Tracing the genetic origin of Europe's first farmers

What could have happened is that some I1 hunter-gatherers copied the farming techniques of their G2a neighbours in Hungary, or that a group of I1 warriors invaded some G2a villages, killed off (most of) the men and took the women for them (or raped them). There descendants would have been mostly I1 men with Near Eastern or Balkanic mtDNA. This new hybrid generation could then have grown and prospered and developed into the LBK culture. That would make sense since LBK mtDNA is overwhelmingly East Mediterranean, while I1 appears to have colonised Germany and Scandinavia late, considering that none of the Mesolithic samples from Scandinavia or Germany/Luxembourg were I1 so far.

Given how much knowledge is required in order to successfully farm and to care for livestock, I'd bet on a peaceful scenario where the more numerous but peaceful farmers traded goods with the more war-like but less numerous hunter/gatherer types and gradually introduced them to farming techniques so that the hunter/gatherer types wouldn't be tempted to hunt their livestock. And, although it's difficult for those of us living in a world of billions of people to wrap our heads around the idea of "not enough people", a small population that was expanding into fertile territory might have been willing to take in any outsiders who wanted to join them. And some of the hunter/gatherers might have envied the food security of the farmers and decided to join them rather than raid them. The Y haplotype I1 that was found could have been a fluke for all we know, but now that I'm thinking about it from that angle, it really does seem to me that it's very difficult to explain the rapid rise of I1 unless I1 people did adopt agriculture fairly early on. The I1 folk appear to have expanded north, for whatever reason, and perhaps that saved them from whatever happened to the G2 farmers.
 
there is something strange about G2a
they seem to have been very succesfull at the early neolithic, being represented in 2 branches : cardium pottery (PF3146 ?) and LBK
yet the only succesfull branch today seems to be P303, which may represent Maykop

In west Caucasus among Adyghe people you will find only G2a3b1a1b1 L1264+ and G2a1a
G2a3b1a1b1 L1264+ is 2700-3000 years old, and there are no other subclades of G2a3b1 P303 among them.
And only some branches of haplogroup J2 are old enough among Adyghe people, reaching ages of 5500 and 5200 years old.
I think that Maykop people were mostly J2
 
While I support most of your post, I cannot see J2 and T involved in metal work. My take is because T in anatolia, Levant, Egypt, africa and arabian peninsula is all too young ....there is a gap of 30000 years from origin of T to any of these areas I noted.
I believe T formed around Azeri lands near caspian sea, they were mostly fishermen/hunters ( lezkins) , moved into North Caucasus as you say, and mostly spread around as hunters, that's why the huge dispersal of T and that's why no bones are found for T.....hunters cannot bury themselves. Otzi ,a hunter was only found due to the ice. Farmers bury or cremate the dead.

It would be interesting to see what % increase there would be for T if all pre-2008 papers looked into the found K ydna . It might change opinions.

J2, IMO formed In the south side of the zargos mountains around mesopotamia, are farmers and I cannot see an earlier migration into the north caucasus.

I would like to know why or how T, G, I, H, L and J sat in the caucasus waiting 40000 years for the creation of R to join them...........seems illogical that people sat around that long in that period of history

there is a gap of 2-3000 years between the 1st neolithic wave and the arrival of IE people in Europe

there was 2nd neolithic wave : http://www.nature.com/news/archaeology-the-milk-revolution-1.13471

the 2nd wave brought J and T IMO, they were not farming the land, they were herders, but also farmers adapted the milk economy : Lyengel , TRB , British neolithic , ..
 
Given how much knowledge is required in order to successfully farm and to care for livestock, I'd bet on a peaceful scenario where the more numerous but peaceful farmers traded goods with the more war-like but less numerous hunter/gatherer types and gradually introduced them to farming techniques so that the hunter/gatherer types wouldn't be tempted to hunt their livestock. And, although it's difficult for those of us living in a world of billions of people to wrap our heads around the idea of "not enough people", a small population that was expanding into fertile territory might have been willing to take in any outsiders who wanted to join them. And some of the hunter/gatherers might have envied the food security of the farmers and decided to join them rather than raid them. The Y haplotype I1 that was found could have been a fluke for all we know, but now that I'm thinking about it from that angle, it really does seem to me that it's very difficult to explain the rapid rise of I1 unless I1 people did adopt agriculture fairly early on. The I1 folk appear to have expanded north, for whatever reason, and perhaps that saved them from whatever happened to the G2 farmers.

