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Jean M
11-21-2013, 08:01 PM
Michael J Sikora, Vincenza Colonna, Yali Xue, Chris Tyler-Smith, Modeling the contrasting Neolithic male lineage expansions in Europe and Africa, Investigative Genetics, 2013, 4:25. http://www.investigativegenetics.com/content/4/1/25/abstract


Background

Patterns of genetic variation in a population carry information about the prehistory of the population, and for the human Y chromosome an especially informative phylogenetic tree has previously been constructed from fully-sequenced chromosomes. This revealed contrasting bifurcating and starlike phylogenies for the major lineages associated with the Neolithic expansions in sub-Saharan Africa and Western Europe, respectively.

Results

We used coalescent simulations to investigate the range of demographic models most likely to produce the phylogenetic structures observed in Africa and Europe, assessing the starting and ending genetic effective population sizes, duration of the expansion, and time when expansion ended. The best-fitting models in Africa and Europe are very different. In Africa, the expansion took about 12 thousand years, ending very recently; it started from approximately 40 men and numbers expanded approximately 50-fold. In Europe, the expansion was much more rapid, taking only a few generations and occurring as soon as the major R1b lineage entered Europe; it started from just one to three men, whose numbers expanded more than a thousandfold.

Conclusions

Although highly simplified, the demographic model we have used captures key elements of the differences between the male Neolithic expansions in Africa and Europe, and is consistent with archaeological findings.

greystones22
11-21-2013, 10:06 PM
Interesting that this paper deals with the basal R1b group, rather than the later expansions under L11. I would have thought the more explosive growth happened more recently than they suggest e.g after P312, thats where the tree starts to get many many more surviving parallel branches, in some regions.

Wing Genealogist
11-21-2013, 10:40 PM
The authors of this study did not have access to Hammer's research (presented at the FTDNA conference, expected to be publish "soon"), which showed the R1b expansion into Europe happened after the Neolithic Revolution. However, that is really a relatively minor point (in terms of what CTS et.al. were studying).

Yes, they could easily have chosen L11/P310 as the start of the expansion, but the average "professional" is still way behind in their understanding of the haplotree.

One of the interesting points of this study (in respect to R1b) is the fact the initial growth was limited to only a couple of male individuals, and within a "few" generations, "expanded more than a thousandfold". This lends more evidence to the thought of the quick cascade of SNPs below L11/P310 and that the origin of the U106 & P312 SNPs may have actually been close cousins. It also makes it clear that dating the clades immediately below U106 & P312 will be very tricky, as they occurred so closely together.

alan
11-22-2013, 01:01 AM
This is not really a shock as it is just a more extreme version of what many people in this hobby had already noted in terms of a real explosion from L11 down with many geographcally pattern early divisions having dates within in century or so of L11 - indicative of both demographic explosion and geographical expansion.

These three men to produce 3000 in a few generations wouldnt have time to do anything else!! I think that is pushing credibility a little. They would need to have had something like 20 kids spread across a number of wives, 10 of them sons, and this pattern continued with every son surviving and the same happening each generation, for them to pull that of in a few generations. Given the birth mortality rate, short lifespans etc they would have had to have had a very large amount of wives and concubines.

Regardless of detail it does tell us something about their social structure. Maybe this is simply the genetic signal of a very steep hierarchical society appearing. For a few men to hog so much of the mating potential its hard to imagine anything else. Clearly that sort of pace is not compatible with either the pace of the spread of farming or the sort of Neolithic societal structure that is generally imagined. There also has to be an element of threat of violence towards the other males and the attraction of wealth to females. However you look at it, a pattern of going from 3 or 4 men to 1000 times that in a few generations is a genetic signal of resource hogging hierarchy of some sort.

The other upshot of this is the difficulty in detecting small groups archaeologically. You would not see the first generation or two and archaeological visibility might only occur around the fourth generation or later. By then some degree of blending in with existing societies will have occurred. However, even if the arrival is blurred by initial invisibility and the effect of small numbers arriving in much bigger populations leading to cultural absorbtion, what should be visible if this demographic explosion was sustained for a long period is the rise of hierachical or warrior type behavours in the archaeological record. Even if this means no 'arrival' horizon is likely to be visible, the extreme change in breeding behavour should become visible after say 4 generations in some indicators of a hierarchical and/or warrior society.

However, as I have often commented, L11 and its early clades would surely not have been in any position to throw their weight about for centuries given that even after a century or so they probably only amounted to several thousand men spread across an entire continent. Just image in say the isles there were 500 of these men spread across the whole area thinly for example. They simply must have been welcomed by the locals. They would presumably have had to have something major to offer that people wanted.

That is why the L11-beaker model has to be seen as very small groups who had some overwhelming attraction to the locals. That can only be something to do with their metal networks. They must have been extremely prestigious to have also led to their influence on societies.

