# Population Genetics > Paleogenetics > Paleolithic & Mesolithic >  More genomes from Mesolithic Romania and Spain (Gonzales-Fortes et al. 2017)

## Maciamo

It's relatively small paper compared to the mastodons published earlier this month, but nevertheless interesting. 

Paleogenomic Evidence for Multi-generational Mixing between Neolithic Farmers and Mesolithic Hunter-Gatherers in the Lower Danube Basin (extended PDF with supplementary materials here)

*Summary*

_"The transition from hunting and gathering to farming involved profound cultural and technological changes. In Western and Central Europe, these changes occurred rapidly and synchronously after the arrival of early farmers of Anatolian origin, who largely replaced the local Mesolithic hunter-gatherers. Further east, in the Baltic region, the transition was gradual, with little or no genetic input from incoming farmers. Here we use ancient DNA to investigate the relationship between hunter-gatherers and farmers in the Lower Danube basin, a geographically intermediate area that is characterized by a rapid Neolithic transition but also by the presence of archaeological evidence that points to cultural exchange, and thus possible admixture, between hunter-gatherers and farmers. We recovered four human paleogenomes (1.1× to 4.1× coverage) from Romania spanning a time transect between 8.8 thousand years ago (kya) and 5.4 kya and supplemented them with two Mesolithic genomes (1.7× and 5.3×) from Spain to provide further context on the genetic background of Mesolithic Europe. Our results show major Western hunter-gatherer (WHG) ancestry in a Romanian Eneolithic sample with a minor, but sizeable, contribution from Anatolian farmers, suggesting multiple admixture events between hunter-gatherers and farmers. Dietary stable-isotope analysis of this sample suggests a mixed terrestrial/aquatic diet. Our results provide support for complex interactions among hunter-gatherers and farmers in the Danube basin, demonstrating that in some regions, demic and cultural diffusion were not mutually exclusive, but merely the ends of a continuum for the process of Neolithization."_

The three Mesolithic southern Romanians belonged to Y-DNA/mtDNA haplogroups R/U5b2c, R1/U5a1c and R1b/K1 + 16362.

The two Mesolithic northwest Spaniards are female and belong to mtDNA haplogroups U5a2a and U5b.

A female Eneolithic sample (5,375 cal BP) from Gura Baciului in central Romania was also tested and belonged to K1a4a.

All Mesolithic samples were pure Mesolithic European autosomally (apparently only WHG). The Chalcolithic sample was about 65% Mesolithic HG, 30% Neolithic farmer and 5% of Steppe/EHG (without CHG).

They also report derived alleles for pigmentation and lactose tolerance. All Mesolithic samples had dark eyes, hair and skin, except one blue-eyed Spaniard. In contrast the Copper Age Romanian had fair skin, blue eyes and possibly lighter brown hair than the others. None of them had derived MC1R mutations for red hair. None of them were able to digest lactose.

Of course the paper would have been much more interesting (for us) if it had been published before the Mathiesen et al. (2017) preprint two weeks ago, but at least this is the definitive paper.

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## Fire Haired14

I guess you were right about K1 in Mesolithic Europe. I always thought mHG K arose in WHG-like people because it belongs to mHG U but I'm surprised to see Mesolithic Serbia/Romania had so much K1(something like 20% of Iron Gate HGs). The K1*(maybe K1c) in Mesolithic Greece now makes sense considering a lot of the Iron Gate HGs had K1c. If Greece was similar to Romania then that could explain R1b V88 in Neolithic Spain. 

But there's no indication any form of H was popular anywhere in Mesolithic but who knows.

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## bicicleur

thanx for your summary, Maciamo

it looks like mesolithic Greece was populated by G2a/K1 HG, and this K1 appears once again in mesolithic Danube Gorge
no EHG in mesolithic Danube Gorge would mean no origin for neither Yamna nor CW, allthough at frist sight, I don't find a clear EHG component nowhere in the paper

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## Maciamo

> thanx for your summary, Maciamo
> 
> it looks like mesolithic Greece was populated by G2a/K1 HG, and this K1 appears once again in mesolithic Danube Gorge
> no EHG in mesolithic Danube Gorge would mean no origin for neither Yamna nor CW, allthough at frist sight, I don't find a clear EHG component nowhere in the paper


As I explained to FireHaired before, I believe that haplogroup K appeared north of the Black Sea as a descendant of Mesolithic European U8b. Some Mesolithic Eastern Europeans eventually made their way to Anatolia and the Caucasus, bringing Y-DNA R1b and I2c and mtDNA K (and probably also U5 and V) lineages with them. 

