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Aberdeen
30-06-14, 19:41
Here's an article published by Nature Communications about a study that suggests Europe's modern mtDNA signature was largely established about 6000 years ago, in the mid Neolithic, by people of an unknown origin who largely replaced the early Neolithic farmers, for reasons that aren't yet clear. Although it does also indicate that Bell Beaker folk expanding out of Iberia did have a significant impact during the late Neolithic.

www.nature.com/ncomms/journal/v4/n4/full/ncomms2656.html (http://www.nature.com/ncomms/journal/v4/n4/full/ncomms2656.html)

Here's the abstract.

Haplogroup H dominates present-day Western European mitochondrial DNA variability (>40%), yet was less common (~19%) among Early Neolithic farmers (~5450 BC) and virtually absent in Mesolithic hunter-gatherers. Here we investigate this major component of the maternal population history of modern Europeans and sequence 39 complete haplogroup H mitochondrial genomes from ancient human remains. We then compare this ‘real-time’ genetic data with cultural changes taking place between the Early Neolithic (~5450 BC) and Bronze Age (~2200 BC) in Central Europe. Our results reveal that the current diversity and distribution of haplogroup H were largely established by the Mid Neolithic (~4000 BC), but with substantial genetic contributions from subsequent pan-European cultures such as the Bell Beakers expanding out of Iberia in the Late Neolithic (~2800 BC). Dated haplogroup H genomes allow us to reconstruct the recent evolutionary history of haplogroup H and reveal a mutation rate 45% higher than current estimates for human mitochondria.

Fire Haired14
30-06-14, 22:32
This is old news,

bicicleur
01-07-14, 08:33
maybe a consequence of the milk revolution, with origins in Anatolia

http://www.nature.com/news/archaeology-the-milk-revolution-1.13471

in Europe : Hamangia culture, Lyengel, Rösen, TRB and British neolithic
Y-DNA , I guess : J1 and T
there was also an expansion into Africa, Y-DNA R1-V88 and T

Maciamo
02-07-14, 08:43
Another paper that completely ignores the Indo-European migrations. Many mtDNA lineages did not spread around Europe until the Bronze Age, as explained here (http://www.eupedia.com/europe/origins_haplogroups_europe.shtml#prehistory).

Fire Haired14
02-07-14, 10:09
Another paper that completely ignores the Indo-European migrations. Many mtDNA lineages did not spread around Europe until the Bronze Age, as explained here (http://www.eupedia.com/europe/origins_haplogroups_europe.shtml#prehistory).

People tend to go with the simple and comforting answer. Nations are constantly moving, mixing, trading, interacting, and so I am sure there were other groups besides eastern Indo Europeans who caused genetic changes in Europe after the initial spread of farming.

Aberdeen
02-07-14, 14:55
Another paper that completely ignores the Indo-European migrations. Many mtDNA lineages did not spread around Europe until the Bronze Age, as explained here (http://www.eupedia.com/europe/origins_haplogroups_europe.shtml#prehistory).

I'm merely trying to generate some discussion about this theory. I haven't seen the background data, so can't argue about how accurate it is. However, when I look at the information available so far from various sources that were collected by Ancestral Journeys, I would have to say that I'm not sure there's enough data available to definitely prove or disprove the idea. Certainly there's more mtDNA H among the 5000 year old Portugese samples and among the Bell Beaker samples than among earlier samples, but I think more detailed data is necessary before one draws definite conclusions about the date when certain subclades of mtDNA H became more dominant in Europe. There does seem to have been some DNA change between the early and later Neolithic, but I don't know if there's enough data to say it all happened at once, as this study seems to be arguing. I suspect it would actually have been ongoing change.

Aberdeen
02-07-14, 20:51
Accidently double posted while editing. See next post.

Aberdeen
02-07-14, 20:53
I was hoping that someone had paid to read the article so they could tell me what it said. When I look at the charts on Ancestral Journeys, I don't see a stable mtDNA structure in Europe between the early Neolithic and the Bronze Age - I think the authors are right to say that there was a significant change between the start of the Neolithic and the start of the Copper Age. But it seems to me that the uptick in mtDNA H really started in Portugal about 3000 B.C. and is also found among Bell Beaker samples after about 2400 B.C. (although H doesn't seem to be as dominant in the earliest Bell Beaker samples). And of course the Bronze Age did make a difference, but more in terms of Y DNA. It looks as if mtDNA remained fairly mixed even in the Bronze Age, with H not being as dominant as it later became. As for this paper and its conclusions about a major mtDNA turnover supposedly taking place about 4000 B.C., I'm having trouble finding out what that conclusion is based on, so if someone has read the whole paper, perhaps they could tell us what it's about.

ebAmerican
02-07-14, 21:35
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3978205/ I believe it is all or most of the paper. What I got out of it was that mtDNA H was brought to Iberia from the near east, and after a natural decline in population of the LBK, and with the spread of the Bell Beaker cultural package mtDNA H merged with the newly developed Corded ware Culture and Central Europe. I believe the author is suggesting that mtDNA H is not Mesolithic/Paleolithic, but was brought to southern Europe during the Neolithic transition. The earlier lines became extinct, and the Iberian lines populated the rest of Europe after the mid Neolithic.

Aberdeen
02-07-14, 22:10
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3978205/ I believe it is all or most of the paper. What I got out of it was that mtDNA H was brought to Iberia from the near east, and after a natural decline in population of the LBK, and with the spread of the Bell Beaker cultural package mtDNA H merged with the newly developed Corded ware Culture and Central Europe. I believe the author is suggesting that mtDNA H is not Mesolithic/Paleolithic, but was brought to southern Europe during the Neolithic transition. The earlier lines became extinct, and the Iberian lines populated the rest of Europe after the mid Neolithic.

Thanks. The time frame is a bit different than indicated in the abstract and fits the actual data better, but the conclusion still seems to me to be a bit of an exaggeration.

polako
03-07-14, 08:26
Here's an article published by Nature Communications about a study that suggests Europe's modern mtDNA signature was largely established about 6000 years ago, in the mid Neolithic, by people of an unknown origin who largely replaced the early Neolithic farmers, for reasons that aren't yet clear.

No, the study doesn't say that. It says that Europe's modern mtDNA signature was largely established about 6000 years ago as a result of migrations into Europe during the early Neolithic.

However, it also says that from the late Neolithic onwards there were new migration waves into the heart of Europe from eastern and western Europe that added new components to this already established gene pool.

Obviously, they're talking about the movements of pan-European cultures like the Bell Beaker and Corded Ware, which makes sense. However, one thing that looks a bit funny in this study is the claim that Indo-European languages spread into Europe during the Neolithic, and then Celtic languages expanded from Iberia with the Bell Beakers.

Of course, any claim that Proto-Indo-European existed during the early Neolithic and sat around in different parts of Europe waiting to expand and differentiate during the Bronze Age can't be taken seriously. It's also at odds with recent ancient DNA evidence which indicates a significant population turnover across Europe during the Copper Age, which is the generally accepted Proto-Indo-European time frame, and a shift from Mediterranean-like to more Northern European-like genetic structure.

Aberdeen
03-07-14, 15:27
No, the study doesn't say that. It says that Europe's modern mtDNA signature was largely established about 6000 years ago as a result of migrations into Europe during the early Neolithic.

However, it also says that from the late Neolithic onwards there were new migration waves into the heart of Europe from eastern and western Europe that added new components to this already established gene pool.

Obviously, they're talking about the movements of pan-European cultures like the Bell Beaker and Corded Ware, which makes sense. However, one thing that looks a bit funny in this study is the claim that Indo-European languages spread into Europe during the Neolithic, and then Celtic languages expanded from Iberia with the Bell Beakers.

Of course, any claim that Proto-Indo-European existed during the early Neolithic and sat around in different parts of Europe waiting to expand and differentiate during the Bronze Age can't be taken seriously. It's also at odds with recent ancient DNA evidence which indicates a significant population turnover across Europe during the Copper Age, which is the generally accepted Proto-Indo-European time frame, and a shift from Mediterranean-like to more Northern European-like genetic structure.

Were you drunk when you wrote that? You "correct" what I said by saying the exact same thing, i.e., that the abstract indicates that Europe's modern mtDNA signature was largely established about 6000 years ago.

Of course, that isn't what the body of the paper is saying. First of all, the paper is partly focussed on one locale but partly addresses the situation throughout Europe, so some of the comments about Europe as a whole are based on the results for that one region, while others are based on the results of Europe as a whole - that was not well thought through. And the paper does conclude that most of the change happened during the Copper and Bronze Ages, although the paper doesn't actually mention the Copper Age. I agree that comment about conflating the Celtic language with Bell Beaker is obvious nonsense. But, no, the Indo-Europeans don't appear to have arrived during the Copper Age, they appear to have arrived during the Bronze Age. It's an interesting question as to why there seems to have been a partial population turnover in Europe during the Copper Age - were those people from an early kind of proto-proto-IE horizon or were they a different but somewhat genetically related group? I think we need more data from Copper Age and Bronze Age Eastern Europe in order to answer that question.

The biggest flaw in the paper, IMO, is that it doesn't really address the mystery of mtDNA H haplotype - while H was present in a small way from early on and it does appear to have increased in frequency all during the period from the mid Neolithic right up to and into the Bronze Age, it still wasn't nearly as dominant as it later became. To me, the real mystery is why H became so much more dominant after the population turnover of the Bronze Age.

polako
03-07-14, 15:39
Were you drunk when you wrote that? You "correct" what I said by saying the exact same thing, i.e., that the abstract indicates that Europe's modern mtDNA signature was largely established about 6000 years ago.

Let me try and explain again.

Your understanding of the study:

Europe's modern mtDNA signature was largely established about 6000 years ago, in the mid Neolithic, by people of an unknown origin who largely replaced the early Neolithic farmers, for reasons that aren't yet clear.

What the study actually argues:

The descendants of early Neolithic farmers weren't replaced about 6000 years ago but were the ones who largely established Europe's modern mtDNA gene pool at this time.

They were then partly replaced by invaders from western and eastern Europe after the final Neolithic (ie. during the Copper Age).

See the difference now?


To me, the real mystery is why H became so much more dominant after the population turnover of the Bronze Age.

Bell Beaker invasion from the west, as mentioned above.

Aberdeen
03-07-14, 17:30
Let me try and explain again.

Your understanding of the study:

Europe's modern mtDNA signature was largely established about 6000 years ago, in the mid Neolithic, by people of an unknown origin who largely replaced the early Neolithic farmers, for reasons that aren't yet clear.

What the study actually argues:

The descendants of early Neolithic farmers weren't replaced about 6000 years ago but were the ones who largely established Europe's modern mtDNA gene pool at this time.

They were then partly replaced by invaders from western and eastern Europe after the final Neolithic (ie. during the Copper Age).

See the difference now?



Bell Beaker invasion from the west, as mentioned above.

Your current comment is somewhat similar to what the paper says, but is not what the abstract says. One of the points I was making is that the abstract doesn't actually reflect what's in the paper. See the difference now? And no, the Bell Beaker "invasion from the west", if it was that, would not explain why the level of dominance by mtDNA H appears to have continued to increase into the Iron Age. As I said, the Copper Age and Bronze Age both seem to have increased the percentage of mtDNA that is H in western Europe, but even at the end of the Bronze Age, H still wasn't yet as dominant as it later became. See my point now?

Fire Haired14
03-07-14, 20:56
Why do you guys group all mtDNA H together? It has over 100 basal clades all with differnt histories.

Greying Wanderer
03-07-14, 22:52
@Aberdeen

"It's an interesting question as to why there seems to have been a partial population turnover in Europe during the Copper Age - were those people from an early kind of proto-proto-IE horizon or were they a different but somewhat genetically related group?"

I think an early IE expansion may have displaced some of the Cucuteni type cultures west of the Black Sea and pushed them west.

"I think we need more data from Copper Age and Bronze Age Eastern Europe in order to answer that question."

yes


"To me, the real mystery is why H became so much more dominant after the population turnover of the Bronze Age."

Yes, my guess it was an adaptation that was only directly advantageous in females e.g. something to do with children.

@Fire-haired

"Why do you guys group all mtDNA H together? It has over 100 basal clades all with differnt histories."

Important point but we'd be having the same argument except referencing specific clades.

polako
04-07-14, 04:05
Your current comment is somewhat similar to what the paper says, but is not what the abstract says. One of the points I was making is that the abstract doesn't actually reflect what's in the paper. See the difference now? And no, the Bell Beaker "invasion from the west", if it was that, would not explain why the level of dominance by mtDNA H appears to have continued to increase into the Iron Age. As I said, the Copper Age and Bronze Age both seem to have increased the percentage of mtDNA that is H in western Europe, but even at the end of the Bronze Age, H still wasn't yet as dominant as it later became. See my point now?


Holy shit. Is English your first language or not?


The paper says exactly what the abstract says. This is the part from the abstract that you're not getting.



Our results reveal that the current diversity and distribution of haplogroup H were largely established by the Mid Neolithic (~4000 BC), but with substantial genetic contributions from subsequent pan-European cultures such as the Bell Beakers expanding out of Iberia in the Late Neolithic (~2800 BC).


So let's break this down:


By 4,000 BC (or 6,000 years ago) the current diversity and distribution of mtDNA H was largely established, and this was among the descendants of early Neolithic farmers, rather than some mysterious population that swept into Europe 6,000 years ago, as you claimed.


However, there were also substantial genetic contributions from around 2800 BC, or the late Neolithic (aka. Copper Age or Chalcolithic), into Central Europe from Western and Eastern Europe that had a significant impact on the modern European mtDNA structure.


The Bell Beakers were one of the most important groups in this respect, because it seems that their Copper Age and early Bronze Age exploits eventually led to later expansions, during the Iron Age, that increased the levels of mtDNA H to around 40% across much of Europe and also upped the frequencies of Atlantic-specific subclades of mtDNA H. So if not for the Bell Beakers, it's likely that Europe today would show lower frequencies of mtDNA H and higher frequencies of Eastern European and Near Eastern derived subclades.

Aberdeen
04-07-14, 04:39
@Aberdeen

"It's an interesting question as to why there seems to have been a partial population turnover in Europe during the Copper Age - were those people from an early kind of proto-proto-IE horizon or were they a different but somewhat genetically related group?"

I think an early IE expansion may have displaced some of the Cucuteni type cultures west of the Black Sea and pushed them west.

"I think we need more data from Copper Age and Bronze Age Eastern Europe in order to answer that question."

yes


"To me, the real mystery is why H became so much more dominant after the population turnover of the Bronze Age."

Yes, my guess it was an adaptation that was only directly advantageous in females e.g. something to do with children.

@Fire-haired

"Why do you guys group all mtDNA H together? It has over 100 basal clades all with differnt histories."

Important point but we'd be having the same argument except referencing specific clades.



You're right. The reason I didn't bother to get into specific subclades is because if we were to discuss specific subclades, we'd have much the same issue to discuss, without getting any better answers. It's not clear why the subclades of H that appear early on in Europe seem to have vanished or become rare, depending on the specific subclade, while other subclades that don't seem to be there earlier later become dominant. I know that strange things happen in genetic history, but I'm sure there are explanations for the changes. I'm just not sure we'll ever have all the answers, although we may get a bit better understanding of the issue once we have more data from the relevant time periods and geographic areas. It still looks as if there was some kind of expansion out of Iberia but we don't yet know how those people got there. And some of the change in H and specific subclades probably came from the east. I don't think the answer is simple.

Aberdeen
04-07-14, 04:45
Holy shit. Is English your first language or not?


The paper says exactly what the abstract says. This is the part from the abstract that you're not getting.





So let's break this down:


By 4,000 BC (or 6,000 years ago) the current diversity and distribution of mtDNA H was largely established, and this was among the descendants of early Neolithic farmers, rather than some mysterious population that swept into Europe 6,000 years ago, as you claimed.