Now I do not lightly dabble in linking Irish mythology to history as its often done very badly, but it is interesting that in later Irish society the highly skilled craftsmen as well as Druids, poets etc were considered Aes Dana and were protected across tribal boundaries. Perhaps an echo of the way beaker people and their successors could once freely penetrate societies across Europe. The name Aes Dana is also linked to the Pantheon of Irish gods, also known as the Tuatha de Danan.

http://en.wikipedia.org/wiki/Tuatha_D%C3%A9_Danann

So, this class was not only ranked among nobles but was seen as sacred and somehow linked to the gods. Its also interesting that several of the gods of this group are characterised by being masters of crafts, arts etc. Finally the name of both the pantheon of Irish (many pan-Celtic) gods and this sacred class of people linked them to the 'people of the goddess Danu'. Is this some kind of very distant echo of the origins of this sacred class. Danu as a goddess appears in Welsh mythology as Don and the name has parallels in Vedic. Could this reflect a distant time when a group of people associated with crafts, skills, learning etc with gods who reflected their values appeared and that this remained echoed in the Celtic gods, especially in Ireland. The beaker people? It is interesting, although not something new, to observe just how sacred people involved in metal crafts and learning were both as a class of people and in the Celtic pantheon. It at least may echo what was originally held in very high esteem. This is not unique. As far as I recall it has been suggested that a similar phenomenon of sacred metalworkers was present in Vucedol culture. Certainly out of the surviving mythologies it does seem that the Celtic pantheon had an usual emphasis on crafts, especially metalwork.

Generalissimo
11-22-2013, 02:20 AM
The authors of this study did not have access to Hammer's research (presented at the FTDNA conference, expected to be publish "soon"), which showed the R1b expansion into Europe happened after the Neolithic Revolution.

They most likely did, because everyone has access to Eupedia. :biggrin1:

Wing Genealogist
11-22-2013, 10:33 AM
They most likely did, because everyone has access to Eupedia. :biggrin1:

While everone does have access to Eupedia, and the maps on that site, they did not have access to Hammer's analysis of the information. Hammer analyzed the ancient DNA data found in Western Europe, and showed where none of the DNA during the Neolithic Period in W. Europe was from Haplogroup R1b, and (from the top of my head, so my percentages may be off a bit) roughly 85% or so was Haplogroup G2a. He thus concluded G2a was likely the haplogroup which entered W. Europe during the Neolithic, and R1b (most likely L11/P310) entered W. Europe sometime after the Neolithic.

The study by Chris Tyler Smith et al. and Hammer's study do mesh quite well. They both predict a late arrival of R1b into Western Europe as well as a quick dominance (as expressed by its very high percentage) over the area. It is possible the origin of L11 pre-dated their push into Western Europe, and their explosive growth was a contributory cause of their migration.

As I stated elsewhere, Hammer did not even speculate when/how/etc. R1b came into Western Europe, but when he questioned the audience on their feelings, the consensus was that it likely occurred during the Bronze age, and the possible benefit of advanced weaponry (as well as military tactics). It should be noted current research suggests mtDNA in Western Europe (primarily Haplogroup H, but also J and possibly others) are believed to have a Neolithic origin, so the spread of R1b later did not also bring in a major change in mtDNA lineages (which also suggests a military conquest, rather than a major migration).

Jean M
11-22-2013, 11:04 AM
This is not really a shock as it is just a more extreme version of what many people in this hobby had already noted in terms of a real explosion from L11 down with many geographically pattern early divisions having dates within in century or so of L11 - indicative of both demographic explosion and geographical expansion.

The rapid expansion of R1b has not just been noted by people in the hobby. It has been mentioned in previous published studies e.g. 1000 Genomes Project Consortium 2010, a map of human genome variation from population-scale sequencing (http://www.nature.com/nature/journal/v467/n7319/full/nature09534.html), Nature, 467, 1061-723:


A striking pattern indicative of a recent rapid expansion specific to haplogroup R1b was observed, consistent with the postulated Neolithic origin of this haplogroup in Europe

Myres et al 2011, A major Y-Chromosome haplogroup R1b Holocene era founder effect in Central and Western Europe (http://www.ncbi.nlm.nih.gov/pubmed/20736979), Eur J Hum Genet, 19(1), 95-101.


The phylogenetic relationships of numerous branches within the core Y-chromosome haplogroup R-M207 support a West Asian origin of haplogroup R1b, its initial differentiation there followed by a rapid spread of one of its sub-clades carrying the M269 mutation to Europe.

Wei et al., 2013, A calibrated human Y-chromosomal phylogeny based on resequencing (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3561879/), Genome Research, 23(2), 388-95.


node representing the major European Y lineage, R1b, to 4000–13,000 yr ago, supporting a Neolithic origin for these modern European Y chromosomes... The third internal node was that of R1b, a well-documented expansion in Europe, but with a much-debated time depth. Here, we estimate a time of 4,300–13,000 yr ago, the most uncertain of the dates. Despite the range of estimates, all these dates favor a Neolithic (Balaresque et al. 2010), more than a Paleolithic (Semino et al. 2000) or Mesolithic, expansion of this lineage. ... Nevertheless, the rapid expansion of R1b ... in Europe

The Sikora et al 2013 model aims to get to grips with detail. How well it does it, I leave others to judge. I am not much of a number-cruncher myself. But I do think it gets across what this means in terms of the migrating numbers.

Generalissimo
11-22-2013, 11:06 AM
It should be noted current research suggests mtDNA in Western Europe (primarily Haplogroup H, but also J and possibly others) are believed to have a Neolithic origin, so the spread of R1b later did not also bring in a major change in mtDNA lineages (which also suggests a military conquest, rather than a major migration).

