Genomes From Verteba Cave Suggest Diversity Within The Trypillians In Ukraine

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https://www.researchsquare.com/article/rs-1044480/v1

Abstract

The transition to agriculture occurred relatively late in Eastern Europe, leading researchers to debate whether it was a gradual, interactive process or a colonization event. In the forest and forest-steppe regions of Ukraine, farming appeared during the fifth millennium BCE, associated with the Cucuteni-Trypillian Archaeological Complex (CTCC, 4800-3000 BCE). Across Europe, the Neolithization process was highly variable across space and over time. Here, we investigate the population dynamics of early agriculturalists from the eastern forest-steppe region based on analyses of 20 ancient genomes from the Verteba Cave site (3789-980 BCE). The results reveal that the CTCC individuals’ ancestry is related to both western hunter gatherers and Near Eastern farmers, lacks local ancestry associated with Ukrainian Neolithic hunter gatherers and has steppe ancestry. An Early Bronze Age individual has an ancestry profile related to the Yamnaya expansions but with 20% ancestry related to the other Trypillian individuals, which suggests admixture between the Trypillians and the incoming populations carrying steppe-related ancestry. A Late Bronze Age individual dated to 980-948 BCE has a genetic profile indicating affinity to Beaker-related populations, detected close to 1,000 years after the end of the Bell Beaker phenomenon during the Third millennium BCE.

Supplementary Tables with Y and mtDNAs:
https://assets.researchsquare.com/files/rs-1044480/v1/be83b4a5450c38c4a77088d7.xlsx
 
The final number they arrive at is still about 80% Anatolia Neolithic with 20% HG for all but two of the samples, although there is some accretion of steppe gradually over time.

Their conclusions in more detail:
"These observations broadly suggest that Eneolithic CTCC descended from the same, or closely related, population that spread the Neolithic across most of Europe and without little or no sign of admixture with earlier Ukrainian Mesolithic or Neolithic groups composed of hunter-gatherer-related ancestry and specifically pointing towards the Baden individuals from Hungary. In fact, most of the Trypillian individuals can be modeled by Eneolithic populations from Europe that have steppe ancestry; however, 4 out of the 20 individuals could be modeled as Moldovan Trypillians. These results in the qpAdm modeling suggest that there were differences in the ancestry composition of the Trypillians of Verteba cave, which could be linked to the proportion of HG in the individuals, although this variability is not substantial enough to differentiate the individuals into different population.

Previous studies of CTCC individuals could not provide a clear origin for the HG component of CTCC-associated groups. Here, although models including Ukranian_N individuals and WHG seem to work, the f4 -statistics suggest that the source of the HG component would be mainly WHG. In addition, not a single qpAdm model using EHG as a source works, which supports that observation. The significant proportion of WHG ancestry found in the Trypillians (up to 18%) might be related to the hunter gatherer resurgence seen in other Middle Neolithic populations of Central Europe, likely due to admixture with groups in the west who already had a higher WHG component derived from Anatolia-related Neolithic groups prior to the origin of the CTCC 8,13,56

This would also indicate that the HG Neolithic populations from Ukraine did not contribute much ancestry to the Trypillians. In addition, we also observed the presence of steppe-related ancestry in these individuals, as was revealed in Moldova 37 , although the proportion in the Verteba individuals was lower, which could correlate with the age of the individuals, suggesting a continuous pulse from the East to the West gradually increasing the Yamnaya-related ancestry during the 4th millennium.

Individual VERT-113, dated to the EBA (1952-1774 BCE), has an ancestry profile that is quite different from the earlier CTCC individuals. There was significantly more Caucasus HG/Yamnaya and EHG ancestry, and thus, this individual was related to Yamnaya expansions. qpAdm results suggest a link between VERT-113 and Corded Ware populations from Poland, pointing to a similarity between this individual and these populations. Additionally, this is the only individual with a higher genetic affinity to Ukraine_N than to WHG, suggesting that the population that originated EBA in the 2nd millennium BC may have shared affinities with the Ukraine_N populations.