However, there were also substantial genetic contributions from around 2800 BC, or the late Neolithic (aka. Copper Age or Chalcolithic), into Central Europe from Western and Eastern Europe that had a significant impact on the modern European mtDNA structure.


The Bell Beakers were one of the most important groups in this respect, because it seems that their Copper Age and early Bronze Age exploits eventually led to later expansions, during the Iron Age, that increased the levels of mtDNA H to around 40% across much of Europe and also upped the frequencies of Atlantic-specific subclades of mtDNA H. So if not for the Bell Beakers, it's likely that Europe today would show lower frequencies of mtDNA H and higher frequencies of Eastern European and Near Eastern derived subclades.

You don't seem to be very bright, but please try re-reading the paper. Nowhere in it is there any evidence given of a major turnover of mtDNA 6000 years ago, mostly because that didn't happen. Regardless of what the authors of the paper said in terms of generalities, when they do get around to discussing specifics, they admit that H was present in small part fairly early on but the first major uptick of mtDNA H seems to have happened with Bell Beaker and another wave of increased H happened during the Bronze Age, as I previously discussed. In order for you to get a better grasp of the facts, you might also want to look at the charts at Ancestral Journeys. Do you see any evidence there that mtDNA H became dominant 6000 years ago? I don't.

kamani
04-07-14, 05:12
Just a hypothesis from the paper: there was E-v13 found in Iberia at 5000 BC and now it is mainly in the Balkans. H also is at 50% in Balkans. So they must have come together from Iberia at 5000 BC.
Then the Turkish farmers came and made everybody R1b, except some well developed areas that kept their ydna makeup.
Then the Indo-Europeans came with R1a and J.

polako
04-07-14, 05:21
You don't seem to be very bright, but please try re-reading the paper. Nowhere in it is there any evidence given of a major turnover of mtDNA 6000 years ago, mostly because that didn't happen.

I never claimed there was a major turnover of mtDNA 6,000 years ago. You did and the evidence is in the posts above.


Regardless of what the authors of the paper said in terms of generalities, when they do get around to discussing specifics, they admit that H was present in small part fairly early on but the first major uptick of mtDNA H seems to have happened with Bell Beaker and another wave of increased H happened during the Bronze Age, as I previously discussed.

No, the authors very obviously state that the major portion of European mtDNA H diversity and distribution was established by the middle Neolithic, around 6,000 years ago. This is stated several times in the posts above, including your own.

And no, there was no "other wave" of mtDNA H into Central Europe during the Bronze Age. The phylogeography of the full mtDNA sequences studied in the paper show that Neolithic farmers and Bell Beakers from the Atlantic facade can explain almost all of the mtDNA H in Europe (the rest can be explained by Corded Ware and other expansions from the east which carried Eastern European-specific mtDNA H subclades).

So what most likely happened was that the Neolithic and Bell Beaker-derived populations, sitting in Western and Central Europe, where population densities were relatively high and could get much higher than in Eastern Europe, experienced large population growth during the metal ages, thereby pushing up the frequencies of mtDNA H and the Atlantic-derived subclades of mtDNA H even further.


Do you see any evidence there that mtDNA H became dominant 6000 years ago?

Yes, it's stated in the paper including in the abstract. Here's that quote again.


Our results reveal that the current diversity and distribution of haplogroup H were largely established by the Mid Neolithic (~4000 BC), but with substantial genetic contributions from subsequent pan-European cultures such as the Bell Beakers expanding out of Iberia in the Late Neolithic (~2800 BC).

Maciamo
04-07-14, 08:52
No, the authors very obviously state that the major portion of European mtDNA H diversity and distribution was established by the middle Neolithic, around 6,000 years ago. This is stated several times in the posts above, including your own.

And no, there was no "other wave" of mtDNA H into Central Europe during the Bronze Age. The phylogeography of the full mtDNA sequences studied in the paper show that Neolithic farmers and Bell Beakers from the Atlantic facade can explain almost all of the mtDNA H in Europe (the rest can be explained by Corded Ware and other expansions from the east which carried Eastern European-specific mtDNA H subclades).

So what most likely happened was that the Neolithic and Bell Beaker-derived populations, sitting in Western and Central Europe, where population densities were relatively high and could get much higher than in Eastern Europe, experienced large population growth during the metal ages, thereby pushing up the frequencies of mtDNA H and the Atlantic-derived subclades of mtDNA H even further.


Although I agree that most major mtDNA H subclades were already present in Europe during the mid Neolithic, I can't agree with this paper that the modern distribution of H subclades was already established back then. As you said, the Corded Ware (+ the Unetice and all subsequent IE cultures) redistributed eastern H subclades across most of Europe, and by doing so changing substantially the mtDNA landscape and the frequencies of H subclades.

I also disagree with the authors' view that mtDNA H was virtually absent from Mesolithic Europeans. Most of the Mesolithic samples are from the northern half of Europe, where indeed hg H was rare (though not completely absent). But southern samples (e.g. Iberia) show a considerable present of hg H at least since the Mesolithic. I wouldn't be surprised if Mesolithic Italians and Southeast Europeans were the true source of most Neolithic H subclades. Only a few subclades appear to be truly Near Eastern in origins, like H2, H5, H7, H13 and H20.

So, in my opinion, in the late Palaeolithic and Mesolithic haplogroup H was already distributed around most of the Mediterranean basin. Early Neolithic farmers only redistributed Near Eastern and Balkanic lineages across Europe. Late Neolithic Bell Beakers redistributed Iberian lineages to central and north-west Europe. Then the Indo-Europeans redistributed eastern European and Caucasian lineages to most of Europe.

Fire Haired14
04-07-14, 09:40
Although I agree that most major mtDNA H subclades were already present in Europe during the mid Neolithic, I can't agree with this paper that the modern distribution of H subclades was already established back then. As you said, the Corded Ware (+ the Unetice and all subsequent IE cultures) redistributed eastern H subclades across most of Europe, and by doing so changing substantially the mtDNA landscape and the frequencies of H subclades.

I also disagree with the authors' view that mtDNA H was virtually absent from Mesolithic Europeans. Most of the Mesolithic samples are from the northern half of Europe, where indeed hg H was rare (though not completely absent). But southern samples (e.g. Iberia) show a considerable present of hg H at least since the Mesolithic. I wouldn't be surprised if Mesolithic Italians and Southeast Europeans were the true source of most Neolithic H subclades. Only a few subclades appear to be truly Near Eastern in origins, like H2, H5, H7, H13 and H20.

So, in my opinion, in the late Palaeolithic and Mesolithic haplogroup H was already distributed around most of the Mediterranean basin. Early Neolithic farmers only redistributed Near Eastern and Balkanic lineages across Europe. Late Neolithic Bell Beakers redistributed Iberian lineages to central and north-west Europe. Then the Indo-Europeans redistributed eastern European and Caucasian lineages to most of Europe.

All we can say about the "H" from Mesolithic Portugal and Magdalenian Spain is that they without almost any doubt had R0 because all of them except the reported H6 had G73A. The testing was so primitive there is no way to know for sure what type of R0 they had(there are many possibilities). There is also for sure 24,000 year old R0 from Gravettian Italy because it had both R0's mutations.

I don't understand how La brana-1 could be 100% the same thing as northwest European hunter gatherers if hunter gatherers from his area had ancestry(maternal lineage R0) the northwest European hunter gatherers lacked. If south European hunter gatherers during the LGM had a high amount of R0 and their descendants resettled northern Europe, then why did northern hunter gatherers completely lack R0? R0 in Italy 24,000 years ago is explainable but R0 in Portugal during the end of the Mesolithic is not. It is an interesting idea and possible but unlikely.

The European hunter gatherer R0 samples could somehow be false, maybe the dates(Neolithic not Mesolithic) are off, who knows. They don't make any sense(especially in Mesolithic Portugal) and at the time of their discovery the mainstream theory was that the first Europeans were full of mtDNA H, and prove that they were not would have been crushing for many people.


But southern samples (e.g. Iberia) show a considerable present of hg H at least since the Mesolithic. I wouldn't be surprised if Mesolithic Italians and Southeast Europeans were the true source of most Neolithic H subclades. Only a few subclades appear to be truly Near Eastern in origins, like H2, H5, H7, H13 and H20.

Would you say the same is true for J1c, T2b, J2b1a, J2a1a, and J1b1a1? To me it seems like mtDNA that the evidence"European" subclades of west Asian haplogroups originated around west Asia were somehow erased after they arrived in Europe, and the only way we know they came from west Asia is ancient DNA.

Maciamo
04-07-14, 11:43
I don't understand how La brana-1 could be 100% the same thing as northwest European hunter gatherers if hunter gatherers from his area had ancestry(maternal lineage R0) the northwest European hunter gatherers lacked.

La Brana is the north of Spain, close to Cantabria and the Basque country, a region that is climatically closer to France and the British Isles, and probably shared more with north-west European hunter-gatherers than with Mediterranean hunter-gatherers.

Wilhelm
04-07-14, 16:24
All we can say about the "H" from Mesolithic Portugal and Magdalenian Spain is that they without almost any doubt had R0 because all of them except the reported H6 had G73A. The testing was so primitive there is no way to know for sure what type of R0 they had(there are many possibilities). There is also for sure 24,000 year old R0 from Gravettian Italy because it had both R0's mutations.

I don't understand how La brana-1 could be 100% the same thing as northwest European hunter gatherers if hunter gatherers from his area had ancestry(maternal lineage R0) the northwest European hunter gatherers lacked. If south European hunter gatherers during the LGM had a high amount of R0 and their descendants resettled northern Europe, then why did northern hunter gatherers completely lack R0? R0 in Italy 24,000 years ago is explainable but R0 in Portugal during the end of the Mesolithic is not. It is an interesting idea and possible but unlikely.

The European hunter gatherer R0 samples could somehow be false, maybe the dates(Neolithic not Mesolithic) are off, who knows. They don't make any sense(especially in Mesolithic Portugal) and at the time of their discovery the mainstream theory was that the first Europeans were full of mtDNA H, and prove that they were not would have been crushing for many people..
There is also a Mesolithic Russian with mtDNA H.

Aberdeen
04-07-14, 17:15
I never claimed there was a major turnover of mtDNA 6,000 years ago. You did and the evidence is in the posts above.



No, the authors very obviously state that the major portion of European mtDNA H diversity and distribution was established by the middle Neolithic, around 6,000 years ago. This is stated several times in the posts above, including your own.

And no, there was no "other wave" of mtDNA H into Central Europe during the Bronze Age. The phylogeography of the full mtDNA sequences studied in the paper show that Neolithic farmers and Bell Beakers from the Atlantic facade can explain almost all of the mtDNA H in Europe (the rest can be explained by Corded Ware and other expansions from the east which carried Eastern European-specific mtDNA H subclades).

So what most likely happened was that the Neolithic and Bell Beaker-derived populations, sitting in Western and Central Europe, where population densities were relatively high and could get much higher than in Eastern Europe, experienced large population growth during the metal ages, thereby pushing up the frequencies of mtDNA H and the Atlantic-derived subclades of mtDNA H even further.



Yes, it's stated in the paper including in the abstract. Here's that quote again.



Your first two statements contradict one another. As for a statement about mtDNA becoming dominant 6000 years ago, I'll explain what's wrong with the statement by using a simple example, because that seems to be necessary if I want you to understand. If researchers found and presented a lot of data showing that donkeys can't fly, but stated in their paper and in their abstract that their evidence shows that donkeys can fly, that would not be proof, it would be an inaccurate statement.

Fire Haired14
04-07-14, 19:05
There is also a Mesolithic Russian with mtDNA H.

But it's only around 7,000 years old and could be explained by farmer admixture unlike 12,000 year old R0 from Spain.

Sile
04-07-14, 22:13
But it's only around 7,000 years old and could be explained by farmer admixture unlike 12,000 year old R0 from Spain.

What are you trying to say?.............that mtDna H advanced along the afrcan coast into iberia before moving into Europe via Anatolia!

Fire Haired14
05-07-14, 00:51
What are you trying to say?.............that mtDna H advanced along the afrcan coast into iberia before moving into Europe via Anatolia!

Farmers had already established themselves in areas where they could have had contact with the hunter gatherers of Uznyi Oleni Ostrov. The east Asian C1 and in later samples Z1a found there are from east Asia, and so why can't the H be from European farmers? Besides it is only one isolated H, in no way is it evidence most H in modern Europe is from European hunter gatherers. It is ridiculous that people focus so much on H when European mtDNA is in subject, H has around 100 basal clades, it was total luck that the diverse and old H lineage became 40% in almost all of Europe.

Sile
05-07-14, 02:15
Farmers had already established themselves in areas where they could have had contact with the hunter gatherers of Uznyi Oleni Ostrov. The east Asian C1 and in later samples Z1a found there are from east Asia, and so why can't the H be from European farmers? Besides it is only one isolated H, in no way is it evidence most H in modern Europe is from European hunter gatherers. It is ridiculous that people focus so much on H when European mtDNA is in subject, H has around 100 basal clades, it was total luck that the diverse and old H lineage became 40% in almost all of Europe.

wasn't mtDna U and its sister K earlier than H in Europe ..............maybe even T2

Fire Haired14
05-07-14, 02:46
wasn't mtDna U and its sister K earlier than H in Europe ..............maybe even T2

U and K are not sister clades, K is another name for U8b2. K was probably not in Europe or at least most of it before the Neolithic. There is an around 30,000 year old U8 sample from the Czech Republic but it's specifically U8c, so not K.

polako
05-07-14, 05:59
Your first two statements contradict one another. As for a statement about mtDNA becoming dominant 6000 years ago, I'll explain what's wrong with the statement by using a simple example, because that seems to be necessary if I want you to understand. If researchers found and presented a lot of data showing that donkeys can't fly, but stated in their paper and in their abstract that their evidence shows that donkeys can fly, that would not be proof, it would be an inaccurate statement.

The fact that most of the modern mtDNA H diversity and distribution was established in Europe by the middle Neolithic doesn't contradict the fact that there was no mtDNA turnover in Europe 6,000 years ago.

That's because this diversity and distribution was established via gradual migrations over thousands of years as well as genetic drift.

Quit embarrassing yourself and move on while you're not too far behind.

polako
05-07-14, 06:05
Although I agree that most major mtDNA H subclades were already present in Europe during the mid Neolithic, I can't agree with this paper that the modern distribution of H subclades was already established back then. As you said, the Corded Ware (+ the Unetice and all subsequent IE cultures) redistributed eastern H subclades across most of Europe, and by doing so changing substantially the mtDNA landscape and the frequencies of H subclades.

They couldn't have changed the character of mtDNA H across Europe too much, because they didn't carry much mtDNA H. Their contribution is better seen in the rising levels of U4, U5, U2, and I in much of Europe after the Neolithic.

epoch
05-07-14, 07:58
I don't understand how La brana-1 could be 100% the same thing as northwest European hunter gatherers if hunter gatherers from his area had ancestry(maternal lineage R0) the northwest European hunter gatherers lacked. If south European hunter gatherers during the LGM had a high amount of R0 and their descendants resettled northern Europe, then why did northern hunter gatherers completely lack R0? R0 in Italy 24,000 years ago is explainable but R0 in Portugal during the end of the Mesolithic is not. It is an interesting idea and possible but unlikely.

The European hunter gatherer R0 samples could somehow be false, maybe the dates(Neolithic not Mesolithic) are off, who knows. They don't make any sense(especially in Mesolithic Portugal) and at the time of their discovery the mainstream theory was that the first Europeans were full of mtDNA H, and prove that they were not would have been crushing for many people.


I have been thinking about that too. The most surprising thing is that Stuttgart is supposedly 25%-30% WHG but there is hardly any mtDNA U5 or U4 in Lineband Beaker settlements found. That itself could mean - I am merely speculating - that they picked it up in the south east of Europa and that Hunter-Gatherers there had different mtDNA.

