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Angela
05-07-14, 13:51
epoch;434955]

it is surprising that current day Swedes aren't similar to early Funnel beaker farmers but show larger affinity to WHG.

Current Swedes are about 40-45% EEF, which may very well have entered their gene pool through Funnel Beaker. They also have significant levels of ANE


The mismatch between mtDNA ancestry and autosomal is quite strange.

Indeed, and one could say the same thing about yDNA.



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.

Perhaps, but we still wind up with central European populations with approximately 50% EEF, the later arriving ANE population contributing about 16% and WHG representing about 33% of the total. I also think it may be possible that the incoming Indo-Europeans, in addition to bringing probably the major portion of ANE, could have brought some of the WHG which is now present in central Europe. The upcoming Samara paper should bring some additional clarity to the issue.

I personally think that Bollongino et al is one of those papers where the data doesn't support the conclusions. That cave was co-used for burials by farmers and fisher/gatherers(a strange thing in and of itself) for a very short period. Those could just as well have been recently arrived or transitory fishers from the Baltic or other northern coastal areas, where these people might indeed have found a refuge.

epoch
05-07-14, 16:12
Current Swedes are about 40-45% EEF, which may very well have entered their gene pool through Funnel Beaker. They also have significant levels of ANE

If I recall correctly partly from Indo-European influx but also because Swedish Hunter-Gatherers also had substantial amount of ANE.



Indeed, and one could say the same thing about yDNA.


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.



Perhaps, but we still wind up with central European populations with approximately 50% EEF, the later arriving ANE population contributing about 16% and WHG representing about 33% of the total.

Since EEF itself is about 20%-30% WHG, if I understand well, that would mean that the 50% EEF consists of 10%-15% WHG added to the 33%. Although EEF possibly picked *that* WHG up in south-east Europe so it wouldn't actually mean local ancestry.

As I stated before: A third local ancestry is quite a substantial amount.



I also think it may be possible that the incoming Indo-Europeans, in addition to bringing probably the major portion of ANE, could have brought some of the WHG which is now present in central Europe. The upcoming Samara paper should bring some additional clarity to the issue.


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.



I personally think that Bollongino et al is one of those papers where the data doesn't support the conclusions. That cave was co-used for burials by farmers and fisher/gatherers(a strange thing in and of itself) for a very short period. Those could just as well have been recently arrived or transitory fishers from the Baltic or other northern coastal areas, where these people might indeed have found a refuge.

Yes, you stated that before. I deliberately used the term "studies suggest". However, we know of more HG's living alongside farmers for a substantial time: Swifterband and Vlaardingen in the Netherlands, Pitted Ware in Sweden.

Also in the same article that spends attention to that Bollongio study Dienekes spends attention to another study of ancient mtDNA, called Brandt en Haak. They see HG mtDNA reappear slightly at the end of LBK, and staronger after LBK. I think that is consistent with my suggestion.

http://4.bp.blogspot.com/-t_KXpJ8f80w/Ulbx5NVzpmI/AAAAAAAAJLE/4ygZg4byqIc/s1600/timeline.jpg

epoch
05-07-14, 16:47
@Angela

Also, there is this:

http://dienekes.blogspot.nl/2008/05/first-contact-between-lbk-farmers-and.html



To conclude, the following model can be put forward. During the 6th Millennium cal BC, major parts of the loess region are exploited by a low density of hunter–gatherers. The LBK communities settle at arrival in locations fitting their preferred physical characteristics, but void of hunter–gatherer activity. Evidently,
multiple processes and contact situations may have occurred simultaneously, but in general the arrival of the LBK did not attract hunter–gatherer hunting activity. Their presence rather restrained native activity to regions located farther away from the newly constructed settlements or triggered fundamental changes in the socio-
economic organisation and activity of local hunter–gatherers. Evidence for the subsequent step in the transition dates to approximately one millennium later (Crombé and Vanmontfort, 2007; Vanmontfort, 2007).

So you could start to think that farmers and HG managed to avoid each other apart from the most Eastern edge of LBK settlement: The Rhine valley. There several mass graves were found indicating very violent clashes and a number of villages were fortified with empalement for a short period.

I recently read a book (partly) about the first contact between Aboriginals and settlers and these contacts also turned very violent, an escalation that certainly wasn't unilaterally done by the settlers. The Rhine valley consists of fertile grounds for farmers as well as good hunting grounds for HG's. At exactly such a spot Aboriginals and early settlers clashed hard in Australia too.

LeBrok
05-07-14, 17:15
@Fire Haired

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.
.
I can easily see that happening, with HG surviving in secluded areas of North into AD era. Perhaps the final mixing happening just in last 200 years due to travel, growth of big cities and rise of nationalism and democracy uniting all people under one country. Once we have genetic resolution showing regional levels, or more samples from villages than cities, we'll see very interesting things. Last Slovenian paper was very symptomatic of things to come.

LeBrok
05-07-14, 17:41
I personally think that Bollongino et al is one of those papers where the data doesn't support the conclusions. That cave was co-used for burials by farmers and fisher/gatherers(a strange thing in and of itself) for a very short period. Those could just as well have been recently arrived or transitory fishers from the Baltic or other northern coastal areas, where these people might indeed have found a refuge. We have to be careful generalizing population genetics by findings in caves. They are excellent places to preserve bones and other artifacts, unlike forests and open fields were average Joe lived.

LeBrok
05-07-14, 18:01
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.

That's pretty high level of "contamination". Can't wait for Ancient Near Eastern sample to clean up this issue. I think there might be some EEF farmer gene flow in Loschbour WHG admixture. It would be a surprise if Loschbour was completely pure Hunter Gatherer at 6k BC, only 150 km and 500 years from Stuttgart farmer.

Angela
06-07-14, 20:09
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.

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.

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."

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.

FrankN
06-07-14, 23:01
@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



EDIT: One more link about pigs genetic history:

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

Thanks for that highly interesting comment and the included links, epoch!

One interesting feature of the LBK expansion is that it stopped in the low mountain foothills, and did not expand further towards the North Sea / Baltic coasts. My (amateurish, not backed up by research papers) interpretation of this phenomenon has been ecological: The North German plain is dominated by alluvial moraines (sandy and rather infertile), intersected by various lakes and swamps - anything but prime agricultural lands, and not very attractive to EEFs. The marshes along the lower Rhine, Weser and Elbe are fertile, but prone to flooding, thus also requiring a fairly different set of technologies to be exploited agriculturally. [In fact, most of the Lower Elbe marshes were only cultivated during the high middle ages by Dutch colonists - the "Alte Land" (-> "Hol-land") SW of Hamburg still testifies the relation in its name]. As such, Ertebolle culture HGs and LBK EEFs seem to have occupied fairly different ecological niches. In that sense, Ertebolle and LBK people probably rather lived nearby, but in separate agro-ecological zones, than "alongside but apart from each other". It is nevertheless likely that part of the trade patterns known from the Medieval was already present during the Neolithic, e.g. peat from the swamps sold to the "farming belt" along the mountain foothills for heating purposes, in exchange for meat, dairy, or linen textiles.

Until the high medieval, domesticated pigs were primarily fed on acorn, chestnut and beechnut. They were sent grazing in so-called "Hütewäldern" (herding forests) close to the villages, which also served local timber supply. One may speculate whether the "sacred forests" and "sacred oak trees" of pagan Germanics were not in fact a religious codification of early environmental regulation geared at protecting such herding forests, which would indicate high relevance and long tradition of pig domestication in the North German plain. In any case, I can imagine that local wild boar proved better adopted to these conditions than "imported" Anatolian domesticates, hence they were increasingly incorporated into the swine gene pool (either intentionally, or by wild boars occasionally 'encountering' sows in such herding forests).

A related note, a bit off-topic: Hunting is still today quite popular among (post-post-Ertebolle) farmers in Holstein. One reason is their interest to control the boar population - an encounter between a wild sow (which like to raise their piglets on maize fields for the food supply and view protection) and a maize harvesting machine is usually fatal for both. But my impression (and I know quite a number of such farmers personally) is also that they really enjoy hunting as leisure activity, and for the contribution to the diet. Boar sausage is actually quite tasty ..

Coming back on the flint mines that I discussed in my previous post - another well-studied example is the Lousberg near Aachen (see photo below). According to the German Wikipedia article
http://de.wikipedia.org/wiki/Lousberg (unfortunately there is no English version), it was intensively mined between 3,500 and 3,000 BC. Based on mining slag and processing remains, the Lousberg is estimated to have supplied 300,000 flint axes over this period. That would be some 500-1,000 axes per year, not a large-scale operation, but nevertheless. Lousberg flint has been found as distant as in Thieusies in Belgium (160 km), Büdingen near Frankfurt (260 km), and Minden on the middle Weser (280 km), and was the main source of flint supply to the LBK communities in the Rhineland (Cologne to Duisburg). I am quite sure it also supplied the southern Netherlands, so you, epoch, might want to check Dutch archaeological papers in this respect.
http://upload.wikimedia.org/wikipedia/commons/thumb/6/69/Aachen%2C_Lousberg.jpg/640px-Aachen%2C_Lousberg.jpg

@Angela: We have already discussed the spread of agriculture from Anatolia into Europe elsewhere. I still think we are talking about two, largely contemporary but nevertheless distinct paths. One was sea-based and originated in the Eastern Mediterranean (whether directly from the Levante or from the Anatolian Lake District north of Antalya is still up for discussion). The other one followed the Danube, and may actually have crossed over from Anatolia by land, or along the (now mostly submerged) Black Sea coast from Armenia / West Georgia. The Paschou et al. paper has attempted to trace the Mediterranean path and, in my view, clearly proved such an expansion to have taken place. However, Pachou et al. can't disprove (and also never intended to do so) the second, Danubian path. In fact, we have ample archaeological evidence of such an expansion having occurred, and Pachou's results demonstrate that genetic links to the Levante along the Danube path are much smaller (though clearly present) than along the Mediterranean coast.
You are nevertheless clearly right that, for all its relevance when it comes to Central Europe, the Middle Elbe-Saale region does not represent all of Europe. It has little to do with the Mediterranean. In the Paris basin, and probably already further east, on the Rhine, the Mediterranean and Danubian paths have apparently converged. In fact, if I read Lazarides correctly, the Stuttgart LBK EEF farmer may have rather originated from the Mediterranean than the Danubian wave. Unfortunately, Lazarides has completely excluded Northern Central Europe, i.e. Belgium, the Netherlands, Germany and Poland, from his study, so it is of little help when discussing the origins of LBK EEFs. The only places within the "LBK core" that Lazarides included were Hungary and the Czech Republic. Both bear comparatively little East Mediterranean EEF admixture. Instead they are majority Mesolithic European with a significant Gedrosian component alongside East Mediterranean EEF, but that mix may of course also have been brought forward by later population movements (Germanics, Huns, Slavs, etc.).