It has now been confirmed that both I2c and R1b were found among Epipaleolithic and Mesolithic Europeans. And yet I2c was also found among Early Anatolian farmers. Mesolithic Europeans all belonged to old subclades of R1b, such as L754, L388 or P297. All evidence points to R1b-M269 having emerged in eastern Anatolia, the southern Caucasus or western Iran (in other words around modern Kurdistan and Armenia) and having returned to the Pontic-Caspian Steppe as R1b-L23 by crossing the Caucasus. So it means that R1b-V88 and R1b-P297 people did migrated from Eastern Europe to the region of Kurdistan during the Mesolithic. This is where R1b hunters would have domesticated cows and later acquired metallurgy from their neighbours, but also how they returned to Europe with 30 to 50% of CHG admixture from millennia of intermingling with West Asians. My point is that if R1b-V88, R1b-P297 and I2c did get to the northern Middle East during the Epipaleolithic or Early Mesolithic, they could easily have brought mtDNA K1 to the region. If they domesticated cattle, their lineages would have quickly blended with other Middle Easterners, so that K1a eventually became a major Neolithic lineage. Let's not forget that the Neolithic lifestyle remained within the confines of the Fertile Crescent for 2500 years (9500 to 7000 BCE) before it started expanding to western Anatolia then Europe. It's more than enough time to spread K1a lineages around. 

Ever since I published my history of haplogroup K, I have maintained that K1c, K2b and perhaps also K2a were Mesolithic East European lineages that were spread by the Indo-Europeans (through their EHG ancestry). Then haplogroup K3 was found in Mesolithic Georgia, which also suggests a Pontic Steppe/North Caucasus origin for hg K. All this to say that K1c in Mesolithic Greece is Mesolithic EHG and not linked to G2a farmers.

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## kingjohn

But there's no indication any form of H was popular anywhere in Mesolithic but who knows.[/QUOTE]

but in the last big paper mtdna h13 was found in mesolithic individual from romania serbia border dated to 7000 bc 
so it was ther and dont forget mtdna pre-hv that was found in magdalenian cantabria iberia ......
so probably h was present in mesolitic southern europe....
i agree that u is older in most europe 
but h was there in mesolitic......
pre- hv is the ancestor of hv and h 
73A i also have this mutation as i am h descendents ... :)

Iron Gates
Romania
Ostrovul Corbului [I4081 / OSTCOR1a+1b / ROM47]
M
7580-7190 calBCE
311701
R1b1a
R1b1a: A702: 10038192G->A; R1b1a: CTS4244: 15510064T->G; R1b1a: CTS8436: 18026855G->A; R1b1a: FGC41: 7900883C->A; R1b1a: L1345: 21558298G->T; R1b1: CTS2229: 14226692T->A; R1: CTS2565: 14366723C->T; R: CTS8311: 17930099C->A; R: FGC1168: 15667208G->C;
H13

Mathieson 2017



Magdalenian
Spain
La Pasiega (Cantabria) [PS-1]





R0 or HV
rCRS in HVRI, G73A, reported as H
Hervella 2012

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## bicicleur

mtDNA K may well have come to an Anatolian LGM refuge along with I2
I would think I2 came to Anatolia after its formation, 27.5 ka, but before its TMRCA 21.9 ka
https://www.yfull.com/tree/I2/
No I2 has been observed in mesolithic Europe prior to the dispersal of the Villabruna clade, so I would suspect it stayed in Anatolia all that time.
Note that the Villabrunans arrived in Europe with microliths made with 'burin' technique and probably also with bow and arrow, something that appeared in the Kabaran in the Levant and in the Zarzian in the Zagros Mts during LGM.

mtDNA K3 was in Satsublia HG, but K1a was in Boncuklu PPN and Tepecik PN, K1c was in mesolithic Greece, while both K1a and K1b were in Barcin neolithic, all in association with G2a

the problem with mtDNA is that there is no reliable dating of the TMRCA's in the pedigree, we don't know when K or K1 originated or split

the recent paper on Indian DNA http://www.eupedia.com/forum/threads...ghlight=Indian claims reliable dating of the mtDNA pedigree based on a 2009 paper, but I have my doubts

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## Angela

> mtDNA K may well have come to an Anatolian LGM refuge along with I2
> I would think I2 came to Anatolia after its formation, 27.5 ka, but before its TMRCA 21.9 ka
> https://www.yfull.com/tree/I2/
> No I2 has been observed in mesolithic Europe prior to the dispersal of the Villabruna clade, so I would suspect it stayed in Anatolia all that time.
> Note that the Villabrunans arrived in Europe with microliths made with 'burin' technique and probably also with bow and arrow, something that appeared in the Kabaran in the Levant and in the Zarzian in the Zagros Mts during LGM.
> 
> mtDNA K3 was in Satsublia HG, but K1a was in Boncuklu PPN and Tepecik PN, K1c was in mesolithic Greece, while both K1a and K1b were in Barcin neolithic, all in association with G2a
> 
> the problem with mtDNA is that there is no reliable dating of the TMRCA's in the pedigree, we don't know when K or K1 originated or split
> ...