We mostly have data of Nothern and north-western WHG's which could be somehow "cherry picking" for U5. Mesolithich HG's are supposed to be the same people as the late Paleolithic (Correct me if I'm wrong) and in the late Paleolithic population was extremely thin. So the omnipresence of U5 in the northern part of WHG's might be due to some sort of founder effect, especially since the area was repopulated only after the LGM. Iberian HG's had Y-DNA had C1 too, another hint that haplogroups absent from WHG in the north and north-west could have been present in the south. Also, apparently the mesolithic finds in Greece point to more mtDNA than just U5. I thought it was X1.

epoch
05-07-14, 08:14
@Aberdeen & Polako



Quit embarrassing yourself and move on while you're not too far behind.


Were you drunk when you wrote that?

Gentlemen. If you left out remarks such as cited yours would actually look like an interesting discussion. I understand how sometimes irkiness enters the discussion but we all should put effort in bending the discussion back to civilized.

Mind you, not to say you aren't civilized gentlemen.

LeBrok
05-07-14, 08:25
@Aberdeen & Polako

Gentlemen. If you left out remarks such as cited yours would actually look like an interesting discussion. I understand how sometimes irkiness enters the discussion but we all should put effort in bending the discussion back to civilized.

Mind you, not to say you aren't civilized gentlemen.
I second that.

Fire Haired14
05-07-14, 09:02
etrdugiuhuohoho

Maciamo
05-07-14, 09:37
They couldn't have changed the character of mtDNA H across Europe too much, because they didn't carry much mtDNA H. Their contribution is better seen in the rising levels of U4, U5, U2, and I in much of Europe after the Neolithic.

It is true that the Bronze Age steppe people carried less mtDNA H than the European average, but based on the few hundreds ancient samples (http://www.eupedia.com/genetics/haplogroups_of_bronze_age_proto-indo-europeans.shtml) from the Yamna, Corded Ware, Catacomb and Unetice cultures, they still had a significant 25% of h H. Typical H subclades of Bronze Age steppe people include H1b, H1c, H2a1, H4a1, H5a, H6, H10 and H11.

epoch
05-07-14, 10:03
@Fire Haired

I have also been suprised that southern Scandinavia has such high WHG affinity where it has low U5/U4 rates compared to northern Scandinavia. Yet la Brana shows great affinity to the Scandinavians. Did they explicitly mean non-southern Swedes by that? In that case it is surprising that current day Swedes aren't similar to early Funnel beaker farmers but show larger affinity to WHG.

The mismatch between mtDNA ancestry and autosomal is quite strange.

I have a hunch that while LBK might be clearly a colonisation of EEF (20% WHG) that managed to stay more or less separate for a thousand of years from surrounding WHG's - as some studies suggest - the cultures that followed were more hybrids. And even then there might be considerable differences in WHG ancestry between different sub-cultures. Rosen culture, Baalberg culture, they might have collected a number of WHG's when colonizing new area's whereas the dutch Swifterband culture might be a slow adaptation of WHG's to husbandry. So in the end we end up with villages across Europa with each different but substantial autosomal WHG ancestry.

Farmer villages in ancient times may have been far more isolated than today. After initial colonization farmer communities may have become quite closed communities, as they are until this day. Even recently one wouldn't marry all that easy into closed communities. In closed communities, due to founder effect en genetic drift, minority haplogroups have a slightly higher chance of disappearing completely so after a while U5 might simply disappear, while autosomically these communities might have considerable WHG ancestry.

It's just a thought, mind you.

Greying Wanderer
05-07-14, 10:36
All we can say about the "H" from Mesolithic Portugal and Magdalenian Spain is that they without almost any doubt had R0 because all of them except the reported H6 had G73A. The testing was so primitive there is no way to know for sure what type of R0 they had(there are many possibilities). There is also for sure 24,000 year old R0 from Gravettian Italy because it had both R0's mutations.

I don't understand how La brana-1 could be 100% the same thing as northwest European hunter gatherers if hunter gatherers from his area had ancestry(maternal lineage R0) the northwest European hunter gatherers lacked. If south European hunter gatherers during the LGM had a high amount of R0 and their descendants resettled northern Europe, then why did northern hunter gatherers completely lack R0? R0 in Italy 24,000 years ago is explainable but R0 in Portugal during the end of the Mesolithic is not. It is an interesting idea and possible but unlikely.



I think the HGs during the LGM are likely to have been split into two related but separate populations mirroring two ecosystems: a southern and coastal population including southern Europe and around the Atlantic coast and a mammoth steppe population extending from northern France to Siberia. If so the resettlement of northern Europe after the end of LGM might easily have been mostly from the mammoth steppe population as they were closer with some distinctive input from the more coastal population along the Atlantic coast.

edit: actually three might make more sense, a southern refuge population, a coastal maritime population up the Atlantic coast (thinking of Eskimo as an example of a coastal population living in a cold environment then possibly quite far north along the Atlantic coast) and thirdly a mammoth steppe population).

Angela
05-07-14, 13:24
There can’t be any rational discussion of the issues raised by Brotherton et al, or any advancement in our knowledge of this period when statements are made concerning it that are totally at odds with a plain reading of the text.


Once again, this is what they say in the abstract: Our results reveal that the current diversity and distribution of haplogroup H were largely established by the Mid Neolithic (~4000 BC), but with substantial genetic contributions from subsequent pan-European cultures such as the Bell Beakers expanding out of Iberia in the Late Neolithic (~2800 BC).


They are thus claiming that already by 4,000 BC, the major part of the current diversity and distribution of H had already been established, and that therefore Bell Beaker, and Corded Ware, and Unetice) did not create the general pattern of diversity and distribution of mtDNA “H” we see in modern populations. It was already in place, or so they claim.


In the body of the paper they then go on to state this, “The combined set of analyses (PCA, Procrustes and Ward clustering) revealed that Mittelelbe-Saale’s earliest farmers (LBK; n=9) cluster with present-day Caucasus, Near Eastern, and Anatolian populations, as previously noted7 (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3978205/#R7). In contrast, individuals from the successor series of regional post-LBK (and Mid Neolithic) Rössen, Schöningen, Baalberge, and Salzmünde cultures (ca. 4625-3025 BC, MNE; n=10) cluster with present-day Central European populations (Figure 2 (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3978205/figure/F2/)


They further state that ,” ENE (Early Neolithic) mt genomes are generally either rare today19 (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3978205/#R19) or have not yet been observed in present-day populations, possibly due to subsequent extinction of these lineages” and that “This suggests that individuals from the Early Neolithic made a marginal contribution to Late Neolithic and present day hg H diversity. Although the relatively small sample numbers from each time period limit detailed analyses of the causes of the distribution shifts, we interpret this phylogenetic pattern as a genetic discontinuity between Early and subsequent Neolithic cultures in Europe, potentially mirroring genetic structure in Neolithic European populations.”


Now, such a change could be caused by either migrations or drift. The authors specifically state that drift was not the cause:

“Genetic drift could also have played a role in generating discrepant hg distributions over time and space. However, if drift was the sole cause we would expect a random distribution across all sub-hgs rather than a clear distinction between ENE and MNE/LNE/Bronze Age mt genomes.”

That leaves migration of peoples bearing other subclades of “H” as the explanation, but they never state that explicitly and so they obviously don’t explain where and when these people arrived in the area. Now, I personally have no problem speculating that there was continuous gene flow during the Neolithic or perhaps a few pulses of gene flow which would bring diversity to the area. I do have a problem with an analysis that leaves that as the only explanation but doesn’t explicitly address the issue. Nor, for that matter, do I think their argument against drift as an explanation is all that convincing. If the sample sizes are too small from each period to figure out how the diversity developed, then why aren't they too small to so definitively rule out drift as one of the factors.


Apologies to Aberdeen for speaking for him, but it seems to me that this must be the same logical trail he followed, a trail which led him to conclude that, “Here's an article published by Nature Communications about a study that suggests Europe's modern mtDNA signature was largely established about 6000 years ago, in the mid Neolithic, by people of an unknown origin who largely replaced the early Neolithic farmers, for reasons that aren't yet clear.”

The authors are clearly proposing that some genetic transition did indeed take place between the LBK or early Neolithic and the mid-Neolithic cultures before the arrival of Bell Beaker, Corded Ware, or Unetice. Now, this may not be supported by the data, it may indeed not be the best explanation of what actually happened in Europe, but it is unambiguously what they did say.


They are most definitely not saying that “ Europe's modern mtDNA signature was largely established about 6000 years ago as a result of migrations into Europe during the early Neolithic" to quote a post by Polako. Given the text of the main paper quoted above, it's also incorrect to maintain that the authors propose that “The descendants of early Neolithic farmers weren't replaced about 6000 years ago but were the ones who largely established Europe's modern mtDNA gene pool at this time." Clearly they are saying that there were indeed changes between the arrival of the first farmers and the Middle Neolithic of 6000 years ago. (4000 BC)


One can disagree with the conclusions of Brotherton et al, but one can’t re-write them; that just leads to frustration, confusion among readers, and further bad conclusions.


In addition to all of the above problems with this paper, I agree with Aberdeen that the authors are making a huge error in taking results from one area in Germany, and very small sample sizes for each specific time period, and presuming to then extrapolate from that to make broad generalizations about the peopling of Europe.

LeBrok
05-07-14, 17:35
That leaves migration of peoples bearing other subclades of “H” as the explanation, but they never state that explicitly and so they obviously don’t explain where and when these people arrived in the area. Now, I personally have no problem speculating that there was continuous gene flow during the Neolithic or perhaps a few pulses of gene flow which would bring diversity to the area.
We can see that with farmers/middle eastern Y-dna pattern too. They don't mirror each other much and not at all in some cases. Suggesting separate entrances into Europe. In other words, few waves of farmers during Neolithic.

Aberdeen
05-07-14, 19:49
There can’t be any rational discussion of the issues raised by Brotherton et al, or any advancement in our knowledge of this period when statements are made concerning it that are totally at odds with a plain reading of the text.


Once again, this is what they say in the abstract: Our results reveal that the current diversity and distribution of haplogroup H were largely established by the Mid Neolithic (~4000 BC), but with substantial genetic contributions from subsequent pan-European cultures such as the Bell Beakers expanding out of Iberia in the Late Neolithic (~2800 BC).


They are thus claiming that already by 4,000 BC, the major part of the current diversity and distribution of H had already been established, and that therefore Bell Beaker, and Corded Ware, and Unetice) did not create the general pattern of diversity and distribution of mtDNA “H” we see in modern populations. It was already in place, or so they claim.


In the body of the paper they then go on to state this, “The combined set of analyses (PCA, Procrustes and Ward clustering) revealed that Mittelelbe-Saale’s earliest farmers (LBK; n=9) cluster with present-day Caucasus, Near Eastern, and Anatolian populations, as previously noted7 (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3978205/#R7). In contrast, individuals from the successor series of regional post-LBK (and Mid Neolithic) Rössen, Schöningen, Baalberge, and Salzmünde cultures (ca. 4625-3025 BC, MNE; n=10) cluster with present-day Central European populations (Figure 2 (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3978205/figure/F2/)


They further state that ,” ENE (Early Neolithic) mt genomes are generally either rare today19 (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3978205/#R19) or have not yet been observed in present-day populations, possibly due to subsequent extinction of these lineages” and that “This suggests that individuals from the Early Neolithic made a marginal contribution to Late Neolithic and present day hg H diversity. Although the relatively small sample numbers from each time period limit detailed analyses of the causes of the distribution shifts, we interpret this phylogenetic pattern as a genetic discontinuity between Early and subsequent Neolithic cultures in Europe, potentially mirroring genetic structure in Neolithic European populations.”


Now, such a change could be caused by either migrations or drift. The authors specifically state that drift was not the cause:

“Genetic drift could also have played a role in generating discrepant hg distributions over time and space. However, if drift was the sole cause we would expect a random distribution across all sub-hgs rather than a clear distinction between ENE and MNE/LNE/Bronze Age mt genomes.”

That leaves migration of peoples bearing other subclades of “H” as the explanation, but they never state that explicitly and so they obviously don’t explain where and when these people arrived in the area. Now, I personally have no problem speculating that there was continuous gene flow during the Neolithic or perhaps a few pulses of gene flow which would bring diversity to the area. I do have a problem with an analysis that leaves that as the only explanation but doesn’t explicitly address the issue. Nor, for that matter, do I think their argument against drift as an explanation is all that convincing. If the sample sizes are too small from each period to figure out how the diversity developed, then why aren't they too small to so definitively rule out drift as one of the factors.


Apologies to Aberdeen for speaking for him, but it seems to me that this must be the same logical trail he followed, a trail which led him to conclude that, “Here's an article published by Nature Communications about a study that suggests Europe's modern mtDNA signature was largely established about 6000 years ago, in the mid Neolithic, by people of an unknown origin who largely replaced the early Neolithic farmers, for reasons that aren't yet clear.”

The authors are clearly proposing that some genetic transition did indeed take place between the LBK or early Neolithic and the mid-Neolithic cultures before the arrival of Bell Beaker, Corded Ware, or Unetice. Now, this may not be supported by the data, it may indeed not be the best explanation of what actually happened in Europe, but it is unambiguously what they did say.


They are most definitely not saying that “ Europe's modern mtDNA signature was largely established about 6000 years ago as a result of migrations into Europe during the early Neolithic" to quote a post by Polako. Given the text of the main paper quoted above, it's also incorrect to maintain that the authors propose that “The descendants of early Neolithic farmers weren't replaced about 6000 years ago but were the ones who largely established Europe's modern mtDNA gene pool at this time." Clearly they are saying that there were indeed changes between the arrival of the first farmers and the Middle Neolithic of 6000 years ago. (4000 BC)


One can disagree with the conclusions of Brotherton et al, but one can’t re-write them; that just leads to frustration, confusion among readers, and further bad conclusions.


In addition to all of the above problems with this paper, I agree with Aberdeen that the authors are making a huge error in taking results from one area in Germany, and very small sample sizes for each specific time period, and presuming to then extrapolate from that to make broad generalizations about the peopling of Europe.

Thank you for that excellent summary of the situation, Angela. This is definitely one of those papers where the facts don't fit the conclusions. The authors state that "Our results reveal that the current diversity and distribution of hg H were largely established by the Mid-Neolithic (~4000 BC), but with substantial genetic contributions from subsequent pan-European cultures such as the Bell Beakers expanding out of Iberia in the Late Neolithic (~2800 BC)." However, that simply isn't the case, as is noted in the paper as well as in various other sources. 6000 years ago, H was less common than it later became, and those examples of H that have been found from 4000 BC and earlier show a substantially different prevalence of subclades than what developed later. And we seem to agree that there's a problem with trying to use the results from one locale as a proxy for all of Europe, since the results in that locale are not in fact representative of Europe. There's a lot wrong with this paper, although I don't have time at the moment to address it all in detail. I just wish I had been able to see the paper before I started this thread on the basis of curiosity about the abstract, since I really don't think this paper should be taken too seriously.

Sile
05-07-14, 20:36
Certainly. However, history shows that few men can father children at a lot of women. The other way around does not seem plausible. So I had the impression that mtDNA should follow autosomical DNA more than Y-DNA.

What I call " dynastic ydna". Men who had a concubine due to their status in society ...................something we still saw in history as late as the 19th century in Zulu culture




If I am right the Baltic states never had significant neolithic immigration that kickstarted the neolithicum there. However, Baltics still do have significant EEF. They also have significant ANE and speak a Indo-European language. This might serve as evidence that there possibly the expanding Indo-Europeans carried an lot of EEF as well as WHG. Allthough possibly the EEF the Baltics show affinity to might be the exact bit in Stuttgart that was derived from WHG. Lazardis discusses that, and suggests a 20% WHG admixture.