Interestingly, VERT-114 (Late Bronze Age) does not show many genetic connections with EBA VERT-113 according to the f3 values, which is clearly associated with Yamnaya pastoralists. The genomic composition of VERT-114 suggests a relationship with Beaker-related populations, despite being almost 1,000 years younger than the end of the Bell Beaker phenomenon 57 , and with a date that would be more coincident with the Cimmerians or Scythians 58 . However, no qpAdm models with these cultures work, and the f4 results seem to confirm the similarity with the Bell Beakers over the Cimmerians. The genetic background of this individual, with its strong Western affinities, supports the evidence shown in Narasimhan et al. 59 of a western influx into the steppe during the Late Bronze Age."

Having read through all the findings, the conclusions seem justified.

Two interesting things:

1)"VERT-113, dated to around the EBA, shows a clear signal of steppe-related ancestry and is the only individual in thedataset that shows a strong influx of this ancestry: f4(Mbuti, Verteba_EBA; Russia_Samara_EBA_Yamnaya, LBK):-0.00398 Z score=-7.848. The same test with individual VERT-114 was not statistically significant (Z=1.382). Relevantly,we observe a major affinity to Russia_Yamnaya over Ukraine_Yamnaya using f4 statistics (Table S4). Furthermore, this isthe only individual who shows a major affinity to Ukranian_N over WHG as the source of HG-related ancestry, as shownby the statistic f4(Mbuti, VERT-113; Ukraine_N, WHG): -0.001276, Z score (Z=-4.202). The distal models of qpAdm usingbasal ancestries reveal that this individual exhibits up to 33% Ukraine_N and 66% CHG, supporting high amounts ofsteppe-related ancestry. When modeled with close chronology populations, the individual requires a single source relatedto the Corded_Ware (Fig. 4, Table S4). We tried to assess whether the signal could, however, correspond to similar geneticpopulations but more contemporary and geographically closer to VERT-113, such as Srubnaya, using the statisticf4(Mbuti, VERT-113; Poland_Southeast_CordedWare, Russia_Srubnaya), but the results (f4=0.00003, Z=0.11) show thatthere is no statistical relationship, which indicates no evidence supporting the Srubnaya origin."

2) There is no sample with the LP allele, and all but two of the samples probably had blue eyes.

I'm sure those with a special interest in pigmentation will pick the data apart, but I suspect that previous descriptions of them as unusually fair will probably be born out, which is interesting in a population basically 80% Anatolia Neolithic and 20% Koros like HG. Something happened.
 
As for the LBA individual:

Individual VERT-114, dated around the LBA, showed a genetic position close to Bell-Beaker populations in PCA and ADMIXTURE. This individual shows a higher influx of ancestry from WHG than from EHG populations f4(Mbuti, VERT-114; WHG, EHG) -0.002, Z score=-8,64), similar to the results obtained for the aggregate group of 22 Verteba individuals. qpAdm results for this individual show that a single model with a Bell_Beaker population as a single source works (Fig. 4). Many of the two-way models involve populations related to Ukraine_Globular_Amphora and to steppe populations, with approximately 60% of ancestry from the former and the rest from the latter.
Click to expand...

https://www.researchsquare.com/article/rs-1044480/v1

I guess it will be closer to G?va and Kyjatice, probably F?zesabony, rather than Beakers, but let's see. They didn't compare with those single samples, but they used Mako too, which being not mentioned and wouldn't fit anyway, because its too WHG. The individual is a female, mtDNA T2.

Looking at which Beakers she is closest too, its clear, its going in a G?va direction:

Verteba_LBA* Poland_Southeast_BellBeaker
Verteba_LBA* Hungary_EBA_BellBeaker
Click to expand...

That's not British Beakers of course...