I think this is in line with what Reich and company have been hinting at in their recent papers.It's not going to sit very well with certain people.

@Maciamo,
I don't think there's any proof it was specifcally R1b people who domesticated cattle, given that cattle came to Europe with the earliest Neolithic, nor do I see how they could have brought metallurgy in the earliest periods given that the earliest samples on the steppe showing a change in ancestry are from a period when their only metallurgy was imported from the Balkans.

However, I think the handwriting is on the wall that all the genetics labs are looking at a movement from the south onto the steppe carrying a new genetic element. In the hobbyist world it's been vociferously argued that this all came via West Asian women. I've argued many times that there are a lot of problems with that theory, not least of which is the fact that there is a lot of mtDNA U5 and U4 on the steppe, and not enough south of the Caucuses mtDNA to account for the large percentage of CHG. Maybe they'll find a big group of J2b somewhere, but if not your hypothesis is a possibility. People forget how quickly the y can become decoupled from it's original autosomal signature.

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## bicicleur

> However, I think the handwriting is on the wall that all the genetics labs are looking at a movement from the south onto the steppe carrying a new
> genetic element. In the hobbyist world it's been vociferously argued that this all came via West Asian women. I've argued many times that there are a lot of problems with that theory, not least of which is the fact that there is a lot of mtDNA U5 and U4 on the steppe, and not enough south of the Caucuses mtDNA to account for the large percentage of CHG. Maybe they'll find a big group of J2b somewhere, but if not your hypothesis makes sense.People forget how quickly the y can become decoupled from it's original autosomal signature.


the strange thing however is that the autosomal got CHG admixed without the Y-DNA being changed
R1b-P297 was all over Eastern Europe before the admixture started and its subclade remained dominant in Yamna, as a matter of facts, no Y-DNA seems to have crossed the Caucasus, except maybe the Khvalynsk newcomer who was R1b but not P297
I hope upcoming papers will clarify this, as I can't come up with a satisfying narrative for this myself.

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## Angela

> the strange thing however is that the autosomal got CHG admixed without the Y-DNA being changed
> R1b-P297 was all over Eastern Europe before the admixture started and its subclade remained dominant in Yamna, as a matter of facts, no Y-DNA seems to have crossed the Caucasus, except maybe the Khvalynsk newcomer who was R1b but not P297
> I hope upcoming papers will clarify this, as I can't come up with a satisfying narrative for this myself.


 I hope I didn't imply that I have this all figured out, Bicycler, because I don't. All I know is that the anthrofora view of the steppe theory has a lot of problems. Like you I'm waiting for the next papers. Rumor has it that the ancient Greek DNA paper is about to come out. Hopefully it will mean more clarity, not more questions.

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## bicicleur

> I hope I didn't imply that I have this all figured out, Bicycler, because I don't. All I know is that the anthrofora view of the steppe theory has a lot of problems. Like you I'm waiting for the next papers. Rumor has it that the ancient Greek DNA paper is about to come out. Hopefully it will mean more clarity, not more questions.


I don't expect clean answers, I'm happy if they bring some hints which tease my imagination.

 :Wink:

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## MOESAN

Maciamo, I'm not sure all your 'go and return' concerning the Y-R1b's is safely established - but I admit I have no certitude, waiting for more ancient Y-DNA of differentes periods for the Anatolia to S-E Caspian regions ; we had so many surprises -

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## Olympus Mons

@Bicicleur,
No Narrative?
.. *iron gates* (R1b), were not much later or at the same time in *Fikirtepe (*R1b?*)* and by 6000bc in South caucasus as *Shulaveri-Shomu* where I M269 was born in the CHG land. by 4900 bc ( as Johannes krause also puts it) is the date I always said that Mentesh tepe fell and the Shulaveri, already with L23, in large numbers because they were many, started the big dispersal. Some run to the mountains, where one still finds lots of R1b in Bagvalins or South Ossetians, some run to the Eastern coast of black sea and moved up to the Kuban river (svodonoe and Mesokho) and later up samara to Become Yamnaya. Some moved back to eastern Anatolia to black sea shores again (Kum6). others moved south to the Aratashen land (armenia) and some even really south to become Merimde and El omari. 

My main point of contentious with people as always because I say merimde by 4000bc start moving via north africa to Portugal to become Bell beakers. We will see.