FrankN
07-07-14, 03:07
Let me draw your attention to another quite remarkable example of cultural exchange between LBK EEFs and HGs further north. Below is the ground plan of an early LBK house, dated to 5,300-4,900 BC, and excavated in Rannersdorf near Vienna:
http://upload.wikimedia.org/wikipedia/commons/d/dd/Rannersdorf_Grundriss_LBK_Sauer_und_Schwarz%C3%A4u gl_2006%2C_19.jpg
The house was a longhouse (29 x 6 m), anchored on three parallel rows of wooden posts. The outer walls were made up of smaller posts, and probably consisted of half-timbered frames with clay infill applied to (willow) tree branches. The roof was most likely covered by straw or reed. The picture below, a reconstruction of a contemporary LBK house from the Frankfurt area, shows how such houses may have looked:
http://www.uni-frankfurt.de/47462545/erlenbach1-1.jpg
Already the early LBK houses show signs of internal differentiation into 2-3 parts: a living (sleeping) area, a working zone, and some stables, probably for milk cows and their calves.
Unsurprisingly, the basic layout was maintained over the following cultural periods. Below, e.g., is the ground plan of an Urnfield culture house (1,250-750 BC) from the Rannersdorf site. Still a longhouse with half-timbered frames - the only difference is larger central posts, which allow for only a single load-bearing, central column line, and placing the central posts further away from each other.
http://upload.wikimedia.org/wikipedia/commons/thumb/e/e2/Rannersdorf_Grundriss_UK_Sauer_2006%2C_50.jpg/609px-Rannersdorf_Grundriss_UK_Sauer_2006%2C_50.jpg

So what? Well, here is a reconstruction of a 4th century BC Germanic farm house from the Aarhus area in central Denmark:
http://upload.wikimedia.org/wikipedia/commons/thumb/b/be/Jernalderhus.JPG/640px-Jernalderhus.JPG
Reed-thatched longhouse with half-timbered frames anchored on a row of central posts. Sounds common? What about this reconstruction of a Wielbark culture (Goths) house from the 2nd to 3rd century AD, in Hrubieszow near Lublin, on the Polish-Ukrainian border:
http://upload.wikimedia.org/wikipedia/commons/thumb/e/eb/Rekonstrukcja_gockiego_d%C5%82ugiego_domu_w_Mas%C5 %82om%C4%99czu.JPG/640px-Rekonstrukcja_gockiego_d%C5%82ugiego_domu_w_Mas%C5 %82om%C4%99czu.JPG
Obviously a bit wider, and probably using 2-3 rows of central posts, but otherwise a reed-thatched longhouse with half-timber framed outer walls that houses farmers and their cattle under the same roof.

Finally, there is the Low German Fachhallenhaus, the typical farmhouse of the North German Plains between the 13th and 19th century. http://en.wikipedia.org/wiki/Fachhallenhaus
http://upload.wikimedia.org/wikipedia/commons/f/fe/Walsrode_Heidemuseum.jpg
I guess you know the pattern by now: A reed-thatched longhouse, anchored on 1-3 rows of central wooden posts, with half-timbered outer walls (initially clay-filled, later filled with bricks) that houses the farmer family and their cattle under the same roof. Here the distribution map:
http://upload.wikimedia.org/wikipedia/commons/5/52/Fachhallenhaus_Verbreitung.png

Intriguingly, in western Germany, the spread corresponds pretty well to what used to be the original Germanic area before they started expanding southwards into Celtic lands around the 1st century BC. Further south, the Low Geman house gives way to the middle German "Ernhaus", a related style (also a longhouse with humans and animals under the same roof) that, however, has load-bearing (and typically higher) outer walls instead of central rows of load-bearing posts.
http://en.wikipedia.org/wiki/Ernhaus
http://upload.wikimedia.org/wikipedia/commons/4/42/Mitteldeutsches_Haus.png

In other words: The original, simple LBK design has survived with Germanics, while areas further south that had gotten under Celtic influence changed it. Shifting the load-bearing function from the central posts to the walls allowed to build higher (two-storey) and better-lighted houses, which also consumed less ground space. Since that innovation never reached the Germanics, they must have adopted the original LBK design rather early, at latest during the Bronze Age.

Angela
07-07-14, 03:12
epoch;434967]If I recall correctly partly from Indo-European influx but also because Swedish Hunter-Gatherers also had substantial amount of ANE.

The Swedish Hunter-Gatherers had 19% ANE. When we get ANE/WHG/EEF data not just from Samara but for more accuracy from the steppe groups at the time period when they would actually have been entering Europe (which may have already been substantially different), we'll know more about possible relative contributions to that, say, 16% ANE that is present in modern day Central Europeans.

There's also another factor to consider, I think. We don't have, and may never have, data sufficiently detailed enough to prove one way or another whether that specific autosomal ANE of the Swedish Hunter Gatherers actually made it into modern peoples, or whether perhaps it was an isolated group that ultimately did not leave much trace any further west. Someone correct me if I'm wrong, but aren't their yDNA clades extinct?


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.

Not only do I agree that mtDNA should correlate better with autosomal DNA, I think it does correlate better; it just isn't a perfect correlation, partly, I think, because of drift, and perhaps partly because of possible selective advantage for certain mtDNA alone, or in combination with certain yDNA lineages.


Since EEF itself is about 20%-30% WHG, if I understand well, that would mean that the 50% EEF consists of 10%-15% WHG added to the 33%. Although EEF possibly picked *that* WHG up in south-east Europe so it wouldn't actually mean local ancestry.

Exactly. I think we can sometimes get caught up in the weeds with this discussion of the percentage of WHG buried within EEF. For anything definitive about the genetic signature of the first farmers who took off from the Levant we’re going to have to wait for ancient samples from that area.


I think that in this discussion we’re trying to figure out what happened to the WHGs who were not among those initially incorporated in, probably, south-east Europe, or even before. Were they all, or substantially all, pushed to the far northeast, far northwest or northern coastal regions that were unattractive to the Neolitic farmers, with only remnants remaining in Central Europe, in a sort of North American Indian model, to filter back in following the Neolithic population crash in Central Europe, which I am persuaded did happen, and perhaps with Indo-European populations? (We can speculate, I think, that some, at least, did remain, and were absorbed, because the Gok samples show additional WHG compared to Stuttgart, or Oetzi, for that matter.)


Or did, alternatively, a substantial number of them remain in Central Europe, living side by side with, but separate from, the Neolithic farmers in their midst. I currently tend toward the first alternative, partly because, as I’ve said before, given the amount of territory it takes to support the hunter-gatherer life style, or even the fisher/gatherer lifestyle, which may be more apropos, and the reproductive advantage the steadily advancing farmers would have had, I think the remaining WHG numbers would likely have been small. How large a population could even the bogs or swampy areas have sustained? Using the American Indian analogy again, imperfect as it is, the Mexican and South American Indians had a much larger population, giving them a greater chance of survival, because they had already experienced their own Neolithic Revolution. Also, I’m far from being a Y DNA expert, but hasn’t there been speculation that yDNA I1, for example, was bottlenecked until it adopted farming? Perhaps one of our more knowledgeable posters can chime in.

I’m keeping an open mind about this whole topic, however, particularly as there might have been many regional variations.




As I stated before: A third local ancestry is quite a substantial amount.

I understand that you believe that; I just don't think the data yet exists to prove it.


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. Although 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.


Are you saying that there was no gene flow from Neolithic groups into the Baltic areas? If the Neolithic spread to the Baltic region from Neolithic cultures to their south, cultures that were already mixed WHG/EEF, then how would that be possible? More importantly, how could that be proved?

As to your comment about Lazaridis et al, I don’t remember the paper concluding that the EEF in Baltic peoples was any different than the EEF in any other Europeans, (which may be, say 20% WHG) or, to say it another way, that they were able to segregate the buried WHG in EEF people, and say that only those genes appear in people in the Baltics. Could you point me to the relevant passage?




Yes, you stated that before. I deliberately used the term "studies suggest". However, we know of more HG's living alongside farmers for a substantial time: Swifterband and Vlaardingen in the Netherlands, Pitted Ware in Sweden.

Also in the same article that spends attention to that Bollongio study Dienekes spends attention to another study of ancient mtDNA, called Brandt en Haak. They see HG mtDNA reappear slightly at the end of LBK, and staronger after LBK. I think that is consistent with my suggestion.

http://4.bp.blogspot.com/-t_KXpJ8f80w/Ulbx5NVzpmI/AAAAAAAAJLE/4ygZg4byqIc/s1600/timeline.jpg[/QUOTE (http://4.bp.blogspot.com/-t_KXpJ8f80w/Ulbx5NVzpmI/AAAAAAAAJLE/4ygZg4byqIc/s1600/timeline.jpg%5b/QUOTE)]


I think Pitted Ware might fit well into a scenario where WHG were mainly isolated to the far northern or northeastern areas, yes? As to Swifterband and Vlarrdingen, I’m assuming you’re basing this on archaeological evidence, because I don’t believe there’s anything available in terms of genetics data.

This is an interesting area for exploration. Do you have any links to papers that show radiocarbon dated HG sites that post-date the arrival of the Neolithic in the area? How many of them are there in relationship to the Neolithic sites? Is there other data that, in your opinion, would indicate that there were substantial numbers of WHG remaining in the area?


Yes, the Brandt et al paper is very interesting. However, the way I read the graphic is that there is, as you say, a slight resurgence in this particular area of HG lineages from around 4600 BC to around 4100 BC (from virtually 0 to about 10% of the total?). However, by 3400 BC it's virtually back to square one, and they only start increasing again with the SMC culture to reach a high of about 30% in 3100 BC? They then see-saw a bit (it looks as if each new culture, CWC, BBC, Unetice, decreases them a bit) to finally level off at about 18%?

Interestingly enough, although there are earlier trickles, the big spike of "new" mtDNA lineages that might specifically be tied to an Indo-European migration (I, U2, T1, R ) really make their appearance around the time of Corded Ware and Unetice, reaching a high of about 20%, but they "settle" at an average of what looks to be less than 10%.

FrankN
07-07-14, 08:50
@Angela: I think you (and possibly a few other posters) are over- or misinterpreting Lazarides findings. Or, maybe I better say: Amidst the various highly interesting results Lazarides has delivered, serious methodical flaws of his work tend to be overlooked. Below is a representation of his admixture results, copied from http://www.eupedia.com/forum/threads/29705-Corded-Ware-Iranic-Aryan-split-of-IE/page11?p=434227#post434227.
http://www.eupedia.com/images/content/Admixtures-Lazaridis.png
Now, look first at the width of each "ethnic" bar, which indicates how many samples from a specific group were included in the analysis. See what I mean?