expansion of I1 must have come after LBK :

1/ I1 expansion is estimated only 4300 years ago
2/ if I1 expanded during LBK , why did G2a not expand at the same rate ; I1 was only a small minority among G2a in LBK
 
In west Caucasus among Adyghe people you will find only G2a3b1a1b1 L1264+ and G2a1a
G2a3b1a1b1 L1264+ is 2700-3000 years old, and there are no other subclades of G2a3b1 P303 among them.
And only some branches of haplogroup J2 are old enough among Adyghe people, reaching ages of 5500 and 5200 years old.
I think that Maykop people were mostly J2

you mean G2a1a P16 1/P16 2 ?
 
Sparkey, I'd really like to know your thoughts on the I1 in LBK Hungary I couldn't find the SNP results, was it positive for IJ and I SNPs, and which I1 SNP(s) was it positive for.

"Data Supplements" > "Supplementary datasets 1-17" > "S5"

You can see that the I1 sample was positive for M89 (F), M170 (I), and M253 (I1) but negative for M438 (I2). No downstream I1 SNPs were tested. IJ was not tested.

Do you think it's possible I1a-Df29 clades have been roaming in central Europe since the Mesolithic or Neolithic? I think it is because although I haven't done any research since pretty much last winter, I remember reading that central Europe has a differnt ratio of I1a-Df29 clades than Scandinavia, plus they have I1b. I1 being in Mesolithic and then Neolithic central Europe doesn't explain how it became so popular in Scandinavia(Motalas didn't have it, Sf11 didn't, The PWC hunters didn't, Gok4 didn't). Weird stuff happened with I1, there had to of been some type of epic rise.

I don't think that I1-DF29 is old enough to be around since the Mesolithic. DF29 represents the primary modern expansion of I1 but it seems to have happened to the north somewhere, since it includes the main subclades of West, East, and North Germanic peoples (Z58, Z63, and L22). I'm guessing that if this ancient sample had been tested for these SNPs, it would have been negative for all of them. If we're supposing that the ancient sample is closer to the true origin of I1, then that means that the I1 of today is mostly from a young displaced branch, while the old I1 branches went extinct (as far as we can tell from samples we have so far). Actually, that scenario would tie up some loose ends, like why modern I1 appears so young, and why Eastern Europe has no native haplogroup that appears to date to the Mesolithic like I2 does in Western and Central Europe. I'm not ready to commit to the hypothesis with only one sample, though.

It'll be interesting to learn how pre-Indo European west European lineages I2a2 and I1 survived so well compared to G2a, I2a1, F*(96?), etc. It's no suprise that all of the Neolithic G2a's so far except Otzi have G2a-P303 like most modern European G2a, and there are still layers of the signature lineage of WHG; I2a1-P37.

Could it be that hunters and hunters-turned-farmers were more likely to be pushed to the north than migrant farmers, and northerners would later experience population expansion at the expense of southerners? Probably too simplistic an explanation, but that could be one of the many effects to contribute to the pattern we see.
 
what strikes me is the Maykop culture area with lowest fst values
i suppose they are not ancestral to STA-LBK culture
does it mean STA-LBK and Maykop have the same genetic origin ?

I lack details but some studies about metric and non-metric traits on skulls and teeth concluded the types of Armenians Highland population was common among AND the North Caucasus ancient populations (Pre-Br and Bronze Age) AND the Cucuteni-Trypillian populations, these types were found too among the supposed I-Ean cultures of the Steppes at same time, but attenuated as distance augmented - "Neolithic" or "bronze age" is a poor adjective to identifygenetically these populations because the regions and their populations which produced, apparently, agriculture are the same which produced diverses stages of metallurgy!
 