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## bicicleur

> @Bicicleur,
> No Narrative?
> .. *iron gates* (R1b), were not much later or at the same time in *Fikirtepe (*R1b?*)* and by 6000bc in South caucasus as *Shulaveri-Shomu* where I M269 was born in the CHG land. by 4900 bc ( as Johannes krause also puts it) is the date I always said that Mentesh tepe fell and the Shulaveri, already with L23, in large numbers because they were many, started the big dispersal. Some run to the mountains, where one still finds lots of R1b in Bagvalins or South Ossetians, some run to the Eastern coast of black sea and moved up to the Kuban river (svodonoe and Mesokho) and later up samara to Become Yamnaya. Some moved back to eastern Anatolia to black sea shores again (Kum6). others moved south to the Aratashen land (armenia) and some even really south to become Merimde and El omari. 
> 
> My main point of contentious with people as always because I say merimde by 4000bc start moving via north africa to Portugal to become Bell beakers. We will see.


do I miss something?
was R1b-M269 detected south of the Caucasus prior to Yamna?
or was his ancestor R1b-P297 detected south of the Caucasus prior to Yamna?

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## Olympus Mons

> do I miss something?
> was R1b-M269 detected south of the Caucasus prior to Yamna?


No. But it will! :)

We were talking about narratives. So follow settlement architecture....

Edited: To me is really important that those samples in Romania and Bulgaria are coming out as R1b. I defend the centrality of Shulaveri in the R1b history (probably the only one who does it) . But I also know that they came from the south shores of black sea, and clearly had something to do with HG from Thrace and eastern balkans. The forest set them apart from the middle and south Anatolian EEF farmers. It doesn't mean there was no interactions.... is just that they remained very pastoral ( as well in Eastern balkans) long after everyone was going full agriculture. The shulaveri arrived to south caucasus highly pastoral... but also evolved agriculturalists.

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## Promenade

> They also report derived alleles for pigmentation and lactose tolerance. All Mesolithic samples had dark eyes, hair and skin, except one blue-eyed Spaniard. In contrast the Copper Age Romanian had fair skin, blue eyes and possibly lighter brown hair than the others. None of them had derived MC1R mutations for red hair. None of them were able to digest lactose.


It seems Blue eyes were much less common in WHG from the Balkans compared to other areas

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## Angela

> It seems Blue eyes were much less common in WHG from the Balkans compared to other areas


You're right. However, even among the new Iberian samples, only one out of three had blue eyes, yes?

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## Promenade

> You're right. However, even among the new Iberian samples, only one out of three had blue eyes, yes?


Who is the third guy you're referring to? Out of the two Iberians here only one has them yes, but so far blue eyes were much more common in WHG elsewhere compared to those we've seen in the Balkans.

I'm wondering about the South Indian "Lilac" component in the older Iberian sample from Chan do Lindeiro now though. She also has a "Peach" component that the study doesnt discuss which she shares with La Brana, but not the girl from Los Canes. The EHG all carry the Lilac component and a few the peach component too. Any thoughts on what the Peach component is or how the South Indian ancestry got there?

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## Angela

> Who is the third guy you're referring to? Out of the two Iberians here only one has them yes, but so far blue eyes were much more common in WHG elsewhere compared to those we've seen in the Balkans.
> 
> I'm wondering about the South Indian "Lilac" component in the older Iberian sample from Chan do Lindeiro now though. She also has a "Peach" component that the study doesnt discuss which she shares with La Brana, but not the girl from Los Canes. The EHG all carry the Lilac component and a few the peach component too. Any thoughts on what the Peach component is or how the South Indian ancestry got there?


Yes, sorry, one out of two, and yes blue eyes are much more common in the WHG compared to the hg population in the Balkans. Also, while the SHG had blue eyes, it seems the EHG were more mixed and in fact leaned toward brown eyes, yes?

I haven't thought about it enough yet to give a complete answer, but off- hand I'd say that SA DNA has been flowing north for thousands upon thousands of years and so it doesn't surprise me some made it's way into the EHG. Nor, given the west/east cline of Hg populations, it doesn't surprise me there would be some flow westward. I have to take a closer look at it.

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## Promenade

> Yes, sorry, one out of two, and yes blue eyes are much more common in the WHG compared to the hg population in the Balkans. Also, while the SHG had blue eyes, it seems the EHG were more mixed and in fact leaned toward brown eyes, yes?
> 
> I haven't thought about it enough yet to give a complete answer, but off- hand I'd say that SA DNA has been flowing north for thousands upon thousands of years and so it doesn't surprise me some made it's way into the EHG. Nor, given the west/east cline of Hg populations, it doesn't surprise me there would be some flow westward. I have to take a closer look at it.


The WHG were all dark skinned but possessed light eyes while the EHG were more likely to have light skin but possessed dark eyes. The SHG were a mix of WHG and EHG and ended up with both light eyes and skin, there was a recent paper about ancient Scandinavia discussing this.

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## Fire Haired14

> Mesolithic Europeans all belonged to old subclades of R1b, such as L754, L388 or P297. All evidence points to R1b-M269 having emerged in eastern Anatolia, the southern Caucasus or western Iran (in other words around modern Kurdistan and Armenia) and having returned to the Pontic-Caspian Steppe as R1b-L23 by crossing the Caucasus.