In search of the "true" Mesolithic HG, Lazarides et al. have massively oversampled Basques, Sardinians, but also French Basques, and Spanyards in general - just to find out that these groups are more like a perfect mix of HG and EEF (Sardinians even very much the prototype of EEF) than typically Mesolithic European. Similarly, they have also massively over-sampled Caucasians, Druze, Palestinians, and various groups of Jews, which is less crucial in the context we are discussing here (though still needs to be kept in mind when interpreting the findings).
Conversely, they haven't sampled Central Europe at all: No Belgians, Dutch, Germans, Poles, Danes, or Swedes. Even English are very much under-represented compared to French, not even speaking of Maltese, who seem to be nearly as numerous in the overall sample as are English. Or, look at the Irish! Oops - no Irish here?
In a first stage, they determined that ancient HG (Loschbour and Motala) weren't that far apart genetically from each other - at least when comparing them to Eskimo, Massai, Japanese, Kalash or Kharia. Fair enough, as long as we are talking about global population movements since Palaeolithic times (which was what they were after). But some people tend to take that statement also being applicable on a Mesolithic Europe scale. It may or may not be so - Lazarides et al. didn't really check for it, they looked at 100 millennia and the whole world! In any case, Loschbour and Motala 12 were both yDNA haplogroup I2a1b*, and mtDNA U5b1a (Loshbour) and U2e1 (Motala 12), respectively, so probably in fact genetically not too far apart.
What they did then was checking whether Loschbour (WHG) or Motala 12 (SHG) represented better what their admixture analysis had indicated to be Mesolithic HG ancestry. Surprisingly, that was Loschbour, which is located only half as far away from the Basque country or from Sardinia, as is Motala in central Sweden. The other interesting point here is that, previous to Lazarides study, mtDNA U2 was believed to only have entered Europe with the Corded Ware culture http://www.eupedia.com/forum/threads/29163-New-map-of-mtDNA-haplogroup-U2?p=418059&viewfull=1#post418059. As it is much rarer than Loschbour mtDNA U5, it is no surprise that Lazarides et al. identified Loschbour (WHG) as the prototypical Mesolithic ancestry.
Unfortunately, they then missed the opportunity to also trace SHG ancestry (i.e. especially mDNA U2) among their population sample. They just concluded that Motala 12 was a mix of 81% WHG and 19% ANE, and left it there The question how, by 6,000 BC, 19% ANE could have been present in the SHG gene pool, wasn't discussed at all.
Finding that a two-way mix of EEF and WHG could not explain Europe's genetic diversity, they introduced a third component, named Ancient North Eurasian (ANE). After a bit of testing, they established that 24,000 year old MA1 from the Lake Baikal area best represented the ANE component in their three-way model. Now, look at their admixture results for MA1 in the diagram above. Doesn't look that pure, doesn't it? So, if ANE equals MA1, which is something like 40% WHG, plus 20% Kalash, plus a bit of Eskimo, Kharia and whatever else, while SHG is 81% WHG plus 19% ANE, that makes the Finns .. well, forget it! You just can't do that kind of admixture arithmetic, definitely not with Lazarides' data that is based on genetic proxies from different points in time and space (and that is not directed against Lazarides et al., they used the best ancient DNA data available, but we have far too little of such data to already arrive at such detailed conclusions).
Finally, they checked whether their 3-way mix could explain most of Europe's genetic make-up (Supplementary Information 12). Result - it couldn't. So they excluded populations "with evidence of additional complex history", i.e. Sicilians, Maltese, Ashkenazi Jews, Finnish, Russians and Mordovians. Result - at least four, probably five components required to explain the genetic make-up. Then, they threw out the Spanish (for which they determined additional North African admixture, especially from Mozabite, possibly during the pre-Roman Iron age). While they state this had improved the fit, they don't give any details on the statistical results - I guess that means they were still not satisfactory. Finally, they employed a "reduced subset" that also eliminated Basques, all Italians expect for Sardinians, all other Mediterraneans that had survived their previous "cleaning", all Balts, and the Ukrainians (and remember, they never included Northern Central Europe nor Ireland, and had already thrown out Russians, Finns, Saami, and other Uralians and Caucasians before). Voila - the make-up of the remaining (12) European populations could be explained from their 3 components. Sorry - this is somewhere between extreme window-dressing and complete bullshit, but has nothing to do with serious scientific work!

Bottom lines:

Lazarides et al. do not tell us anything about the fate of Mediterranean HGs - for them, they did not exist. Especially when looking at the Sardinian results, and also at North African Bedouins (Bedouins 1), I suspect their genes could be hidden somewhere inside the EEF component (Remember- they could not fit their EEF/ WHG / ANE mix to Mediterranean populations!).
Lazarides et al. do not tell us anything about the fate of Scandinavian HGs - after establishing that WHG (Loschbour) better fit their sample of current Europeans (which is heavily skewed away from northern Central Europe), they did not look into SHG further (Remember here that their 3-way WHG/EEF/ANE model only worked after they had "thrown out" Finns, Russians, Ukrainians, and Balts).
Lazarides et al. do not tell us anything about the fate of Central European HGs - because they hardly looked at Central Europe in their study.
They pretend to tell us a lot about WHG, though. WHG took over Russia and the Baltic states, account for the majority of British and French genetic heritage, and could by a hair also have overtaken EEF in the Basques. Believe it, or not!
We do learn other interesting things: EEF, e.g., appear to have left more genetic traces on Icelanders than on Iranians or Chechens (Yeah, early farmers obviously spread by sea!). And while only few of them made it into Russia, a couple settled among the Northern Ethiopian Afar, which looks like an -ehm - interesting place for early farmers to settle http://en.wikipedia.org/wiki/Danakil_Desert
Finally if you look at the aDNA admixture analysis in their Annex, you will note a significant WHG / EEF component among Native American Algonquin, Cree and Ojibwa (apparent from k=3, splits into the two components at k=19). That finding in itself is actually very remarkable and deserves a separate, in-depth discussion. But it confirms my suspicion that, whatever they have been actually measuring, it doesn't really correspond to "pure" WHG and EEF heritage, respectively.


To remain fair: Lazarides et al. themselves state in the main body of their study that all three "source populations" may already have been admixed. For EEF they estimate "44 +/-10% basal Eurasian ancestry", and a WHG-related ancestry of 0-45%. They also caution that finding three admixture sources could be consistent with a larger number of admixture events (though the main study stays rather tacit on the questionable way they have concluded on such a three-way mix, other than noting that this finding was statistically not significant). The more often I read their study, the more it seems to me that the real story they wanted to tell was that of a significant trans-Atlantic gene flow, but somehow, somebody convinced them to rather focus on Europe and the Near East instead.

Vedun
07-07-14, 14:06
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.

There was no Slavic migration in early middle ages at all... No archeological discoveries haven't proved that (yet). That's why we do not exist at all - according to these theories (no N haplogroup migration from Asia in Slovenia, Croatia, Slovakia,...) Secondly the Hallstatt cultures were clearly proto Celtic and not Germanic.

Nori, Aedui, Volcae, Boii, Helvetii (Kelveti; probably a derivative of "Kelti") weren't Germanic.

FrankN
07-07-14, 15:50
There was no Slavic migration in early middle ages at all... No archeological discoveries haven't proved that (yet). That's why we do not exist at all - according to these theories (no N haplogroup migration from Asia in Slovenia, Croatia, Slovakia,...).

I have made my points on Slavic migration in several threads:
http://www.eupedia.com/forum/threads/26341-who-were-Slavic-people?p=434359&viewfull=1#post434359
http://www.eupedia.com/forum/threads/29386-Thracians-spoke-Balto-Slavic-language?p=433040&viewfull=1#post433040
http://www.eupedia.com/forum/threads/29632-Poland-more-Germanic-or-Slavic?p=434521&viewfull=1#post434521
Feel free to comment on them in the threads concerned. However, as this thread is already dealing with enough issues that are only loosely related to the original topic (and I confess, I have also partly gone off-topic), I suggest we keep anything related to Slavs out of here.


Secondly the Hallstatt cultures were clearly proto Celtic and not Germanic.
I never stated otherwise. Please read my post again.

LeBrok
07-07-14, 16:59
Let me draw your attention to another quite remarkable example of cultural exchange between LBK EEFs and HGs further north. Below is the ground plan of an early LBK house, dated to 5,300-4,900 BC, and excavated in Rannersdorf near Vienna:

The house was a longhouse (29 x 6 m), anchored on three parallel rows of wooden posts. The outer walls were made up of smaller posts, and probably consisted of half-timbered frames with clay infill applied to (willow) tree branches. The roof was most likely covered by straw or reed. The picture below, a reconstruction of a contemporary LBK house from the Frankfurt area, shows how such houses may have looked:


n other words: The original, simple LBK design has survived with Germanics, while areas further south that had gotten under Celtic influence changed it. Shifting the load-bearing function from the central posts to the walls allowed to build higher (two-storey) and better-lighted houses, which also consumed less ground space. Since that innovation never reached the Germanics, they must have adopted the original LBK design rather early, at latest during the Bronze Age.
Great history of long houses, thanks. I had no idea they go so far back till 5th millenium BC! The EEF Stuttgart fellow possibly lived in one of these?

Angela
07-07-14, 17:23
@Epoch
A few additional thoughts:

As to Brandt et al, the correlation between the various cultures and the mtDNA lineages studied in that paper really deserve a detailed discussion.

Brandt et al and Bollongino et al are addressed in general terms in this eupedia thread: http://www.eupedia.com/forum/threads/29148-Two-major-new-papers-on-Early-Neolithic-to-Early-Bronze-Age-mtDNA-in-Central-Europe?highlight=Brandt

Unfortunately, the discussion got bogged down in the quagmire of mtDNA "H" dating, which is also the subject of a current, sometimes heated discussion on anthrogenica.

Perhaps the thread could be revived. (I have something of a personal interest in this discussion, as I'm mtDNA U2e myself. :))

My post number 19 on that thread is not one of my best efforts. :( It illustrates the danger of accepting other people's analyses of a paper instead of carefully reading the whole thing oneself. The last paragraph contains what I now know after my own reading of the paper to be an obvious error about mtDNA "H" and Brotherton's claims about it, but alas, it's too late to add an Ed. statement.

I provided a link in that thread to a paper about the spread of the Neolithic in the Balkans. For ease of discussion, here it is: http://www.pnas.org/content/110/9/3298.full

As I stated in that thread, I'd welcome critiques of it from other people. What I do gather from it is that the HGs there were rather quickly absorbed. The other interesting tidbit is that there seems to have been relatively more movement of Neolithic women into the HG cultures than the other way around. Perhaps initially the HG women were a bit less attractive to Neolithic settlers because of their lack of knowledge of farming and animal tending technology? Could we also speculate that it was the Neolithic women who "Neolithicized" the HG bands by bringing the knowledge of the Neolithic lifestyle to them?

Apropos of this discussion, take a look at the Pitted Ware mtDNA sequences here at Jean Manco's page on Ancient DNA haplogroups. which is an invaluable resource : http://www.ancestraljourneys.org/mesolithicdna.shtml
Although very much minority haplogroups, you already have present an HV, aT2b and a K1a mtDNA. (The mtDNA frequencies of the Neolithic cultures including LBK and Rossen etc. are on a separate page for those unfamiliar with that site.)

Speaking of Jean Manco, I sometimes take a look at what's being posted at Anthrogenica on subjects that interest me. Here is her latest take on Bollongino et al:
http://www.anthrogenica.com/images/BittenFruit_fluid/misc/quote_icon.png Originally Posted by Jean M http://www.anthrogenica.com/images/BittenFruit_fluid/buttons/viewpost-right.png (http://www.anthrogenica.com/showthread.php?p=44421#post44421)
I've been searching for my previous comments on the Bollongino 2013 paper ...

