either goat or sheep), but also seals, and wild roe deer. Besides, they were fishing too (bones of lagoon fish were found: pike, pikeperch, bream) So the CWC economy could be different, and in some locations domestic animals were raised alongside seal hunting and fishing. No remains of cultivated plants were found only some burnt hazelnut shells and fruit of an unidentified species of sorrel (the settlement dates to 2600 BC)

@Dagne: That was a position I didn't share, but which was possible to argue for before the new data came in. Now we see that in Southern France the Neolithic clans were not able or willing to replace the local HG clans, which in turn adopted Neolithic techniques and customs, took farmer wives and formed their own Neolithic community and culture. And it seems to be quite clear that the increased HG ancestry and I2a lineages spread from there, to other regions in which first farmers were established already and replaced them.
So while individual cases of mixture and increased HG ancestry certainly took place, this big shift we are talking about happened because the more HG dominated groups expanded and partly or fully replaced the first farmer lineages in many regions of Europe.

This mixture and takeover by local forager paternal lineages happened early on, in the early phase of the Neolithic expansion. Not later. That's what the study is proving.
 
Somehow, it is difficult for me to imagine HG adapting to farmer ways of life, or, moreover, taking over farmer livelihood by force.
HGs need forest, rather that farmer "property". Perhaps these were descendants of HG, who had farmers as mothers who merged with farmers, in a slow and peaceful (?) way, generation after generation.
I could see well however, how HG could have had advantage over farmers during the years of famine. In Baltics it were more CWC who would adapt to HG ways of life, as in some conditions it is very easy to fish and but very difficult to cultivate plants (sandy dunes) or even to raise cattle (no pastures).
The HG share increased with time, however, we still don't say that HG took over CWC in the Baltics. No, it was still the same CWC, only it developed by absorbing different peoples and traditions (more HG and a newly introduced farming culture (via CWC Western contacts).
 
I guess what Dagne is proposing, and she can correct me if I'm wrong, is a possible scenario in which they firstly mixed normally, but at certain point something happened* (climate change, for example); something that affected more farmers than those who kept some of the HG lifestyle. If so, it would not have been a replacement according to her, but just one group almost dying out, whereas the other (already mixed) "continued". It naturally implies the two groups were a bit different, but an explanation for the bias towards farmer autosomal would still lack.

*
41467_2018_4375_Fig1_HTML.jpg
 
@Dagne: That was a position I didn't share, but which was possible to argue for before the new data came in. Now we see that in Southern France the Neolithic clans were not able or willing to replace the local HG clans, which in turn adopted Neolithic techniques and customs, took farmer wives and formed their own Neolithic community and culture. And it seems to be quite clear that the increased HG ancestry and I2a lineages spread from there, to other regions in which first farmers were established already and replaced them.
So while individual cases of mixture and increased HG ancestry certainly took place, this big shift we are talking about happened because the more HG dominated groups expanded and partly or fully replaced the first farmer lineages in many regions of Europe.

This mixture and takeover by local forager paternal lineages happened early on, in the early phase of the Neolithic expansion. Not later. That's what the study is proving.

Are we speaking about the same paper?
original-1590693135.png

original-1590693135.png


In Hungary the proportions of "HG" actually seem to have gone down a bit with time in one area over time.

In 5000 BC there's very little in Central Europe or the Pyrenees. Some, maybe 15% in southern Spain. France becomes involved from 5000 BC to 4500 BC, but the levels are about what they were in Southern Spain, or maybe it's now 20%. By 4000-3500 BC, rather than an advance where more and more WHG is picked up, I see a British Isles proportion extremely close to those of Spain and Southern France and the northern France and southwestern Germany of the prior 500 years. The only area which sees a marked increase is what looks like the border of the Benelux countries and northwestern Germany. Any movements to the area from the north or east? that would have to be investigated. Even in east Germany it looks like a mixed bag; one group has less than a quarter HG. As I said, we see it in the yDna too.

It makes perfect sense for me. You get away from the Loess areas and get some wetter, colder weather, and someone who still does a lot of fishing and hunting might start to look more attractive.

It looks as if the only farmer groups which really didn't want to mix much were the groups from just north of the Balkans, but from what I remember, there weren't many hunter-gatherers there to start with, and Mathiesen said a while ago that the groups there didn't seem to spread northward much, and not westward. It makes sense, the admixture is going to take place on the frontier, not back in the "homeland".

That's what I see from the yDna and mtDna of the various cultures too. Proportions could vary from settlement to settlement; lots of founder effect, drift, and patrilocality from what I can see. The Neolithic didn't need the steppe people to bring that in.


According to this, the far northeast actually saw the first admixture. Anyone have data as to whether it's farmer mtDna? If so, where did they get the women? That part of Europe is not my area of expertise.