I definitely think that possible but it still makes more sense to me R1b M269 emerged in Europe. Samara_HG had pre-R1b M73 and many of the Latvian HGs had pre-R1b M73. If a R1b M73 rich population existed in the Baltic then a R1b M269 population could have existed somewhere else in Eastern Europe. 

But a West Asian homeland for R1b M269 wouldn't surprise me since EHG-WHG and CHG had common ancestry. 




> My point is that if R1b-V88, R1b-P297 and I2c did get to the northern Middle East during the Epipaleolithic or Early Mesolithic, they could easily have brought mtDNA K1 to the region.


K1 looks pretty old in the Middle East. It was pretty popular in the Anatolia Neolithic(20%+?) and Levant Neolithic. It has a weak presence in India, Pakistan, and Afghanistan which could mean Iran Neolithic had some K1. 

The non-Basal stuff in ancient Middle Easterners could be native to the Middle East. Natufians had a bunch of WHG-type stuff but no confirmed K1 yet. The Mesolithic Balkans apparently also carried a large dose of K1. Yeah, it makes sense a K1-rich Mesolithic European group migrated to the Middle East to help form EEF but it also makes sense that K1's presence in both the Mesolithic Balkans and Middle East has more complex origin. Maybe a common ancestor between Balkan HGs and EEF not 




> Ever since I published my history of haplogroup K, I have maintained that K1c, K2b and perhaps also K2a were Mesolithic East European lineages that were spread by the Indo-Europeans (through their EHG ancestry). Then haplogroup K3 was found in Mesolithic Georgia, which also suggests a Pontic Steppe/North Caucasus origin for hg K. All this to say that K1c in Mesolithic Greece is Mesolithic EHG and not linked to G2a farmers.


Well that was a good call by you. But the old ways of using modern mtDNA/Y DNA is begging to look less and less able to deceiver the origin of lineages. The old assumption is that origin of a haplogroup can be determined by which region holds the most variety in the haplogroup is looking less and less able to determine the origin of a haplogroup.

Because modern day mtDNA/Y DNA diversity is not the result of gradual small scale migrations in which only some of the mtDNA/Y DNA diversity in one region goes to another. Instead it is the result of rapid mass migrations which brought most of the mtDNA/Y DNA diversity in one region to another region. 

The migration of Anatolian farmers to Europe is a grand example of this. Because of tit lots of Middle Eastern mtDNA haplogroups look like they originated in Europe. This is why old studies argued mtDNA H1, T2b, J1c, V, and other typical EEF lineages originated in Paleolithic Europe. 

I'm inclined to be skeptical of any argument that haplogroups other than U5/U4/U2 in Europe derive from European hunter gatherers. Because of the new K1 results from Mesolithic SouthEast Europe I'm more open to the idea but I think it's just as likely most K1c and K2a in modern Europe derive from Anatolian farmers. 

Unless Balkan hunter gatherers admixed a lot into Anatolian farmers who later moved into Northern and Western Europe why would K1 outside of the Balkans derive from Balkan hunter gatherers? Other European hunter gatherers could have had lots of K1 I guess. Also Anatolian farmers could have received their K1c from Balkan hunter gatherers.

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## Fire Haired14

> Yes, sorry, one out of two, and yes blue eyes are much more common in the WHG compared to the hg population in the Balkans. Also, while the SHG had blue eyes, it seems the EHG were more mixed and in fact leaned toward brown eyes, yes?


It's interesting that Mesolithic Europeans had variable pigmentation which didn't always correlate with ancestry. There was also some variation in Neolithic farmers. Ones in Hungary and Germany were more fair than ones in Iberia. It could help understand the difference in pigmentation between Andronovo and Yamnaya and variation in modern Europeans.

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## Maciamo

> I definitely think that possible but it still makes more sense to me R1b M269 emerged in Europe. Samara_HG had pre-R1b M73 and many of the Latvian HGs had pre-R1b M73. If a R1b M73 rich population existed in the Baltic then a R1b M269 population could have existed somewhere else in Eastern Europe.


What's pre-M73? Do you mean L388, which is ancestral to both M73 and P297? No M73 has been found in Mesolithic Europe yet. It looks increasingly like old R1b clades like L754 and L388 could have been present from the Balkans and East Baltic all the way to Central Asia, including all the Pontic-Caspian Steppe and North Caucasus region. In that case M73 could represent a Central Asian branch of hunter-gatherers.



