More DNA from stone age European(Swedish) farmers and hunter gatherers

[Angela]

I did check the alleles for immune function for the oldest sample (not the 6000 B.C. ones who were living right alongside the farmers) to see if my thinking on the prior post was correct. I see two A's, one D, and two D*, or provisional derived states because of post mortem damage. The rest are all no calls, so I think it's difficult to say whether or not this old, pre-Neolithic contact European would have had the protective alleles. Perhaps it's still plausible that they were introduced through admixture? Some obviously had gone on even in terms of these isolated fisher/gatherers bands, since they already had a T2b mtDNA among them.

 

[LeBrok]

Mind you, the notion that North-Europeans have one third to two fifths of WHG genetic material brings a number of problems with it. This study specifically claims that farmers absorbed WHGs as they spread rather than the other way around. But that still leaves one of the problems that arises.

From the study:

The study was not able to show whether the gene flow into the immigrant communities of farmers was the result of indigenous women or men intermarrying with farmers – or people of both sexes.

I'm still suspecting it was mostly by women. We need more samples from farmer communities to see some U5s or 4s.

A while ago another study, I think it was also by Skoglund, claimed that the genetic investigation of Gotland Pitted Ware Culture showed that at the onset of the neolithic the population was completely replaced. [1] They showed tables of the rate of mtDNA in current day as compared to the baltics and the PWC finds to prove that point. However, when la Brana was published however, it clearly showed close relation to Northern Europeans rather than Iberians. How can both those assumptions be true?

I have been wondering about this a lot. Where has the mtDNA gone? Swedes have rather a high WHG rate, yet very low U5 (or U4) mtDNA. If Skoglund is right and farmers mobbed up WHGs we would expect rather a high number of U5/4.

I wouldn't read too much by Mt or Y DNA frequency of today. It's been so long that ordinary competition between them, and drifting will mud the picture completely. That's why it is so hard ot make sense of it.
When we look at H1 and H3 map, that are suppose to be the farmer haplogroups, try to figure out how farming spread in Europe?

Right.
By this distribution map I would guess that H wasn't original farmer hg but was picked up by them in western Europe and then becoming successful spreading with them NE.

 

[LeBrok]

I did check the alleles for immune function for the oldest sample (not the 6000 B.C. ones who were living right alongside the farmers) to see if my thinking on the prior post was correct. I see two A's, one D, and two D*, or provisional derived states because of post mortem damage. The rest are all no calls, so I think it's difficult to say whether or not this old, pre-Neolithic contact European would have had the protective alleles. Perhaps it's still plausible that they were introduced through admixture? Some obviously had gone on even in terms of these isolated fisher/gatherers bands, since they already had a T2b mtDNA among them.

Even though these European HGs are more isolated by their nature than farmers, I don't think we can expect such big immune disparity as per Europeans meeting Native Americans. After all they lived on same continent with farmers and there was always some contact and cultural exchange, to go through same infectious diseases gradually one by one.

 

[LeBrok]

Although I think we should be cautious about making broad generalizations, it's interesting that while you had a "farmer" mtDNA in the hunter-gatherers, you didn't yet have the hunter-gatherer mtDNA in the farmers. The same thing happened in the Balkans according to a prior study. In the earliest stages of the encounters, the mtDNA gene flow seemed to go only in one direction, although of course that changed later. Is it possible that it was the "farmer" women who brought agriculture to the natives? Of course, in the Balkans, they did also incorporate some I2a yDNA lineages. I wonder if that was because, given the climate in the Balkans, the agricultural package was even more obviously successful there.

I would be afraid to claim which mtDNA had farmer origin unless we go to deep deep subclades. At low resolution level they all have paleolithic origin, therefore HG, and were already scattered around Eurasia in secluded communities.
If it comes to I2a becoming farmers, (judging by eagerness of Prairie Indians (HGs) to farming), I would expect rather late Neolithic/Copper Age for this to happen. In a way of very slow trickling farmer's genetic components to I2a HG communities.