I think Dagne is on the right track, although the ultimate proportions were still heavily skewed toward Anatolian Neolithic, with only certain areas reaching as high as 30% HG.

Oh, and there are no samples from either the Tyrennhian or Adriatic side of Italy, so all of that is complete speculation on the part of the writers.


The British changed a lot; southern Europeans less...anyway, this is the trail. (Cassidy et al)
Bally.PNG
 
Somehow, it is difficult for me to imagine HG adapting to farmer ways of life, or, moreover, taking over farmer livelihood by force.
HGs need forest, rather that farmer "property". Perhaps these were descendants of HG, who had farmers as mothers who merged with farmers, in a slow and peaceful (?) way, generation after generation.
I could see well however, how HG could have had advantage over farmers during the years of famine. In Baltics it were more CWC who would adapt to HG ways of life, as in some conditions it is very easy to fish and but very difficult to cultivate plants (sandy dunes) or even to raise cattle (no pastures).
The HG share increased with time, however, we still don't say that HG took over CWC in the Baltics. No, it was still the same CWC, only it developed by absorbing different peoples and traditions (more HG and a newly introduced farming culture (via CWC Western contacts).

They were HG lineages, clans in Southern France/Northern Italy, but they became farmers, Neolithics themselves, but sticked to their own clan. That's the point. They were not staying foragers, they became farmers, but it was a HG clan which did so in this area and kept other farmers, the original Neolithics, away from its area. The transition seems to have been coming from the female side more than the male. Early farming was, to a large degree, a female business. So what I'm proposing is that a HG clan took farmer wives, one way or another, and some techniques, by some sort of contact, and by doing so adopted a lot from the Cardial, but with their own adaptation, in their own style. They expanded over other, probably unmixed foragers and later replaced even a lot of their Neolithic competitors. The early Cardial people, from which they had their Neolithic culture and wives, tried to avoid them it seems, and moved around them on to Iberia.

@Angela:
France becomes involved from 5000 BC to 4500 BC, but the levels are about what they were in Southern Spain, or maybe it's now 20%. By 4000-3500 BC, rather than an advance where more and more WHG is picked up, I see a British Isles proportion extremely close to those of Spain and Southern France and the northern France and southwestern Germany of the prior 500 years. The only area which sees a marked increase is what looks like the border of the Benelux countries and northwestern Germany. Any movements to the area from the north or east? that would have to be investigated. Even in east Germany it looks like a mixed bag; one group has less than a quarter HG. As I said, we see it in the yDna too.

The original HG clan based Southern French Neolithics had a significantly higher proportion of WHG ancestry and what the paper is stating is that the WHG ancestry and I2a's expansion on a large scale in Western Europe can be attributed to these Southern French tribes moving out and expanding in different directions. Let's assume that the original clan had about 30-40 percent WHG ancestry and was 100 percent I2a, what you see in many places where they moved to is an increase of WHG ancestry and a strong dominance, if not total replacement, of other farmer lineages, like G2/H2 in particular, while we have later in Michelsberg an alliance with E1b.

Just assume most of the HG ancestry is indeed from the South, with the PWC being on a low level, and can be explained like you said, by individuals assimilated here and there, we deal with a replacement rate of 40-50 percent of the original ICC and LBK people, where they were affected by these expansion. That's actually pretty close to the BB expansion in Iberia, its almost the same pattern.

I repeat how the paper sums it up:
In accordance with the established chronology of first Neolithic settlements in the French territory, the overlapping/synchronous date estimates obtained for southern ICC sites are consistent with the signal of a first HG contribution in the south of France, followed by a subsequent northward expansion of groups carrying this HG legacy

They tested for where this additional HG ancestry which spread in the Middle Neolithic was coming from and they concluded it came from the deviating, HG-dominated groups in the South of France. The local HG input was minimal, only significant in the North with a minor PWC contribution. But that's not it, at least not in the West. There we deal with a large scale expansion and replacement, most likely from this Southern French I2a clan.
 
^^Sorry, all speculation and supposition unsupported by data for each point, and in fact contradicts the graph from the paper.

Not my style; not into fantasizing.
 
G-L497 is one of the best to check imo in BAM files because L497 SNP has lots of equivalents. It helps.
Obviously I'm talking here on full developed G-L497 (based on current modern people tested) and descendants.