P297 tribes would have been found from the North Caucasus to the Baltic. Some would have crossed the Caucasus during the Late Glacial or immediate postglacial period, before agriculture emerged in the Near East. It would have been just one of numerous HG haplogroups found in the region, although possibly the only tribe with an ancestral experience of hunting mammoths and other big game in Siberia and Eastern Europe. That is what I believe gave them an advantage in taming aurochs (who would have looked pretty inoffensive compared to mammoths, but still daunting for local HG used to hunt gazelles or wild goats). 

R1b-P297 or M269 wouldn't turn up in Early farmers from the Levant or Anatolia since it was confined to Kurdistan/Armenia/Azerbaijan/NW Iran and lived in patriarchal pastoral societies where only women intermarried with neighbouring tribes. That's how their original female lineages (such as K1 and K2, but also obviously U5a and U5b1, and probably also U2e, U4, U8, V and some H subclades) found their way in the Early Neolithic gene pool in the northern Middle East.




> K1 looks pretty old in the Middle East. It was pretty popular in the Anatolia Neolithic(20%+?) and Levant Neolithic. It has a weak presence in India, Pakistan, and Afghanistan which could mean Iran Neolithic had some K1.


Pretty old? According to Soares et al. (2009), haplogroup K itself is no older than 12,000 years old. It might be a bit older than that as the K3 from Satsurblia in Georgia is about 13,000 years old. The oldest known K1 sample is a 10,500 year old K1a from Boncuklu in central Anatolia, which is actually pretty near from the place where cattle were domesticated in SE Anatolia/N. Syria. That was probably one of the very first K1a. If K1 was found in the Mesolithic Balkans, it could easily have spread to Anatolia. There were no borders in those days and HG were very mobile. If even Neolithic farmers could move into Europe and live side by side from the local HG for centuries, what is to prevent European HG to do the same in Anatolia? That being said, I have no idea if K1 came to Anatolia via the Balkans or via the Caucasus, but I think that European HG moved to the northern Middle East from both sides, each bringing different clades, or each bringing K1* that developed into different clades in the Middle East. 




> The non-Basal stuff in ancient Middle Easterners could be native to the Middle East. Natufians had a bunch of WHG-type stuff but no confirmed K1 yet. The Mesolithic Balkans apparently also carried a large dose of K1. Yeah, it makes sense a K1-rich Mesolithic European group migrated to the Middle East to help form EEF but it also makes sense that K1's presence in both the Mesolithic Balkans and Middle East has more complex origin. Maybe a common ancestor between Balkan HGs and EEF not


Not possible of K is only 12,000 years old. That's the same age of the beginning of the Mesolithic and early agriculture in the Fertile Crescent.




> Well that was a good call by you. But the old ways of using modern mtDNA/Y DNA is begging to look less and less able to deceiver the origin of lineages. The old assumption is that origin of a haplogroup can be determined by which region holds the most variety in the haplogroup is looking less and less able to determine the origin of a haplogroup.
> 
> Because modern day mtDNA/Y DNA diversity is not the result of gradual small scale migrations in which only some of the mtDNA/Y DNA diversity in one region goes to another. Instead it is the result of rapid mass migrations which brought most of the mtDNA/Y DNA diversity in one region to another region. 
> 
> The migration of Anatolian farmers to Europe is a grand example of this. Because of tit lots of Middle Eastern mtDNA haplogroups look like they originated in Europe. This is why old studies argued mtDNA H1, T2b, J1c, V, and other typical EEF lineages originated in Paleolithic Europe.


Actually my explanation for J1c and several T2 clades is that they were present since the Late Glacial period in the Balkans *and* the northern Middle East. It was almost certainly the case of Y-haplogroups C1a2 and I2c as well. I originally thought J2b, but I have to revise that to C1a2 and I2c instead.

As for H1 it has been found among Mesolithic Sweden, but I don't remember if that individual had any EEF admixture or not.




> I'm inclined to be skeptical of any argument that haplogroups other than U5/U4/U2 in Europe derive from European hunter gatherers. Because of the new K1 results from Mesolithic SouthEast Europe I'm more open to the idea but I think it's just as likely most K1c and K2a in modern Europe derive from Anatolian farmers. 
> 
> Unless Balkan hunter gatherers admixed a lot into Anatolian farmers who later moved into Northern and Western Europe why would K1 outside of the Balkans derive from Balkan hunter gatherers? Other European hunter gatherers could have had lots of K1 I guess. Also Anatolian farmers could have received their K1c from Balkan hunter gatherers.


The K1 HG from the Balkans had no Near Eastern admixture whatsoever.

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## Maciamo

> the strange thing however is that the autosomal got CHG admixed without the Y-DNA being changed
> R1b-P297 was all over Eastern Europe before the admixture started and its subclade remained dominant in Yamna, as a matter of facts, no Y-DNA seems to have crossed the Caucasus, except maybe the Khvalynsk newcomer who was R1b but not P297
> I hope upcoming papers will clarify this, as I can't come up with a satisfying narrative for this myself.