We might have a good view on farmer/HG mixing from Cucuteni and Yamna cultures. There was obvious contact between both so different cultures, and most likely this fusion of both giving start to Corded-ware farming culture of early Bronze age. Judging by the speed of new papers coming and dna testing we might learn this in next 3 years. Maybe 6 giving economic status of Ukraine and Romania.

 

[Angela]

Even though these European HGs are more isolated by their nature than farmers, I don't think we can expect such big immune disparity as per Europeans meeting Native Americans. After all they lived on same continent with farmers and there was always some contact and cultural exchange, to go through same infectious diseases gradually one by one.

I didn't expect it. Epoch did.

I do think, however, that the Neolithic lifestyle subjected humans to more pathogens, given what a filthy business keeping animals was in those days, and even now if the proper precautions aren't taken. One would expect, I think, that farmers who were descendents of people who had been practicing that lifestyle for thousands of years would have more immunity to those pathogens than hunter-gatherers being introduced to it for the first time. However, some admixture, even if it didn't occur at very high levels in the beginning, would have spread those immunity alleles to the hunter gatherers. That's why I was interested to see whether the oldest specimen, from a period before the first farmers reached the area, carried the same number of immunity alleles. The results are such that I don't think a conclusion can be drawn either way.

 

[epoch]

It's easier to retain your genetic cohesiveness when no one wants the land you're sitting on, or at least is at the limits of what is desired.Even in central Europe,there was a climate collapse, and perhaps an environmental crisis created deforestation. All of these factors come into play it seems to me, as well as the fact that there was a later movement of peoples into Europe who were not part of the original encounter at all.

Dienekes had a link to a study which suggested that even in the middle of LBK area hunter-gatherers persisted [1]. That reminds me of a PDF I read that tried to assess the first contact between farmers and hunter-gatherers. It tried to make some estimates about the importance of Loess grounds for mesolithic hunter-gatherers. With no mesolithic camp site ever found on Loess grounds the paper tries to make a distinction between flint tools by hunter-gatherers and farmers. While the method seems a bit uncertain it stated that Loess grounds were hardly used by mesolithic hunter-gatherers, who were far more attracted to river valleys. So, that might be the second stroke of luck for European hunter-gatherers.

The only place farmers and hunter-gatherers might become competitors might be the largest river valleys, where the deposits of river clay may both attract wild life and be used as fertile ground. It may be no coincidence that Stuttgart showed HG admixture. The Rhine and Meuse valleys are not only places where there is evidence for violence between the group [2] but also the area where some form of burial practise continuity has been suggested [3].

[1] http://dienekes.blogspot.nl/2013/10/ancient-central-european-mtdna-across.html
[2] https://www.academia.edu/215068/Beating_Ploughshares_back_into_Swords
[3] http://arheologija.ff.uni-lj.si/documenta/pdf34/DPlenneis34.pdf

 

[LeBrok]

I didn't expect it. Epoch did.

I do think, however, that the Neolithic lifestyle subjected humans to more pathogens, given what a filthy business keeping animals was in those days, and even now if the proper precautions aren't taken. One would expect, I think, that farmers who were descendents of people who had been practicing that lifestyle for thousands of years would have more immunity to those pathogens than hunter-gatherers being introduced to it for the first time. However, some admixture, even if it didn't occur at very high levels in the beginning, would have spread those immunity alleles to the hunter gatherers. That's why I was interested to see whether the oldest specimen, from a period before the first farmers reached the area, carried the same number of immunity alleles. The results are such that I don't think a conclusion can be drawn either way.

I was writing this in context that Farmers didn't cause extreme die off of HG in Europe, unlike in America.
However, if it comes to pathogens I would expect farmers to have more immunological mutations. They lived in more numerous and crowded communities by factor of 5-10, when compared to HGs, and by sheer numbers it will speed up mutations and combinations. Statistically speaking farmers should have had more "advanced", or complicated I should say, immune system than HGs. We can expect a direction of pathogen gene flow to be from farmers to HGs, and rarely reversed.