I add I regreat ancien anthropology because it was focalized on superficial traits (and typology) which inabled to differenciate cousin or sister (close) populations in a way very more precise than can do autosomals studies or distances metric modern comparisons, sometime confusing - it's true that genetical heritability of some features are in discussuin but I think the exercice was of some worth, nevertheless, when added to other ways of investigation.
a survey and compilation of ancient works concludes there were changes in people features at the beginning of Neolithic in the Balkans (around Donau river,in Serbia and Croatia (with the eternal considerations of new types and crossings opposed to in situ evolution: evolution I find a bit rapid, all that is new official science like the "no-move-theories" in archeology - we know now what value this last theory has) - the old science said that two population among local HGs existed at first: one almost "pure" cromagnoid, the other a mix of cromagnoids and brunnoid types (I have to find it back) whan Neolithic send a pannel of so called mediterranoid subtypes, what is not too surprising - crossings appeared a few later, if I don't mistake-
the apparition of brachycephally in Armenia and North Mesopotamia ddin't precede the 2000's BC according to old scholars: what would be their aurosomals and haplo's today affiliation? I wonder... the same phenomenon appeared (in less)in the steppes almost at the same time: from Carpathians or from East Caspian??? it would be interesting concerning the metal ages and the I-Ean question knowing what other means of rechearch could tell us about all that???
 
there is a gap of 2-3000 years between the 1st neolithic wave and the arrival of IE people in Europe

there was 2nd neolithic wave : http://www.nature.com/news/archaeology-the-milk-revolution-1.13471

the 2nd wave brought J and T IMO, they were not farming the land, they were herders, but also farmers adapted the milk economy : Lyengel , TRB , British neolithic , ..

Have you read the June 2014 Karafet papers?

Its, states IIRC, GHIJTLNO formed within 6000 years of each other , and P formed in modern Malaysia .........R and Q came from this P and R-207 has 8% in Java, Borneo etc....there is nearly 30000 years between this GHIJTL group and the creation of R.
Why would this west-asian/southasian group that formed in and around Iran , Pakistan and India wait 30000 years for R ?

Below is the new age of markers from July 2014 as what is now accepted


I am not saying all the main R formed in South-East Asia...I am saying the wait time for creation of R makes no sense for the other markers to wait around and do nothing

K' = the new K1 which is T and L
K'' = the new K2a which is N and O
and R and Q belong to K2b
 
Given how much knowledge is required in order to successfully farm and to care for livestock, I'd bet on a peaceful scenario where the more numerous but peaceful farmers traded goods with the more war-like but less numerous hunter/gatherer types and gradually introduced them to farming techniques so that the hunter/gatherer types wouldn't be tempted to hunt their livestock. And, although it's difficult for those of us living in a world of billions of people to wrap our heads around the idea of "not enough people", a small population that was expanding into fertile territory might have been willing to take in any outsiders who wanted to join them. And some of the hunter/gatherers might have envied the food security of the farmers and decided to join them rather than raid them. The Y haplotype I1 that was found could have been a fluke for all we know, but now that I'm thinking about it from that angle, it really does seem to me that it's very difficult to explain the rapid rise of I1 unless I1 people did adopt agriculture fairly early on. The I1 folk appear to have expanded north, for whatever reason, and perhaps that saved them from whatever happened to the G2 farmers.


I agree for the most part. From everything in recent papers about the culture of the foragers at the time of the Neolithic transition, they were not the mobile mammoth hunters of the popular imagination. Rather, whether it was the people living around the Danube Gorges, in Germany (Bollongino et al) or in Gotland, they seem to have been relatively sedentary fisher folk. More like the Indians of the Pacific northwest than the Dakota. (See Boric et al about mobility at this period: Our study unequivocally proves that in the earliest phases of the Neolithic in southeastern Europe, perhaps paradoxically, farming communities were much more mobile than local foraging populations, which in the case of the Danube Gorges remained tied to the exploitation of particular ecological niches since the beginning of the Holocene up until ∼6200 cal B.C. ")

According to the same paper, there seems to have been some absorption of local hunter gatherers,at least in the Danube Gorges area. We also know that from the fact that some samples found in a Neolithic context in Hungary were I1, and in other Neolithic contexts were I2a.

From the paper: "Strontium isotope ratios indicate a dramatic increase in the numbers of nonlocal, first-generation migrants buried in this region at two sites—Lepenski Vir and Ajmana—during the course of Mesolithic–Neolithic transformations in the Danube Gorges (∼6200–6000/5950 cal B.C.) in the sample of 25 securely dated individuals from three sites. This trend continues into the period of the Early/Middle Neolithic after ∼6000/5950 cal B.C. in the sample of 26 securely dated individuals from two sites. Strontium isotope ratios associated with individuals dated to these periods occur outside both the upper and lower boundaries of the defined local strontium range (Figs. 2 and 3), suggesting at least two or more geologically distinct regions of origin for these incomers."