I would interpret all the results as mounting evidence for a local continuity of a WHG heavy Neolithic population around the Carpathians and Pannonia, which was later fusing with Epi-Corded/Beaker groups to create the North Pannonian clusters which emerged with G?va, Kyjatice and F?zesabony/late Otomani.
 
As expected the three I2 samples don't belong to I2-L621 subclade. In the Eupedian article about Haplogrup I2 > Haplogroup I2a1b (M423) > Haplogroup I2a1b-L621 is claimed:

The high concentration of I2a1b-L621 in north-east Romania, Moldova and central Ukraine reminds of the maximum spread of the Cucuten > Trypillian culture (4800-3000 BCE). No Y-DNA sample from this culture has been tested to date, but as it evolved as an offshoot from the Starčevo–Kőrös–Criş culture, it is likely that I2a was one of its main paternal lineages, and a founder effect could have increased considerably its frequency. The Cucuten > Trypillian culture was the most advanced Neolithic culture in Europe before the Indo-European invasions in the Bronze Age and seems to have had intensive contacts with the Steppe culture before the expansion of Yamna to the Balkans and Central Europe (see histories of R1a and R1b). From 3500 BCE, at the onset of the Yamna period in the Pontic-Caspian Steppe, the Cucuten > Trypillian people started expanding east into the steppe of what is now western Ukraine, leaving their towns (the largest in the world at the time), and adopting an increasingly nomadic lifestyle like their Yamna neighbours. It can easily be imagined that Cucuten > Trypillian people became assimilated by the Yamna neighbours and that they spread as a minority lineage alongside haplogroups R1a and R1b as they advanced toward the Baltic with the Corded Ware expansion.
Click to expand...

This is completely wrong and should be removed since the theory is not based on any evidence or even worse is outdated for some time now. The most probable spread of I2-L621 > I-CTS10936 > I-S19848 > I-CTS4002 > I-CTS10228 > I-Y3120 was from central Europe by some Proto-Celtic population associated with Urnfield culture which moved to Eastern Europe and their descendants prospered in the Pre-Slavic & Proto-Slavic ethnogenesis.
 
from the paper :

It is also interesting to remark that, except for two individuals, the majority of individuals from Verteba cave have the variant of SNP rs12913832 associated with blue eyes and the other two associated with dark eyes​




1-s2.0-S0002929719301922-gr3.jpg


i am A/G eyes are hazel
 
kingjohn said:
from the paper :

It is also interesting to remark that, except for two individuals, the majority of individuals from Verteba cave have the variant of SNP rs12913832 associated with blue eyes and the other two associated with dark eyes​




1-s2.0-S0002929719301922-gr3.jpg


i am A/G eyes are hazel
Click to expand...

What was with the LBA sample?
 
Miroslav said:
As expected the three I2 samples don't belong to I2-L621 subclade. In the Eupedian article about Haplogrup I2 > Haplogroup I2a1b (M423) > Haplogroup I2a1b-L621 is claimed:



This is completely wrong and should be removed since the theory is not based on any evidence or even worse is outdated for some time now. The most probable spread of I2-L621 > I-CTS10936 > I-S19848 > I-CTS4002 > I-CTS10228 > I-Y3120 was from central Europe by some Proto-Celtic population associated with Urnfield culture which moved to Eastern Europe and their descendants prospered in the Pre-Slavic & Proto-Slavic ethnogenesis.
Click to expand...

Ok "Mr. completely wrong". There are only 11 Y-DNA samples in this study, including one I2a2a1, one I2a1a2a and one I2c. That's a lot of I2 variety. The study does not mention the SNPs tested, but if they follow the latest phylogeny I2a2a1 should be I2-CTS616 and I2a1a2a should be I2-Y3104, which is just upstream of (and ancestral to) I2-L621. This means that there is still a good chance to find I2-L621 in other samples from the Cucuteni-Trypillian culture.
 