I guess what Dagne is proposing, and she can correct me if I'm wrong, is a possible scenario in which they firstly mixed normally, but at certain point something happened* (climate change, for example); something that affected more farmers than those who kept some of the HG lifestyle. If so, it would not have been a replacement according to her, but just one group almost dying out, whereas the other (already mixed) "continued". It naturally implies the two groups were a bit different, but an explanation for the bias towards farmer autosomal would still lack.
I don't know... If something such climate change happened and affected farmers even before Steppe invasion, explaining this supposed replacement of G by I, it remains the question on why G2a would have survived in some other areas such Central Europe and "Old Europe". But they remained G, T etc. till the arrival of Steppe folks, no? So... Perhaps some "boundaries" were substantially affected, as Angela suggested (if I got it right). In LN Switzerland - and possibly in parts of Germany - G2a was replaced directly by R1b, who at the end also replaced the I2 guys in the more Western parts after Neolithic. The replacement by R1b is pretty clear, since we have the "before" and the "after", but I still cannot visualize it with I2 vs G2a (say, a place once richer in hg G2a that at the end became richer in hg I2). If they exist, I cannot remember. Perhaps Iberia? Anyway, if it's true, I2 rich folks could have "infiltrated" farmer communities heavily till certain point, but could not go further for whatever reason.
Either way, as Riverman suggested, lifestyle could have been a plus for certain groups in certain places and times.
The reality must have been way more complex than what we can imagine though.
 
They were HG lineages, clans in Southern France/Northern Italy, but they became farmers, Neolithics themselves, but sticked to their own clan. That's the point. They were not staying foragers, they became farmers, but it was a HG clan which did so in this area and kept other farmers, the original Neolithics, away from its area. The transition seems to have been coming from the female side more than the male. Early farming was, to a large degree, a female business. So what I'm proposing is that a HG clan took farmer wives, one way or another, and some techniques, by some sort of contact, and by doing so adopted a lot from the Cardial, but with their own adaptation, in their own style. They expanded over other, probably unmixed foragers and later replaced even a lot of their Neolithic competitors. The early Cardial people, from which they had their Neolithic culture and wives, tried to avoid them it seems, and moved around them on to Iberia.

Ok this is your hypothesis.
I could think of other ideas too how to explain HG expansion.
I know more about HG who lived in the Baltic so I take examples from these type of HG, but perhaps French HG might have been similar?


Excavations in the Šventoji settlement [5, 6] revealed three beautiful ritual bone staffs with she-elk head tops (Fig. 1) [5]. Such staffs may have been used for performing pre-hunting rites. In eastern Lithuania and in Latvia numerous deer figurines have been found. From analogy with other mythologies, we can suppose that people of Nemunas and Narva cultures considered the Goddess-elk or Goddess-deer to have specific power, such as life-, fertility- and birth-giving. Even the present Lithuanian Advent songs mention a she-deer with nine horns. Some European myths reveal two she-elks, women, birth-givers of the world [7].
Fig. 1. One of the ritual bone staffs with a she-elk head found in the Šventoji settlement (2400 B.C.).
It is also probable that Neolithic people worshiped the grass-snake which is often represented by bone and horn figurines and frequently in pottery decoration [8]. Primeval worship of gods and demons in the shape of animals expressed the idea of human identification with them after death, metempsychosis. An important place in cultic rites was given to fire [9, 10].In the Neolithic, anthropomorphic gods appear as evidenced by a two-meter high wooden sculpture found in the Šventoji settlement (Fig. 2.) [5], by amber figurines of the Juodkrantė (Schwarzort) settlement [11], by a bone figurine found near the Kretuonas lake [12] and the Nida pottery decorations [13]. A number of bone and clay figurines of antropomorphic beings has been found in the Neolithic settlements in eastern Latvia [14].There is no doubt that in the Early Neolithic the people of Nemunas and Narva cultures lived in a matriarchal community.
http://www.lithuanian.net/mitai/cosmos/baltai.htm


So overall, it does not seem that HG decided whom to take like wives, not to speak about going and killing farmers to take their wives. Eskimo like communities in the Siberian North were very peaceful up until historic times. In order to kill a seal they had the shaman to perform a ritual, asking for seal's spirit forgiveness and begging the seal god to send one of its children to sacrifice itself for people. Only after praying like this they would go to hunt, and use the hunted seal for the benefit of all community, spreading the hunted spoils equally to those who needed rather than developing the property of families like farmers did. So I would not compare HG lifestyle to that of farmers.

We don't know how the HG spread to farmer communities. I could create a story like like this - in a time of famine some farmer women, instead of perishing from hunger with their clan, would go to forest and cling to hunter communities, then after living in close proximity for let's say a long winter they would get pregnant, but with spring time they would want to leave back to farmer communities. (HG did not have slaves, so there was no one stopping them from going where ever they wanted).
The lifestyles or farmers and hunters must have differed drastically, and one must have detested/found unacceptable the other. After some time, back in farmer communities, farmer women who returned from forest gave birth to half farmer half hunter babies who were raised like farmers. And - voila - we have an increase in hunter gatherer genes among farmers.
 

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