The amber trade/migration trail and the jutland to Genoa migration/trade trail where already in action by the late-neolithic period

FrankN
06-07-14, 04:39
I need a few more looks at the paper itself to comment on its content, but it seems to me that some posters here have problems to place it into the correct historical setting:
Let's start with the Elbe-Saale region that has been analysed in the paper. Of course it is only one Central European region, and as such not necessarily representative of other regions. However, if there is a single region that captures most of Central European population dynamics, it is exactly that area along the Middle Elbe and Saale:

Central part of the LBK EEF expansion during the early Neolithic from the Lower Danube north-westwards (LBK arrived at the Middle Elbe a couple of centuries before it reached the middle and lower Rhine);
Part of the area where lactase persistency developed first. Recent genetic research puts the origin somewhere into Western Hungary, i.e. the starting point of the LBK expansion, in the second half of the 5th millennium BC. Lactase persistency seems to have spread strongest along the central LBK expansion path - through Moravia, Bohemia and along the Upper Elbe - and less intensively / slower along the western path (Upper Danube / Rhine). The Middle Elbe/ Saale region is still today part of the area where lactase persistency is highest within Europe;
Part (actually the western edge) of the first stage of IE expansion into Central Europe, marked by the Globular Amphora Culture, and the place where quite a number of linguists assume Proto-Germanic to have been formed;
Eastern border of the Bell Beaker expansion out of Iberia, and the only region where co-existence of Bell Beakers and (IE- influenced) Global Amphora / Corded Ware people is archeologically documented;
Convergence zone of (Proto-Celtic) Urnfield & Hallstatt cultures, and (Proto-Germanic) Jastorf culture;
Western border of the Slavic expansion during the early middle ages.

LBK EEFs settled on the loess plains along the major rivers (Danube, Morava, Elbe, Vltava, Oder, Rhine, Main, etc.). Archaeological research in the Rhine-Main area around today's Frankfurt, and by Czech archaeologists along the Upper Elbe and around Prague shows that the LBK heartland was quite densely populated. The mountain ranges between the main rivers, however, were left to HGs. This may, especially when looking at older maps, create the impression of isolated settlements. But in fact, as recent excavations have shown, the settlement pattern included larger "towns" every 50-60 km, and smaller settlements as well as individual "farms" in-between and reaching into secondary valleys. Not a pattern that necessarily promotes genetic isolation and drift.
Below is a recently published map by the Archeological Service of Saxony Anhalt, which indicates all Unetice sites (2,700 - 2,100 BC) that have been identified/ documented to date. The indicated settlement density is actually higher than today (though many of the places may not have been settled continuously, but only for some 3-5 generations before the soils were exploited and settlers moved on to a nearby place).
http://vanaland.files.wordpress.com/2013/05/aunjetitz_fundstellen.jpg

Entrance of EEF into HG areas has obviously lead to conflicts, in particular as LBK farming seems to have been strongly based on cattle herding, with field crops (grain, linen) only playing a secondary (but nevertheless important) role. What is more tempting to hunters than large herds of well-fed cattle? However, aside from the "Australian Aborigines" / "Native Americans" way to "solve" such conflicts, there is another possibility: Farming and cattle-herding communities along the Niger in West Africa employ Saharans (Tuareg) for their salt supply. Salt trade, while it surely existed already during the Neolithic, is difficult to trace archeologically. But we have ample evidence of medium-distance flint and tool trade. One well documented case is the flint mines near Kehlheim in Upper Bavaria, which supplied their flint as far as Lake Constance and the Middle Rhine to the West, and Dresden, Prague and Linz to the (south-)east. The "flint road" from Kehlheim to Pilsen (from where it continued to Prague) is archeologically well documented from a series of camps in the Bavarian Forest where evidence of small-scale stone processing has been found. The area is still scarcely populated today, and, with one exception (near the town of Cham), no traces of Neolitihic agriculture have been found along the "flint road". As such, it is assumed that traders, following the rivers and creeks, just used their spare time on the evening campfire to get rid of a bit of weight by doing some stone processing. And I am pretty sure that these traders didn't carry sandwiches with them, but hunted (or, more likely, fished) for their food along the way. In other words - some LBK EEF communities found a way to economically integrate HGs from the periphery into their economy, just as Sahel farmers have done with the Tuareg. I furthermore assume that most (all?) of the flint miners and processors were originally HGs - they probably were already regularly (outside hunting season) visiting such mines and preparing replacement spear and arrow heads before farming created additional demand for flint tools. EEFs marrying women from HG communities may have helped to promote such, more peaceful ways of coexistence, and could have become quite frequent once regular trade connections were established.

I am not aware of genetic studies of LBK graves. However, later graves from the extended Middle Elbe / Saale region (Eulau, Lichtenstein cave) clearly indicate a patrilocal culture, whereby women move towards the husband's residence. In both cases, strontium analyses yielded that the adult women were born at least 60 km away from their burial place. As such, a mtDNA "travelling speed" of 200-250 km/ century across Central Europe is well possible even in the absence of mass migrations.

John Doe
06-07-14, 11:11
Haplogroup U was present in Europe before the Neolithic, is it possible that some carriers of U migrated to Western Asia where K, the subclade of U originated, and then some West Asian carriers of K (such as Jews) migrated into Europe?

epoch
06-07-14, 11:14
@FrankN

The genetic history of pigs is quite an interesting one. It shows that early LBK settlers took their own pigs with them, as the earliest remains of pigs show affinity with Anatolian wild boar. There are a number of cultures, if I understand correctly that are considered surviving hunter-gatherers, one of them being the Ertebolla culture, that started to keep pigs in their villages. The remains of the oldest pigs found at Ertebolla sites also show affinity with Anatolian wild boar. That makes it very clear that trade between LBK and HG's existed.

http://archaeology.about.com/od/domestications/qt/pigs.htm
http://www.nature.com/ncomms/2013/130827/ncomms3348/full/ncomms3348.html

(The latter of the two links also states that Ertebolla and LBK lived alongside each other but apart from each other for almost thousand years.)

Strangely enough remains of more recent neolithic pigs shows them to be descendant from local wild boar.

https://www.academia.edu/1949053/Genetic_aspects_of_pig_domestication



The same Near Easternhaplotype was also identified in four specimensfrom the 8th millennium Linear Bandkeramik (LBK) site of Eilsleben in northern Germany,and in two samples from the mid Neolithic(very early 6th millennium Chasséen culture)site of Bercy in the Paris Basin. At Bercy,mtDNA sequences of European origin werealso extracted from archaeological specimensidentified by zooarchaeological criteria asdomestic swine, making it the earliest sitewhere both Near Eastern and Europeandomestic swine have been identified together.


EDIT: One more link about pigs genetic history:

http://geknitics.com/2007/09/ancient-pig-dna-and-the-neolithic-transition/


It appears that domesticated swine came into Europe with Neolithic farmers. Interestingly, the archaeological sequences suggest that these Near Eastern lineages were replaced by indigenous wild boar which were quickly domesticated.

Angela
06-07-14, 18:57
Thank you for that excellent summary of the situation, Angela. This is definitely one of those papers where the facts don't fit the conclusions. The authors state that "Our results reveal that the current diversity and distribution of hg H were largely established by the Mid-Neolithic (~4000 BC), but with substantial genetic contributions from subsequent pan-European cultures such as the Bell Beakers expanding out of Iberia in the Late Neolithic (~2800 BC)." However, that simply isn't the case, as is noted in the paper as well as in various other sources. 6000 years ago, H was less common than it later became, and those examples of H that have been found from 4000 BC and earlier show a substantially different prevalence of subclades than what developed later. And we seem to agree that there's a problem with trying to use the results from one locale as a proxy for all of Europe, since the results in that locale are not in fact representative of Europe. There's a lot wrong with this paper, although I don't have time at the moment to address it all in detail. I just wish I had been able to see the paper before I started this thread on the basis of curiosity about the abstract, since I really don't think this paper should be taken too seriously.

Thanks for the kind words, Aberdeen. I just thought that before even getting to the merits of the conclusions, we needed to know precisely what, in fact, the authors were claiming in terms of the populating of Europe from a mtDNA "H" perspective.

LeBrok
06-07-14, 19:36
Thanks for the kind words, Aberdeen. I just thought that before even getting to the merits of the conclusions, we needed to know precisely what, in fact, the authors were claiming in terms of the populating of Europe from a mtDNA "H" perspective.
It was a great summary Angela, I gladly enjoyed it not having time to read research papers these days.

epoch
06-07-14, 22:33
GailT, who posts at Anthrogenica makes the point far better than I can, and coming from a far better grasp of mtDNA than I have:

"My guess is that there were not just a few waves of migration - the concept of early farmers partially replacing hunter-gatherers, and late Neolithic immigrants partially replacing earlier farmers is much too simplistic. There were very likely many waves of migration that varied in different parts of Europe. For example, much of the U5 found in Finland today is not a Paleolithic remnant, rather, a large part of it appears to be a Neolithic migration that arrived via eastern Europe. So I think we need many more ancient samples from a much wider region to even begin to understand the complexity of past migrations. The new studies from Brandt et al and Bollinger et al. are fascinating, but they focus on a small geographical area. We need that level of analysis from many different areas."

Until very recently - In the Netherlands that was 1980 - wild areas in Europe were drained and cultivated. Or, in the case of undrainable lands, tranformed to meadows for cattle. Agriculture and the spread into wild areas are intimately connected. That means that agricultural cultures are *constantly* colonizing. Taking my country as an example again, one third of the Netherlands consisted of peatbogs. They all are transformed to agricultural grounds, apart from small remnants. One could call this something like "migration within".

Couldn't it be that *some* changes in haplogroups somehow reflect the uptake of people met during this "migration within"?




I would just add that given Lazaridis et al and now the Paschou et al paper, it seems to more probable that although there might have been different pulses of the Neolithic into Europe, the autosomal signature shows a departure for Europe from the Levant area primarily by sea and then splitting and differentiation after that, but further testing will clarify matters.

Could you elaborate on this?

epoch
06-07-14, 22:49
Indeed, that's why the Haak group has been mining this area for years, not just in this paper but in the prior Haak et al papers and the Brandt et al paper referred to by Epoch.

I just don't think that changes the fact that, setting aside any other issues with the paper, such as the number of ancient samples per culture, and their determinations concerning drift just as two examples, it is injudicious to extrapolate from this area to make broad generalizations about all of Europe.

I understand this hesitation, mind you. The answer to it is the answer that has been given on many occasions: "We need more data" :)

Angela
07-07-14, 02:19
epoch;435031]Until very recently - In the Netherlands that was 1980 - wild areas in Europe were drained and cultivated. Or, in the case of undrainable lands, tranformed to meadows for cattle. Agriculture and the spread into wild areas are intimately connected. That means that agricultural cultures are *constantly* colonizing. Taking my country as an example again, one third of the Netherlands consisted of peatbogs. They all are transformed to agricultural grounds, apart from small remnants. One could call this something like "migration within".

Couldn't it be that *some* changes in haplogroups somehow reflect the uptake of people met during this "migration within"?

I absolutely think that's possible. What I think is a mistake is to imagine that the present day haplogroup distribution in European countries, especially in terms of yDNA, but even to some extent for mtDNA, is an exact mirror of what it was even after the arrival of the Indo-Europeans, for example. When climate change and the Huns pushed the Germanic tribes west and south, tribes that ultimately toppled the Empire, there had to have been some "mixing" of genes. Then the Slavic invasions further "stirred the pot". (That's not to say that the affect was the same for all countries. I don't see much evidence for influence in Italy from the Slavic tribes, and even the "Germanic" influence seems pretty minor if we are to go by the y-DNA sub-groups that I think most people would see as markers of those migrations, like R1b U106 and I1.)

Industrialization is another huge factor. Just in terms of Italy, which I know best, there has been an extremely large internal migration from the areas south of Rome to the north, and in particular to the Lombardia (Milano), Piemonte (Torino), and Liguria (Genova) triangle, as these were the areas that were first industrialized. The majority of that migration took place starting in the mid 1950's and continues to this day, but there was migration to Genova in the late 19th century, for example, from the hinterlands of Liguria itself, from the Veneto, and from northern Toscana. *

I think it's probable that such "internal" migrations, particularly of women, would have taken place in Neolithic Europe as well, and may go some way toward explaining the changes from the early Neolithic to the Middle Neolithic which Brotherton et al purport to see, in addition differentiation within Europe, and to drift.




Could you elaborate on this?

The comment I made to the effect that, "although there might have been different pulses of the Neolithic into Europe, the genetic signature shows a departure for Europe from the Levant area primarily by sea and then splitting and differentiation after that" is based on my interpretation of the findings of Paschou et al. (and Lazaridis et al) FrankN and I are going to have to agree to disagree about it. :) Of course, a paper could come out tomorrow showing there were two autosomally different waves of the Neolithic, one from the Levant and one from Anatolia; it's just that I don't think that's what Paschou shows.

You can see the discussion here:

http://www.eupedia.com/forum/threads/30129-The-Mediterranean-route-into-Europe-%28Paschou-et-al-2014%29?highlight=Paschou

*Edited to add "in the late 19th century".

Aberdeen
12-07-14, 17:49
I think this thread has gone badly off course. Is there anyone who's actually read the study I originally referenced and who has an opinion as to whether it makes sense?

FrankN
13-07-14, 09:52
I think this thread has gone badly off course. Is there anyone who's actually read the study I originally referenced and who has an opinion as to whether it makes sense?
While I understand that some of the discussion has gone off topic, and actually myself proposed to shift some posts to a new thread, as you did, I feel you done a bit too much of re-arranging. For the points I was to bring forward (and I have been working on more than one answer to the original question posed), the following posts that I feel are pertinent to the discussion would greatly help to understand my arguments (numbers relate to the new thread):

#8 Flint mining as essential part of Neolithic interchange in Central Europe
#17 Background on the LBK culture that, according to Brotherton, has been completely overturned in the middle & late Neolithic
#20 MK as a possible culprit (I plan to go into further detail on that)
#22 Epoch on Swifterblade culture (I intend to further comment on the Swifterblade-MK relation outlined in his links)
#29 If it is possible for you to split the post, the second half should go here, the first half is fine where it is now. Otherwise, I might just repost the second half here.
#30 Epoch on Dutch land losses (I intend to look into the issue of population pressure from drowning lands in more detail. Short answer is - it's probably not an issue, but the point is nevertheless interesting enough for a few more paragraphs).

I also think that the housing piece may still become relevant, but at the moment I haven't progressed beyond 4,000 BC, the initial genetic turnover found by Brotherton. If I understood you correctly, your interest is more on BB/ Corded Ware, and housing might tell us something about them as well. However if you prefer to keep it outside this thread, I'll accept it (though I don't really see it in any way related to "limitations of genetic models").

Once you have shifted the above posts back, I will delete this post. If you feel they should stay in the other thread I will have quite some problems to contribute further in a meaningful way, which would be regrettable as I think the study, and the region in question are important to better understand Central European Neolithic.

Aberdeen
13-07-14, 15:00
While I understand that some of the discussion has gone off topic, and actually myself proposed to shift some posts to a new thread, as you did, I feel you done a bit too much of re-arranging. For the points I was to bring forward (and I have been working on more than one answer to the original question posed), the following posts that I feel are pertinent to the discussion would greatly help to understand my arguments (numbers relate to the new thread):

#8 Flint mining as essential part of Neolithic interchange in Central Europe
#17 Background on the LBK culture that, according to Brotherton, has been completely overturned in the middle & late Neolithic
#20 MK as a possible culprit (I plan to go into further detail on that)
#22 Epoch on Swifterblade culture (I intend to further comment on the Swifterblade-MK relation outlined in his links)
#29 If it is possible for you to split the post, the second half should go here, the first half is fine where it is now. Otherwise, I might just repost the second half here.
#30 Epoch on Dutch land losses (I intend to look into the issue of population pressure from drowning lands in more detail. Short answer is - it's probably not an issue, but the point is nevertheless interesting enough for a few more paragraphs).