 

[LeBrok]

Dienekes had a link to a study which suggested that even in the middle of LBK area hunter-gatherers persisted [1]. That reminds me of a PDF I read that tried to assess the first contact between farmers and hunter-gatherers. It tried to make some estimates about the importance of Loess grounds for mesolithic hunter-gatherers. With no mesolithic camp site ever found on Loess grounds the paper tries to make a distinction between flint tools by hunter-gatherers and farmers. While the method seems a bit uncertain it stated that Loess grounds were hardly used by mesolithic hunter-gatherers, who were far more attracted to river valleys. So, that might be the second stroke of luck for European hunter-gatherers.

I wouldn't be surprised to see small HGs communities lingering in secluded places deep to the end of Bronze Age. I forgot the name of the site in Germany where of HG/herding village, of 1 millenium BC, with all males composed of I2a and R1a type.

 

[epoch]

I didn't expect it. Epoch did.

I do think, however, that the Neolithic lifestyle subjected humans to more pathogens, given what a filthy business keeping animals was in those days, and even now if the proper precautions aren't taken. One would expect, I think, that farmers who were descendents of people who had been practicing that lifestyle for thousands of years would have more immunity to those pathogens than hunter-gatherers being introduced to it for the first time. However, some admixture, even if it didn't occur at very high levels in the beginning, would have spread those immunity alleles to the hunter gatherers. That's why I was interested to see whether the oldest specimen, from a period before the first farmers reached the area, carried the same number of immunity alleles. The results are such that I don't think a conclusion can be drawn either way.

http://eurogenes.blogspot.nl/2014/01/mesolithic-genome-from-spain-reveals.html

La Brana 1 is 7000 years old. Perhaps diseases originating from the Middle- and Near-East or even European LBK area already had reached Iberia. However, the mesolithic HG from Sweden also seem to have at least some of these adaptive variants. I am not sure, as the information is thin. However, the confirmation of these genes is very interesting.

Moreover, we provide evidence that a significant number of derived, putatively adaptive variants associated with pathogen resistance in modern Europeans were already present in this hunter-gatherer.

 

[Angela]

I wouldn't be surprised to see small HGs communities lingering in secluded places deep to the end of Bronze Age. I forgot the name of the site in Germany where of HG/herding village, of 1 millenium BC, with all males composed of I2a and R1a type.

Yes, I think that's what the Bollongino et al paper showed, although they might also have been a group that moved down from the Baltic at a certain point.

 

[epoch]

I wouldn't be surprised to see small HGs communities lingering in secluded places deep to the end of Bronze Age. I forgot the name of the site in Germany where of HG/herding village, of 1 millenium BC, with all males composed of I2a and R1a type.

We still have fishing villages in Europe, don't we? The most famous one in the Netherlands is Urk, which used until recently be a small village sized island. The only thing is almost all males from Urk - which even if it is currently part of reclaimed land is still very much on its own - have Y-DNA haplogroup G2

 

[Fire Haired14]

What mesolithic F-96 is there? I know of F-96 LBK neolithic.

F* was found in LBK.

 

[Angela]

Dienekes had a link to a study which suggested that even in the middle of LBK area hunter-gatherers persisted [1]. That reminds me of a PDF I read that tried to assess the first contact between farmers and hunter-gatherers. It tried to make some estimates about the importance of Loess grounds for mesolithic hunter-gatherers. With no mesolithic camp site ever found on Loess grounds the paper tries to make a distinction between flint tools by hunter-gatherers and farmers. While the method seems a bit uncertain it stated that Loess grounds were hardly used by mesolithic hunter-gatherers, who were far more attracted to river valleys. So, that might be the second stroke of luck for European hunter-gatherers.

