Tracing the genetic origin of Europe's first farmers

Eldritch

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Farming was established in Central Europe by the Linearbandkeramik culture (LBK), a well-investigated archaeological horizon, which emerged in the Carpathian Basin, in today's Hungary. However, the genetic background of the LBK genesis has not been revealed yet. Here we present 9 Y chromosomal and 84 mitochondrial DNA profiles from Mesolithic, Neolithic Starčevo and LBK sites (7th/6th millennium BC) from the Carpathian Basin and south-eastern Europe. We detect genetic continuity of both maternal and paternal elements during the initial spread of agriculture, and confirm the substantial genetic impact of early farming south-eastern European and Carpathian Basin cultures on Central European populations of the 6th-4th millennium BC. Our comprehensive Y chromosomal and mitochondrial DNA population genetic analyses demonstrate a clear affinity of the early farmers to the modern Near East and Caucasus, tracing the expansion from that region through south-eastern Europe and the Carpathian Basin into Central Europe. Our results also reveal contrasting patterns for male and female genetic diversity in the European Neolithic, suggesting patrilineal descent system and patrilocal residential rules among the early farmers.

Three STA individuals belong to the NRY haplogroup F* (M89) and two specimens can be assigned to the G2a2b (S126) haplogroup, and one each to G2a (P15) and I2a1 (P37.2) (Dataset S3, S5).

http://biorxiv.org/content/early/2014/09/03/008664
 
To me, the most interesting detail was this.

"The two investigated LBKT samples carry haplogroups G2a2b (S126) and I1 (M253). Furthermore, the incomplete SNP profiles of eight specimens potentially belong to the same haplogroups; STA: three G2a2b (S126), two G2a (P15), and one I (M170); LBKT: one G2a2b (S126) and one F* (M89) (Dataset S5).

So, while most of the Y DNA was typical of Neolithic farmers from the Middle East, Y haplotype I and specifically I1 was also present.
 
Interesting to see I1 there; it shouldn't be too surprising though as we don't know a lot about its early history but this is very interesting! And another case against R1b being from farmers.
 
What does this map mean from the paper?
Y.jpg
 
It seems that in addition to I2a1, I1 was one of the y lineages that were assimilated by the Neolithic farmers very early on. This is consistent with papers showing that the assimilation of hunter-gatherers in the Balkans happened within a few generations, whatever might have been the case in central Europe.
 
What surprised me was to see in neolithic farmers a very large percentage of F* which they say was very related to G2a. Mind that most current ydna in Eurasia descends from F* but very far down the tree.
 
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.
 
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.

I never thought about it before in quite that light, but the age of this I1 and its presence in a Neolithic farming setting in the Carpathian Basin may indicate that, after being assimilated by Neolithic farmers whose ancestors came from the Middle East, I1 folk entered Scandinavia as Neolithic farmers, and that's why I1 is so common there, even though it may have evolved in Europe from IJ. So, instead of I1 being proof of a high rate of survival of pre-Neolithic hunter/gatherer type folk in Scandinavia after the advent of farming in that area, I1 could actually be proof of farmers displacing hunter/gatherer types in Scandinavia. Perhaps the I1 farmers pushed out the I2 hunter/gatherers.
 
Interesting to see I1 there; it shouldn't be too surprising though as we don't know a lot about its early history but this is very interesting! And another case against R1b being from farmers.

Given the time frame, this does not in fact tell us anything about whether R1b could have entered Europe during the late Neolithic. IMO, that's a separate issue.
 
It seems that in addition to I2a1, I1 was one of the y lineages that were assimilated by the Neolithic farmers very early on. This is consistent with papers showing that the assimilation of hunter-gatherers in the Balkans happened within a few generations, whatever might have been the case in central Europe.
I wouldn't jump into these conclusions too fast Angela. If farmers had slaves, from what populations do you think these slaves would come from?
 
I never thought about it before in quite that light, but the age of this I1 and its presence in a Neolithic farming setting in the Carpathian Basin may indicate that, after being assimilated by Neolithic farmers whose ancestors came from the Middle East, I1 folk entered Scandinavia as Neolithic farmers, and that's why I1 is so common there, even though it may have evolved in Europe from IJ. So, instead of I1 being proof of a high rate of survival of pre-Neolithic hunter/gatherer type folk in Scandinavia after the advent of farming in that area, I1 could actually be proof of farmers displacing hunter/gatherer types in Scandinavia. Perhaps the I1 farmers pushed out the I2 hunter/gatherers.

Probably not since I count a grand total of maybe 4-5 distinct G2a male lineages in Scandinavia. I don't believe the LBK farmers ever made a push into far north Europe - the society collapsed long before then. Based on the dates, 'farming' pushed into Scandinavia during the copper or bronze age, at which point R1b/R1a had already been introduced to Europe. Lo and behold, the 2nd and 3rd most common lineages in Scandinavia are R1b and R1a respectively. These later immigrants had farming, but they were probably not the "first" European farmers. The LBK left a very minimal imprint on far northern Europe.
 