Have you seen this current research? Because I don't think Hammer has.

http://www.sciencemagazinedigital.org/sciencemagazine/11_october_2013?pg=134#pg131

Don't worry about logging in. Just get rid of that pop up and then use the arrows to move from page to page.

What this study shows is that there were moderate to significant shifts in mtDNA with every archeological culture that appeared in Central Europe. These shifts were most significant during the time when R1b is supposed to have entered Europe, which means that R1b had to come with a relatively distinct mtDNA gene pool. So Hammer is wrong when it comes to that, and he's probably wrong about other things too.

Jean M
11-22-2013, 11:46 AM
What this study shows is that there were moderate to significant shifts in mtDNA with every archeological culture that appeared in Central Europe. These shifts were most significant during the time when R1b is supposed to have entered Europe, which means that R1b had to come with a relatively distinct mtDNA gene pool.

Not exactly. What we see in the Brandt study is


The background (shown by previous studies). LBK farmers arrived in the Mittelelbe-Saale region with people carrying almost exclusively mtDNA haplogroups new to that region and with origins in the near East: H, J etc not seen in Mesolithic Europe, which was dominated by U5 and U4.
The clever bit. These farmers were not the direct and complete ancestors of farmers of the Late Neolithic cultures which followed the LBK in the same region. This is big news for most archaeologists, who had assumed that once farming arrived in a particular place, the farmers just stayed put and bred more farmers. It is only now percolating through academia that there was a boom and bust sequence in the LBK. So some new influx with Funnel Beaker, which fits my proposal in AJ that Funnel Beaker came from the Balkans.
Yet more new arrivals with Corded Ware and Bell Beaker, who carry a mixture of Mesolithic and Neolithic mtDNA haplogroups. Brandt et al make guesses about where these people came from. Some of said guesses are fine. They link CW to a "genetic influx into Central Europe from the East, likely influenced by Kurgan cultures." But other guesses are based on the old ideas (undermined by recent papers) that subgroups of H spread from Iberia, which they tie to Bell Beaker, because they have got the message from radiocarbon dates that the first BB pottery came from Portugal. But BB pottery was made by the descendants of people who had brought copper-working to Iberia a bit earlier from eastern Europe. Brandt et al declare that the "BBC is distinguished from the CWC by the absence of haplogroup I and U2 and an
overwhelmingly dominant genetic signature of haplogroup H". But they admit that today H dominates all over Europe. That includes the whole area once Corded Ware, which was certainly not all taken over by Bell Beaker! Fact is that the higher level of H in the Bronze Age is a Europe-wide feature, according to another study which combined all aDNA studies available at the time.



Everything will get clearer with more ancient Y-DNA.

greystones22
11-22-2013, 12:15 PM
The rapid expansion of R1b has not just been noted by people in the hobby. It has been mentioned in previous published studies e.g. 1000 Genomes Project Consortium 2010, a map of human genome variation from population-scale sequencing (http://www.nature.com/nature/journal/v467/n7319/full/nature09534.html), Nature, 467, 1061-723:



Myres et al 2011, A major Y-Chromosome haplogroup R1b Holocene era founder effect in Central and Western Europe (http://www.ncbi.nlm.nih.gov/pubmed/20736979), Eur J Hum Genet, 19(1), 95-101.



Wei et al., 2013, A calibrated human Y-chromosomal phylogeny based on resequencing (http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3561879/), Genome Research, 23(2), 388-95.



The Sikora et al 2013 model aims to get to grips with detail. How well it does it, I leave others to judge. I am not much of a number-cruncher myself. But I do think it gets across what this means in terms of the migrating numbers.

Thanks for this summary of the published material. To be honest the number of samples all 3 papers have used are not sufficient to address with precision the tree structure and phases of population growth. This latest paper applies a new modelling approach to the problem, but the modelling can only be used to fit the data in their tree, which is based on only 6 (!!!) R1b Y chromosomes.

I am also not a mathematician, but my feeling is that the data which is used to build the model influences the models that are generated and validated by simulation to match the tree.

So my feeling is that if they had a tree with >100 L11+ chromosomes (as found in the 1000 genomes project, albeit with lower coverage) they would have built a very different model, with a different result.

Generalissimo
11-22-2013, 12:18 PM
Brandt et al declare that the "BBC is distinguished from the CWC by the absence of haplogroup I and U2 and an
overwhelmingly dominant genetic signature of haplogroup H". But they admit that today H dominates all over Europe. That includes the whole area once Corded Ware, which was certainly not all taken over by Bell Beaker! Fact is that the higher level of H in the Bronze Age is a Europe-wide feature, according to another study which combined all aDNA studies available at the time.

mtDNA H didn't reach modern levels in Eastern Europe before the Iron Age. You should know that. It actually only became increasingly more common there due to secondary Indo-European expansions to the east from Central Europe and the Baltic, close to where the eastern Bell Beaker phenomenon was present.

alan
11-22-2013, 12:43 PM
I think though the possibility of a middle Neolithic wave should have been considered by archaeologists as soon as Eversheds work on the spread of dairying emerged. I cant remember how long ago that was but I was at a conference at qub where he presented it before he published them and it seemed probably that there was some secondary input from Anatolia.