The only place farmers and hunter-gatherers might become competitors might be the largest river valleys, where the deposits of river clay may both attract wild life and be used as fertile ground. It may be no coincidence that Stuttgart showed HG admixture. The Rhine and Meuse valleys are not only places where there is evidence for violence between the group [2] but also the area where some form of burial practise continuity has been suggested [3].

[1] http://dienekes.blogspot.nl/2013/10/ancient-central-european-mtdna-across.html
[2] https://www.academia.edu/215068/Beating_Ploughshares_back_into_Swords
[3] http://arheologija.ff.uni-lj.si/documenta/pdf34/DPlenneis34.pdf

Sorry, I missed this initially.

As far as Bollongino et al is concerned, it's a study whose data doesn't logically lead to its conclusions, in my opinion, or at least doesn't support the claims that have been made based on the study. The period in question, when the two groups were simultaneously in the same area, amounted to only a few hundred years if I remember correctly. It's just as likely that this group moved down from the Baltic shores.

Regardless, neolithic peoples are always going to out breed hunter-gatherers, a point made in the Skoglund study itself. It was only with the Neolithic that some distance was put between humanity and extinction.

I'm still leaning toward the idea that in the southern areas where the original Neolithic package was a good "fit", the hunter-gatherers were indeed "wiped out" very early as a distinct group, not, from what we can see in the Balkans through any kind of genocide, (death through disease we don't know, but I doubt it) but from being outbred and incorporated.

In central Europe that probably happened as well, if the population figures I've seen in prior papers are accurate, with only scattered communities of hunter-gatherers remaining, as was alluded to by LeBrok. In the far northern regions, the " Neolithic package" just wasn't optimum for a long time, and so they were left relatively undisturbed for a long time. I think it has to be remembered that even now Scandinavia, Finland, the eastern Baltics etc. are very low in population compared to the densities we see in central and southern Europe.

Then, there seems to have been a climate collapse in central Europe, or an environmental collapse, which caused a population collapse as well. We don't know how many people of which groups survived.

At some point, you have a new group of people entering central Europe, in unknown numbers, containing an unknown amount of EEF, WHG and ANE, to refer to the discussion on another thread. Later still, you have the migrations after the fall of Rome affecting central Europe and northwestern Europe, and even the Balkans later on, which could have brought more WHG and ANE for that matter.

That's how I think we have gotten to the current genetic landscape in Europe.

All speculation I know, but I think there's solid evidence for some of it. We may get more soon.

Ed. I don't remember the specifics of those two papers. It's been eight years, and my memory isn't what it was.... I'm going to glance through them when I have a chance.

 

[Fire Haired14]

I wouldn't read too much by Mt or Y DNA frequency of today. It's been so long that ordinary competition between them, and drifting will mud the picture completely. That's why it is so hard ot make sense of it.
When we look at H1 and H3 map, that are suppose to be the farmer haplogroups, try to figure out how farming spread in Europe?

Right.
By this distribution map I would guess that H wasn't original farmer hg but was picked up by them in western Europe and then becoming successful spreading with them NE.

Lebrok, do you know that one of the Swedish TRB farmer samples had mtDNA H1c like you? If H1, H3, and whatever H subclades in western Europe were native why does every study keep finding only U5, U4, U2, U8, and U*. H1 and H3 are not very popular in the strange mtDNA gene pool of LBK farmers but they are more frequent than U5(which we know took up the majority of hunter mtDNA) and there haven't been any U4 and U2 samples found in Neolithic central Europe period. The fact is that near eastern farmer mtDNA is dominate in Europe even if European hunter gatherer ancestry is over 50% for some northern Europeans like Finns.

 

[epoch]

As far as Bollongino et al is concerned, it's a study whose data doesn't logically lead to its conclusions, in my opinion, or at least doesn't support the claims that have been made based on the study. The period in question, when the two groups were simultaneously in the same area, amounted to only a few hundred years if I remember correctly. It's just as likely that this group moved down from the Baltic shores.