I never thought about it before in quite that light, but the age of this I1 and its presence in a Neolithic farming setting in the Carpathian Basin may indicate that, after being assimilated by Neolithic farmers whose ancestors came from the Middle East, I1 folk entered Scandinavia as Neolithic farmers, and that's why I1 is so common there, even though it may have evolved in Europe from IJ. So, instead of I1 being proof of a high rate of survival of pre-Neolithic hunter/gatherer type folk in Scandinavia after the advent of farming in that area, I1 could actually be proof of farmers displacing hunter/gatherer types in Scandinavia. Perhaps the I1 farmers pushed out the I2 hunter/gatherers.

Exactly. That also explains why Scandinavians have so much Mediterranean admixture in the Dodecad K12, and even more Early European Farmer(EEF)admixture (>30%) in the Lazaridis 2014 paper. But that doesn't mean that Pre-Indo-European Scandinavians were already a blend of I1 and G2a lineages. The current evidence rather suggests that I1 entered Scandinavia late, perhaps with R1a during the Corded Ware period, or just before that.

What seems quite likely is that Neolithic Germans and Poles were a blend of Mesolithic I1 from central or southeast Europe and Neolithic farmers from the eastern Mediterranean. This is the population that the R1a people of the Corded Ware would have met advancing from Ukraine and Belarus westward to Germany.

The oddest thing so far is the high percentage of F* in Central European Neolithic samples. Either they didn't test all the subclades or that means that this male population underwent a severe pruning, possibly when the Indo-Europeans invaded, considering that 60 to 70% of Polish and German paternal lineages are now either R1a or R1b.
 
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.

finding I1 is maybe the biggest surprise of the year, indeed
I1 expansion was estimated some 4300 years old
the sample is 4900 - 5600 old, so this might be the single source from where all present-day I1 expanded

the Balkans south of the Danube and north of Greece were uninhabited during the mesolithic

this suggests I2a1 - P37.2 originated in the Greek ice age refuge , from where the split north (starcevo) and west along with cardium pottery (Sardinia, Southern France)

I1 may have been in the Italian/Adriatic refuge, which expanded into the Carpathian basin some 17000 years ago

what happened to F* ? isn't it very rare in Europe?
 
G2a wins again, but it's the elusive I1 sample that will get a lot of play out of this. This particular I1 sample may predate the TMRCA of (at least most) modern I1, so it's perhaps no surprise that it's not very close to the area of the highest modern frequency of I1. Actually, it's not too far from an outlier subclade of modern I1, which has presence in the Czech Republic IIRC. So although this doesn't prove much about I1's past, it's certainly evidence that it could have spread from that direction.

I wish they had tested for the subclade of I2a1. As is, it could be I2a-M26 (consistent with other Neolithic farmer samples) or an I2a-Din relative (which would be very notable!) or it could be something else. Oh well.
 
I wouldn't jump into these conclusions too fast Angela. If farmers had slaves, from what populations do you think these slaves would come from?

where did they find the skeletons? was it in a burrial site
i don't think slaves got proper burrial at that time
 
About those F*'s:

Based on the supplementary datasets, here are the haplogroups the first two F*'s could be:
F*, F1, F2, F3, H, IJ*, IJK*, HIJK*, GHIJK*

The second two F*'s got no-calls for J* (but negative for J1 and J2) so they could be any of those as well, or J*.
 
I wouldn't jump into these conclusions too fast Angela. If farmers had slaves, from what populations do you think these slaves would come from?

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.
 
What does this map mean from the paper?
View attachment 6604
Figure 4. Genetic distance map of the STA-LBK Y chromosomal data.
Y chromosomal genetic distances (Fst) were computed between the STA-LBK samples and 100 present-day
populations of Eurasia and North Africa and visualized on a geographic map. Grey dots denote the
location of present-day populations. Color shadings indicate the degree of similarity or dissimilarity of
Neolithic samples to the modern-day populations. Short distances and great similarities to present-day
populations are marked by red areas. Fst values were scaled by an interval range of 0.01. Fst values higher
than 0.21 were not differentiated (grey areas). The map shows remarkable affinities of the STA-LBK
Downloaded from http://biorxiv.org/ on September 4, 201436
samples to present-day populations of the northwest and south Caucasus. Population information and Fst
values are listed in table S15.
 
About those F*'s:

Based on the supplementary datasets, here are the haplogroups the first two F*'s could be:
F*, F1, F2, F3, H, IJ*, IJK*, HIJK*, GHIJK*

The second two F*'s got no-calls for J* (but negative for J1 and J2) so they could be any of those as well, or J*.

quite a broad definition of F*
which one is most common among todays Europeans?
isn't that F2?
 
Figure 4. Genetic distance map of the STA-LBK Y chromosomal data.
Y chromosomal genetic distances (Fst) were computed between the STA-LBK samples and 100 present-day
populations of Eurasia and North Africa and visualized on a geographic map. Grey dots denote the
location of present-day populations. Color shadings indicate the degree of similarity or dissimilarity of
Neolithic samples to the modern-day populations. Short distances and great similarities to present-day
populations are marked by red areas. Fst values were scaled by an interval range of 0.01. Fst values higher
than 0.21 were not differentiated (grey areas). The map shows remarkable affinities of the STA-LBK
Downloaded from http://biorxiv.org/ on September 4, 201436
samples to present-day populations of the northwest and south Caucasus. Population information and Fst
values are listed in table S15.

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 ?
 

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