I also think that the housing piece may still become relevant, but at the moment I haven't progressed beyond 4,000 BC, the initial genetic turnover found by Brotherton. If I understood you correctly, your interest is more on BB/ Corded Ware, and housing might tell us something about them as well. However if you prefer to keep it outside this thread, I'll accept it (though I don't really see it in any way related to "limitations of genetic models").

Once you have shifted the above posts back, I will delete this post. If you feel they should stay in the other thread I will have quite some problems to contribute further in a meaningful way, which would be regrettable as I think the study, and the region in question are important to better understand Central European Neolithic.

I can't move posts - only a moderator can do that. I'm just saying that my personal opinion is that the thread is now badly off topic in many respects. I originally posted a reference to this study because I only had the abstract to look at, since the rest of the paper was behind a paywall, and I was curious as to how they arrived at their conclusions. Once ebAmerican posted the whole study for us to look at, I felt that the conclusions of the study failed to match the data. While the mtDNA haplotype H was present in Europe from an early date (and was found in at least one LBK site), it had not reached its present dominance in Europe by 6000 BP, which is what the authors indicated. And while subclade H1 was present by then, H5 seemed to still have been the most common subclade at that point. And I don't see any point in trying to decide how the present structure of mtDNA H came to be established 6000 years ago when that is not in fact the case.

As for housing styles, you would have to show consistency of styles over a very long period of time for housing styles to be relevant. Showing that some barns that currently exist follow a style somewhat similar to a really ancient building doesn't, in itself, prove continuity. Nor does it address the question of whether different ethnic or tribal groups may have used a similar style of building construction and/or layout. There have been a lot of population movements in that part of the world. And I'm not sure what housing styles in one part of Europe have to do with the present predominance of mtDNA H, particularly H1 and H3, in the whole of Europe.

FrankN
13-07-14, 17:42
The Brotherton study has yielded several remarkable results that, however, could definitely have been presented in a clearer and less misunderstandable way. In order to create an overview on their actual findings, I try to put together a cultural timeline, based on the cultural epochs used by them, and describe their findings (including those only documented in the annex) within that timeline. I furthermore add Wikipedia information on the cultures themselves, and on the development of metallurgy in the region as described here:
http://www.eupedia.com/forum/threads/29077-New-map-of-the-diffusion-of-the-Copper-Age-in-Europe?p=432246&viewfull=1#post432246



LBK (5,500 - 4,800 BC): The share of mtDNA H increases from virtually zero to 20%. In the PCA analysis, the LBK sample clusters together with present-day populations from the Near East and the Caucasus, especially Lebanese and Ossetians. Brotherton et. al interpret this as confirmation for the spread of farming via substantial immigration from these regions, and not just by cultural exchange as supposed by other theories.
Rössen (4,800 - 4,200 BC): According to their Supplementary Material, the phase actually contains several, partly temporally overlapping or regionally distinct cultures. Aside from Rössen, there is the ornamented ware (STK) culture that precedes Rössen, and the Gatersleben culture as regional phenomenon in the late Rössen phase. Increasing regional differentiation during this phase has also been observed elsewhere, e.g. the Großgarlach, Hinkelstein, Bischheim and early Michelsberg cultures that have preceded or paralleled the Rössen culture along the Rhine. The beginning of this period is marked by the construction of several large earthworks with astronomical functions such as the Goseck circle (http://en.wikipedia.org/wiki/Goseck_circle), a phenomenon that originated in the Danubian Lengyel culture and apparently spread along the Elbe. Along the Rhine, Rijkholt flint (Maastricht area) is increasingly being replaced by upper Bavarian flint. This corresponds to an expansion of the Michelsberg Culture (MK) from the Paris Basin towards the Rhine, and most likely also into the upper Danube basin.
The share of mtDNA H increases to 40%. Changes in the sub-composition are not assessed. Visual inspection of the phylogenetic network suggests a relatively constant composition. Brotherton's demographic simulation yields a substantial increase in absolute mtDNA H population, within a population that is growing as well, but apparently at a lower rate. Unfortunately, they have only published their skyride plot, not the underlying data. In any case, they seem to interpret this as a sign of relative demographic success of early farmers vis-à-vis the original HG population. I personally think that additional immigration from the LBK "homeland" around the central Danube would plausibly explain both the emergence of astronomical earthworks, and at least part of the increase in mtDNA H frequency.
Schöningen (4,100 - 3,950 BC): A rather obscure culture that is neither covered in their supplementary materials nor by Wikipedia. It seems to be commonly regarded either as final phase of the Gatersleben culture, or as initial phase of the subsequent Baalberge culture. The period is nevertheless remarkable, as it shows the first evidence of copper imports from the Western Carpathians (Slovakia). The westerly adjacent Michelsberg Culture MK expands into the Scheldt and Rhine-Meuse basins, replacing the Mesolithic Swifterbank culture there, and also gains cultural influence on the Elbe-Saale region. Under influences of Elbe-Saale cultures and MK, the Mesolithic Ertebolle culture in Northern Germany, Denmark and Southern Sweden transforms into the Funnelbeaker Culture. The shift in Northern Germany is fast, in Denmark abrupt.
Brotherton et al. are apparently themselves not sure where to place this phase. For statistical purposes, they treat it as Middle Neolithic, but in their phylogenetic analysis it appears as Early Neolithic. The fact that mtDNA H frequency drops to below 20% suggests significant demographic change. However, as there are only two samples from this phase, it is hardly possible to assess whether this demographic change relates to additional gene inflow (immigration), mtDNA H gene outflow (migration to other regions, e.g. due to overpopulation), and/or a major farming crisis. The period in question marks the transition from the wet and hot Atlantikum to the cooler and dryer Boreal. The local climate may have furthermore been affected by the Littorina Transgression, i.e. the flooding of the (previously dry) Western Baltic Sea basin through the Great Belt, which started in 5,500 BC and reached its maximum extent by 2,500 BC. Brotherton's demographic simulation displays stagnation, but not decline of the absolute mtDNA H population.


As I have a football match to watch tonight, and will be visiting friends, I leave it here for the time being, but will continue next week.

LeBrok
13-07-14, 18:22
Off toppic posts were moved yesterday to this new thread: http://www.eupedia.com/forum/threads/30234-Limitations-of-current-genetic-models

I'm not sure why this new thread didn't show up on Forums main page? Sorry, I should have mentioned it in a new post here.

MOESAN
13-07-14, 18:35
Mt DNA autosomals in some regions




I precise the geographic distribution does not imply any climatic rapid selection of mt DNA-H1-

I'm aware I base that on the today distribution -

the founding survey of this thread is very unprecise concerning geography

the debate concerning mt H is veryinsteresting, even if personally I lack more detailed data about thegeographical distribution of all the mt-H subclades
-H1 I found very interesting : itsdistribution peaks among Berbers, Atlantic Europe and Finland/Estoniadoesn't seem illustrating an I-Ean East to West introgression OR wewould imagine a first vawe followed and partly erased by othersteppic people, but the presence in North Africa (of which LybianBerbers) even if exagerated maybe by founder effects, doesn't suggestthis I-Ean origin –
but on the other hand, the strongenough presence in not-I-Ean populations as Finland (18,0%) andEstonia (16,7%), finnic speaking and TOO among other finnic speakersof the Oural region (13,6%), if not typical of I-Eans, could seemweird for Neolithic peasants from the Near-East or even Anatolia -confirmation of the russian position close to finnic speakingpopulations, here too; by the fact, the Near-Eastern regions, as theCaucasus as a whole, are poor enough for mt-H1 (from 0,0% to 4,2%,the most being around Lebanon) – the BBs influence was evocatedconcerning the spreading in Atlantic and central peri-danubianregions of Europe (here at poor enough levels from 8% to 12% with alittle peak in Slovakia – but the presence among western finnicspeaking people is still a question ! -
the Eupedia map of mt-H1+H3 doesn'tchange this global sketch, in complement we can add the British Islesand Scandinavia as hotspots – in fact, the western steppes Norththe Caucasus are less poor for H1/H3 than southeastern Europeregions as Italy, Greece and most of all Albania and CarpathianRomania Moldavia (echo of Starcevo and then Cucuteni-Tripoljehere?) ; this seems excluding even more the FIRST genuineneolithic peasants from Near-Eastern and farther South than theI-Eans -
first consequence, knowing agricultureseem having been born until us by populations with a strong'south-western asian' component, we can say the H1+H3 element didn'tcome with it – maybe at a close enough time, BUT NOT withit – the strong enough density on the northern side of North Africais very confusing – one of the BBs theory (not stupid) imagining amaritime East to West road until Iberia and N-Africa requires morebasis – the Long Barrows or other megalithic people possiblecolonizations of northwestern and northern Europe doesn't explain thenorthwestern eurasian steppes and Finland presence, even if someonescould object time passed since... -
As the today Basques seem the focus ofthis grouping of mt-DNA and as the presence in Dodecad K12 of'basque' autosomals component in northern Europe and even Finland(even if the densities are not comparable in absolute) are noticeablecompared to southeastern Europe and Near-East we could imagine therethe signal of an old enough colonization by these « females » ?a survey about the respective distributions of 'south-east-farmers'EEF , 'west-hunters-gatherers' WHG and 'ANE' showed roughly a highenough WHG in northern Europe as for ANE, even if this two componentsare not parfectly parrallelic – the Spain Basques are in some wayricher for EEF + ANE (!) than the France Basques who are richer forWHG : I 'm almost sure the french Basques are the most typicallybasque, as a whole – the ANE component in Iberia is confusing butit seems the Spain Basques share there something with theCantabrians ? (whose mt DNA has something special recording someparts of northern Europe, according to some scholars, but I've notthe details) – we would need more regional data about autosomals– the ANE would recall some eastern colonizations, I-Ean or not, orboth ; I think Portugal W+S Spain WHG would be more linked tothe mt-H1+H3 complex, the N-Spain and other WHG more linked to mt-U(5the most) – U5 and ANE could be arrived in western Europefrom East just after the 9000 BC (Solutrean people from W-Asia asbelieved the old scholars? More brutal phenotypes ?) -
all that is speculations because noneof the criteria Y-DNA, mt-DNA and autsomals are evolving at the verysame speed – (even the categorizations of autosomals is cause ofinternal autosomals differences) and we have not the sufficientsamples for every place and period -


now I've the problem of elderness ofmt-H as a whole and H1-H3 in particular, and also the way taken toreach western and northern Europe : I'm tempted to think H1-H3were born by the first mt-H already present in W or S Iberia at lastMesolithic... their presence of their ancetors Haplo's in southernEurope could trace back very earlier (Paleolithic, before LGM) -
hypothesis : a) E >> Wthrough Mediterranea (middle or late Neolithic ? lateMesolithic?) – b) W >> E through N-Europe (late Neolithic ?calcholithic-bronze age?) - mt-U5 seemed the only or almost only mtDNA among the HGs of Europe – we lack dense data about theMesolithic compared to Neolithic (and yet, in this last case too thedensity is weak in fact) – is mt-U so old as Palolithic in westernEurope ? Nothing tell us for sure - Was Cro-Magnon of mt-Ustock ??? We (I only?) lack every kind of DNA from recentPaleolithic in Europe -
WHG with already some mt-H amongmt-U when mt-U was more exclusive in ANE ? Or H1-H3 ancestorsalready present at two ends of Europe in late Mesolithic and andjunction helped by Atlantic Bronze with the help of BBs boosting ???very imaginative this last one! -
whatever thehypothesis we can construct, it seems to me the « explosion »of mt-H in Europe, for me already before metal ages, as conclude somesurveys, even it at this last date new mt-H could have been arrivedthere – the surveys about the mtDNA in the post-Danubianagricultural cultures in Hungary show clearly a very bigger weight ofmt-H West the Tisza (LBK : ) compared to East the Tisza (ALPC),being the opposite for mt-J – other mtDNA haplogr's show somedifferences of distribution, but very less strong – I 'm sure theH1-H3 are not the descendants of the mt-H rare yet among theNear-East agricultors and who gave birth surely to other subclades ofmt-H –the strong presence in Finland, Scandinavia and the light butnoticeable presence among Balts and some Russians compared to thepoorness among S-E Europe confirms that – but at the same time, thestronger weight in far North seems eliminatating an I-Ean« importation » at first sight – so mt -H1/H3 is OLD inW and N Europe – the presence in N-Africa could could have beenmagnified by the BBs colonization or exploitation (even if alreadythere) + the reinforcing of this presence in Atlantic and CentralEurope at immediate pre-Bronze-Age is surely linked to BBscolonization and later to a demographic increase linked to theprogress due to previous BBs intervention in other subesquentcultures -
a W to E or E to Wsingle travel at the daybreak of metal ages is too simplistic :but males mediated colonizations more or less local or ongreat scale (1° Long Barrows and assimilated, early : 4200/4000BC ??? – 2° protoceltic or partly celtic Y-R1b-<<L11,later : 2500 BC ?) could have reinforced AND modified theprevious mt-H1/H3 geographic distributions locally, principally inCentral Europe -
so : a firstW>>E move at middle-to-last Neolithic time with a preferencefor N-coasts (what Y-Haplos?) and then at pre-metal and metal agescomplicated intrication of diverse moves, where Celts played a bigrôle, after the help send by BBs of unknown origin to me – Iadd the presence in Finland and around is older : just after theLGM -
western females anddiverse southern and eastern males + demographic encrease – thesefemale (maybe with a strong 'basque' + light 'sardinian' autosomalelement present among West-HGs, these last maybe corresponding to the« SECOND Mesolithic » trapeze microliths) mixed more thana time with male mediated autosomals from South-East('west-mediterranean' as 'sardinian' + 'east-mediterranean' and maybe'southwest-asian') and some East ('northsea' or 'northwesterneuropean' autosomal elements +others ???: more linked toproto-Celts-Italics-Germanics) –
I'm not sureEurogenes thinks mt-H1 among Trichterbecher Culture in Sweden is asign of southeastern mediterranean agricultors ? - it could verywell be an error if it were the case: I red somewhere theTrichterbecher (Funnelbeaker) culture was born in a crossing place(peri-Danmark area and great rivers around) with previous megalithic'Long-Barrows'like, Neolithic agricultors (more bovins) cultures andlater Globular Amphores cultures influence - the last one, lateneolithical, could have had I-Eans influences and had comprised someeastern influences (?) - but for me the mt-H1 elements in theTrichter-B culture of Scandinavia came more surely from themegalithical elements -the mt-H bearers population could even hadbeen there before the megalithers, being after reinforced by the mixcalled 'Trichterbecher' where finally the megalithers (LongBarrows-like, tombes à couloir) seems the impulsing element -whatever the case I recall my opinion about what I consider the'long-barrow' anthropological mix where western pre-neolithicalelements from atlantic France-Iberia dominated eastern mediterraneanelements – concerning autosomals understanding, the Eurogenesthought that the mix 'Europeans' + Near-Easterners took place on thecontinent and then was transmitted to Scandinavian by Trichterbecherpeople is not stupid – all the way it says nothing about datationof mt-H1 pr pre-H1 presence in North...
the presence inFinland and near the Baltic could be due to W >> E earlier(Ertebölle?) and later (I-Ean corded?) movements implicating diverseY-DNA bearers (among them Y-I1 people, first, and later Y-R-U106 +Y-R1a) : it would be interesting to know the estimated age ofthe mt-H1/3 in Finland an around -
Sorry for thisunprecise enough post concerning dates – it is just a try to applygood sense to very unlevel data (for time, localisation, homogeneity,sample size) = for the fun of betting -