I have no idea if that links to r1b. In some ways its tempting but in other ways it does work as its too early and dairying even reached Britain and northern Europe by 4000BC. It does raise the question if it wasnt R1b then what was it. Just more G?

alan
11-22-2013, 12:52 PM
I do think it lazy of some of these studies to link beaker and R1b and paint P312 as an out of Iberia thing while glossing over the fact that Iberia is at the wrong end of Europe when you look at the upstream bits. Jean has suggested how that might have happened but everyone else publishing just seems to skip over the awkward aspect of much of the length of Europe separating upstream R1b from P312.

I just wish there was a bit more discussion in papers about the older forms of R1b in eastern Europe. Also M73 is crying out for some sort of study. If we understood it a bit better we would understand the possibilities for M269 a lot better too IMO. I dont even recall any attempt at a Eurasian map for M73. I could be wrong but I dont think anyone has done one. Maybe that is a project for Macciamo. The population studies are building up now so it must be getting closer to be able to make a map.

Jean M
11-22-2013, 01:20 PM
mtDNA H didn't reach modern levels in Eastern Europe before the Iron Age.

I think that's more or less right, though not just in Eastern Europe. It seems to rise gradually everywhere, though we don't have enough samples yet to be making firm statistical statements. It is possible that Iron Age migrations had an impact, but mtDNA H confers an advantage in recovery from sepsis, which may well have gradually increased its levels in Europe over the millennia, particularly after the various rounds of plague that started with the Justinian one.

MtDNA H is not linked to just one archaeological culture as you know very well. H arrived in Europe in the Neolithic. It was all over the place. There was H in the Dnieper-Donets Culture, which we can deduce was absorbed as this culture took on farming wives along with farming. We can further deduce that this was one part of the mixture that spread Indo-European languages. The statistical level of H in this or that sample is not vitally important. I can see the temptation to make much of it, but we should try to resist. Breaking down H into subclades and trying to track back to the origin of specific subclades in aDNA will be more informative as to origins.

Jean M
11-22-2013, 01:39 PM
I am also not a mathematician, but my feeling is that the data which is used to build the model influences the models that are generated and validated by simulation to match the tree.


Couldn't agree more! It makes me wary of models.


So my feeling is that if they had a tree with >100 L11+ chromosomes (as found in the 1000 genomes project, albeit with lower coverage) they would have built a very different model, with a different result.

Interesting. Any guesses as to outcome?

newtoboard
11-22-2013, 02:49 PM
I think that's more or less right, though not just in Eastern Europe. It seems to rise gradually everywhere, though we don't have enough samples yet to be making firm statistical statements. It is possible that Iron Age migrations had an impact, but mtDNA H confers an advantage in recovery from sepsis, which may well have gradually increased its levels in Europe over the millennia, particularly after the various rounds of plague that started with the Justinian one.

MtDNA H is not linked to just one archaeological culture as you know very well. H arrived in Europe in the Neolithic. It was all over the place. There was H in the Dnieper-Donets Culture, which we can deduce was absorbed as this culture took on farming wives along with farming. We can further deduce that this was one part of the mixture that spread Indo-European languages. The statistical level of H in this or that sample is not vitally important. I can see the temptation to make much of it, but we should try to resist. Breaking down H into subclades and trying to track back to the origin of specific subclades in aDNA will be more informative as to origins.

The Dnieper-Donets mtDNA looked pretty Eastern overall though. Btw do you know if anyone is ever going to test Y-DNA from Dnieper Donets?

alan
11-22-2013, 04:10 PM
I think for those interested in R1b and exploring its potential hiding place in the Neolithic, testing a whole range of pre-Yamnaya cultures from the western end of the steppes between the Don and Bug is important: Dnieper Donets, Bug Dniester, Sredny Stog and several other spring to mind. I am not saying it has to have been there but its a possible zone that needs to be checked out. It could have come from somewhere else. The western steppes, the north Caucasus, NW Iran (the last two were linked) are all zones where farming arrived late enough to correlate. What we really ideally would want to find in ancient DNA is something like P297* which is the state the ancestor of most Eurasian R1b other than was in during the Neolithic. As none of this had been found despite it spanning 9000-5000BC or so, it clearly wasnt doing well and I imagine it would be very hard to stumble on.

Jean M
11-22-2013, 05:22 PM
.. do you know if anyone is ever going to test Y-DNA from Dnieper Donets?

All the ongoing studies including ancient DNA that I know about are listed (with links) on http://www.ancestraljourneys.org/migrationprojects.shtml

I really wish that I could answer these periodic pleas for more information on various unpublished studies. I would like to know more myself. :)

newtoboard
11-22-2013, 05:58 PM
All the ongoing studies including ancient DNA that I know about are listed (with links) on http://www.ancestraljourneys.org/migrationprojects.shtml

I really wish that I could answer these periodic pleas for more information on various unpublished studies. I would like to know more myself. :)

I was curious if there was any Y-DNA available at all.

What do you make of their mtDNA? From what was listed on your site it looked very eastern especially with the mtDNA T and mtDNA C.