Maciamo said:
Ok "Mr. completely wrong". There are only 11 Y-DNA samples in this study, including one I2a2a1, one I2a1a2a and one I2c. That's a lot of I2 variety. The study does not mention the SNPs tested, but if they follow the latest phylogeny I2a2a1 should be I2-CTS616 and I2a1a2a should be I2-Y3104, which is just upstream of (and ancestral to) I2-L621. This means that there is still a good chance to find I2-L621 in other samples from the Cucuteni-Trypillian culture.
Click to expand...

No, as other contemporary studies they used 2016/17 ISOGG phylogeny tree which still had I2c hence I2a1a2a is I2-L1286 which is not and cannot be ancestral to I2a1b2-L621.


We have three samples from 2018 study all G2a; numerous samples from Neolithic-Chalcolithic archaeological cultures directly or indirectly related to Cucuteni-Trypillian culture and in all of them there is absence of I2-L621 - because it was not part of an Southeastern European culture. Until the Roman period we have evidence for I2-L1286 and I2-CTS616 but not I2-L621 (> I-CTS10936 > I-S19848 > I-CTS4002 > I-CTS10228 > I-Y3120+) because it arrived only in the migration period. Through I2-L1286 & I2-CTS616 and on other side I2-L621 we follow and differentiate population until the end of Roman period from the migrating mainly Slavic population from the Early Middle Ages. Using variety as an argument is highly misleading in this case.
 
Miroslav said:
...it was not part of an Southeastern European culture.
Click to expand...

I2-L621+ was not part of Neolithic culture in both Southeastern and Eastern Europe because it migrated from Central European late Bronze Age culture to Eastern Europe and from there in the Middle Ages migrated to Southeastern Europe as well as partly Central Europe.
 
They had a round face with little longer lower part, a broad and high forehead, narrow eye sockets, wide set eyes. A straight or upturned long nose bridge, nose tip points down, little broader nostrils. Broad and big mouth. European eyelids.

They had very likely blue eyes and light skin.

Wavy hair, brown to blonde.

https://www.wertpapier-forum.de/uploads/post-2022-1192022240_thumb.jpg

https://external-content.duckduckgo...NxB37HJgM3lpLpNx8lQtLL4CKqL8g=s0-d&f=1&nofb=1

They looked different from Anatolian Farmers or typical Linear Pottery

Matching with today's populations based on trait SNPs:

United Kingdom 81%
Swede 80%
Danish 78%
Estonian 76%
Ukrainian 70%
Basque 66%
Netherlands 64%
French 63%
Middle East 62% Sardinian 62%
Italian 58%
Iberian 55%
Russian 54%
Hungarian 50%
 
Last edited:
Maciamo said:
Ok "Mr. completely wrong". There are only 11 Y-DNA samples in this study, including one I2a2a1, one I2a1a2a and one I2c. That's a lot of I2 variety. The study does not mention the SNPs tested, but if they follow the latest phylogeny I2a2a1 should be I2-CTS616 and I2a1a2a should be I2-Y3104, which is just upstream of (and ancestral to) I2-L621. This means that there is still a good chance to find I2-L621 in other samples from the Cucuteni-Trypillian culture.
Click to expand...

they don't follow the latest phylogeny. "i2c" was last used in the 2017 isogg tree.

according to that tree:

I2a2a1- I-CTS616 https://www.yfull.com/tree/I-CTS616/
I2a1a2a- I-L1286/I-Y4192 https://www.yfull.com/tree/I-Y4192/
I2c- I-L596 https://www.yfull.com/tree/I-L596/
 
Doggerland said:
They had a round face with little longer lower part, a broad and high forehead, narrow eye sockets, wide set eyes. A straight or upturned long nose bridge, nose tip points down, little broader nostrils. Broad and big mouth. European eyelids.

They had very likely blue eyes and light skin.