Aberdeen
13-07-14, 21:08
The Brotherton study has yielded several remarkable results that, however, could definitely have been presented in a clearer and less misunderstandable way. In order to create an overview on their actual findings, I try to put together a cultural timeline, based on the cultural epochs used by them, and describe their findings (including those only documented in the annex) within that timeline. I furthermore add Wikipedia information on the cultures themselves, and on the development of metallurgy in the region as described here:
http://www.eupedia.com/forum/threads/29077-New-map-of-the-diffusion-of-the-Copper-Age-in-Europe?p=432246&viewfull=1#post432246



LBK (5,500 - 4,800 BC): The share of mtDNA H increases from virtually zero to 20%. In the PCA analysis, the LBK sample clusters together with present-day populations from the Near East and the Caucasus, especially Lebanese and Ossetians. Brotherton et. al interpret this as confirmation for the spread of farming via substantial immigration from these regions, and not just by cultural exchange as supposed by other theories.
Rössen (4,800 - 4,200 BC): According to their Supplementary Material, the phase actually contains several, partly temporally overlapping or regionally distinct cultures. Aside from Rössen, there is the ornamented ware (STK) culture that precedes Rössen, and the Gatersleben culture as regional phenomenon in the late Rössen phase. Increasing regional differentiation during this phase has also been observed elsewhere, e.g. the Großgarlach, Hinkelstein, Bischheim and early Michelsberg cultures that have preceded or paralleled the Rössen culture along the Rhine. The beginning of this period is marked by the construction of several large earthworks with astronomical functions such as the Goseck circle (http://en.wikipedia.org/wiki/Goseck_circle), a phenomenon that originated in the Danubian Lengyel culture and apparently spread along the Elbe. Along the Rhine, Rijkholt flint (Maastricht area) is increasingly being replaced by upper Bavarian flint. This corresponds to an expansion of the Michelsberg Culture (MK) from the Paris Basin towards the Rhine, and most likely also into the upper Danube basin.
The share of mtDNA H increases to 40%. Changes in the sub-composition are not assessed. Visual inspection of the phylogenetic network suggests a relatively constant composition. Brotherton's demographic simulation yields a substantial increase in absolute mtDNA H population, within a population that is growing as well, but apparently at a lower rate. Unfortunately, they have only published their skyride plot, not the underlying data. In any case, they seem to interpret this as a sign of relative demographic success of early farmers vis-à-vis the original HG population. I personally think that additional immigration from the LBK "homeland" around the central Danube would plausibly explain both the emergence of astronomical earthworks, and at least part of the increase in mtDNA H frequency.
Schöningen (4,100 - 3,950 BC): A rather obscure culture that is neither covered in their supplementary materials nor by Wikipedia. It seems to be commonly regarded either as final phase of the Gatersleben culture, or as initial phase of the subsequent Baalberge culture. The period is nevertheless remarkable, as it shows the first evidence of copper imports from the Western Carpathians (Slovakia). The westerly adjacent Michelsberg Culture MK expands into the Scheldt and Rhine-Meuse basins, replacing the Mesolithic Swifterbank culture there, and also gains cultural influence on the Elbe-Saale region. Under influences of Elbe-Saale cultures and MK, the Mesolithic Ertebolle culture in Northern Germany, Denmark and Southern Sweden transforms into the Funnelbeaker Culture. The shift in Northern Germany is fast, in Denmark abrupt.
Brotherton et al. are apparently themselves not sure where to place this phase. For statistical purposes, they treat it as Middle Neolithic, but in their phylogenetic analysis it appears as Early Neolithic. The fact that mtDNA H frequency drops to below 20% suggests significant demographic change. However, as there are only two samples from this phase, it is hardly possible to assess whether this demographic change relates to additional gene inflow (immigration), mtDNA H gene outflow (migration to other regions, e.g. due to overpopulation), and/or a major farming crisis. The period in question marks the transition from the wet and hot Atlantikum to the cooler and dryer Boreal. The local climate may have furthermore been affected by the Littorina Transgression, i.e. the flooding of the (previously dry) Western Baltic Sea basin through the Great Belt, which started in 5,500 BC and reached its maximum extent by 2,500 BC. Brotherton's demographic simulation displays stagnation, but not decline of the absolute mtDNA H population.


As I have a football match to watch tonight, and will be visiting friends, I leave it here for the time being, but will continue next week.

No, you are mistaken, IMO. HV may have been present in Spain and Italy during the Paleolithic, and H was definitely present in Russia, the Ukraine and Spain over 7,000 years BP, i.e. prior to Rosen. However there were a variety of mtDNA types, and U was the most common. If I look at the results reported in Ancestral Journeys, it indicates that the Rossen results show 4 out of 15 have mtDNA H (and one HV) but only one of those four is H1 and none are H3, so we don't have a pattern similar to the modern H distribution in Europe. And H is much less apparent in the large number of samples from Schoningen and Baalberge at a slightly later date, right around 4,000 B.C. or a bit later for Baalberge. So H seems to have been at least marginally present in Europe since at least the Mesolithic but only shows up in large numbers in Iberia 5000 years BP and later, and the modern mtDNA structure of Europe doesn't really seem to have developed until the Iron Age. IMO, there's no point in trying to discuss what the conclusions of this paper mean for DNA in Europe when the conclusions seem to be quite wrong.

FrankN
15-07-14, 01:51
No, you are mistaken, IMO. HV may have been present in Spain and Italy during the Paleolithic, and H was definitely present in Russia, the Ukraine and Spain over 7,000 years BP, i.e. prior to Rosen. However there were a variety of mtDNA types, and U was the most common. If I look at the results reported in Ancestral Journeys, it indicates that the Rossen results show 4 out of 15 have mtDNA H (and one HV) but only one of those four is H1 and none are H3, so we don't have a pattern similar to the modern H distribution in Europe. And H is much less apparent in the large number of samples from Schoningen and Baalberge at a slightly later date, right around 4,000 B.C. or a bit later for Baalberge. So H seems to have been at least marginally present in Europe since at least the Mesolithic but only shows up in large numbers in Iberia 5000 years BP and later, and the modern mtDNA structure of Europe doesn't really seem to have developed until the Iron Age. IMO, there's no point in trying to discuss what the conclusions of this paper mean for DNA in Europe when the conclusions seem to be quite wrong.
I was simply reporting their findings. Supplementary Fig. 3 in the Annex states clearly that none of the 13 Mesolithic samples from the region contained mtDNA H, and I recall having read an earlier paper from some of the co-authors where they already reported these results in more detail. Among the 102 LBK samples (and this is quite a sample size, better than what we have for most countries' present population!), however, 20% had mtNA H. Of course, mtDNA H must already have been present somewhere else in Europe in the Mesolthic, but those places weren't in the Elbe-Saale region. Their LBK samples date from around 7,000 BP, that rather seems to be contemporary or even older than the samples from Russia, Ukraine and Spain that you mention (I hope you realise that Rössen was about 500-1,000 years after the period in question here).

The more important point is that the mtDNA H they found in the LBK samples is hardly present any more in Central Europe.

Early Neolithic (and in particular LBK) mt genomes are either rare today (H16, H23, H26), extinct or have not yet been observed in present-day populations (H46b, H88, H89). In sharp contrast, most of the later H sub-hgs are more common in present-day European populations (e.g. hg H3, H4, H6, H7, H11, and H13)12 (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3978205/#R12),14 (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3978205/#R14)-16 (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3978205/#R16). Of the 39 haplotypes detected, only three (within the common, basal, sub-hg H1) were shared between ENE and MNE/LNE cultures. Since the observed gene diversity is high, we might expect the number of shared haplotypes within and between cultures to be low. However, since the MNE/LNE haplotypes are on different sub-hg branches from the ENE haplotypes, these patterns combined show minimal local genetic continuity over this time period.
(..)
Our genetic distance data also indicate minimal local genetic continuity between the Early Neolithic and the Mid/Late Neolithic in Central Europe (Figure 1 (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3978205/figure/F1/); Table 1 (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3978205/table/T1/)), again suggesting that Early Neolithic lineages were largely superseded during the Mid/Late Neolithic (~4100-2200 BC) in a previously unrecognised major genetic transition. This pronounced genetic changeover between Early and Mid/Late Neolithic cultures is comparable to other known major genetic transition thus far revealed by ancient DNA and coalescent simulations (between indigenous European hunter-gatherers and incoming early farmers from the Near East during the initial Meso-Neolithic transition from ~7500 BC in Central Europe).
I invite you to demonstrate why this conclusion is wrong. Distribution maps of H16, H23, H26, H46b, H88 and H89 could be of great help in this respect to show to which places in Europe the offspring of all those EEF females migrated. Oh, they didn't migrate anywhere - 66% of the EEF female gene pool just disappeared. Well, that certainly hasn't any relevance to Europe as a whole.

It would, btw, also be nice if anybody could explain to me how those other 3 LBK samples, which all carry mtDNA H1, made it from their presumed Franco-Cantabrian LGM refugium to the Middle Elbe just in time to join their distant Anatolian cousins at the LBK party.

This, btw, is another interesting finding by Brotherton et al. that so far hasn't been discussed:

Another major advantage of the temporal calibration points provided by ancient hg H mt genomes is that the data allow a relatively precise estimate of the evolutionary substitution rate for human mtDNA. The temporal dependency of evolutionary rates predicts that rate estimates measured over short timespans will be considerably higher than those using deep fossil calibrations, such as the human/chimpanzee split at ~6 million years29 (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3978205/#R29). The rate calibrated by the Neolithic and Bronze Age sequences is 2.4×10−8 substitutions/site/year (1.7 to 3.2×10−8; 95% High Posterior Density) for the entire mt genome, which is 1.45× (44.5%) higher than current estimates based on the traditional human/chimp split (e.g. 1.66×10−8 for the entire mt genome30 (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3978205/#R30) and 1.26×10−8 for the coding region31 (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3978205/#R31)). Consequently, the calibrated ‘Neolithic’ rate infers a considerably younger coalescence date for hg H (10.9 to 19.1 kya) than those previously reported (19.2 to 21.4 kya for HVSI10 (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3978205/#R10), 15.7 to 22.5 kya for the mt coding region31 (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3978205/#R31) or 14.7-22.6 kya when corrected for purifying selection30 (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3978205/#R30) .
I am anything but an expert when it comes to these kinds of calculations, but I suppose that finding is not only relevant to mtDNA H, but to most of the coalescence dates that we have been using so far.

Aberdeen
15-07-14, 04:18
IMO, you have a marvellous capacity to miss the point, FrankN. While the results from Rossen show more mtDNA H than has been found at any Euopean site previously, it also shows more mtDNA H than has been found at European sites that are many centuries later than Rossen, so those results cannot be taken as a proxy for Europe around 4,000 B.C. (and they're actually a bit earlier than that). The Rossen results can't even be taken as a proxy for Germany around 4,000 B.C., since the many samples from Schoninger, which do date from around 4,000 B.C., have much less mtDNA H, as do the somewhat later results from Baalberge. The conclusion of the study was that "Our results reveal that the current diversity and distribution of haplogroup H were largely established by the Mid Neolithic (~4000 BC), but with substantial genetic contributions from subsequent pan-European cultures such as the Bell Beakers expanding out of Iberia in the Late Neolithic (~2800 BC)." That's clearly not correct for Europe, or even for Germany as a whole. I would argue that if you actually look at the Rossen results and the modern DNA results, such a conclusion is only slightly valid for that one small set of samples from Rossen. One cannot extrapolate too much from that, since the results are not replicated in subsequent finds for Neolithic Europe.

Since we don't seem to be getting anywhere with this conversation, I will leave you to grind out your long posts, even though I personally believe that they generally obscure the point at issue in most threads where you post. But maybe that's just my perspective.

FrankN
15-07-14, 05:16
IMO, you have a marvellous capacity to miss the point, FrankN. While the results from Rossen show more mtDNA H than has been found at any Euopean site previously, it also shows more mtDNA H than has been found at European sites that are many centuries later than Rossen, so those results cannot be taken as a proxy for Europe around 4,000 B.C. (and they're actually a bit earlier than that). The Rossen results can't even be taken as a proxy for Germany around 4,000 B.C., since the many samples from Schoninger, which do date from around 4,000 B.C., have much less mtDNA H, as do the somewhat later results from Baalberge. The conclusion of the study was that "Our results reveal that the current diversity and distribution of haplogroup H were largely established by the Mid Neolithic (~4000 BC), but with substantial genetic contributions from subsequent pan-European cultures such as the Bell Beakers expanding out of Iberia in the Late Neolithic (~2800 BC)." That's clearly not correct for Europe, or even for Germany as a whole. I would argue that if you actually look at the Rossen results and the modern DNA results, such a conclusion is only slightly valid for that one small set of samples from Rossen. One cannot extrapolate too much from that, since the results are not replicated in subsequent finds for Neolithic Europe.

Since we don't seem to be getting anywhere with this conversation, I will leave you to grind out your long posts, even though I personally believe that they generally obscure the point at issue in most threads where you post. But maybe that's just my perspective.
The way the study conclusion has been formulated is obviously bullshit, and doesn't represent their findings. I actually don't understand why they are making such a statement that is discrediting an otherwise highly valuable and interesting piece of research. I still try to figure out what they actually uncovered, maybe that's the reason for long posts.

I think, the best way to rephrase their findings is:
"The genetic structure established by EEF didn't survive the climatic transition from Atlanticum to Boreal. The current diversity and distribution of haplogroup H in Central Europe were largely established by the Mid Neolithic during the 4th millennium BC, but with substantial genetic contributions from subsequent pan-European cultures such as the Bell Beakers expanding out of Iberia in the Late Neolithic (~2800 BC).".
Your comment on their Rössen findings is legitimate. As I already said before (unfortunately in a post that somebody decided to be "off topic"), all four Rössen samples come from a single gravefield, and may thus not be representative. However, their LBK samples have been taken from 3 different locations, as have been those for MNE (Schöningen / Baalberg / Salzmünde), and the above statement remains valid even if the Rössen samples are discarded.

That was obviously not what they expected to find. Otherwise, I am sure, they would have done more MNE sampling, and especially have included samples from the late 4h millennium Bernburg culture so we would have known the genetic situation before BB and Corded Ware arrived.

FrankN
17-07-14, 02:08
As Angela had asked for it - here a map of documented LBK settlements in the Elbe-Saale region (NW of map), Upper Franconia, Bohemia, Moravia and Western Silesia. The westward "bay" in the Elbe-Saale settlement zone is the Harz. The Lichtenstein cave (itself not LBK!) is just west of the Harz.
http://denkstroeme.de/heft-6/s_137-167_stock/content-pic_137-167_stock-3.jpg/@@download/src/content-pic_137-167_stock-3.jpg
Settlement concentrated on regions that are today characterised by "Steppenheide" (heath steppe), i.e. widely unforested, and covered with loess. The picture below is from Thuringia, south of the Harz.
http://www.thueringen.de/imperia/md/images/homepagethringen/tmlnu/naturschutz/steppenrasen/202_11_23_236_steppenrasen_grosses_federgras.jpg

FrankN
18-07-14, 15:28
I have looked a bit deeper in the Rössen results reported by Ancestral Journeys. For the Elbe-Saale region, ancient mtDNA is available from four locations. Two of them (Hettstedt-Oberwietderstedt, and Wittmar near Wolfenbüttel) have a >=50% share of mtDNA H/HV0 (5 out of 9 samples in Oberwiederstedt, 2 out of 4 in Wittmar). The locations are 90 km distant from each other. From each of the two other locations (Halberstadt, Esperstedt) there is only one sample available. Halberstadt is V, Esperstedt T2e. If we take all those 15 samples, we get the following distribution of mtDNA:



mtDNA HG


n

%



HV0

2

13.3 %



V

1

6.7 %



H1

1

6.7 %



H5 / H5b

2

13.3 %



other H (rare/ extinct)

2

13.3 %



Subtotal H/V/HV0


8


53.3%



U5b / K

3

20.0%



T2

2

13.3%



X2c

1

6,7%



N1a1a

1

6,7%



Total

15

100 %





I don't have an explanation for the results yet. Hettstedt-Oberwiederstedt (silver, copper) and Wittmar (salt) are historical mining towns, but that doesn't apply to Halberstadt, where one mtDNA V sample was found. I think, for the time being, we have to accept the fact that the Elbe-Saale region, btw, the so-far best sampled European region, displays elevated frequencies of mtDNA H/V/HV0 during the middle of the 5th millennium BC.