A change in culture is dramatically apparent not only in the old Mesolithic settlements like Lepinski Vir, but, of course, in the totally new centers established by the newcomers.The authors conclude that "this pattern could be interpreted as suggesting a reciprocal mating network between the Danube Gorges foragers and the earliest farming communities in the surrounding areas, with largely nonlocal women being buried at the central forager site of Lepenski Vir during these phases."

They further go on to state that "important changes were taking place in these forager societies that came under pressure from the growing Neolithic presence in the adjacent areas after ∼6200 cal B.C. (14, 16, 17). This process of asymmetrical acculturation ended up in a complete absorption of forager specificity in the first several centuries of the sixth millennium B.C."

Furthermore, they state that...As for the Danube Gorges foragers, judging by, at first, a rather organic appropriation and incorporation of new elements of social and material existence as well as biologically “new blood” into the existing modes of being, and, consequently, the abandonment of forager cultural specificity, the mentioned process might have had a predominantly positive connotation or at least represented the only viable solution in the face of the emerging demographics."

That isn't to say that violence didn't occur;the authors also document the discovery of a couple of severed heads. However, the transition seems to have been relatively peaceful.

Given this model, how then are we to make sense of the mtdna findings from the site in Hungary which is the subject of this thread? In Hungary, at least, the U5 lineages which the scholars have been associating with the Mesolithic fisher/foragers form only 10% of the total. Does that mean the wife exchanges were asymmetrical? I don't know. Perhaps forager women, totally foreign to the Neolithic lifestyle, would not have been the first choice for a Neolithic village, while a farmer mate could provide a source of technological expertise in the forager communities? Or, perhaps it was simply a case where the Neolithic farmers just outbred them.

Also, I think we have to keep in mind that the I1 and even the I2a forager lineages are decidedly in the minority. So perhaps we have to think in terms of some absorption of the foragers, but it might be that the majority of them were pushed further to the northeast. There is an analogy in what happened to the North American Indians.

Finally, I think that one has to keep in mind the papers which have argued for a major population crash in central Europe, a population crash which didn't affect southern Europe. How many of the LBK (and related cultures) people remained to encounter the "Indo-Europeans" I don't know, nor do I know the autosomal make-up of this third wave. I would be very surprised, however, if they didn't carry a large EEF component themselves.
 
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you mean G2a1a P16 1/P16 2 ?
Yes G2a1aP16_1, P16_2, FGC645/Z6616
But P16_1, P16_2 are unstable and unreliable
Better if we say G2a1a FGC645
 
I don't think that I1-DF29 is old enough to be around since the Mesolithic. DF29 represents the primary modern expansion of I1 but it seems to have happened to the north somewhere, since it includes the main subclades of West, East, and North Germanic peoples (Z58, Z63, and L22). I'm guessing that if this ancient sample had been tested for these SNPs, it would have been negative for all of them. If we're supposing that the ancient sample is closer to the true origin of I1, then that means that the I1 of today is mostly from a young displaced branch, while the old I1 branches went extinct (as far as we can tell from samples we have so far). Actually, that scenario would tie up some loose ends, like why modern I1 appears so young, and why Eastern Europe has no native haplogroup that appears to date to the Mesolithic like I2 does in Western and Central Europe. I'm not ready to commit to the hypothesis with only one sample, though.
Thanks for this elegant explanation. Always admire your ability to word your ideas in so easy to comprehend style.



Could it be that hunters and hunters-turned-farmers were more likely to be pushed to the north than migrant farmers, and northerners would later experience population expansion at the expense of southerners? Probably too simplistic an explanation, but that could be one of the many effects to contribute to the pattern we see.
I don't think it was that easy for HGs to break into farmers society. When we look at LBK culture and samples that's around 2,000 years after first farmers took first step in Europe around Bulgaria and Greece. At the time of LBK we should be able to start seeing first HGs among farmers. More precisely HGs paternal haplogroups in farmers population. When this I1 individual is tested autosomally, sample should show predominantly EEF admixture. If it shows mainly WHG that I would swear he was most likely a slave. ;)

Do you have any hypothesis what happened to E1b folks? Are they still deep in South Balkans at this time?
 