It isn't strange that the Y-DNA should remain stable if they belonged to a patriarchal society, like all pastoralists. 

If you look at the phylogeny, M269 has no less than 105 defining mutations, which took about 7000 years to accumulate, from the time the first one arose (13,300 years ago according to Yfull) until the TMRCA of 6,300 years. This means that M269 lived in isolation during all that time, from the Late Glacial period until the Chalcolithic. Since M269 has never been found in Mesolithic or Neolithic Europe, and it only shows up as its subclade L23 in Yamna with CHG admixture, the evidence is indeed very strong for R1b-M269 having lived as a small tribe of pastoralists somewhere south of the Caucasus range (probably moving around quite a bit over the millennia, between eastern Anatolia and western Iran).

But that is not all. Where do we find M269* people today? In northwestern Iran and Armenia. Grugni et al. (2012) found R1b-M269* among Teheran Zoroastrians, in Mazandaran Province and among Iranian Azeri. They are all located on the south coast of the Caspian Sea! However M269* only makes up 1.4% of the population in that region (except Teheran Zoroastrians who have an amazing 15%, but that's a small sample size). However there is really a lot of R1b-L23* in Iran, found at high frequencies (23% to 55%) among the Assyrians, Armenians and in Lorestan (in the Zagros), and at reasonable frequencies (11-19%) among the Azeri, and in the western provinces of Gilan and Fars. But there is no R1b in Balochestan or most of the eastern provinces, except Palaeolithic subclades like M343.

Where do we find the PF7562 subclade of M269, the only other known brother clade of L23? In Trabzon, NE Turkey.

This strongly suggests that M269 and L23 evolved somewhere between NW Iran and NE Turkey, in the southern Caucasus perhaps around or on either sides of Armenia and Kurdistan. I did not come up with that theory based on a whim. I analysed all the data available and envisaged all the possible scenarios. I still cannot find a better explanation.

You can see in my migration maps that already many years ago I hypothesised R1b (M269 or L23) migrated from the South Caucasus along the Caspian Sea to reach the Pontic-Caspian Steppe some time between 6000 and 5000 BCE. That may have been a bit too early, unless they did found Khvalynsk around 5000 BCE, which seems entirely possible.

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## Maciamo

> @Maciamo,
> I don't think there's any proof it was specifcally R1b people who domesticated cattle, given that cattle came to Europe with the earliest Neolithic, nor do I see how they could have brought metallurgy in the earliest periods given that the earliest samples on the steppe showing a change in ancestry are from a period when their only metallurgy was imported from the Balkans.


The strongest evidence that R1b people domesticated cattle is that R1b-L23 Yamna people were cattle pastoralists AND R1b-V88 tribes in Africa are also cattle pastoralists to this day. Obviously domesticated animals and crops were readily exchanged between tribes in the two and a half millennia (9500-7000 BCE) preceding the Neolithic expansion beyond the Fertile Crescent. Farmers couldn't have come to Europe with cereals, legumes, goats, sheep, pigs, cattle and also pottery if each of them were domesticated (or invented for pottery) by separate tribes at various locations in the Fertile Crescent. And we know that there were many independent tribes (E1b1b, G2a, H2, J1, J2a, J2b, L1a, R1b, R2) in the Middle East at the time, and that cereal cultivation arose in the southern Levant, while cattle were domesticated in northern Syria and goats in the Zagros (western Iran), and the first pottery came from northern Iraq. There must have been a lot of exchanges between tribes. Well, a lot is relative as it took them 2500 years to get the full Neolithic package relatively evenly distributed over that region. It doesn't matter how quickly it happened. My point is that whoever domesticated cattle in northern Syria, those cattle could easily have ended up in all Neolithic tribes before the Neolithic expansion. What is more surprising is that R1b tribes decided to remain primarily cattle pastoralists wherever they went (Europe, Africa, Central Asia) and did not really care for land cultivation, nor other animals (they did keep some sheep in Yamna, but mostly for the wool, as they still do in Scotland and Ireland). 

As for metallurgy, I don't understand why you are telling me that. I have never said that R1b people invented metallurgy. Only *bronze* metallurgy, and that was probably in Maykop, where R1b-L23 would have started associating with G2a-U1. So it could have been the G2a-U1 who were the metal workers who originally invented bronze too. If you check this thread from early 2010, I already proposed that G2a (what was then G2a3b1 but is now known as L140) were the metallurgist accompanying the R1b stockbreeders in the PIE migrations. It's a well established fact that copper metallurgy arose in the Balkans and central Anatolia, two regions where G2a was dominant at the time. 