I checked the Ancestral Journeys' data for other samples from that period:


Tisza Culture (Hungary), 5,000-4.500 BC: 67% mtDNA H (2 out of 3 samples)
Trentino / South Tirol, 4.500-4,000 BC: 50% mtDNA H (1 out of 2)
St. Pau de Camp (Spain), 4,250-3,750 BC: 33% mtDNA H (1 out of 3)
Navarre (Spain), 4,185-3,185 BC: 39% mtDNA H/HV (9 out of 23)
Verteba Cave (Ukraine), 4th millennium BC: 67% mtDNA H/HV (4 out of 6)
Gokhem (Sweden), 3,500-2,500 BC: 50% mtDNA H (3 out of 6)


Opposite to what Aberdeen claims, to me this looks like quite a pan-European pattern (of course with temporal variation).

FrankN
18-07-14, 16:53
Here for comparison ancient mtDNA data for the late 6th/ early 5th millennium BC. The increase in H/HV frequency during the mid-/late 5th millennium was obviously not only a local phenomenon in the Elbe-Saale region, but also occurred in Ukraine, the central Pyrenees, and, to the extent the limited number of samples allows for conclusion, in Hungary.


Culture


Location


Time (BC)


n


hgH


%



Cardial Pottery

Spain/ Portugal (Can Sadum, Chaves)


5,500-5,000

9

3

33




Dnieper-Donets

Ukraine (Yasinovatka, Nikolskoje)

5,600-5,000

11

4

36




LBK

Morava (Vedrovice)

5,300

5

1

20




Alföld

Hungary

5,250-5,000

3


1

33




LBK

Elbe-Saale (Brotherton study)

5,500-4,800

102


~17




LBK

Elbe-Saale (Anc. Journ. data)

5,300-4,900

43

6

14




LBK

SW Germany (Stuttgart, Vaihingen, Flomborn)

5,100-4,800

6

0

0




Epi-Cardial

Spain (Avellaner Cave)

5,000

5

1

20





Interestingly, in SW Germany, home to the prototypical EEF, no mtDNA H has so far been found in LBK graves.

Edit: I checked the Hungary data. While the main article is behind a paywall, the abstract reports "a remarkable genetic discontinuity" between early Neolithic and current populations in Hungary, just as Brotherton reports for the Elbe-Saale region. The Hungarians study concludes "It is worth further investigation as a non-descendant theory, instead of a continuous population history, supporting genetic gaps between ancient and recent human populations." Maybe someone with access to the full study wants to check for details..
http://www.nature.com/jhg/journal/v56/n11/full/jhg2011103a.html

FrankN
21-07-14, 17:49
Based on the Ancestral Journeys data, I have compiled a time series on the pre-historic mtDNA structure of the Middle Elbe-Saale region for the periods investigated by Brotherton. Note that Ancestral Journeys counts close relatives, i.e. siblings or mother & child found in the same grave, as a single sample. As such, the number of samples for each period tends to be slightly lower than the numbers given in the Brotherton study, which appears to count each individual. Note also that in several cases, partially defective DNA did not allow to identify the specific subclade of mtDNA H.




Rössen

Schöningen

Baalberge

Salzmünde

Bernburg

Corded_Ware


Bell_Beaker


Unetice



Locations


4

1

5

1

1

6

7

6




mtDNA HG

n

%

n

%

n

%


n

%

n

%

n

%

n

%

n

%



HV0

2

13,3%

1

4,3%

1

5,9%

1

5.0%




1

2,7%



2

5,0%



V

1

6,7%





1

5.0%


1

7,7%





1

2,5%



H1

1

6,7%

2

8,7%

1

5,9%



1

7,7%

1

2,7%

2

7,7%





H3







2

10.0%






2

7,7%

2

5,0%



H5 / H5b

2

13,3%





1

5.0%


1

7,7%

1

2,7%

2

7,7%





other H (rare/ extinct)

2

13,3%

2

8,7%

1

5,9%





4

10,8%

2

7,7%

5

12,5%



H (no subclade)





3

17,6%

2

10.0%


1

7,7%

4

10,8%

4

15,4%

2

5,0%



Subtotal

8

53,3%

5

21,7%

6

35,3%

7

35.0%


4

30,8%

11

29,7%

12

46,2%

12

30,0%



U / K

3

20,0%

8

34,8%

2

11,8%

5

25.0%


6

46,2%

14

37,8%

8

30,8%

13

32,5%



I











1

2,7%



4

10,0%



J



3

13,0%

1

5,9%

4

20.0%



2

5,4%

1

3,8%

1

2,5%



T

2

13,3%

3

13,0%

5

29,4%

1

5.0%

1

7,7%

6

16,2%

3

11,5%

6

15,0%



X

1

6,7%

1

4,3%

2

11,8%

1

5,0%

1

7,7%

2

5,4%



2

5,0%



W



2

8,7%





1

7,7%

1

2,7%

2

7,7%

2

5,0%



N1a1a

1

6,7%

1

4,3%

1

5,9%

2

10.0%











Total

15

100%

23

100%

17

100%

20

100%

13

100%

37

100%

26

100%

40

100%



Results on the Schöningen and Salzmünde cultures need to be treated with some caution, They relate to only one location, the Salzmünde plateau, which is characterised by specific, rather unusual burial habits. More on that in a subsequent post.

Let me furthermore add that Corded Ware and Bell Beaker samples in many cases stem from the same grave field and period, and have been assigned to the respective culture based on the specific artefacts that accompanied each burial. To a lesser extent, this also applies to Bell Beaker vs. Unetice. As such, the obvious genetic differences should be interpreted as signals of two separate cultures that simultaneously lived in the Elbe-Saale region. Their coexistence appears to have been relatively peaceful - in several cases, both cultures used existing megalithic graves from the Bernburg culture, and left older burials undisturbed. Brotherton's study clearly identifies the Bell Beakers as genetically Iberian that immigrated into the already existing Corded Ware / Globular Amphora culture. How sizeable that immigration has been is difficult to estimate. Overall, the Unetice mtDNA structure is more resembling the Corded Ware than the Bell Beaker structure, but certain BB lineages (e.g. H4) continue into the Unetice culture.

The rarer H subclades for Corded Ware are H10e, H2a1, H6a1a and H4a1. H4a1 was a female from the period of CW/BB coexistence that shares all SNP markers with a somewhat younger BB sample from the same site. The other rare BB subclade is H13a1a2c. Rarer subclades for Unetice are H2a1a3, H4a1a1a2, H7h, H11a, and H82a. The Unetice H4a1a1a2 appears to be a descendant of the CW/BB H4a1 individuals (identical basic SNP markers, plus a few more recent mutations). It was found in the Eulau, grave some 100 km SE of Quedlinburg, where those other two H4a1 individuals were buried.

Edit: Table corrected for Salzmünde. To maintain consistency with Brotherton's timeline, only Bell Beaker data from 2,500 BC onwards is included. An older BB grave (2,600-2,500 BC) from Kromsdorf / Thüringen shows a quite different genetic structure that is more in line with Corded Ware and, most notably, doesn't include mtDNA H. The respective study also reports a genetic break between the early and late Neolithic, in this case concerning sub-groups of mtDNA U5. The study identifies 8 agro-pastoralist, and 4 hunter-gatherer sub-groups of U5 that have been observed in ancient DNA but are extinct today. Outside the basal clades, only one sub-group of EEF U5 is still observed in present populations.
http://www.academia.edu/1596369/Emerging_genetic_patterns_of_the_European_Neolithi c_perspectives_from_a_Late_Neolithic_Bell_Beaker_b urial_site_in_Germany

FrankN
21-07-14, 19:11
As addendum to my previous post:
mtDNA T up to Schöningen is exclusively T2; this also applies to the LBK samples from the region. Baalberge includes a single T1, Salzmünde and Bernburg are again excusively T2. Corded Ware has two T1 (5.4% of total) vs. four T2, for Bell Beakers it is 2:2. Among the Unetice samples there is only a single T1.

In looking at the T data, I have realised that the table in my previous post may miss a few samples. Please wait with interpretation and comments until I have checked and updated the table.

Sile
21-07-14, 20:38
As addendum to my previous post:
mtDNA T up to Schöningen is exclusively T2; this also applies to the LBK samples from the region. Baalberge includes a single T1, Salzmünde and Bernburg are again excusively T2. Corded Ware has two T1 (5.4% of total) vs. four T2, for Bell Beakers it is 2:2. Among the Unetice samples there is only a single T1.

In looking at the T data, I have realised that the table in my previous post may miss a few samples. Please wait with interpretation and comments until I have checked and updated the table.

I see no correlation between Schöningen and the other 2 highly % T2 mtdna areas of North-east italy ( 19% ) and coastal Romania ( 14% )...............is there one?

what was the german area! , the ancient WHG basin for deer hunters ?...................I thought the basin was near hamburg

FrankN
21-07-14, 22:58
I see no correlation between Schöningen and the other 2 highly % T2 mtdna areas of North-east italy ( 19% ) and coastal Romania ( 14% )...............is there one?

what was the german area! , the ancient WHG basin for deer hunters ?...................I thought the basin was near hamburg
Well, the ancient German WHG basins are now called North Sea, but the surrounding hills were in fact close to Hamburg (and just where I grew up) - the Ahrensburg culture. But that was some 5,000 years before the time in question here.

mtDNA T has apparently survived the last Ice Age somewhere near the Black Sea. Maciamo speaks about an Antatolian refuge; to me the distribution maps point more towards the Bessarabia / Crimea area (possibly lands a bit further south that in the meantime have been flooded). The earliest ancient mtDNA T dates to around 5,000 BC, and has been found in Russia, Ukraine, the Avellaner Cave in the Spanish Pyrenees, and LBK settlements in the Czech Republic, the Saale-Elbe region, and SW Germany (Stuttgart). Russian and Ukrainian samples contained both T1 and T2, the samples from further west were always T2.
T2 is apparently linked to the spread of farming. It may have been first distributed by Mesolithic HGs across Europe, to be "picked up" by these early farmers, or it was from the beginning part of the EEF gene pool. In the second case, it has almost certainly expanded along the Danube, and LBK farmers carried it into the Elbe-Saale region and SW Germany (Problem here: How did it get into the Avellaner cave?). T2 is still quite frequent in the LBK "homeland" (the Budapest-Szombathely-Vienna triangle), and there are dozens of possibilities how it could have reached North-East Italy from there (Baden Culture, Illyrians, Celts, Romans, Goths, etc.).

The much more interesting question is about the spread of T1, which so far hasn't been found among EEF, but is typically associated with Steppe cultures and the spread of Indo-Europeans. Since I assumed somebody might anyway ask for the T1/ T2 relation to speculate on IE incursions, I prepared and posted the respective count in advance. The bottom lines here are:

T1 isn't present in large enough quantities to base much conclusions on its frequencies over time;
The first appearance during the Baalberge Culture (early 4th millennium BC) predates the commonly assumed time line of IE expansion into Central Europe;
Corded ware isn't any more likely to have spread T1 into the Elbe-Saale region than are the Bell Beakers, or population inflow down the Elbe from Bohemia / Moravia / Hungary.

FrankN
22-07-14, 02:55
While I am still reading more on Salzmünde, here already a video introduction:

http://www.youtube.com/watch?v=0k2BZjn1hzw
More videos on the excavations and subsequent analysis:
http://www.sciencemovies.de/en/08_was_it_murder?c=3

The burial practice appears less unusual as initially assumed - it is now also reported from a large Boleraz / Baden culture settlement at Lake Balaton in Hungary:
http://www.academia.edu/2169947/Halle_2012_Oktober_18_20_Salzmunde_-_rule_or_exception_T._Horvath_Intrasite_human_buri als_at_the_site_of_Balatonoszod-Temetoi_dulo

FrankN
24-07-14, 05:38
Apparently, while nobody has anything to comment, people are still reading, so I continue with putting together some puzzle pieces. Whether they get us closer to solving the question why, as Brotherton's study has found out, most of the early Central European farmers died out, and who replaced them when - I don't know yet.

The European Neolithic experienced massive climate change. While after the end of the Ice Age, the climate had been similar to today, around 8,000 BC Europe got warmer and moister. By around 7,300 BC, average temperatures in Central and Northern Europe were some 0.5-2.5 °C higher than today. It is no coincidence that this climate optimum marks the beginning of the agricultural expansion from the Near East into Europe. Around 6,200 BC, the climate got markedly colder, but returned to the previous temperature levels around 6,000 BC. Furthermore, glacier melt had resulted in substantial sea level rise. Reduced glacier weight caused the underlying areas to rise, while those further away lost elevation. Combination of both factors resulted in flooding of the North Sea, the coastline of which reached today's West and East Frisian islands by around 6,000 BC. This also resulted in the Central European climate becoming more maritime, i.e. with milder winters and relatively humid springs and autumns. Nevertheless, overall higher temperatures are believed to have resulted in relatively long, hot summers, so the climate in the Middle-Elbe Saale region would have been more "Mediterranean", possibly comparable to today's climate in Hungary and Serbia. This second warm phase corresponds to the LBK expansion into Central Europe.

The northern periphery of the Elbe-Saale region at that time looked quite different from today. Melt water from Scandinavian glaciers had created a giant freshwater lake, the Ancylus Lake, in the area of today's Baltic Sea. A land bridge extended from East Holstein via the Danish isles to Southern Sweden. Ancylus Lake was connected to the North Sea via the Svea river, Lake Vanern and the Göta river. Motala, where a 6,000 BC grave of HGs has been found (Lazarides' NHG, 6x mtDNA U5a*, plus one U2e1) is located slightly south of the eastern end of this outlet.
http://upload.wikimedia.org/wikipedia/commons/thumb/5/5f/Baltic_History_7500-BC.svg/353px-Baltic_History_7500-BC.svg.pnghttp://upload.wikimedia.org/wikipedia/commons/thumb/e/e5/Baltic_History_5000-BC.svg/353px-Baltic_History_5000-BC.svg.png

During the 6th millennium BC, further sea-level rise caused the land bridge to break, gradually connecting the Ancylus lake to the North Sea - first as brackwater lake (Litorina Sea), then gradually developing into today's Baltic Sea. Recent investigations have clarified the timing and sequence of this transgression:http://www2008.io-warnemuende.de/documents/mebe67_2006-roessler.pdf
New cores were drilled west and east of the Darss Sill, the Mecklenburg Bay and the Arkona Basin, to investigate the transition from the fresh water conditions of the Ancylus Lake to the marine-brackish conditions of the Littorina Sea in a high spatial and temporal resolution.In general, Arkona Basin sediments display more abrupt shifts in the proxy parameter at the transition from Ancylus Lake to Littorina Sea stage material than those from the Mecklenburg Bay. Radiocarbon dating on calcareous material indicates that marine waters entered the Mecklenburg Bay first at c. 7,500 14C yr BP and the Arkona Basin approximately 1,000 years later at c. 6,500 14C yr BP. These results therefore suggest a transgression pathway via the Great Belt into the Mecklenburg Bay and then into the Arkona Basin.
In a yet not fully understood way, flooding of the North and Baltic Sea, and associated changes in Atlantic currents, had profound effects on the North-West European climate. A short global cooling around 5,200-5,100 BC is observable in Greenland ice cores and the Near East, but not in Scandinavia. It may, however, have affected areas further south. There is indication of late LBK societies as far apart as the Paris Basin, the middle Rhine, Saxony and Upper Austria reverting to ritualised cannibalism by the end of the 6th millennium, interpreted as symptom of a deep crisis that shook societies during the late LBK period.http://www.academia.edu/1403683/Cannibalism_in_the_Linear_Pottery_culture_at_Herxh eim_Palatinate_Germany_

Following a short recovery, temperatures on the northern hemisphere started to gradually decline from around 4,700 BC on due to a slight tilt in the earth axis. In addition, major volcanic eruptions (e.g. Mt. Hudson 4,750 BC, Kikai Caldera 4,400 BC) may have played a role. Some authors place the transition from the Atlantic to the Subboreal climate period here, others at 4,000 or even 3,710 BC. This again is related to obvious strong regional differentiation. Scandinavia, e.g., became only slightly cooler but substantially dryer after 4,800 BC (though volcanic winters around 4,400 are apparent), while Alpine glaciers start to grow, and sediment analysis from the upper and middle Vistula basins indicates a substantial increase in flooding frequency and intensity between 4,650 and 4,050 BC. This generally corresponds to the Central European archaeological record of post-LBK cultures (Rössen, Lengyel, Michelsberg etc.) having retreated from the river areas towards hill settlements. Forest coverage gradually decreases, possibly due to intensified pastoral and agricultural use.http://hol.sagepub.com/content/5/1/34.abstract

Around 4,100 BC, Scandinavia gets much cooler (though still 1°C warmer than today) and slightly wetter, and Central Europe warmer and dryer again. Possibly triggered by two major volcanic eruptions (Masaya, Pago), around the turn of the 4th millennium Central Europe enters a cool and dry phase that lasts for around 300 years, and corresponds to massive deforestation. Swiss glaciers grow to an extent similar to the 16/17th century "little ice-age".

