European mtDNA Signature Established in the Mid Neolithic

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.
 
Aberdeen said:
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.
Click to expand...
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.
 
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.
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.
202_11_23_236_steppenrasen_grosses_federgras.jpg
 
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).
 
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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
 
Timeline of the mtDNA structure of the Elbe-Saale region

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/Eme..._Neolithic_Bell_Beaker_burial_site_in_Germany
 
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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.
 
FrankN said:
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.
Click to expand...

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
 
Sile said:
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
Click to expand...
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:
  1. T1 isn't present in large enough quantities to base much conclusions on its frequencies over time;
  2. The first appearance during the Baalberge Culture (early 4th millennium BC) predates the commonly assumed time line of IE expansion into Central Europe;
  3. 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.
 
Climate change during the Neolithic

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.
353px-Baltic_History_7500-BC.svg.png
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.
Click to expand...
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/Can...ttery_culture_at_Herxheim_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).
Bond-events2.png


Further reading (nice diagrams):http://www.dandebat.dk/eng-klima7.htm
http://www.academia.edu/3584928/Men..._in_Mitteleuropa_in_Vergangenheit_und_Zukunft
 
FrankN said:
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%
HV0213.3 %
V16.7 %
H116.7 %
H5 / H5b213.3 %
other H (rare/ extinct)213.3 %
Subtotal H/V/HV0
8
53.3%
U5b / K320.0%
T2213.3%
X2c16,7%
N1a1a16,7%
Total15100 %

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).
Click to expand...

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
 
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._2012._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.
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The study had two shortcomings:
  1. 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.
  2. 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:

  1. 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.
  2. 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):
  1. LBK: HV/V (4.5%) + J (9%) + K (6.6%) + T2 (7.8%) + W (1.7) = 29.6%
  2. Mid-Neolithic: T1 (2.8%) + U2/5b/8 (6%?) + X (1.3%) = 10.1%
  3. Corded ware: I (2.1%) + U4 (2.9%) + U5a (4%?) = 9%
  4. 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.
 
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.
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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.
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:
  1. 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.
  2. 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.
  3. 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?).

 
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