Limitations of current genetic models

Angela

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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.
 
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
 
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@Angela

Also, there is this:

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

Dienekes Article said:
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.
 
@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.
 
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.
 
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.
 
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:
  1. 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);
  2. 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;
  3. 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;
  4. 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;
  5. Convergence zone of (Proto-Celtic) Urnfield & Hallstatt cultures, and (Proto-Germanic) Jastorf culture;
  6. 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).
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

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.
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.).
 
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:
Rannersdorf_Grundriss_LBK_Sauer_und_Schwarz%C3%A4ugl_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:
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.
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:
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:
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
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:
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
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.
 
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]


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%.
 
@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/thread...-Aryan-split-of-IE/page11?p=434227#post434227.
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?

  1. 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).
  2. 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?
  3. 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.
  4. 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/thread...-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.
  5. 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.
  6. 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).
  7. 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:
  1. 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!).
  2. 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).
  3. 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.
  4. 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!
  5. 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
  6. 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.
 
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:
  1. 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);
  2. 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;
  3. 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;
  4. 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;
  5. Convergence zone of (Proto-Celtic) Urnfield & Hallstatt cultures, and (Proto-Germanic) Jastorf culture;
  6. 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).
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.
 
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/thread...lavic-language?p=433040&viewfull=1#post433040
http://www.eupedia.com/forum/thread...anic-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.
 
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?
 
@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/thread...-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:
quote_icon.png
Originally Posted by Jean M
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.
 
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.
 
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_Börde
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
 
  1. 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: 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).
 
This map may help to put some (though not all) of Brotherton's findings into context:
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).
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.
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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..
 
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