If this study was only piece of evidence pointing to this I would be far more cautious. But there is the site of Ostorf, where hunter-gatherers lived in a Funnelbeaker environment, having funnelbeaker pottery - prehaps imitation - but living off hunting. Their mtDNA shows this:

Forager/FB	Germany	Ostorf [SK 28a]	M	3200 BC			K	16224C, 16311C	Bramanti 2009
Forager/FB	Germany	Ostorf [SK 8d]	M	3200 BC			U5	16270T, +12308 HinfI, +7028 AluI	Bramanti 2009
Forager/FB	Germany	Ostorf [SK 35]	F	3100 BC			U5	16270T, +12308 HinfI, +7028 AluI	Bramanti 2009
Forager/FB	Germany	Ostorf [SK 12a and 18]	M	3000 BC			T2e	2 samples 16093, 16126C, 16153A, 16294T	Bramanti 2009
Forager/FB	Germany	Ostorf [SK45a]	?	3000 BC			J	16069T, 16126C	Bramanti 2009
Forager/FB	Germany	Ostorf [SK19]	M	2950 BC			U5a	16168T, 16192T, 16256T, 16270T, 16302G, +12308, HinfI +7028 AluI	Bramanti 2009

There is more archeological evidence collected by German, Danish and Dutch archeology that suggest continuation. Seen in this light I think the Bollongino article may simply add to the idea.

 

[Angela]

If this study was only piece of evidence pointing to this I would be far more cautious. But there is the site of Ostorf, where hunter-gatherers lived in a Funnelbeaker environment, having funnelbeaker pottery - prehaps imitation - but living off hunting. Their mtDNA shows this:

Forager/FB	Germany	Ostorf [SK 28a]	M	3200 BC			K	16224C, 16311C	Bramanti 2009
Forager/FB	Germany	Ostorf [SK 8d]	M	3200 BC			U5	16270T, +12308 HinfI, +7028 AluI	Bramanti 2009
Forager/FB	Germany	Ostorf [SK 35]	F	3100 BC			U5	16270T, +12308 HinfI, +7028 AluI	Bramanti 2009
Forager/FB	Germany	Ostorf [SK 12a and 18]	M	3000 BC			T2e	2 samples 16093, 16126C, 16153A, 16294T	Bramanti 2009
Forager/FB	Germany	Ostorf [SK45a]	?	3000 BC			J	16069T, 16126C	Bramanti 2009
Forager/FB	Germany	Ostorf [SK19]	M	2950 BC			U5a	16168T, 16192T, 16256T, 16270T, 16302G, +12308, HinfI +7028 AluI	Bramanti 2009

There is more archeological evidence collected by German, Danish and Dutch archeology that suggest continuation. Seen in this light I think the Bollongino article may simply add to the idea.

Thank you for providing the data. Interesting how much of what I would consider "farmer" mtDNA is in them.

I didn't mean to imply, by the way, that there were no remaining settlements of hunter-gatherers in Europe even at this late date. There obviously were. It was the number of such settlements which I was questioning, and the total number of people who would have been practicing this life style, particularly in central Europe. From what I remember of the archaeological studies that dealt with this issue, scholars have proposed very large numbers for the "farmers", and very small numbers for the hunter-gatherers, which makes sense, because that kind of lifestyle just won't sustain large populations, especially when so much of the territory is progressively, year after year, being claimed by farmers. The same thing has happened all over the world. I don't see anything in the European archaeological record that contradicts that. That means that the largest number of hunter-gatherers would have been found in the far north and the far northeast, and to some extent in the far northwest, which were not desirable or optimal locations for the "farmer" lifestyle of the time.

I think the genetic evidence supports that conclusion, especially if we factor in subsequent movements into Europe from the east and then from the north after the fall of Rome.

I'll try to see if I can find some of those papers that discuss the relative population sizes.