Given this model, how then are we to make sense of the mtdna findings from the site in Hungary which is the subject of this thread? In Hungary, at least, the U5 lineages which the scholars have been associating with the Mesolithic fisher/foragers form only 10% of the total. Does that mean the wife exchanges were asymmetrical? I don't know. Perhaps forager women, totally foreign to the Neolithic lifestyle, would not have been the first choice for a Neolithic village, while a farmer mate could provide a source of technological expertise in the forager communities? Or, perhaps it was simply a case where the Neolithic farmers just outbred them.

Also, I think we have to keep in mind that the I1 and even the I2a forager lineages are decidedly in the minority. So perhaps we have to think in terms of some absorption of the foragers, but it might be that the majority of them were pushed further to the northeast. There is an analogy in what happened to the North American Indians.
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Well, 10% might be just right. Roughly speaking, population density between farmers and HGs is 10 to 1. If farmers settled and mixed right away with HGs that is the proportion of haplogroups or admixtures one might expect coming from density factor. I don't think it was the case, but surprisingly the numbers added up, lol.

To bad they didn't use Lazarides autosomal admixture "standards", we would know right away much EEF mixed with HGs.
 
To bad they didn't use Lazarides autosomal admixture "standards", we would know right away much EEF mixed with HGs.

They didn't here, but the Lazaridis paper tested the autosomal DNA of one LBK individual from Germany (Stuttgart) and the proportion was also 90% East Mediterranean (EEF) and 10% Mesolithic European (WHG).

That makes it even more difficult to understand how I1 became so overwhelmingly dominant compared to G2a (3:1) in Germany and especially in Scandinavia (15:1 in Denmark, 30:1 in Sweden/Norway). I suppose it has to do with a founder effect in the northward expansion of farmers from Germany to Scandinavia. After all, even if the Neolithic farmers that colonised Scandinavia were 90% EEF and 10% WHG, if it was just one extended family and all the men were I1 (by chance), then I1 ended up introducing Mediterranean genes into Scandinavia. That would explain why Mesolithic Scandinavians had 0% EEF, but modern Scandinavians have 40% EEF. That would also explain why a mere 15% of mtDNA lineages in Sweden and Norway are Mesolithic U4 and U5, and only 8% in Denmark. In fact we cannot even be sure that these are direct descendants from the Mesolithic inhabitants of Scandinavia since the Indo-Europeans definitely had U4 and U5 among their lineages too.

Ironically most of the 45% WHG admixture in modern Scandinavians may have come from R1a invaders from the Corded Ware (and some from the subsequent R1b invaders in the Late Bronze Age).
 
To bad they didn't use Lazarides autosomal admixture "standards", we would know right away much EEF mixed with HGs.

The idea is to find results independently. If they then match you can be relatively sure it is right.
 
I think we have to keep in mind here that there isn't a perfect correlation between the percentages for uniparental markers and autosomal results. While the "hunter/fisher/forager" mtdna might only account for 10% of the lineages, their autosomal contribution might be greater. There is then the y lineage contribution. So, you could easily reach the 20% 'hunter-fisher-gatherer" component hypothesized for the EEF signature. I don't know if that's coincidence or not, but the numbers fit.

The authors also come down in favor of a scenario were most of the prior inhabitants evacuated the area, presumably for the northeast regions. From the paper:
Residual Neolithic hunter-gatherer isolates, as reported from Central Europe by Bollongino et al. [30], have not yet been observed in our study region. According to the low proportion of hunter-gatherer mtDNA lineages in the LBK gene pool, we assume that admixture between hunter-gatherers and colonizing LBK farmers was negligible in Central Europe.
Considering the relative size and speed of the LBK expansion, we have to assume a substantial population growth during the earliest LBKT, which might have resulted in a population pressure and led to emigration from Transdanubia [55]. While such a radical
population increase was not palpable from the Early Neolithic archaeological records [7], but recent extensive archaeological excavations have provided new insights into large-scale early LBKT settlements in western Hungary [9,56,57], which suggest larger source communities for a possible colonization than previously assumed.