> However, I think the handwriting is on the wall that all the genetics labs are looking at a movement from the south onto the steppe carrying a new genetic element. In the hobbyist world it's been vociferously argued that this all came via West Asian women. I've argued many times that there are a lot of problems with that theory, not least of which is the fact that there is a lot of mtDNA U5 and U4 on the steppe, and not enough south of the Caucuses mtDNA to account for the large percentage of CHG. Maybe they'll find a big group of J2b somewhere, but if not your hypothesis is a possibility. People forget how quickly the y can become decoupled from it's original autosomal signature.


I don't understand what U4 and U5 have to do with CHG admixture. IMO, CHG admixture came in two waves to the Steppe. The first was an early Neolithic migration of J2b from NW Iran. The second was a middlle Neolithic migration of R1b-L23 from Armenia/NW Iran/Azerbaijan to the Steppe (see above post). The first wave would have brought lineages like N1a (or I), and perhaps also T1a, T2 and W. The second would have brought J1b1a, H2a1, H4, H8c, H15, T1a, T2c, as well as I and W.

I don't support the theory that R1b men from the Steppe crossed the Caucasus to steal West Asian women. That is ridiculous. If R1b-M269 evolved for 7000 years around the South Caucasus (see above post), there is no way they wouldn't have intermarried with West Asian women.

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## Olympus Mons

@Maciamo.
Speak clearly. That second migration into steppe was the shulaveri shomu with r1b L23 . right?

Second...
How long have you been stating that? Show me your earlieast (2015/2016) comments and posts saying that.

I predicted long ago that once it starts to be obvious the role of my shulaveri in the r1b history....many would start to post as if they knew all along. Are you one of those?

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## Maciamo

> @Maciamo.
> Speak clearly. That second migration into steppe was the shulaveri shomu with r1b L23 . right?


I don't know yet, but I'd say it was just before Shulaveri Shomu. However that doesn't preclude that L23 could also have been present in Shulaveri Shomu, as it has been present in the region until this day.




> Second...
> How long have you been stating that? Show me your earlieast (2015/2016) comments and posts saying that.
> 
> I predicted long ago that once it starts to be obvious the role of my shulaveri in the r1b history....many would start to post as if they knew all along. Are you one of those?


I predicted in 2009 the migration of R1b from the South Caucasus circa 6000-5000 BCE. Use the search engine if you want to find those posts.

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## LeBrok

I'm fascinated by this GB1 individual. It is Late Neolithic in Balkans, almost Bronze Age, and she has 3/4 of genome WHG! Remaining quarter is EEF. From Bronze Age Hungarian genome we know that in Bronze Age there was explosion of WHG admixture in this area. A little bit of Steppe too, but mostly WHG, up to 50% of total genome. 
Where is the source of WHG at this time?
My guess for quite some time is West Yamnaya, but they might have been coming from farther North like Belarus, and we know WHG lasted in Latvia till BA.
Looks more and more like the EHG/Steppe by Bronze Age pushed remaining WHGs into Balkans.

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## Fire Haired14

The Bronze age genomes from Hungary which carry a lot of WHG share the most IBD with Poles and other Slavs. Maybe WHG-rich populations from SE Europe closely related to those Bronze age Hungarians are important to Slavic genesis or at least Eastern European genesis.

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## Angela

When I suggested, right after Haak et al came out, I think, that there was perhaps a reservoir of WHG somewhere in eastern or northeastern Europe which might have gone into the genesis of these LNBA populations, and that perhaps even SHG might be involved, didn't you, following Eurogenes, say that was impossible? I'm pretty sure I have that right.

It's always good to look at different points of view and to be open minded.

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## LeBrok

> The Bronze age genomes from Hungary which carry a lot of WHG share the most IBD with Poles and other Slavs. Maybe WHG-rich populations from SE Europe closely related to those Bronze age Hungarians are important to Slavic genesis or at least Eastern European genesis.


Absolutely they are. Modern Poles are more related and mostly descendents from Hungarian BA than from CW and Unetice.

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## Maciamo

Just an addendum to post #23 above, Nasidze 2009 found 48% of R1b among Azeri Talysh, 14% among Iranian Talysh, 22% among the Gilaks and 14% among the Mazandarans. Based on Grugni 2012, this is all R1b-L23 (xP312, U106).

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## MOESAN

It's to be believed that a pop in N-Carpathians took advantage from close contacts with Late CTC (Tripolye)... and took part in the earlier Slavs genesis (among them a lot of Y-I2a1); already it seems that Y-I2a2 (males!) took the strong side upon Neolithic Y-G2a males or exchanged females (neolithic mt-DNA) with them, what could be seen in ALP culture (Hungary); surely they were not the same I2a lineages but it shows that exchanges had already taken place with ex-Mesolithic HG's playing a big role at the mergins of the LBK world, after their apparent "dispariton" in Early Neolithic - some metric anthropology extracts seems confirming the mixture in some place of Hungary even before BA or Pit Grave introgressions -

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