I found this study puzzling, as two separate groups would not normally share a burial place. A better explanation just occurred to me. The fishing people could have been just occasional visitors from the coast, delivering fish. Occasional deaths in situ would be dealt with by the locals burying the body in the local burial spot that they used themselves.
Had a nagging feeling that I was remembering the paper incorrectly. I was. The later bunch of foragers were eating freshwater fish. This still distinguishes them from the earlier Mesolithic burials at the Blätterhöhle, who were meat-eaters. The DNA of the fishermen is also too different from that of the earlier Mesolithic group for confidence of direct descent, though the paper was keen not to rule it out:

At first sight, this would seem to cast doubt on a direct genetic continuity between them. However, coalescent simulations do not reject a model of population continuity under a very wide range of demographic parameters..


Also there were U5b2a2 individuals among both the farmers and fishermen. So intermarriage had actually taken place, though the paper plays this down. (There was also a U5b2a5 among the farmers, though not matched among the fishermen.)

So what we have now looks to me like a case of in-laws arriving periodically, or perhaps being cared for by daughters in old age/injury. That would explain the burial in the farmer's usual burial site. It would also explain the complete lack of any sign of Mesolithic life surviving in the surrounding area. The original contact could still have come from trade, but not of marine fish.

I think this partly speaks to Le Brok's point that it might be injudicious to make broad generaliztions about the peopling of even Central Europe based on the somewhat strange practices at one cave burial site.

Finally, I'm pressed for time, so I didn't do an exhaustive search, but I couldn't find a map of radiocarbon dated HG sites in Central Europe either pre-or-post the arrival of the Neolithic settlers.

Which reminds me, does anyone have a link to a study with a reliable estimate for the total number of hunter-gatherers in Central Europe around the time of the arrival of these new settlers, and one that guesstimates the number of farmers in LBK settlements, or Neolithic settlements in general prior to the population collapse? Those kinds of findings might provide further insight.

LeBrok
07-07-14, 17:46
Which reminds me, does anyone have a link to a study with a reliable estimate for the total number of hunter-gatherers in Central Europe around the time of the arrival of these new settlers, and one that guesstimates the number of farmers in LBK settlements, or Neolithic settlements in general prior to the population collapse? Those kinds of findings might provide further insight. As you mentioned yourself before, we should be able to observe a see-saw effect going between Farmers and HG populations in Central and North Europe. With every collaps (cold weather spells) creating HGs population recovery, and Farmers dominating in warm and moist periods. In this case whoever studies HG populations of Northern Europe have to carefully go through all millenia till pretty much Iron Age.

FrankN
07-07-14, 21:00
Which reminds me, does anyone have a link to a study with a reliable estimate for the total number of hunter-gatherers in Central Europe around the time of the arrival of these new settlers, and one that guesstimates the number of farmers in LBK settlements, or Neolithic settlements in general prior to the population collapse? Those kinds of findings might provide further insight.
The Brotherton study itself (Figure 1, inset in bottom right corner) includes some graphs. They put the population density of Central Europe (however it is defined) after the LBK expansion at 0.6 persons/ km². The combined area of today's Benelux, Germany, Switzerland, Austria, Hungary, Slovakia, the Czech Republic and Poland is around 1.1 million km² - that would mean some 660,000 inhabitants during the LBK period.

The LBK farmer population might be guesstimated based on the following paper, which describes the three LBK settlements that provided data to the Brotherton study in detail: https://openaccess.leidenuniv.nl/bitstream/handle/1887/19045/05.pdf?sequence=13. But remember - its only one specific region, and not necessarily representing all of Central Europe :innocent:
Two of the three sites, Halberstadt and Derenburg, are located less than 10 km apart. The Halberstadt settlement consisted of four longhouses; Derenburg of 11 longhouses from at least three different settlement periods. The third settlement, Karsdorf, 100 km further south, consisted of 24 longhouses from three settlement stages. From another paper that I read, but unfortunately did not bookmark, I recall that in the Rhineland, the structure actually consisted of individual longhouses every 1-2 km, small villages in-between, and somehow larger settlements every 50-60 km. I take Karsdorf, with maybe 10 longhouses during its biggest extension, as such a larger settlement, and Halberstadt and Derenburg as small villages. Derenburg and Halberstadt are located on the Magdeburg Börde, an extremely fertile area, so they may actually be a bit closer to each other than smaller villages would usually be. Nevertheless, we might talk about something like one longhouse / km² in average along the fertile loess plains and major rivers.
http://en.wikipedia.org/wiki/Magdeburg_B%C3%B6rde
This leads to the question how many people lived in each longhouse. Some studies assume several families, up to 25 people in total, per longhouse. However, in Halberstadt, 6-8 graves were found around each longhouse. Derenburg had a central cemetery with 44 (excavated) graves in total. This pattern rather suggests one family per longhouse (and possibly a group of several related families clustering in and around one small village). I suppose we are talking a 3-generation family here, which possibly includes one unmarried 2nd-generation adult alongside his or her married brother, so something like 4-5 adults in average. 19 of the 97 skeletons that were analysed (20%) were below 6 years of age, which suggests substantial infant mortality, even if not all adults may have been buried close to their home, [Note in that respect that several studies have shown that domestication of animals has substantially increased infection risks - the resulting higher mortality could at least partly, if not almost completely have offset the nutritional advantage of early farmers vis-à-vis HGs] Bone tissue analysis on the Halberstadt, Derenburg and Karsdorf skeletons suggests that the infants were breast-fed to the age of three, though the authors warn that their sample size is fairly limited in this respect. Anyway, if furthermore accounting for still births, a couple may have only reproduced every 4-5 years, and would thus be unlikely to have had more than five children in average, 1-2 of which may have died before reaching six years of age. All in all, 8 persons per longhouse look like a fair estimate, so the LBK population density within the core agricultural area may have ranged around 8 persons / km².
If you ask, however, how large that LBK agricultural area may have been - I have no idea! Let's say we are talking 5% of all Central Europe, which is probably too much, since the coastal plains, and the mountain ranges, weren't covered at all. That would yield something like 1.1 million km² * 5% * 8 persons/km² = 440,000 LBK farmers. The remainder (220,000) would be HGs. Roughly one HG per two farmers. At 6% LBK coverage, the ratio goes down to 1:4 (530,000 : 130,000).

A few other takeaways from the a/m study:

Lactose tolerance was tested on three individuals. All were negative
The diet was quite balanced between animal protein and grain / vegetables. Compared to Neolithic Britain, animal protein consumption was slightly lower in the Middle Elbe / Saale region. (Sweetwater) fish was hardly consumed. Of the 32 animal remains found in Karsdorf, 12 were bovine, 8 pigs, 8 sheep or goat, and 4 wild animals (3 Aurochs, 1 deer).
In general, there was no discernible difference between male and female diet. One older man from Halberstadt, however, showed significantly higher meat consumption than average. He was found in a separate grave, distant from the longhouses, "buried in an extremely crouched position". A HG who had found acceptance in the LBK community, without becoming part of any nuclear family? The other extreme is one adult woman from Karsdorf "that could be classified as a vegan."


I assume you are already aware of the excellent work done by Walter Scheidel at Stanford University. Although he mainly focuses on demographic and socio-economical aspects of the Roman empire and the early migration period, you might also find some comparison with other historical periods in his works. His publication list (mostly available as PDF free of charge) is here: http://www.princeton.edu/~pswpc/papers/authorMZ/scheidel/scheidel.html

JS Bach
08-07-14, 04:20
Finally if you look at the aDNA admixture analysis in their Annex, you will note a significant WHG / EEF component among Native American Algonquin, Cree and Ojibwa (apparent from k=3, splits into the two components at k=19). That finding in itself is actually very remarkable and deserves a separate, in-depth discussion. But it confirms my suspicion that, whatever they have been actually measuring, it doesn't really correspond to "pure" WHG and EEF heritage, respectively.



@FrankN: Would you happen to know where I might find the actually breakdowns of the Algonquin, Cree and Ojibwa for the k=20 level? I can't tell from the chart if they have any of the Caucaso-Perso-Gedrosian component, in addition to the WHG- and EEF-related components. There have been some claims about the Basque and Algonquin languages having similarities. Basques don't seem to have any of the Caucaso-Perso-Gedrosian component there.

FrankN
08-07-14, 06:39
@FrankN: Would you happen to know where I might find the actually breakdowns of the Algonquin, Cree and Ojibwa for the k=20 level? I can't tell from the chart if they have any of the Caucaso-Perso-Gedrosian component, in addition to the WHG- and EEF-related components. There have been some claims about the Basque and Algonquin languages having similarities. Basques don't seem to have any of the Caucaso-Perso-Gedrosian component there.
Unfortunately, no. All that I have is the Lazarides study with the Supplemental Information. I browsed it again, but it seems they only published admixture estimates (WHG/EEF/ANE) for Europeans. There are quite a number of hints on the transatlantic gene flow scattered across the various supplements, but none of them has easily-interpretable data (at best some coefficients from various statistical models). Anyway, they seem to hint at a "Nordic" connection via Iceland, in addition to the well-known Siberia-Alaska path. Your comment on Basque language suggests that you may be more looking at the Solutrean hypothesis, but I don't feel one can draw much from Lazarides in this respect.
.
We should open a separate thread on it - feel free to start it (I will be busy for some more time with digging further into the prehistory of the Elbe-Saale region, as my ancestry is from there).

FrankN
08-07-14, 10:37
This map may help to put some (though not all) of Brotherton's findings into context:
http://p12897.typo3server.info/fileadmin/gruppen/rgzm/2010/Neolithische_Gesellschaften/Enceintes_MK_EAA_copie.jpg
It comes from an on-going joint French-German research project on the Michelsberg Culture (MK), ca. 4,300-3,600 BC, which followed on the Rössen culture in Western Germany and Northern France. The MK core is shaded bluish, its contact zone with neighbouring cultures (including Middle Elbe/ Saale to the east, marked yellow as MK/TRB) is highlighted. In Brotherton's chronology, MK corresponds to late Rössen - Schöningen - early Baalberge [Central European Neolithic chronology is an absolute mess, every author and every regional school appears to use a specific terminology and time scale, which makes it quite difficult to identify common patterns on a larger geographical scale].

Several (all?) of the settlements (triangles) may be interpreted as early cities - one of the best known and researched is the Glauberg north of Frankfurt. The Michelsberg near Karlsruhe, a 10 ha summit plateau, after which the culture was named, is another example.
http://en.wikipedia.org/wiki/Michelsberg
http://en.wikipedia.org/wiki/Glauberg

Two other large and fortified settlements were Halle-Döhlauer Heide, and nearby Salzmünde. They are represented by the two yellow triangles inside the dotted square around Halle, near the eastern end of the map. The Salzmünde earth walls encircled an area of 40 ha. As already indicated by the names (old German "Hall"=salt, see also "Hallstatt"), both places were important centres of salt extraction and trade. The same, btw, applies to other places that have lent their names to local cultures such as Bernburg and Schöningen (Schöningen is most likely the black "Rössen" star NNW of Halle, just on the border between Lower Saxony and Saxony-Anhalt). The earliest evidence for salt boiling north of the Alps (ceramic evaporation vessels, so-called briquetage, see photo below) was found in Halle-Döhlener Heide. Northern Saxony Anhalt is still today the most important salt production area inside Germany. Salt springs are indicated in white. The map also shows important Neolithic flint mines, including Lousberg near Aachen, and the recently discovered Bottmersdorf mine near Mansfeld in the Elbe-Saale region (though it does not include the Upper Bavarian mines near Kelheim, a bit SW of Regensburg).
http://www.museum-digital.de/san/images/201108/25154659913.jpg

The on-going French-German research project tries to establish whether the MK evolved around a hypothesized central axis "Ile de France" - "Mosel" - "Rhein-Main-area (Glauberg)" - "Erfurt" - "Halle/ Saale" (shaded in pink on the map). It will pay special attention to tracking longer-distance trade, especially of salt, in this respect. Let's see what they will find out...