Just generally, it seems pretty clear to me, and apparently to the Skoglund et al researchers, although they don't specifically say so, that old Cavalli Sforza may have been generally correct, and that what happened in Europe was that there was a demic diffusion in terms of the spread of agriculture. The fact that the cline in terms of genetics is more south/north east than the south-east/northwest cline that explains the actual movement of the farmers and of agriculture itself has to do, I think, with those later population movements.

However, that's just my opinion as things stand today. I'm not one of those people who has a theory and holds to it no matter what. My thinking on all of this is very fluid. My thoughts on the matter change as new data becomes available. I'm still particularly interested, for example, in what the analysis of the mesolithic peoples of the Balkans will show, and an analysis of the earliest farmers of Anatolia as well.

 

[epoch]

From what I remember of the archaeological studies that dealt with this issue, scholars have proposed very large numbers for the "farmers", and very small numbers for the hunter-gatherers, which makes sense, because that kind of lifestyle just won't sustain large populations, especially when so much of the territory is progressively, year after year, being claimed by farmers.

My idea is that both the adaptation of the HG's and the restriction of the EEFs to the most fertile grounds during the initial phase could have made those differences in population growth less spectacular. I have hunch these numbers were partly assumed because of assessments of other contact zones. In my current idea European HG's are an exception to that rule. Hence the importance of the disease resistance allels I suspect. And, yes: That is a bias.

I suspect we underestimate the resilience of the HG's because we misinterpret a number of semi-neolithic cultures as non-HG.

The same thing has happened all over the world.

Again, I think Europe is an exception.

I don't see anything in the European archaeological record that contradicts that. That means that the largest number of hunter-gatherers would have been found in the far north and the far northeast, and to some extent in the far northwest, which were not desirable or optimal locations for the "farmer" lifestyle of the time.

I think the genetic evidence supports that conclusion, especially if we factor in subsequent movements into Europe from the east and then from the north after the fall of Rome.

 

[epoch]

I'm still leaning toward the idea that in the southern areas where the original Neolithic package was a good "fit", the hunter-gatherers were indeed "wiped out" very early as a distinct group

So, the idea would be that another semi-population replacement afterwards would bring back the WHG part into current day European genome, originating from the east? But I fail to see that if HG's population growth lost so badly from the farmers here, it somehow did not in those parts from which those population movements originated. Shouldn't these areas have experienced a far larger survival rate in order to enhance the WHG rate in Western Europa to present day levels?

 

[motzart]

I am Y DNA haplogroup I2a1 (-M26, -M423). Looks like they found my Grandpa, always thought I was English but looks like I'm Swedish now. I'm also mtDNA H1c3, let me lock myself in a closet with some opium and do a little past-life regression. Hopefully I'll have an explanation for you guys soon.

 

[Fire Haired14]

I am Y DNA haplogroup I2a1 (-M26, -M423). Looks like they found my Grandpa, always thought I was English but looks like I'm Swedish now. I'm also mtDNA H1c3, let me lock myself in a closet with some opium and do a little past-life regression. Hopefully I'll have an explanation for you guys soon.

There are none I2a1-P37.2 subclaees besides M26 and M423. I2a1a-CTS95, I2a1c-L1294, and I2a1b-M423 are the known subclades of I2a1-P37.2, M26 is one of three known I2a1-CTS95 subclades, the two less popular ones according to Eupedia are most popular in north-western Europe. I2a1c-L1294 is the most popular form of I2a1-P37.2 in the British isles, so i bet that's what you have. Pre-I2a1b has been found in Mesolithic north-west Europe, they had 7 of 9 defining mutations of I2a1b. Ajv58 a Neolithic Gotland hunter gatherer who is I2a1-P37.2, probably belonged to a brother lineage of modern I2a1b, or some rare I2a1 lineage that may or may not have been found in modern Europeans.

Since your M423- that means you don't belong to the same lineage as did the pre-I2a1b Mesolithic European samples since they were postive they had derived alleles in SNO M423 but were missing two other I2a1b defining mutations.

Your paternal lineage though is for sure descended of stone age north-west European hunter gatherers, like my maternal lineage(U5b2a2).