By around 3,650 BC, Central Europe and Scandinavia revert to a warmer and wetter climate. The African humid period in the Sahara, however, comes to an end, and the Aegean encounters one-hundred years of drought. While climatic patterns apparently vary, it is tempting to speculate that this drought could also have affected the northern Black Sea, and played a role in the replacement of the Dnieper-Donetsk culture by the Yamna culture around 3,500 BC.
The last major climate change in the period in question here was the Piora cold period between 3,200 and 2,900 BC. Alpine glaciers grew suddenly and massively, and the Eifel (NW Germany) saw a sudden drop of average temperature by 1°C, coupled with milder winters and increasing precipitation. Flooding frequency on the Vistula increased. Elsewhere, the water level of the Dead Sea rose by 100m, some historians also link the fall of Uruk in Mesopotamia to this climatic anomaly. Scandinavia again appears to have been hardly affected. Note that the Salzmünde culture, with its specific burial practices that may have included human sacrifice and ritual cannibalism falls into this period. http://www.eupedia.com/forum/newreply.php?do=postreply&t=30209

After the end of the Piora cold phase, Central Europe seems to have enjoyed a relatively stable climate similar, though slightly warmer to the one today that lasted until around 1,800 BC, with temperatures gradually decreasing and precicipation / flooding increasing after 2,500 BC.
The diagram below displays cyclical temperature changes in the Atlantic (Bond cycles), the red is the temperature read from Greenland ice core. Yellow lines mark the different cycles, but they also correspond pretty well to our understanding of historic phase. Note the Greenland temperature drops around 5,200 BC (LBK crisis), 3,700BC (Aegean drought) and 3,200 BC (Piora cold period).
http://www.skepticalscience.com/pics/Bond-events2.png

Further reading (nice diagrams):http://www.dandebat.dk/eng-klima7.htm
http://www.academia.edu/3584928/Mensch_Klima_und_Landschaft_in_Mitteleuropa_in_Ver gangenheit_und_Zukunft

MOESAN
31-07-14, 14:41
I have looked a bit deeper in the Rössen results reported by Ancestral Journeys. For the Elbe-Saale region, ancient mtDNA is available from four locations. Two of them (Hettstedt-Oberwietderstedt, and Wittmar near Wolfenbüttel) have a >=50% share of mtDNA H/HV0 (5 out of 9 samples in Oberwiederstedt, 2 out of 4 in Wittmar). The locations are 90 km distant from each other. From each of the two other locations (Halberstadt, Esperstedt) there is only one sample available. Halberstadt is V, Esperstedt T2e. If we take all those 15 samples, we get the following distribution of mtDNA:



mtDNA HG

n
%


HV0
2
13.3 %


V
1
6.7 %


H1
1
6.7 %


H5 / H5b
2
13.3 %


other H (rare/ extinct)
2
13.3 %


Subtotal H/V/HV0

8

53.3%


U5b / K
3
20.0%


T2
2
13.3%


X2c
1
6,7%


N1a1a
1
6,7%


Total
15
100 %




I don't have an explanation for the results yet. Hettstedt-Oberwiederstedt (silver, copper) and Wittmar (salt) are historical mining towns, but that doesn't apply to Halberstadt, where one mtDNA V sample was found. I think, for the time being, we have to accept the fact that the Elbe-Saale region, btw, the so-far best sampled European region, displays elevated frequencies of mtDNA H/V/HV0 during the middle of the 5th millennium BC.

I checked the Ancestral Journeys' data for other samples from that period:


Tisza Culture (Hungary), 5,000-4.500 BC: 67% mtDNA H (2 out of 3 samples)
Trentino / South Tirol, 4.500-4,000 BC: 50% mtDNA H (1 out of 2)
St. Pau de Camp (Spain), 4,250-3,750 BC: 33% mtDNA H (1 out of 3)
Navarre (Spain), 4,185-3,185 BC: 39% mtDNA H/HV (9 out of 23)
Verteba Cave (Ukraine), 4th millennium BC: 67% mtDNA H/HV (4 out of 6)
Gokhem (Sweden), 3,500-2,500 BC: 50% mtDNA H (3 out of 6)


Opposite to what Aberdeen claims, to me this looks like quite a pan-European pattern (of course with temporal variation).

the problem is that
1- a lot of places lack serious sample
2- we're mixing here in the same bag different mt DNA H with different points of diffusion even if their far origin is the same (as for all Europeans!) - a survey about the current mt H in Europe and Eurasia shows a big difference in within distributions of mt-H

FrankN
06-08-14, 06:41
The Brotherton study is actually a sequel, and, as all sequels, difficult to understand if you have missed the previous part (and it is not recalled in the introduction). In 2012, Christine Adler, one of Haak's students and co-author of the Brotherton paper, had amended the set of previously known ancient DNA from the Elbe-Saale region, and compared the mtDNA structure of LBK, Rössen, BB, Corded Ware and Unetice samples with the structure found today in Central Europe (Germans, Poles, Austrians, Swiss, Czech?, Slovaks?, Hungarians?). She concluded:
http://www.academia.edu/3306830/Ancient_DNA_studies_of_human_evolution._Adler._201 2._PhD_thesis

Population continuity from the introduction of agriculture in the Early Neolithic until today was found to be an unlikely model of demographic history (..) Internal population changes in Europe between the Early Neolithic and Late Neolithic/Early Bronze Age appear to have contributed substantially to the population structure of extant Central Europeans, as all the Late Neolithic/Early Bronze Agecultures examined (Bell Beakers, Corded Ware or Unetice) were found to be more
likely ancestors of modern Central Europeans than either of the Early Neolithic cultures (LBK and Rössen) investigated. Haplogroup distributions suggest that Palaeolithic mtDNA haplogroups which were infrequent in the Early Neolithic, such as haplogroups H and U, became more frequent during the Late Neolithic. (..)
The frequency of haplogroup N1a decreases (..) from 25% among the LBK and 10% among the Rössen culture to 0% in the Late Neolithic/Early Bronze Age cultures The frequency of N1a haplogroup remains extremely rare (0.2%) in modern Central European populations.

The study had two shortcomings:

mtDNA H had not been broken down into sub-clades. For its high overall share among Bell Beakers as well as in extant populations, statistical models pointed at the Bell Beakers being the most important contribution to Central Europe's current population structure, a result that contradicts the archaeological record.
The middle Neolithic, i.e. the 3rd millennium BC, had not been covered. This made it impossible to determine when and in which stages the EEF (LBK) mtDNA structure had been so profoundly overturned.

The Brotherton study has obviously aimed at overcoming these shortcomings, and delivered the following results:


Bell Beakers have contributed to the extant Central European mtDNA diversity, but weren't a major factor. More specifically, they are not required to explain the substantial presence of H1, H3, and H5 in current Central Europeans, though they contributed H5a.
Transition to the extant Central European mtDNA structure, i.e. of dominating mtDNA H sub-clades, occurred already in the Rössen (H5), Baalberge (H7) and Salzmünde (H3) cultures. Corded Ware contributed H1b (secondary peak in Elbe-Saale region), H2 (peak in Germany reported elsewhere, but not seen in the Brotherton data), H4 and H6. Basal H1 had already been introduced by the LBK. In that sense, I think for mtDNA H it is fair to state that the main shift occurred in the mid-Neolithic, with further contributions from BB and Corded Ware.

Looking at other major mtDNA hgs:

J was introduced by the LBK. Fluctuations appear to be mostly statistical noise, overall its share appears to have remained rather constant.
I was introduced by Corded Ware.
K and HV/V have already been present during the LBK. They retreated to their current frequencies of 5-10% during the Schöningen (HV/V) and Baalberge (K) cultures, respectively.
N1a1a, which made up 12% of unrelated LBK samples, disappears together with the Salzmünde culture (3,300-3,100 BC).
T2 (17% share in LBK) retreats to current levels with the Salzmünde culture. T1 first appears with the Baalberge culture.
(Mesolithic) U5b was boosted with the post-Nordic TRB Bernburg culture, but at its current level with Corded Ware. Corded Ware newly introduced U5a and U4. In this context, it should be noted that local archaeologists interprete Bernburg, Corded Ware and Bell Beakers as overlapping, partly contemporary cultures that are distinguished by burial style and the ceramics found therein. During the early copper age, the Elbe-Saale region was literally multi-cultural. Differing DNA structures reflect this and shouldn't be interpreted as population replacement (though the subsequent Unetice culture allows some judgement on which cultures/ genes ultimately prevailed).
W was already present in the LBK (5%) and has been regularly found thereafter if sampling size was large enough.
X wasn't present in LBK samples, but has from the Rössen culture onwards been more or less constantly found at low levels.


So, let's do the math on all other hgs based on Maciamo's list of current mtDNA frequencies per country (for simplification, I restrict myself to Germany):

LBK: HV/V (4.5%) + J (9%) + K (6.6%) + T2 (7.8%) + W (1.7) = 29.6%

Mid-Neolithic: T1 (2.8%) + U2/5b/8 (6%?) + X (1.3%) = 10.1%
Corded ware: I (2.1%) + U4 (2.9%) + U5a (4%?) = 9%
Unallocated: L (0.3%) + U3 (1.1%) + other U (0.6%) + others (4.5%) = 6.5%

In fact, it seems the paper has played down Corded Ware a bit too much. Still, the conclusion remains that the Neolithic revolution, even when H1 is added to the figure above, can explain less than 40% of Germany's current genetic diversity. A lot of things have happened afterwards, especially during the middle Neolithic that so far has received little attention in this respect.

FrankN
10-08-14, 14:51
I have come across a highly interesting paper that may help to shed some more light on Brotherton's findings:
http://www.nature.com/ncomms/2013/131001/ncomms3486/full/ncomms3486.html

We show that, in contrast to the steady population growth usually assumed, the introduction of agriculture into Europe was followed by a boom-and-bust pattern in the density of regional populations. We demonstrate that summed calibrated radiocarbon date distributions and simulation can be used to test the significance of these demographic booms and busts in the context of uncertainty in the radiocarbon date calibration curve and archaeological sampling. We report these results for Central and Northwest Europe between 8,000 and 4,000 cal. BP and investigate the relationship between these patterns and climate. However, we find no evidence to support a relationship. Our results thus suggest that the demographic patterns may have arisen from endogenous causes, although this remains speculative.

Here are the demographic trends they have produced for various regions (you might want to check the study itself for larger pictures, the Supplementary Materials include higher-resolution graphs for each region). The dotted lines display a hypothetical steady growth, blue arrows mark the onset of farming. Significant (p>0.95) deviations from the "steady growth scenario" are marked in red (overpopulation) and light blue (underpopulation), respectively.
https://forwhattheywereweare.files.wordpress.com/2014/02/b3d40-regdemographyannotated.png

While I highly appreciate their work, I have a few, minor technical issues with it, which I feel need to be considered when looking at their diagrams:

The population estimates have been extrapolated from archaeological findings. There is reason to assume that HGs and EFs concentrate on the coasts (fishery, trade contact, fertile marshland etc.). Considerable amount of coastland, especially along the North Sea and the English Channel, has of course been flooded in the meantime, and submarine archaeology is still in its beginnings. As such, I think that their diagrams for Jutland, possibly also Northern Germany, Wessex/Sussex, the Danish Isles and Paris Basin/ Normandy, tend to underestimate population densities before 5,000-4,500 BC. Early increases prior to the onset of farming may reflect re-location from flooded areas to higher ground that is better covered archaeologically.
Some of their sub-regions could have been delineated better. Specifically, "Central Germany", which is essentially the Elbe-Saale region, extends too far north into the plains and covers areas that weren't neolithisised by LBK, but only during the Funnelbeaker period (Altmark etc.). As such, the neolithisation push there doesn't become apparent. The upper Weser & Westphalia have culturally always been linked more to the Rhine than the Elbe, and would have been better placed with the Rhineland (Michelsberger Culture) than with Central Germany. As the area is somewhat a Neolithic late-comer (early 5th millennium), here again the EEF population push on the Elbe-Saale gets underestimated, that in the Rhineland exaggerated. "Southern Germany" includes the upper Rhine and Alsace; a split between the Rhine and Danube basins could have been more appropriate here.
The climate-related analysis is exclusively using data from the Atlantic (Greenland, Ireland, Norway etc.). There is substantial indication that flooding of the North and Baltic Seas affected the Central European and Mediterranean climates and de-coupled them from NW European climates (see my post further above). As such, I question the validity of their finding of no relationship between population patterns and climate as concerns Rhone-Languedoc, Southern and Central Germany. Note also that they actually have found a statistically significant relation between North Atlantic water temperatures and demographic patterns for the Paris Basin, the Rhineland, and Jutland.

I will zoom in onto the Elbe-Saale region in the next post. For the time being, let me point out some general patterns:
A. Migration seems to be apparent in several cases: Paris Basin->Ireland and Rhineland->England around 6,200 BP, Rhineland->Southern Germany around 6,200 and 5,200 BP, Scania/ Danish Isles/ Jutland->Northern & Central Germany around 5,500 BP, Scotland/ Ireland->Wessex/Sussex at about the same time. I am sure more of such migrations can be detected. Here, it would of course have been nice if the study had also covered Bohemia, Silesia and the Polish lowlands.

B. The onset Bronze Age brings about substantial population shifts from about 3,500 BC (brownish arrow). Most regions encounter substantial population declines. Exceptions are Ireland (copper mining?), Southern England (Cornish tin) and Central Germany (Harz copper, Erzgebirge tin). Unfortunately, we don't have demographic information for Brittany, Northern Iberia, and the Central/Eastern Alps (Tyrolia & Mondsee copper), but I would assume substantial population growth there as well. Jutland is holding up surprisingly well, which might indicate substantial maritime trade and east-west connections (British tin to the Mediterranean via the Baltics & Vistula/Dnieper?, Batic amber?).