Edit: The recently discovered flint mine in the Mansfeld area is not the Bothmersorf mine recorded on the map (I actually wasn't aware of that one), but another one further south. some 20 km west of Halle. For lack of organic material, the mine could so far not be dated exactly. However, mine refill includes sherds of middle Neolithic ceramics (whatever middle Neolithic means in this context). The local press has been interpreting it as "6,000 years of mining in the Mansfeld area". Detailed excavation report (in German, but with photos): http://www.lda-lsa.de/landesmuseum_fuer_vorgeschichte/fund_des_monats/2014/juli/
To understand better local enthusiasm on the finds, I should add that copper and silver mining has been the main economic activity in the Mansfeld area since the 12th century, and mine closure after 1990 has sent the region in economic depression. Has Mansfeld copper and silver already been mined in pre-historic times? Many suppose so, but so far no evidence has been found (and, after at minimum 800 years of open pit and underground mining, one shouldn't expect many archaeological traces to have survived). Let's also add that just around Halle, a major lignite deposit starts, part of which is still today exploited in open-pit mining (see map below). Now, if you are regularly boiling substances, e.g. salt lake, lignite deposits nearby might come in handy (just saying). Finally, in Bitterfeld, 20 km NE of Halle, amber was mined industrially between 1974 and 1993 from alluvial deposits.
http://www.hs-zigr.de/schule/scholl/faecher/eru/fue_04/KrisKath/Bilder/bodenschatzkarte.jpg

What I am trying to say here: Instead of interpreting the genetic history of the Middle Elbe - Saale region along EEF vs. HG lines, one should rather regard the region as a kind of late Neolithic / Chalcolithic Ruhr area or Midlands. This implies that specific locations may have had a very specific economic focus other than agriculture. All Brotherton samples for the Rössen period, e.g., have been taken from a grave field close to where medieval copper mining in the Mansfeld area started in 1189 AD. The grave field is around 1 km away from a place called "Silbergrund", where a silver refinery was operated during the late medieval, and 2.5 km away from Hettstedt-Kupferberg ("copper mountain"). All samples for the Schöningen and Salzmünde phases, OTOH, come from the Salzmünde fortified area described above.
I am still in the process of digging deeper into the individual locations, trying to figure out what Brotherton's results might actually mean, but I clearly agree to other poster's statements that broader (and more geographically spread) sampling would have desirable. However, let's not be unfair - 37 ancient samples is better than nothing..

Sile
08-07-14, 11:25
This map may help to put some (though not all) of Brotherton's findings into context:
http://p12897.typo3server.info/fileadmin/gruppen/rgzm/2010/Neolithische_Gesellschaften/Enceintes_MK_EAA_copie.jpg
It comes from an on-going joint French-German research project on the Michelsberg Culture (MK), ca. 4,300-3,600 BC, which followed on the Rössen culture in Western Germany and Northern France. The MK core is shaded bluish, its contact zone with neighbouring cultures (including Middle Elbe/ Saale to the east, marked yellow as MK/TRB) is highlighted. In Brotherton's chronology, MK corresponds to late Rössen - Schöningen - early Baalberge [Central European Neolithic chronology is an absolute mess, every author and every regional school appears to use a specific terminology and time scale, which makes it quite difficult to identify common patterns on a larger geographical scale].

Several (all?) of the settlements (triangles) may be interpreted as early cities - one of the best known and researched is the Glauberg north of Frankfurt. The Michelsberg near Karlsruhe, a 10 ha summit plateau, after which the culture was named, is another example.
http://en.wikipedia.org/wiki/Michelsberg
http://en.wikipedia.org/wiki/Glauberg

Two other large and fortified settlements were Halle-Döhlauer Heide, and nearby Salzmünde. They are represented by the two yellow triangles inside the dotted square around Halle, near the eastern end of the map. The Salzmünde earth walls encircled an area of 40 ha. As already indicated by the names (old German "Hall"=salt, see also "Hallstatt"), both places were important centres of salt extraction and trade. The same, btw, applies to other places that have lent their names to local cultures such as Bernburg and Schöningen (Schöningen is most likely the black "Rössen" star NNW of Halle, just on the border between Lower Saxony and Saxony-Anhalt). The earliest evidence for salt boiling north of the Alps (ceramic evaporation vessels, so-called briquetage, see photo below) was found in Halle-Döhlener Heide. Northern Saxony Anhalt is still today the most important salt production area inside Germany. Salt springs are indicated in white. The map also shows important Neolithic flint mines, including Lousberg near Aachen, and the recently discovered Bottmersdorf mine near Mansfeld in the Elbe-Saale region (though it does not include the Upper Bavarian mines near Kelheim, a bit SW of Regensburg).
http://www.museum-digital.de/san/images/201108/25154659913.jpg

The on-going French-German research project tries to establish whether the MK evolved around a hypothesized central axis "Ile de France" - "Mosel" - "Rhein-Main-area (Glauberg)" - "Erfurt" - "Halle/ Saale" (shaded in pink on the map). It will pay special attention to tracking longer-distance trade, especially of salt, in this respect. Let's see what they will find out...

Glauberg is the burial places for the "royal" celtic chiefs ......IIRC this place has been excavated for more than 10 years

Nobody1 earlier in the year mentioned a new post from glauberg...IIRC 27 skeletons where found

epoch
08-07-14, 16:57
@Angela



Do you have any links to papers that show radiocarbon dated HG sites that post-date the arrival of the Neolithic in the area? How many of them are there in relationship to the Neolithic sites? Is there other data that, in your opinion, would indicate that there were substantial numbers of WHG remaining in the area?


I haven't had a lot of time and this thread fills up with interesting responses. However, I feel I have an obligation to answer this as I, as an amateur, so pretentiously have called for attention to these cultures. I initially learned about from Dutch archeology books, books that I have all currently packed as I am moving. The original Vlaardingen finds, from a culture that seems related if not similar to Swifterband, are on display in Leidens museum of antiquity. They consist of a number of well preserved fish traps and a lot of freshwater fish remains. This is therefore not inconsistent with continuing mesolithic HG's using freshwater fisheries for sustenance.

The original Swifterband find is described in the link below and in the first map with an overview of Swifterband finds you again see them concentrated around freshwater systems.

http://dissertations.ub.rug.nl/FILES/faculties/arts/2013/i.devriendt/thesis_Izabel_Devriendt___proe_1.pdf

But you want dates off course, in order to check the impact of coexistence. This might be interesting: Hoge Vaart being a typical Swifterband site. Actually, on slightly more than a day's walk distance from the Swifterband site.

www.academia.edu/1231539/Radiocarbon_dating_of_Mesolithic_open-air_sites_in_the_coversand_area_of_the_north-west_European_plain_problems_and_prospects (http://www.academia.edu/1231539/Radiocarbon_dating_of_Mesolithic_open-air_sites_in_the_coversand_area_of_the_north-west_European_plain_problems_and_prospects)

There is quote in it that indicates something happened to HG's though:


The reliability of these samples for dating Mesolithic sites is corroborated by more than 404 dates for the Netherlands (Niekus 2005–6) and about 25 for Belgium (Van Strydonck andCrombé 2005). Except for a few isolated dates, all belong to the Mesolithic, with a sharp decrease during the Final Mesolithic (Swifterbant Culture) from c. 5800 bp onwards (Niekus2005–6). From then onwards, and during the entire Neolithic, hearth pits occur only very incidentally (Groenendijk 1987; Hamburg and Louwe Kooijmans 2006).

epoch
08-07-14, 19:44
@FrankN




In other words: The original, simple LBK design has survived with Germanics, while areas further south that had gotten under Celtic influence changed it. Shifting the load-bearing function from the central posts to the walls allowed to build higher (two-storey) and better-lighted houses, which also consumed less ground space. Since that innovation never reached the Germanics, they must have adopted the original LBK design rather early, at latest during the Bronze Age.

Dutch farms are similar. But there are local differences

A farmhouse called "Saksische boerderij", Saxon farmstead. The difference between the examples you give and these is that these regularly have the frontdoor on the small side.

http://upload.wikimedia.org/wikipedia/commons/0/06/Vragender_dorp.JPG

The Frisian type, which seems to be a worked out version of the former but is not considered a "hallenhouse":

http://upload.wikimedia.org/wikipedia/commons/thumb/d/d2/West-Beemster%2C_Jisperweg_130.jpg/800px-West-Beemster%2C_Jisperweg_130.jpg

and

http://upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Stjelp.jpg/1024px-Stjelp.jpg

The Frankish type, a clear "hallenboerderij":

http://upload.wikimedia.org/wikipedia/commons/thumb/4/4b/Gevel_-_Oirschot_-_20171901_-_RCE.jpg/800px-Gevel_-_Oirschot_-_20171901_-_RCE.jpg


And, just as in Germany, in the original LBK settlement area the farmhouses have changed considerably. Frams consist of houses around a central yard. The "carréboerderij":

http://upload.wikimedia.org/wikipedia/commons/thumb/e/ed/Sibbe-Dorpstraat_45_%284%29.JPG/800px-Sibbe-Dorpstraat_45_%284%29.JPG

Angela
08-07-14, 19:47
@FrankN,
I don't think this is quite the thread for a detailed discussion of Lazaridis et al, and I'm very pressed for time as I'm about to go on “sabbatical” for about a six to eight week period, but I don’t want it to appear as if I am ignoring your post, and, frankly, I am loathe to let what I consider serious misunderstandings of the paper stand for those who may not be very familiar with autosomal analysis.

(I want to extend my apologies to Aberdeen and the moderators for continuing this hijacking of this thread. Please feel free to remove both of these extended posts about Lazaridis et al to a more appropriate venue.)


So, to begin, you stated that I misrepresented the findings of Lazaridis et al. Having said so, it would have been helpful if you had pointed out specific examples where I did that. I think I have been pretty scrupulous about quoting specifically from the text, and rather cautious about extrapolating from it.