Wavy hair, brown to blonde.

https://www.wertpapier-forum.de/uploads/post-2022-1192022240_thumb.jpg

https://external-content.duckduckgo...NxB37HJgM3lpLpNx8lQtLL4CKqL8g=s0-d&f=1&nofb=1

They looked different from Anatolian Farmers or typical Linear Pottery

Matching with today's populations based on trait SNPs:

United Kingdom 81%
Swede 80%
Danish 78%
Estonian 76%
Ukrainian 70%
Basque 66%
Netherlands 64%
French 63%
Middle East 62% Sardinian 62%
Italian 58%
Iberian 55%
Russian 54%
Hungarian 50%
Click to expand...

Please post a picture of a typical person from the British Isles who has those traits, please.
 
https://i2-prod.walesonline.co.uk/i...st-women-in-wales-20-11-gallery-666309537.jpg

https://hips.hearstapps.com/hmg-prod.s3.amazonaws.com/images/emily-atack-at-film-premiere-2018-1544970617.jpg?crop=1.00xw:0.669xh;0,0.0139xh&resize=480:*

https://upload.wikimedia.org/wikipedia/commons/thumb/2/25/Cara_Delevingne_by_Gage_Skidmore.jpg/1200px-Cara_Delevingne_by_Gage_Skidmore.jpg

https://www.newdvdreleasedates.com/images/profiles/sam-heughan-06.jpg

https://vistapointe.net/images/hugh-dancy-1.jpg

https://hydra.mediasearch.verizon.com/ResourcesFiles/ImageLibrary/Celebrities/V/34_Lrg/238249.jpg
 
As I suspected. None of the men approach the look you're describing, imo. Long faces with angular jaws are not round. Cara Delavigne and the first girl have somewhat round faces and reasonably wides-paced eyes, but neither, in fact not one of your examples, has a long nose, much less a long nose which points downward. They have snub noses, which have quite a different shape.

Sorry, not convinced.

What you're describing sounds to me like Alpines from eastern and southeastern Europe, which makes sense given where the Cave in question is found.

The plates are easily accessible, but here is an example.

troe141.jpg
 
Angela said:
As I suspected. None of the men approach the look you're describing, imo. Long faces with angular jaws are not round. Cara Delavigne and the first girl have somewhat round faces and reasonably wides-paced eyes, but neither, in fact not one of your examples, has a long nose, much less a long nose which points downward. They have snub noses, which have quite a different shape.

Sorry, not convinced.
Click to expand...

The guy you posted has a flat and narrow forehead, doesn't match much I would suggest.
But you would never find someone who matches 100% to a description. 80% for example means still 20% not matching with the sample of a population.

More examples of Brits who where similar to them:

https://ilarge.lisimg.com/image/8592111/740full-emma-rigby.jpg

https://fr.web.img3.acsta.net/pictures/17/02/10/15/26/425395.jpg

They don’t looked like today's southeastern Europeans, but northern Europeans and Brits. They where far from looking southeastern European in any way.
Southern Europeans of today don’t really match any ancient European population from Neolithic or Bronze Age in traits, because they had many population changes in the past. Sardinias who where thought to carry most neolithic ancestry, doesn't match much to Linear Pottery in traits. It is hard to find any population from those time periods matching the Iberian people too. Cardial Pottery for example only matches 50% to today's Iberians.

Neolithic Brits also had blue eyes and fair skin, but only matching 74% with today's United Kingdom people.
The reason could be that Tripillians where one of the main components of the people that later invaded the Isles.

And there is another thing. Non of the people I posted looks long faced in my opinion. But it could be because we are raised in a different social environment, we don’t rate traits of people similar.
This would I consider long faced Brits:

https://www2.pictures.zimbio.com/gi/Premiere+Of+Rogue+Red+Carpet+qoDjej635XMx.jpg

https://bestofcomicbooks.com/wp-content/uploads/2018/06/kaya-scodelario-cleavage.jpg
 
Miroslav said:
As expected the three I2 samples don't belong to I2-L621 subclade. In the Eupedian article about Haplogrup I2 > Haplogroup I2a1b (M423) > Haplogroup I2a1b-L621 is claimed:



This is completely wrong and should be removed since the theory is not based on any evidence or even worse is outdated for some time now. The most probable spread of I2-L621 > I-CTS10936 > I-S19848 > I-CTS4002 > I-CTS10228 > I-Y3120 was from central Europe by some Proto-Celtic population associated with Urnfield culture which moved to Eastern Europe and their descendants prospered in the Pre-Slavic & Proto-Slavic ethnogenesis.
Click to expand...