Jean M
11-22-2013, 06:19 PM
What do you make of their mtDNA? From what was listed on your site it looked very eastern especially with the mtDNA T and mtDNA C.

There seem to be three elements in the mtDNA of Dnieper Donets II people, which we can date like this:


Mesolithic -U5a1a. I assume this was predominant in the hunter-gatherers who settled around the Dnieper and Donets.
c. 6000 BC - C and C4a2. I'm assuming that C arrived with pottery. Dnieper Donets I people made the kind of pottery that arrived in Europe from Lake Baikal. It would have been made by women. It probably did not come all in a rush in one big journey from Lake Baikal. Picture it travelling in stages from one hunter-gatherer band to another as they hunted across the Asian steppes. Now and then hunter bands would meet each other, which was a good way to acquire partners from another band and avoid inbreeding. So pottery-making could gradually spread westwards. It reached the Volga about 7000 BC and went from there to Dnieper Donets.
c. 5200 BC - H, T, U3. I assume these were acquired from the neighbouring farmers, as the Dnieper Donets people adopted animal breeding as so turned into the Dnieper Donets II culture. I don't know why you see T as eastern particularly. It seems to have originated in the Near East and spread with farming.

newtoboard
11-22-2013, 06:58 PM
There seem to be three elements in the mtDNA of Dnieper Donets II people, which we can date like this:


Mesolithic -U5a1a. I assume this was predominant in the hunter-gatherers who settled around the Dnieper and Donets.
c. 6000 BC - C and C4a2. I'm assuming that C arrived with pottery. Dnieper Donets I people made the kind of pottery that arrived in Europe from Lake Baikal. It would have been made by women. It probably did not come all in a rush in one big journey from Lake Baikal. Picture it travelling in stages from one hunter-gatherer band to another as they hunted across the Asian steppes. Now and then hunter bands would meet each other, which was a good way to acquire partners from another band and avoid inbreeding. So pottery-making could gradually spread westwards. It reached the Volga about 7000 BC and went from there to Dnieper Donets.
c. 5200 BC - H, T, U3. I assume these were acquired from the neighbouring farmers, as the Dnieper Donets people adopted animal breeding as so turned into the Dnieper Donets II culture. I don't know why you see T as eastern particularly. It seems to have originated in the Near East and spread with farming.



Doesn't T has its highest frequency and diversity in Eastern Europe, Central Asia and Iranian plateau ? I could be wrong about that though.

Jean M
11-22-2013, 07:11 PM
Doesn't T has its highest frequency and diversity in Eastern Europe, Central Asia and Iranian plateau ?

It might for all I know, but that would be where it went to, not where it came from. T2b was present in the pre-pottery Neolithic of Syria. Then we see T2 in the LBK and so on. Actually those two T samples in Dnieper Donets II were excluded from Nikitin 2012. Presumably they were among the results which failed to replicate. So perhaps we shouldn't make too much of them anyway.

GailT
11-22-2013, 09:44 PM
Breaking down H into subclades and trying to track back to the origin of specific subclades in aDNA will be more informative as to origins.

One of the challenges of interpreting mtDNA H origins and migrations is that there are nearly 100 named subclades of H, and almost as many unnamed subclades, many of which are represented by a single sample.

If we focus only on the major subclades, there is still a challenge with the poor time resolution of mtDNA. H5 has an age estimate of around 10,000 years, right around the time that agriculture and herding cultures were expanding from southwest Asia. We have 20 named subclades of H5 and at least 55 H5* samples that represent 42 unnamed subclades.

There are five H5 ancient FMS samples:
H5b, Rossen 6400 ybp (KC553992, Brotherton et al)
H5 Blatterhohle, farmer, 5600 ybp (KF523405, Bollinger et al)
H5* Blatterhohle, farmer, 5600 ybp KF523402, Bollinger et al)
H5a3, Bell Beaker 4500–4200 and 4050 ybp, ( KC554002 & KC554005, Brotherton et al)

There are also H5 samples (not tested at the full genome) in Syria around 8400 ybp and Minoan Greece around 4000 ybp.

The H5* Blatterhohle sample KF523402 has 3 extra mutations, (3197C, 73G, 11719A) only one of which (3197C) is shared with one modern sample from Italy (JQ703228). So the common maternal ancestor of KF523402 and JQ703228 might have been thousands of years before 5600 ybp, and they could have their origins in different ancient cultures.

So we very clearly need full genome sequencing of ancient samples, and we probably need a very large number of ancient samples. If may turn out that H5 was present in many different Neolithic cultures. It will be interesting to see if specific mtDNA fingerprints can be developed for different ancient cultures, but I wonder if the precursor Neolithic population was already a mix of many different haplogroups, such that it may difficult to get very good precision for the mtDNA fingerprint of ancient cultures.

Jean M
11-22-2013, 10:20 PM
but I wonder if the precursor Neolithic population was already a mix of many different haplogroups, such that it may difficult to get very good precision for the mtDNA fingerprint of ancient cultures.

I think you are right. MtDNA is pretty much a mish-mash from the Neolithic onward. But I really thank you for your analysis here. Table 1 in my book already needs updating, with all the new aDNA samples we have.