As to the validity of the analysis itself, Lazaridis et al is a ground breaking study of the peopling of Europe. I hold it in high esteem, and it seems I am in good company, as 72 of the world's leading population geneticists have signed on as "contributing" authors, including Brenna Henn, Mark Thomas, George Busby, Christian Capelli, Toomas Kivisild, Joachim Burger, Wolfgang Haak (from the group that produced the paper we are discussing) to Qiaomei Fu and Svante Paabo. I hardly think they would have put their names to it if they thought there were serious methodological problems with it. Then, of course, there are the listed co-senior authors, Johannes Kraus and David Reich, although I’m sure as senior authors they have no problems with the conclusions of their own paper. If you wish to pit your expertise in population genetics, the new statistical algorithms that have been developed, and the new statistical tools created to assist in this analysis, that’s of course your prerogative. Far be it from me to suggest that the validity of a study should be determined by an appeal to authority.


The following are just a few thoughts from my non-professional population geneticist’s, “ lay person's” , perspective on the paper and the various tools used in the paper, a lay person who has, however, been reading autosomal analyses and papers since the publication of The History and Geography of Human Genes by Luca Cavalli-Sforza in 1994, (a book that is still invaluable as a primer in this area), so I have, I hope, some familiarity with the issues.

You state that you are concerned that the results from the ADMIXTURE software analysis are not valid because there are not dozens of samples from each population group. More samples are always better. However, autosomal DNA is not like uniparental DNA. There has been so much "mixing" that, from what I have seen, you don't need a lot of samples to get a statistically reliable number for a certain area. There are analyses that address this issue, but I don't have the time to search for them right now. Just as a simple example, there are numerous sample sources for Tuscans. One has 7 samples (HGDP), one has 20 (Hap Map 3) and one has 100 individuals in it (1000 Genomes Project). There is virtually no difference in the autosomal analysis results. For another example, again from the Italian population, the results for the people in the Bergamo HGDP sample are almost identical to the garden variety Northern Italian participants in Dienekes' Dodecad sample.

(Also, to your point about uniparental markers, they are good for tracking individual migration movements, but they are very poor indicators, particularly in the case of yDNA, of autosomal make-up, as I would think you would know. There is the often cited example of some pastoralists in western Sub-Saharan Africa who carry very significant amounts of R1b, and yet they are autosomally practically indistinguishable from surrounding non R1b tribes. Particularly as to your point about the ancient genomes, Loschbour is I2a, and La Brana is “C”, a haplogroup we now associate with eastern Asia. Yet, they form an autosomal cluster. )

Further, as to your point about the analysis with ADMIXTURE, if you re-read the paper and the supplement, it might refresh your recollection as to the fact that the Admixture software is just one of the tools they used to reach their conclusions. They also used PCA analysis, f-statistical analysis and ADMIXTUREGRAPH, unsupervised Mix Mapper and Tree Mix software, plus IBD analysis and chromosome painting analysis to arrive at their conclusions.
.
Now, would it have given the consumers of the paper more information if more populations were tested? The answer is yes, of course. That doesn't invalidate the results, which state broadly that "these analyses allow us to infer that EEF ancestry in Europe today ranges from ~ 30% in the Baltic region to ~90% in the Mediterranean, a gradient which is also consistent with patterns of identity by descent sharing (IBD)(S118) and chromosome painting (S119)”

Those patterns would indicate, in my opinion, that the area from Czechoslovakia (which was tested-.495 EEF), west through Germany and the Netherlands to Britain(which was also tested and had exactly the same EEF score-.495) hovers around a 49-50% rate on average, a fact that is borne out in the results from the calculator, with the number dropping a few points in far northern Germany from what I recall. (Although I have criticized that calculator, Central Europe, going by the correlation between the calculator results and the study results, is not one of the geographic areas of concern. ) Ukrainians score .46, by the way.


Of course, it’s unfortunate that more western and northwestern European populations were not included. I’m sure that they will be included in the future, since, as the authors speculate, that may be why their PCA does not exactly duplicate those of some prior studies. However, that is one, and not a particularly good tool, so this doesn’t in any way invalidate the findings of the paper or the percentages of these ancient genomes that show up in modern populations. It just means we don’t have the figures for some modern European populations.

From the figures that are provided, there seems to be a very narrow range in terms of these scores across the northern European plain and into at least part of the British Isles, which goes from about .46 for Ukrainians for EEF to about .49 in Britain. The number for EEF starts to increase in France, where a more northerly sample is .554 , but France_South is already .675 EEF. The EEF number of about 50% in Central Europe jumps to .715 in northern Italy, .713 in Spanish_North, .712 in Bulgaria and then to .745 in Tuscany, .81 in the Spanish etc. and so on. This all makes total sense in terms of prior autosomal analyses of modern Europeans.

I would be absolutely shocked if there is much difference in the scores for people from different areas on the Northern European plain. That would be contrary to every result I've ever seen from personal or academic autosomal genetic analysis of these groups. There is a great deal of homogeneity in this area, which crosses national borders. Even since the period of the Indo-European migrations, you have the “mixing” that took place in this region as the result of the Germanic migrations, the Slavic migrations, the numerous wars like the Thirty Years War, the migrations brought about by industrialization, and then the dislocations of the First and Second World Wars. Many of these events did not impact populations in Iberia and the Italian peninsula to the same degree. If you've read all the pertinent papers, and followed the personal results as well, I don't understand how you can fail to see this pattern.

For those who have not seen them, these are some graphics of PCA’s based on autosomal results for Europeans that I think nicely illustrate the point:

http://upload.wikimedia.org/wikipedia/commons/2/2b/European_genetic_structure_%28based_on_SNPs%29_PC_ analysis.png

http://2.bp.blogspot.com/_Ish7688voT0/R6Nt4XlfrwI/AAAAAAAAAFA/NzaQHAnUOvI/s1600-h/pc300k.jpg (http://2.bp.blogspot.com/_Ish7688voT0/R6Nt4XlfrwI/AAAAAAAAAFA/NzaQHAnUOvI/s1600-h/pc300k.jpg)


http://s21.postimg.org/6pawkhkjb/russiangwafig3.png (http://s21.postimg.org/6pawkhkjb/russiangwafig3.png)


You also make some statements that I think are very off the mark, given the reading that I have done on this subject. Why on earth, for example, would it be surprising that Icelanders have an EEF component? They are a mixture of settlers from Norway and Ireland, both areas that received genetic input from Neolithic farmers. (Norway has an EEF score of .41 and Scotland has an EEF score of .39. This makes the Icelandic score of .394 perfectly plausible.) The Neolithic farmers also spread to South Asia (a recent paper explored it in depth) and, yes, Africa, in the form of pastoralism, with some speculation that it's connected to the Cushitic speakers, as per your comment about the Afars of Ethiopia, who are pastoralists who indeed speak a Cushite language. Perhaps you haven't gotten around to reading the papers showing a large Southwest Asian autosomal component in Horner populations, some of which could indeed have entered their genomes with such a migration? Ukrainians also have, of course, an EEF component, as do far NWestern Russians, but the latter so called “ Volga Russian” population is a heavily Siberian admixed population, so their numbers will, of course, be quite different. You may also not be aware of the detail that the Algonquin, Cree and Ojibway are some of the most heavily European admixed populations among all Amerindian groups. There is speculation as well that they have a somewhat differential gene flow as shown by their levels of mtDNA “X”. These are all known relationships genetically speaking.

I also don't quite know how to respond to your complaint that Lazaridis et al doesn't tell us the fate of the WHG in the various areas of Europe. (Although they do, contrary to your assertion, speculate that the perhaps 20% WHG in EEF is attributable to the Mediterranean HGs. They obviously need ancient samples from them to be sure. Also, they did indeed try to fit the SHG into the model, but the statistical analysis refuted it.)


This is an incremental process...as more ancient data becomes available, from Mediterranean HGs, from Ancient Near Eastern farmers, from the first farmers in the Balkans, indeed, from Mesolithic samples in the Balkans, from Samara and from the new groups entering the Balkans around 3500 BC, I’m sure this group will adjust their analyses, just as they adjusted their findings as published in Moorjani et al in the Patterson et al paper.

The fact remains that, properly understood, this is a quantum leap forward in terms of genetics population studies, and, from what I have seen, is accepted as such in the academic and hobbyist communities.

For those who haven’t been tracking the analyses produced by the Reich Lab, I would suggest reading the following papers in the listed order:

Moorjani et al: http://www.plosgenetics.org/article/info%3Adoi%2F10.1371%2Fjournal.pgen.1001373
Patterson et al: http://www.genetics.org/content/early/2012/09/06/genetics.112.145037.full.pdf
Lipson et al: http://arxiv.org/pdf/1212.2555v2.pdf

The following improvements in statistical analysis appeared in these papers:



Reich et al, 2009 (Indian Cline; introduction of f-statistics)
Green et al, 2010 (Neandertal genome; introduction of D-statistics)
Durand et al 2011 (elaboration on D-statistics)
Meyer et al, 2012 (high coverage Denisova; enhanced D-statistics)
Pickrell et al, 2012 (Khoisan origins; use of admixture LD with 1-ref)
Reich et al. (2012 (Native American origins; multiple waves of admixture)


The following tools were used in the following papers: Patterson et al (ADMIXTOOLS), Loh et al (ALDER), and Moorjani et al (updates to ROLLOFF)


I would also recommend a careful reading of Ralph and Coop:

http://www.plosbiology.org/article/info%3Adoi%2F10.1371%2Fjournal.pbio.1001555

To explore in detail what happened to the hunter-gatherers in each area of Europe, their number, where and when the admixture with farmers took place, whether it was from HG's who remained in place, or from later repopulation from the north, or with the Indo-European migrations, wasn't the purpose of this paper. On the most basic level, what it is showing is that the genes of the WHGs (Loschbour and La Brana) appear in the genomes of modern Europeans in certain percentages. The same applies to the genes of the Stuttgart sample and the ANE sample. (And the Gok sample and Oetzi himself cluster with Stuttgart) The purpose of the paper is to show that the populations of Europe, not the Middle-East, can best be modelled as a mix of three ancient genome clusters. Other admixtures, when more ancient samples are available, are possible. I can't see how they can be faulted for not answering every detailed question we have about every minor as well as major migration flow and extinction that has taken place in a particular area of Europe over the last thousands of years.

Your comment about them "throwing out" certain groups also shows a fundamental misunderstanding, in my opinion, of this analysis. First of all, the Ukrainians can indeed be fit with the three population model. The Finns, the Mordovians, and the Volga Russians cannot, because they have, in the case of the Finns, for example, according to other autosomal analyses, at least 5-6% "Eastern" admixture. (Siberian/East Asian, the percentages differing slightly by analysis) They weren't "thrown out". Nor were the Sicilians and the Maltese thrown out. They're all still Europeans. It's just that unlike all other Europeans, they don't fit the model because they have too much "other" admixture, which in the case of the Sicilians and Maltese is not yet properly understood, by the way. Said another way, they are not a statistical fit as a combination of Neolithic farmers from the Near East, Western Hunter Gatherers, and an ancient North Eurasian population that is also ancestral to Amerindians. The fact that the model doesn't fit a few groups at the periphery of Europe doesn't mean that the model is invalid. It is valid for the vast majority of Europeans.

I don't have the time to address all your points, nor is this the thread for it, but I think I've posted enough to show that I strongly disagree with your analysis. People will have to make up their own minds after reading all the pertinent papers.