I don't see any argument in favour of a proto-Celtic or Celtic pop rich for Y-I2a1-L621, and no more in favour of a measurable Celtic input into the Slavic ethnogenesis, to date.
Concerning Y-I2a1b, I am not a specialist, but a strong presence in Eastern Carpathians at the margins with CTC doesn't seem so amazing even if it was not the typical haplo of CTC core. It could have furnished human elements in Daco-Thracian tribes at the fringes of the proto-Slavic world of future, whose some have been integrated later in the Slavs expansion;
 
Miroslav said:
No, as other contemporary studies they used 2016/17 ISOGG phylogeny tree which still had I2c hence I2a1a2a is I2-L1286 which is not and cannot be ancestral to I2a1b2-L621.


We have three samples from 2018 study all G2a; numerous samples from Neolithic-Chalcolithic archaeological cultures directly or indirectly related to Cucuteni-Trypillian culture and in all of them there is absence of I2-L621 - because it was not part of an Southeastern European culture. Until the Roman period we have evidence for I2-L1286 and I2-CTS616 but not I2-L621 (> I-CTS10936 > I-S19848 > I-CTS4002 > I-CTS10228 > I-Y3120+) because it arrived only in the migration period. Through I2-L1286 & I2-CTS616 and on other side I2-L621 we follow and differentiate population until the end of Roman period from the migrating mainly Slavic population from the Early Middle Ages. Using variety as an argument is highly misleading in this case.
Click to expand...

It's annoying when studies don't mention the actual SNPs. Even if you were right, though, we'd still have three different I2 lineages out of three I2 samples. Who is to say that if they test 1000 Cucuteni-Trypillian samples there won't be any I2-L621.

One thing I don't understand is why you write L621 (> I-CTS10936 > I-S19848 > I-CTS4002 > I-CTS10228 > I-Y3120+) ? I was talking about L621, not its subclades. I-Y3120 is 3400 years old, so it's impossible that it is belongs to the Cucuteni-Trypillian culture, which started 7500 years ago and ended almost 5000 years ago.
 
MOESAN said:
I don't see any argument in favour of a proto-Celtic or Celtic pop rich for Y-I2a1-L621, and no more in favour of a measurable Celtic input into the Slavic ethnogenesis, to date.
Concerning Y-I2a1b, I am not a specialist, but a strong presence in Eastern Carpathians at the margins with CTC doesn't seem so amazing even if it was not the typical haplo of CTC core. It could have furnished human elements in Daco-Thracian tribes at the fringes of the proto-Slavic world of future, whose some have been integrated later in the Slavs expansion;
Click to expand...

https://www.yfull.com/tree/I-CTS10228/

The Slavic subclade is I-Y3120, it split 3400 years ago from I-Y81696 which is found in Bas-Rhin and Baden-Wurttemberg.
3400 years ago is just when the mutation occured, the migration into Eastern Europe probably wasn't immedieately but some generations after.

More widely speaking these are subclades of the 14,100 year old I-M423, which was found in mesolithic and neolithic samples from Scandinavia, British Isles, Spain, etc.

https://www.yfull.com/tree/I-M423/

Even today I-M423 is found only in western Europe. Out of 100s of modern and ancient samples, in Eastern Europe there is only this rare subclade : https://www.yfull.com/tree/I-Y13336/
and the Slavic one which I already mentioned, which splits from the West European ones 3400 years ago.

With such deep roots, I can hardly imagine what kind of samples could be found which would shift the origins of I-L621 away from Western Europe.
 
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