Generalissimo
11-23-2013, 02:58 AM
I think that's more or less right, though not just in Eastern Europe. It seems to rise gradually everywhere, though we don't have enough samples yet to be making firm statistical statements. It is possible that Iron Age migrations had an impact, but mtDNA H confers an advantage in recovery from sepsis, which may well have gradually increased its levels in Europe over the millennia, particularly after the various rounds of plague that started with the Justinian one.

Eastern Europe didn't suffer from plague to the same extent as Western and Southern Europe. That's probably why the Slavs managed to expand so rapidly at the time into the Balkans.

So if the Proto-Slavs had the same levels of mtDNA H as the Dnieper-Donets, Corded Ware and Unetice cultures, Scythians from Rostov. etc. until 500AD, then it's a real puzzle how they acquired much higher levels so quickly without being in primary contact for a long time with former populations of the Bell Beaker Culture range, like Urnfield for instance, which appears to have had an mtDNA H frequency of around 47%.


MtDNA H is not linked to just one archaeological culture as you know very well.

What I'm saying is that modern levels of mtDNA H across Europe are linked to events which started in Copper Age Portugal (PCA_). You can see that clearly on this PCA.

http://img199.imageshack.us/img199/8420/euadnapca12.png

http://img829.imageshack.us/img829/2051/euadnapca12l.png

Jean M
11-23-2013, 01:10 PM
Eastern Europe didn't suffer from plague to the same extent as Western and Southern Europe. That's probably why the Slavs managed to expand so rapidly at the time into the Balkans.


Yes that is true. But the point is that H may have gradually worked its way up through natural selection.


So if the Proto-Slavs had the same levels of mtDNA H as the Dnieper-Donets, Corded Ware and Unetice cultures, Scythians from Rostov. etc. until 500AD, then it's a real puzzle how they acquired much higher levels so quickly without being in primary contact for a long time with former populations of the Bell Beaker Culture range, like Urnfield for instance, which appears to have had an mtDNA H frequency of around 47%.


We don't really know what level of H the Mid-Dnieper Culture had. There is no guarantee that it was exactly the same as the samples we have from neighbouring or distantly related cultures.

We can easily separate out the Scythians from Rostov -on-Don, as they were the descendants of people who had spread far into Asia, and evidently mixed with Central/East Asian groups, from whom they had acquired mtDNA haplogroups A4, C, D and F1b. They migrated to the European steppe around 700 BC. They had some contact with the Slavs. It looks as though a few descendants of Scythians (if we see the Serbi as such) took refuge from the Huns with the Slavs, and then travelled with at least one Slavic group in the great migrations, if I'm reading things aright. As I recall an Iranian name crops up among the early Serbs/Sorbs. But they made next to no impact on the present-day Slavic gene pool, as is easily seen from both Y-DNA and mtDNA.

Generalissimo
11-23-2013, 02:44 PM
But the point is that H may have gradually worked its way up through natural selection.

All non-Beaker pre-Iron Age Central and Eastern European cultures tested to date show low mtDNA H frequencies by modern standards. This would have to mean that natural selection started working suddenly in Eastern Europe during the Iron Age to push up H, even though the plague didn't have much of an impact on the early Slavs. That doesn't sound very convincing.

I'm betting the real cause was a migration, or a series of migrations. That's probably why Russians from near the Volga show a lower genetic distance to Spaniards these days than to North Caucasians.


We don't really know what level of H the Mid-Dnieper Culture had.

We can extrapolate, and it's already obvious that it was low, like in all pre-Beaker European cultures, except those in Portugal and Sardinia, and probably parts of Spain and North Africa.

By the way, on a related note, which uniparental markers do you think are associated with this autosomal component modal in Basques and the Irish? And which archeological culture expansion might it represent?

http://img43.imageshack.us/img43/3213/opv1.png

Silesian
11-23-2013, 03:12 PM
I just wish there was a bit more discussion in papers about the older forms of R1b in eastern Europe. Also M73 is crying out for some sort of study. If we understood it a bit better we would understand the possibilities for M269 a lot better too IMO. I dont even recall any attempt at a Eurasian map for M73. I could be wrong but I dont think anyone has done one. Maybe that is a project for Macciamo. The population studies are building up now so it must be getting closer to be able to make a map.

It would also be nice to see where a cluster consisting of { R1b-M343*_R1b-M73_R1b-M269[L23]_R1b-L23[51] } exist among Iranian groups.

alan
11-23-2013, 03:43 PM
This is all very well and unclear but this thread is about R1b and there is basically no case for R1b either

1. Somehow hiding in the farming world in the Neolithic only to spring up

2. R1b spreading from Iberia (except perhaps the local DF27 clade -and even then its not certain that clade originated there)

There is too much simplification of the idea of beaker being attached to P312 and having to closely mirror the spread of a type of pot. The pot may not have been spread most widely by the people who earliest made it.

There is no evidence of entire population spreads involved in beaker. Its impact just doesnt look like that because it usually takes on local characteristics of the local pre-beaker cultures in many places. So, I dont think its likely that a dual y and mt DNA signal is going to exist on the simple basis of P312 and H.

The real story of beaker is networking and ability to penetrate into existing societies. The key is networking. However, there is nothing to say it was like a stream or wave. It was probably more like a chain and anywhere in this chain another male line could have taken up part of the cultural package and then moved it on.