In other words, previous scenarios underestimated the numbers of farmers moving into Central Europe, and the Bollongino results seem to be anomalous. Indeed, I never thought the interpretation of that find ever made much sense. A strange use of a burial cave by two different cultures for a couple of hundred years should not have been, in my opinion, extrapolated to mean that there were two, co-equal in numbers, cultures inhabiting Central Europe at the same time. (Fisher/forager and Farmer)

An obvious take away from the results is the extreme diversity of the mtDNA package in comparison to the y dna results. The authors explain it in terms of patrilocality, which makes sense. What I find more interesting is the question of where that mtDNA package was formed, especially in light of the so far unpublished results from an analysis of mesolithic Greek mtdna, which found no U5 or U4, but did find what are usually called "Neolithic" or "Near Eastern" lineages. There are also the recently published mtDNA samples from the Near Eastern Neolithic, Fernandez et al 2014, which can be found here: http://www.plosgenetics.org/article/info:doi/10.1371/journal.pgen.1004401

This is a table of the Fernandez results:
http://www.ephotobay.com/image/picture-32-55.png

As to this last find, this is what the authors of the current paper have to say:

Recent aDNA study from 8000 BC Near Eastern farmers raises the question whether modern Near Eastern mtDNA can be used as a proxy for the Near Eastern Neolithic variability [44]. In our opinion, these newly described seven different incomplete HVS-I haplotypes (np 16095-16369) only provide a limited basis for comparative aDNA analyses, and we thus still consider modern-day Near Eastern genetic data sufficient proxies, when tracing the origin of the first European farmers.

The fact that prior papers found an affinity between the mtDna of Crete and the Near East on the one hand, with that of Europe on the other, also has to be factored into the equation.

So, I don't know where we stand. Maybe the supposedly Mesolithic dna found in Greece which is not U5 or U4 is misdated. Or, the movement of mtDNA from the Near East into Europe began during the Mesolithic. There's something to be said for the fact that population increases in the Near East which coincided with improving climatic conditions actually propelled the development of this new technology. It could also have propelled migration into similar climatic regions in the Greek Islands and mainland proper.

We know that the Neolithic transition occurred in the Near East. However, when these farmers moved into Europe, they might have encountered people already related to them at least through some mtDNA lines.

Btw, speaking of the Fernandez paper, I think their map is a great way to visualize the spread of the Neolithic:

View attachment 6605
 
When trying to make sense of the population movements, I think it's important to remember that we're talking about the Carpathian Basin, a relatively flat area of grassland with a river running through it. During the Early Neolithic, farmers with stone axes would congregate in an area like that and avoid the more forested area, so they could graze their livestock and till the soil without first having to clear the land of trees, so the population of Hungary during the Early Neolithic could have been much higher than elsewhere in Central Europe. Of course, once people acquired copper and perhaps bronze axes, clearing the forest would have been a little easier, but I doubt the population would have shifted off the plains that quickly. And the Carpathian Basin would have also seemed highly appealing to the Bronze Age IE pastoralists, so I suspect the population replacement rate as a result of the later Indo-European invasions would have been much higher in this area than in some other parts of Central Europe, specifically those heavily forested areas to the north. If the G2 folk remained primarily on the Hungarian Plain, they would have felt the full impact of the IE invasions. However, if I1 folk did learn farming for G2 people, then decided to move north or were pushed north into more forested areas, they would have initially been much less vulnerable to Indo-European pastoralists with bronze weapons, simply because forest lands with farms carved out of them are much less appealing to pastoralists than open plains.
 
I'm not quite sure I understand, but I'll take a stab at it. :) Are you thinking that the "I" lineage males were slaves? I have to read the paper more carefully. Is there any indication from the archaeology that the "I" males were of a lower social status?

I also don't know that we have any indication that very early farmers had slaves. I would think that slavery would involve a level of stratification in society that would be more typical of the very late Neolithic if not the Metal Ages, although that's just my best guess. Perhaps an analogy might be the difference between the more advanced Aztecs and Incas, who indeed had slavery (and ritual sacrifices using slaves) and the Plains Indians, who often, particularly with young people, adopted them into the tribe.