Angela
08-07-14, 19:52
I haven't had a lot of time and this thread fills up with interesting responses. However, I feel I have an obligation to answer this as I, as an amateur, so pretentiously have called for attention to these cultures. I initially learned about from Dutch archeology books, books that I have all currently packed as I am moving. The original Vlaardingen finds, from a culture that seems related if not similar to Swifterband, are on display in Leidens museum of antiquity. They consist of a number of well preserved fish traps and a lot of freshwater fish remains. This is therefore not inconsistent with mesolithic HG continuing freshwater fisheries

The original Swifterband is described here, and in the first map you also see it concentrated around freshwater systems.

http://dissertations.ub.rug.nl/FILES/faculties/arts/2013/i.devriendt/thesis_Izabel_Devriendt___proe_1.pdf

But you want dates off course, in order to check the impact of coexistence. This might be interesting: Hoge Vaart being a typical Swifterband site. Actually, on slightly more than a day's walk distance from the Swifterband site.

www.academia.edu/1231539/Radiocarbon_dating_of_Mesolithic_open-air_sites_in_the_coversand_area_of_the_north-west_European_plain_problems_and_prospects (http://www.academia.edu/1231539/Radiocarbon_dating_of_Mesolithic_open-air_sites_in_the_coversand_area_of_the_north-west_European_plain_problems_and_prospects)

There is quote in it that indicates something happened to HG's though:

Thanks for the links, Epoch, and I will definitely read them carefully. Unfortunately, I won't be posting much, if at all, for the next 6-8 weeks. Perhaps when I return, if this is still being discussed, I can add my two cents. :)

Aberdeen
08-07-14, 20:53
Once people start discussing the more academic aspects of autosomal analysis, etc., I find it best to sit back and read, rather than showing my lack of background in the more esoteric aspects such stuff by commenting too much. But I think that in critiquing such material, it's useful to go back and remind oneself of what the author was and wasn't addressing. And I think it's also useful to remember what certain terms mean - for example, although the early European farmers came to Europe from the Middle East during the Neolithic, we all know that EEF doesn't equate to modern Middle Eastern Farmers, because of genetic changes in that area since the Neolithic. However, knowing that and keeping it in mind are sometimes two different things.

I do think that some of the discussion of Lazaridis et al that has appeared in this thread so far is useful and on topic up to a certain point, but any further analysis of that material might be more useful in a separate thread devoted to that subject. And while I certainly find the lesson material on modern German and Dutch farmhouses to be fascinating, I'm not sure how relevant it is to the thread. A similarity of design with Neolithic structures may mean continuity of of design, but it could also simply be a demonstration of the fact that people who have similar problems to address often create similar designs without continuity. (I think only the Atlantis crowd would argue that the similarity between Egyptian and Mesoamerican pyramids showed a flow of cultural continuity between the two areas.)

The main issue I want to address is that, even if one accepts the view of the authors of the paper that the specific locale shows that the current pattern of mtDNA H in the area was already primarily set 6000 years ago and was only somewhat affected by Bell Beaker and the Bronze Age folk (and I don't agree), that still tells us nothing about Europe as a whole. And yet the paper does appear to be extrapolating on that basis. So even if you agree with its conclusions for that specific area, so what?

I hope you enjoy your vacation, Angela. But I'll miss your learned and well written comments.

epoch
08-07-14, 23:16
Thanks for the links, Epoch, and I will definitely read them carefully. Unfortunately, I won't be posting much, if at all, for the next 6-8 weeks. Perhaps when I return, if this is still being discussed, I can add my two cents. :)

Go away. There is a lot more in life than posting on the internet: Go and enjoy it :)

JS Bach
09-07-14, 06:40
Unfortunately, no. All that I have is the Lazarides study with the Supplemental Information. I browsed it again, but it seems they only published admixture estimates (WHG/EEF/ANE) for Europeans. There are quite a number of hints on the transatlantic gene flow scattered across the various supplements, ...

I don't doubt it. That topic has piqued my interest for some time now. I read Stanford and Bradley's book "Across Atlantic Ice" and have since made lots of posts on the Solutrean Hypothesis thread further down here in this "Ancient DNA Studies" sub-forum. My current thinking is that there may have been three separate waves across the Atlantic in prehistoric times. There are lots of hints out there in this DNA data world. For instance, in this ADMIXTURE breakdown for k=13: https://docs.google.com/spreadsheet/ccc?key=0ArJDEoCgzRKedGR2ZWRoQ0VaWTc0dlV1cHh4ZUNJR UE#gid=24 it lists a sample of 17 Mayans as being: 96.9% Amerindian, 2.1% Mediterranean, 0.8% North European, and 0.2% Siberian. There's also this paper: http://dienekes.blogspot.ca/2013/08/y-chromosome-haplogroup-q-and-native.html that has found ydna Q1b in Central and South America. There's an interesting discussion in the comments on that article, where a commenter says that the closest STR match is from Morocco. It might be a stretch to suggest that a Q1b once had those Mediterranean/North European/Siberian admixture proportions, but you never know - La Brana was ydna C. Sorry to go off topic there.

FrankN
09-07-14, 11:07
@Angela: Here is what I said:

@Angela: I think you (and possibly a few other posters) are over- or misinterpreting Lazarides findings. Or, maybe I better say: Amidst the various highly interesting results Lazarides has delivered, serious methodical flaws of his work tend to be overlooked.
And you are correct - I shouldn't have used the word "misinterpreting", it put a wrong notion to my post. I also shouldn't have addressed that remark specifically to you - it was actually geared towards a number of "conclusions" from Lazarides drawn by various posters, and I lumped your position together with others, which was unfair. My apologies!

I nevertheless stand to the second sentence. There is neither a need to be a genetic expert for that, nor for in-depth technical discussion - Lazaris et al. are pretty open themselves on various methodological shortcomings, as put forward in their Supplemental Materials No. 10, 12, and 14. I just intended to sum up these shortcomings for those here that haven't taken the time to look through all the Annexes.

The main problem is the model itself. A three-way mixture model implies the following:

There has only been one North-Eurasian population inflow. As you state yourself, that is against everything we know about Russians, Chuvash, Finns etc.
There has only been one population inflow from the Near East. Well, their admixture analyses clearly point at two such inflows, one Levantine, and one Caucaso-Gedrosian. Existence of these two, different flows seemed to be generally accepted before they published their study.
There only has been one European LGM refuge. That would actually have been worthwhile to be examined. Unfortunately, they haven't. Instead, they overloaded their population sample with Iberians and Western Mediterraneans, which automatically pulled the European HG towards the Cantabrian refuge [The problem with the sampling sizes is not the reliability of the results for each country/ region. The problem is that all of the various methods that are used try to find the best fit to all samples. And if over 25% of all samples are from Spain or Sardinia, but 0% from Central Europe (figures taken from p.77 of their Supplemental Analysis), you get a result that fits well in the Western Mediterranean, but not necessarily elsewhere.]

But, didn't they prove that their three-way mixture works? Look at their Supplementary Information 12 (p. 88):


The finding of at least 4 ancestral populations is seemingly at odds with our modeling approach which assumes 3 populations, so we sought to determine the cause of the added complexity. We removed each of the populations of R in turn and repeated the analysis over all 23 subsets. If the 4th ancestral population has largely affected only one of the populations in R, the evidence for four populations should disappear or greatly weaken when one of the affected population is removed.
We find that the P-value for rank 2 remains <10`\ for 22 subsets, but for the subset R-{Spanish} it becomes 0.019, which is not significant after correcting for multiple hypothesis testing.
(..)
A different approach is not to start with the full set of populations, but to choose a “small” R as:
R = {Belorussian, Bulgarian, Croatian, Czech, English, French, Hungarian, Icelandic, Norwegian, Orcadian, Sardinian, Scottish}
This set of populations includes members of the main south-north European cline (Fig. 1B), and avoids most Mediterranean and Baltic populations where there may be more complex history involving Near Eastern, African, or East Eurasian ancestry.
(..) We find that rank 1 is excluded (P < 10-12), and thus there must be at least 3 source populations related to the outgroups even for this restricted set of European populations (Table S12.3)
What they have proved is that one can fit the 12 populations above into a 3-way WHG / EEF / ANE split. For these 12, the model is valid. For all the others, it is at best a statistical not significant hypothesis, at worst not tested at all.

So, what happens when you try to squeeze six (maybe seven, a bit from North Africa seems to also have reached the Western and Southern Mediterranean Europe) admixtures into three groups? You end up with some strange proxies that try to combine 2-3 fairly unrelated components into one "hyper component". And suddenly, Sardinians are the prototypical EEF. Stupid as I am, I always thought yDNA I2a1a to be an indicator of WHG from the Cantabrian refuge...

Why do I think that neglecting Central Europe might be problematic? Well, wasn't it you who just told us a few days ago that Tuscans are quite different genetically from other Italians? The map below (btw pertinent to the original focus of the thread) indicates where, in the absence of autosomal data, I would expect to find elevated HG concentrations - for example in the Middle Elbe- Saale region, or in northern Sweden. Both are places that are not too far away from Motala!
http://cache.eupedia.com/images/content/Haplogroup-I2b.gif
If there had been a Black Sea LGM refuge, I would also expect some autosomal traces of it to still pop up, maybe in Moldova or Romania (both not sampled). Getting from the Black Sea by boat to the Baltic Sea and then into Central Sweden shouldn't have been that difficult for a Mesolithic HG (actually, the Baltic Sea wouldn't even have been there initially). In fact, it is probably much easier to get into Central Sweden from the Black Sea than from Cantabria. Unfortunately, Lazarides et al didn't investigate whether there may have been more than one European LGM refuge, so this is all speculative.


I would be absolutely shocked if there is much difference in the scores for people from different areas on the Northern European plain. That would be contrary to every result I've ever seen from personal or academic autosomal genetic analysis of these groups. There is a great deal of homogeneity in this area, which crosses national borders.
Do you have a link to these analyses (especially, of course, to any relating to the Elbe-Saale region and its periphery)? What I have seen so far for Germany (but data is pretty scarce) doesn't at all point to homogeneity, but to strong regional differentiation (and mind you, not all of Germany, and also not all of the Elbe-Saale region, is on the Northern European Plain).

Anyway, enjoy your holiday. I don't expect an answer now, we can continue discussion when you are back.

@Aberdeen:
And while I certainly find the lesson material on modern German and Dutch farmhouses to be fascinating, I'm not sure how relevant it is to the thread. A similarity of design with Neolithic structures may mean continuity of of design, but it could also simply be a demonstration of the fact that people who have similar problems to address often create similar designs without continuity. (I think only the Atlantis crowd would argue that the similarity between Egyptian and Mesoamerican pyramids showed a flow of cultural continuity between the two areas.)
I think we are trying to establish whether there has been some kind of population inflow between LGM and Bronze Age, and from where. Our knowledge of the middle- and late Neolithic is pretty thin - just look at Maciamo's migration maps for that period (hint: There aren't any!) Wikipedia is also a mess in that respect - sometimes the region in question is MK, sometimes stroke-ornamented culture, sometimes Lengyel, sometimes TRB, and when you start checking German papers, it gets even messier (among others as the region was divided by the Iron Curtain, and West and East German archaeologists obviously didn't communicate much with each other during that period) So the main message of the study seems to be: "Hey guys, something important has been going on during that period that we all have overlooked so far, but we also don't really have a clue what that could have been".
I am trying to figure out what might actually have happened. And since I am better in looking at houses than in looking at pot sherds (and also think houses may tell us more about how the people lived and where they might have come from), I thought I place a kind of baseline here. Whether housing changed, and in which way, and if any of that will really be meaningful - no idea yet..