A typical method of cementing networks was marriage. That could IMO have a lot to do with the amount of H in the beaker phase. Not because H moved in some sort of wave but because once elite marriages started with some H women of importance, marriage of important people's daughters could have meant the lineage passed on with further alliances.

Another aspect of this is that women likely made pottery. Women also are likely to have been behind a number of the other aspects that Harrison and Heyd call the proto-Beaker package. The evidence of patrilocal pattern in now well established to have existed long before the beaker period and while marriage networks involving alliance marriages moving women is easy to envisage, its less clear how male lineages would have been welcomed, especially into areas where they already had equally if not superior metallurgy, mining etc traditions going back to 3000-3500BC such as Italy and the southern/western Alps, southern France etc. They simply had no need or motive to allow male groups to take over something they already had and controlled.

In fact one of the mysteries of beaker is its unique ability to penetrate into other societies, often taking on local characteristics although bringing new traits too. That is not such a mystery if you consider a wide network that was largely cemented through elite marriage networks. That makes complete sense in a patrilocal society who wanted to join this new network.

The motives to join such a network probably varied. In some areas there would have been no motive and probably horror at the idea of letting new male lines in (who after all would have been tiny numbers) to take over their long established turf. I think especially of the pre-beaker network of long duration that used Remedello dagger symbolism and dominated north Italy and part of Austria, Switzerland and SE France. However, RC dates show that the Ligurian mines run down at exactly the time beaker appeared so the likely motive for new exotic links would be importing metal and one of the obvious targets for putting our feelers from new sources would have been Iberia which appears to have been so copper rich that it was of little status in Iberia but had long been used as an export item to secure other exotica. I do not think there is a logical scenario for those Alpine areas to allow Iberian groups to settle and take over the metal trade around the Alps and north Italy but there would have been a logical scenario to obtain copper from Iberia perhaps by sending middlemen to Iberia and establish alliance marriages.

Southern France is unclear. The Languedoc group (apparently G people) had copper sources and a metal working tradition but it has been observed that they did not appear to want to trade it out and apparently used it as a purely local thing. That would have left part of southern France potentially without supply. They may have had a motive to allow actual male metallurgist settlers because they didnt have a previous history of doing it themselves.

Many areas though like the Alps and central Europe didnt have any logical incentive to allow new groups in IMO as the sources were already there and the same metal types etc existed in pre-beaker times.

For me the main scenario where actual beaker male settlers being allowed in, rather than just females moving for alliance marriages, are

1. Where local metallurgy was undeveloped

2. Where no ore sources existed locally

3. Where ore existed locally but there was previous tradition or expertise to exploit it

4. Where the above conditions applied and middlemen would help establish a link to distant copper sources

In a large chain like that with different motives for contact with the beaker culture, I think the female chain may have been more extensive than the male chain and that anywhere along the chain beaker traits could have been taken up by local male lineages. I think this is very plausible as beaker traits tends to be absorbed and local traits often retained. In other words the female chain may have been wide and unbroken but the male aspect of beaker culture may have not had a continous chain of a single male lineage. That in my opinion is a far more rational fit to the archaeology.

I will post this and elaborate in another post shortly



All non-Beaker pre-Iron Age Central and Eastern European cultures tested to date show low mtDNA H frequencies by modern standards. This would have to mean that natural selection started working suddenly in Eastern Europe during the Iron Age to push up H, even though the plague didn't have much of an impact on the early Slavs. That doesn't sound very convincing.

I'm betting the real cause was a migration, or a series of migrations. That's probably why Russians from near the Volga show a lower genetic distance to Spaniards these days than to North Caucasians.



We can extrapolate, and it's already obvious that it was low, like in all pre-Beaker European cultures, except those in Portugal and Sardinia, and probably parts of Spain and North Africa.

By the way, on a related note, which uniparental markers do you think are associated with this autosomal component modal in Basques and the Irish? And which archeological culture expansion might it represent?

http://img43.imageshack.us/img43/3213/opv1.png

alan
11-23-2013, 05:02 PM
To follow on, I think there are a number of suggestions that the male chain involved in the beaker cultural spread was less continuous than the female

1. The localised aspect of the way beaker traits were encorporated.

2. The lack of rational motive to allow male groups to spread like that in a patrilocal society many areas that I have outline above.

3. Evidence at Sion that the people who took up beaker pot (who had previously used the Remedello dagger symbol) were the same people.

4. The clear distinction between what look like Cardial descended people in early beaker Iberia and the evidence for distinct groups with different craniology and dental traits elsewhere.

5. The fact that the brachycephalic beaker type known in many areas of central, NW Europe and the isles is different from the Iberian beaker types and in fact better resembles the pre-beaker Remedello and related groups in Italy as well as Balkans groups. This type did not spread from SW Europe where it is unknown until later beaker phases. This type is also unknown in pre-copper age Anatolia.