This is the study to which I was referring about assimilation of hunter-gatherers in the north-central Balkans around Lepinski Mir:
http://www.pnas.org/content/110/9/3298
"Strontium Isotopes document greater human mobility at the start of the Balkan Neolithic"


This is the abstract:
Questions about how farming and the Neolithic way of life spread across Europe have been hotly debated topics in archaeology for decades. For a very long time, two models have dominated the discussion: migrations of farming groups from southwestern Asia versus diffusion of domesticates and new ideas through the existing networks of local forager populations. New strontium isotope data from the Danube Gorges in the north-central Balkans, an area characterized by a rich burial record spanning the Mesolithic–Neolithic transition, show a significant increase in nonlocal individuals from ∼6200 calibrated B.C., with several waves of migrants into this region. These results are further enhanced by dietary evidence based on carbon and nitrogen isotopes and an increasingly high chronological resolution obtained on a large sample of directly dated individuals. This dataset provides robust evidence for a brief period of coexistence between indigenous groups and early farmers before farming communities absorbed the foragers completely in the first half of the sixth millennium B.C.

Following is a link to the supplementary information. It's well worth a read.
http://www.pnas.org/content/suppl/2013/02/06/1211474110.DCSupplemental

Ed. Perhaps this is the source of the "hunter-gatherer" component of EEF.

it makes sense, at first there was place for 2 complimentary economies at Lepenski Vir : the fisher/hunter/gatherer and the new neolithic one
both economies could benefit from mutual trade, as their products (and resources) were complimentary
but when the neolithic population kept growing, it must have created problems
a tough choice for the original fisher/hunter/gatherers
 
Finally an interesting study this summer ! There hadn't been much happening for several months.

These new samples come from the Early Neolithic Starčevo and LBK cultures. The most valuable results here are the ancient Y-DNA. Haak 2010 and Brandt 2013 had already tested three Y-DNA samples from the LBK culture in Germany and yielded haplogroups F* and G2a2b. Identically the same haplogroups were found in these Hungarian samples, with the addition of haplogroups I2a1 and I1. I2a1 was also found among Neolithic farmers in France and probably represents the Mesolithic European population assimilated by Near Eastern farmers in the Balkans before their expansion across Europe. I1 is by far the most interesting for two reasons:

1) It is the oldest attested existence of I1 and it suggests that I1 may have been far more widespread in the Mesolithic than the Baltic region.

2) Since I1 hasn't been found yet among Mesolithic Scandinavians (who so far were found to belong to I*, I2* and I2a1), it could mean that I1 was also among the first lineages of Mesolithic Europeans assimilated by Neolithic farmers, and that I1 actually entered Scandinavia during the Neolithic/Chalcolithic. In other wors, I1 could have been living in the Balkans in the Mesolithic, then spread to Germany with the LBK culture, then to Scandinavia afterwards.
"We sampled one Mesolithic, 47 Starčevo and 61 LBKT skeletons, excavated in Croatia and western Hungary.

I1 was found in western Hungary not in the Balkans.
The STA expanded from present-day Serbia to the western part of the Carpathian Basin, encompassing the regions of today’s northern Croatia and south-western Hungary (ca. 6,000- 5,400 BC) [7,8] (Figure 1), and resulting in the formation of the Linearbandkeramik culture (LBK) [9]."

 
Journal of Language Relationship • 9 (2013) • Pp. 69–92 • © Dybo A., 2013
Anna Dybo
Russian State University for the Humanities (Moscow)
Language and archeology: some methodological problems.1. Indo-European and Altaic landscapes
The article is the first part of a larger work that represents an attempt to systematize ourideas on the natural environment and material culture of the Proto-Indo-Europeans. It is based on a more or less complete selection of reconstructed words from the appropriate semantic areas and on their comparison with a similar selection performed for a proto-language of similar time depth, whose speakers evidently inhabited a territory that was notin contact with the Proto-Indo-European one — Proto-Altaic. In this part, only the words that belong to the semantic field of landscape terms are analyzed. The main conclusion is that thehypothesis of a steppe environment is more applicable for the Proto-Altaic population,whereas for Proto-Indo-Europeans a mountainous region seems more appropriate. As forthe water bodies, for Proto-Indo-Europeans we should suppose the existence of a sea (or of avery big lake), and for speakers of Proto-Altaic, the existence of very big rivers with seasonfloods.
http://jolr.ru/files/(108)jlr2013-9(69-92).pdf
 

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