The main issue I want to address is that, even if one accepts the view of the authors of the paper that the specific locale shows that the current pattern of mtDNA H in the area was already primarily set 6000 years ago and was only somewhat affected by Bell Beaker and the Bronze Age folk (and I don't agree), that still tells us nothing about Europe as a whole. And yet the paper does appear to be extrapolating on that basis. So even if you agree with its conclusions for that specific area, so what?
First of all, the study never claimed to speak about Europe at a whole, they speak about Central Europe. But if some major, so far largely unnoticed change has been going on in a region that somehow is being linked to all other major cultures from Normandy to Eastern Poland, and down to the Danube, some of that change should also have affected those related cultures. And those people with new and additional mtDNA H must ultimately have come from somewhere..

@epoch: Thanks for the links. The first paper seems to indicate that the main sites near the Ijsselmeer were submerged sometimes around 4,300 BC. You don't happen to by chance have seen any maps on changes of the Dutch North Sea coast over that period? Those Doggerland maps only date to 9,000- 5,000 BC, and are anyway a bit speculative and with little detail about the continent. While there is quite good German cartography tracing land loss since the 2nd century AD, I haven't yet found any research or modelling on the times before.
http://log.doggerland.net/wp-content/uploads/2010/10/doggerland5.png

epoch
09-07-14, 21:48
@epoch: Thanks for the links. The first paper seems to indicate that the main sites near the Ijsselmeer were submerged sometimes around 4,300 BC. You don't happen to by chance have seen any maps on changes of the Dutch North Sea coast over that period?


5000 year ago (Purple is peat):

http://www.historischekringdebilt.nl/bodemonderzoek/geologie/Paleografie%20Laat-atlanticum%205100%20j%20gel.jpg

3000 year ago (A lake starts to appear. The name in the Middle Ages of the expanding lake was "Almere", all lake. After a while it turned to a estuary, then a sea, the Zuiderzee. After this was dammed parts were reclaimed which is why they made the finds. Currently one of the new cities in these polders is called "Almere")

http://www.historischekringdebilt.nl/bodemonderzoek/geologie/Paleografie%20Subboreaal%203000%20j%20gel.jpg


EDIT: http://www.historischekringdebilt.nl/bodemonderzoek/geologie/nederland.html

Greying Wanderer
11-07-14, 05:24
"I would be absolutely shocked if there is much difference in the scores for people from different areas on the Northern European plain. That would be contrary to every result I've ever seen from personal or academic autosomal genetic analysis of these groups. There is a great deal of homogeneity in this area, which crosses national borders."

The I2a2 hotspot on the DNA map above would be the Harz mountains so just south of the Northern European plain.

edit: above may be an unwarranted assumption, see post below from FrankN

FrankN
12-07-14, 00:07
"I would be absolutely shocked if there is much difference in the scores for people from different areas on the Northern European plain. That would be contrary to every result I've ever seen from personal or academic autosomal genetic analysis of these groups. There is a great deal of homogeneity in this area, which crosses national borders."The I2a2 hotspot on the DNA map above would be the Harz mountains so just south of the Northern European plain.
That's where the problem starts. The hotspot on the map would actually be south of the Harz, in Thuringia, somewhere around Erfurt. Several non-scientific German texts talk about the hotspot extending from the Harz northwards to around Luneburg. I have been looking for more details for quite some time already, but it seems this is al based on the old study by Rootsi et al (2004), which also (at least not the main body) doesn't give specific information on the sampling area, but instead just comes up with a map showing the European hotspot somewhere in Central Germany. Harz of course sounds good, because the Lichtenstein cave is located nearby (though neither to its south, nor to its north, but to its west).http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1181996/

Anyway, after Rootsi, it appears that nobody has again looked systematically into the I2a2 distribution inside Germany. The regionalised studies that I have seen typically lump all yDNA I together, which isn't really helpful, as I1 is of course very common in Northern Germany (and, btw, also not really homogenically distributed over an east-westerly cline, say Central Ukraine to Paris basin). A recent comparison between German border regions and Poland shows I2a2 at 2% in Bavaria and 5% in Mecklenburg, that's about it.

I don't know whether it is any better in the Netherlands - Rootsi has them at 10% I2a2, Underhill (2007) at 5.4%, but Maciamo in his map appears to have only used the second figure. Now, one might think that such a difference between two studies could indicate quite a differentiation inside the Netherlands, but I am not sure if anybody ever looked into it...

For the Czech republic, I have seen figures ranging from 1% (Rootsi) to 6.4% (Underhill). I guess it all depends how near or far away your sampling locations are from the "big river" (Elbe, Rhine) and/or loess plains, but that's just a guess...

Greying Wanderer
12-07-14, 01:52
@FrankN

"The hotspot on the map would actually be south of the Harz, in Thuringia, somewhere around Erfurt. Several non-scientific German texts talk about the hotspot extending from the Harz northwards to around Luneburg."

Ah, that was just me assuming then.


"Now, one might think that such a difference between two studies could indicate quite a differentiation inside the Netherlands, but I am not sure if anybody ever looked into it..."

Yes I wonder if prior assumptions effect this more generally? If a researcher's premise is this dna migrated from somewhere else (and maybe a lot of it did) then the exact current distribution might not seem to matter much. However if a researcher is looking for possible indigenous survival in specific likely areas then the precise location of hotspots of specific clades matters a lot.

FrankN
12-07-14, 18:17
@FrankN

"The hotspot on the map would actually be south of the Harz, in Thuringia, somewhere around Erfurt. Several non-scientific German texts talk about the hotspot extending from the Harz northwards to around Luneburg."

Ah, that was just me assuming then.


"Now, one might think that such a difference between two studies could indicate quite a differentiation inside the Netherlands, but I am not sure if anybody ever looked into it..."

Yes I wonder if prior assumptions effect this more generally? If a researcher's premise is this dna migrated from somewhere else (and maybe a lot of it did) then the exact current distribution might not seem to matter much. However if a researcher is looking for possible indigenous survival in specific likely areas then the precise location of hotspots of specific clades matters a lot.



That's exactly the question I wanted to bring up. As British and Irish are much more into individual DNA testing than Germans (or central Europeans in general), we now have a pretty good picture of yDNA differentiation inside the British isles, and it has turned out that patterns are far from being homogenous or following simple geographical clines. The same is becoming apparent in the Alps. As to I2a-Din, what was once assumed to be a West Balkan pattern has now been shown to have specific concentrations on Dalmatian islands and in some valleys in Herzegovina and SW Bosnia, while I2a-Din is far less present along the Danube. If researchers started to look into the Carpathians, or the Massif Central, at a geographic resolution similar to the one that has already been used in Spain and Italy, I wonder what they would find.

That leads of course into a general problem. Big cities tend to be excluded for methodological reasons (you ideally want your test sample to have been resident in the area for at least three generations). Depending on the country in question, this implies that something like 50-70% of the current population are a priori excluded. That's ok as long as one assumes that mobility is unrelated to genetics, but is that really the case? Mobility has to do with education, with has to do with parental professional background, which may relate to socio-economic patterns that already have been established long ago. Then there is the issue of mining, which has a millennium-long tradition in several of the areas that have been studied intensively (Sardinia, NW Spain, Bosnia-Herzegovina), or not that intensively (Harz, Erzgebirge, Carpathians). Field-crop farming along the rivers vs. specialised tree-crops (viticulture, olives etc.) vs. transhumating cattle-herding vs. fishing? We are starting to getting some glimpses of a possible differentiation, especially on the Balkans, but so far, systematic evaluation hasn't been done. However, when looking at the archaeological record, the assumption that EEF would rather go into field-crop farming, while HGs might switch towards cattle herding or specialise on fishing is anything but implausible.

We are rightfully happy about any ancient DNA coming in. But do we really know which "culture" that DNA may be assigned to? All available evidence points to Ötzi the Iceman not being an EEF, but engaged in metal exploration and/or trade. A spectacular "royal grave" with lots of precious artefacts is uncovered next to simple collective burials. Surely the local leader. But, take for example the Amesbury Archer, who was born somewhere in the NW Alps. Was he really the local leader, or rather a long-distance trader, who happened to die far away from his home, and was buried together with all of his belongings since there were no relatives nearby to claim the heritage? Unlikely that a stranger will be given an exquisite burial? Check out this case from the High Medieval:
http://en.wikipedia.org/wiki/Nicholas_the_Pilgrim

In summary - while we all agree that more DNA testing, and especially of ancient DNA, will be required, we may still be much too confident about the state of knowledge that apparently has already been achieved. What we have so far consists of relatively regionalised DNA data around the Mediterranean and on the British Isles, but comparatively poor sampling of central & eastern Europe, which excludes cities (including those that have been existing already for millenniums). The data is analysed under the assumption of homogenous regional clines, without paying attention to (pre-)historic economic specialisation of the sampled regions. That analysis is combined with ancient DNA that is again just grouped according to simplified geographical and economic criteria (Med / West/ North; HG vs. EEF). And ultimately, we are applying the results of this genetic analysis to an archaeological record that is far from being complete, especially when it comes to the middle and late Neolithic.

I realise that I have been getting a bit off topic again - maybe we should open a new thread for a general discussion on the limitations of current genetic models. Let me also add that I in general prefer an incomplete model to not having a model/analysis at all. I am grateful for all those new studies, including the Lazarides one, but we need to be aware that much more work may be needed until we arrive at a half-way complete picture.

Greying Wanderer
13-07-14, 12:28
"So I had the impression that mtDNA should follow autosomical DNA more than Y-DNA."

These kind of mismatches can provide clues to what happened. IIRC there are countries in South America (Argentina?) where the mtdna is largely native while the autosomal dna is largely European. This is a result of multiple waves of mainly male migration so the first wave marry native women resulting in 100% native mtdna and 50/50 autosomal but then as further male migrants arrive some of the mixed women marry mixed men and some marry the new arrivals. Over time the European autosomal percentage keeps going up while the percentage of native mtdha stays the same. In theory a situation like that could eventually lead to 100% native mtdna and 90%+ European autosomal.

MOESAN
18-07-14, 15:54
That's pretty high level of "contamination". Can't wait for Ancient Near Eastern sample to clean up this issue. I think there might be some EEF farmer gene flow in Loschbour WHG admixture. It would be a surprise if Loschbour was completely pure Hunter Gatherer at 6k BC, only 150 km and 500 years from Stuttgart farmer.

why would it be so surprising, Lebrok?
6000 BC is old enough - the 2 pop's didn't melt together so easily - agriculture didn't take a huge place in N and NW Europe before about 5000/4500 -
let's remember Michelberger-"neolithical"-s had seemingly a strong remnant of mesolithic genes