6. As Harrison and Heyd point out, many of the beaker cultural traits and artifacts were not known in the early SW European phase and only appear later. It is interesting to note how many of the developed beaker traits, both physical and cultural are actually not present in Iberian and earliest beaker in general but do have strong echoes in pre-beaker cultures like Remedello around the southern Alps. Remedello not only had the cranial types but had the whole dagger, archery etc symbolism already as well as fascinating hierarchical looking burials. This contrasts with the collective burials in Iberia and the west Med. before and during the early beaker phase.

7. SW Europe, especially Atlantic Iberia, appears to have been totally dominated by one downstream clade, DF27. This by all logical standards points to it as an arrival point rather than an origin point for P312. There is a far greater P312 clade mix further east around France, the Rhine etc. Also, despite some reports simplifying the R1b story, there is a trail of higher variance and phylogenically upstream branchings that points to an east to west move. L23xL51 and M269* show a Balkans concentration in Europe while L51* has a concentration that is not a bad fit for pre-beaker Remedello dagger culture and zone of influence.

So, all in all Iberia does not look like a source point for the spread of anything above DF27 in the R1b story. The overall picture for European R1b fits a first fall in the farming area of Europe in the Balkans area followed by a trail across the Alpine area and finally a move into Iberia in DF27 form. While this does not nicely fit the chonology of beaker pot, it is clear to me that tying the spread of beaker pot to P312 in a simplistic way simply does not work. If I had to guess how P312 got to Iberia, one rational scenario would be that it got there c. 2800BC or so along with a prototype of the beaker pot which has no believable local predecessor but has parallels in central Europe. The mysterious appearance of AOO beaker in Iberia with its very different distribution to Maritime beaker in Iberia and early dates could be a signal of contact with areas of Europe to the east of Iberia. The other scenario, which may not be entirely unrelated, is west and south Alpine groups who had an interesting combination of Remedello dagger symbolism, the running down of the Ligurian mines that had supplied that group around the period of beaker contact and who also neighboured corded ware groups who had just arrived in the western Alps and where only a pass away to the north from c. 2800BC. They had an interesting combination of circumstances and contacts that might have led to contact with Iberia as well as a prototype beaker. I would suggest that they may have initiated the contacts with Iberia and that they were also the people where R1b in its L51-L11-P312 phase had resided.

That raises the question of how these Remedello dagger imagery using groups originated themselves. Well the culture and the sudden appearance of developed local metallurgy and mining appeared in north Italy c. 3500BC, were mining in Liguria at the same time and soon influenced the whole Alpine area from Austria to SE France. The outstanding charachteristic other than metallurgy, mining etc was the dagger symbol which in combination with the remarkable Remedello burials and the appearance of an element with the bracycephalic skulls does suggest something major was happening in the area.

This is a very tentative suggestion. Recent reports have placed the origin of the copper dagger as a symbol in the copper working Bodrogkeresztúr culture. They had been controlling what was an important copper source when the other Balkans sources collapsed. Just before this culture emerged the area had featured a small Suvorovo type steppe intrusion of people who were clearly interested in the copper and had no other reason to settle in what was an atypical area for steppe settlers. They were a local component around the time of the emergence of the Bodrogkeresztúr culture. Although it has a lot of continuity with the previous Tiszapolgár culture there are changed such as very ephemeral settlements indicative of mobile pastoralism and hierarchy is indicated in the graves. They were major networkers with significant contacts in many directions including Cucuteni who probably relied on them for metals. This culture is described as located in 'the Alföld, with most settlements located at the river Tisza and the eastern border of the lowlands. The area between the rivers Danube and Tisza is hardly settled. The north-eastern border is made up of the rivers Tisza and Someş. Single finds occur far away from the centre zone, in south Poland as well as in Transylvania, here especially at the river Mureş (for Romania cf. especially Luca 1999). The southern border is disputed; however, we find cultural remnants as far as the Serbian Banat. The finds from the Belgrade region, however, seem to be imports.The Budapest region is a special case. It was attributed to the distributional area of the culture by Patay, but some researchers – particularly N. Kalicz, J. Pavúk, J. Bátora and Zs. M. Virág – argue conclusively against this. The finds from the middle Copper Age from this region greatly resemble those from the Bodrogkeresztúr culture'.

The date of origin is around 4000BC around the same time as the Suvorovo element was added to this area. The date of origin also corresponds rather well with the M269 variance date and perhaps within the bounds of L23. Interestingly the Bodrogkeresztúr culture and its enormous metal wealth collapses c. 3500BC about the sort of time metallurgy, Remedello etc appears in Italy.

This is just a tentative suggestion. However, there are many other scenarios which could involved a link to the Balkans where the cultural sequence c. 4500-3500BC is fiendishly complex. Regardless it seems virtually certain that the Balkans c. 4000-3000BC was the stepping stone for R1b into central and western Europe.

alan
11-23-2013, 05:11 PM
I would not tend to mix clades like P343, P25 or V88 with ones like L23, M73 etc. They hadnt shared a certain common ancestor with L23 and M73 since perhaps 14000BC. However, M269 and M73 shared a P297 SNP c. 9000BC and both would have remained in the P297* state until after 5000BC. That is essentially the period of the Neolithic and this lineage was barely surviving given the lack of any modern representatives so far recorded.


It would also be nice to see where a cluster consisting of { R1b-M343*_R1b-M73_R1b-M269[L23]_R1b-L23[51] } exist among Iranian groups.