Human Paternal Lineages, Languages and Envrionment in the Caucasus

This is the link to the paper:
http://digitalcommons.wayne.edu/cgi/viewcontent.cgi?article=1053&context=humbiol_preprints

It's open access...

Here is the abstract:
"Publications that describe the composition of the human Y-DNA haplogroup in diffferent ethnic or linguistic groups and geographic regions provide no explicit explanation of the distribution of human paternal lineages in relation to specific ecological conditions. Our research attempts to address this topic for the Caucasus, a geographic region that encompasses a relatively small area but harbors high linguistic, ethnic, and Y-DNA haplogroup diversity. We genotyped 224 men that identified themselves as ethnic Georgian for 23 Y-chromosome short tandem-repeat markers and assigned them to their geographic places of origin. The genotyped data were supplemented with published data on haplogroup composition and location of other ethnic groups of the Caucasus. We used multivariate statistical methods to see if linguistics, climate, and landscape accounted for geographical diffferences in frequencies of the Y-DNA haplogroups G2, R1a, R1b, J1, and J2. The analysis showed significant associations of (1) G2 with wellforested mountains, (2) J2 with warm areas or porrly forested mountaina, and (3) J1 with poorly forested mountains. Rib showed no association with environment. Haplogroups J1 and R1a were significantly associated with Daghestanian and Kipchak speakers, respectively, but the other haplogroups showed no such simple associations with languages. Climate and landscape in the context of competition over productive areas among diffferent paternal lineages, arriving in the Caucasus in diffferent times, have played an important role in shaping the present-day spatial distribution of patrilineages in the Caucasus. This spatial pattern had formed before linguistic subdivisions were finally shaped, probably in the Neolithic to Bronze Age. Later historical turmoil had little influence on the patrilineage composition and spatial distribution. Based on our results, the scenario of postglacial expansions of humans and their languages to the Caucasus from the Middle East, western Eurasia, and the East European Plain is plausible. "


I've only quickly skimmed it, but I'm not sure if I buy all of their conclusions. If I understand them correctly, while they acknowledge that the "G" y Dna lineages might have been pushed to forested mountain refugia, they also believe that their presence in these mountains can be explained by the fact that yDna "G" originated in the Caucasus during the LGM and deliberately chose and kept the forested mountains because as a specific type of hunter gatherer they were accustomed to that terrain. I'm not sure I buy that. What then about all the yDna "G" men who became farmers and brought farming into Europe?

They also seem to give a nod to what I think might be the out of the Armenian highlands theory of the Indo-European languages, but I couldn't find either of the papers they cited.
"Archaeologists suggest that the first Indo-Europeans expanded to the Southern Caucasus from the southwest in the 3rd millennium BC (Melikishvili, 1959; Melaart, 1970). Currently, relatively high proportion of the haplogroup R1b is found in Indo-European speaking Armenians, but also in Georgians from the areas south of the Lesser Caucasus (this study) and in some Daghestanian ethnic groups (Yunusbayev et al. 2012).


I did find this interesting in light of some of the recent discussions on this Board:

"Expandingtribes (most likely dominated by haplogroups J1 and J2) that had survived the Ice Age south of the Caucasus probably started settling in the Caucasus both before emerging early agricultural settlements in theFertile Crescent about 9.5 KY ago (Allaby et al. 2008) and after that."

It also provides a nice summary of the connection, or disconnection, between y lineages, language and religion in the region. I think it might serve as a cautionary tale for those who tend to see Indo-European and certain y lineages as indissolubly linked.

Okay, if we try to find a trend by looking at this paper, the other one you referenced recently and what's been leaked about the upcoming Lazardis paper concerning a "Karelian" and "Armenian" mixture in the IE homeland, I don't know what conclusions one might reach, other than it's looking more and more as if R1b was one of the Y haplotypes in the IE homeland, which is not what I was expecting. And I guess one might conclude that the "Karelian" and "Armenian" components adopted a common language and culture without a lot of genetic mixing, given the lack of R1b in the Baltic (which was the main reason I was skeptical about R1b in the IE homeland).

Aberdeen said:

Okay, if we try to find a trend by looking at this paper, the other one you referenced recently and what's been leaked about the upcoming Lazardis paper concerning a "Karelian" and "Armenian" mixture in the IE homeland, I don't know what conclusions one might reach, other than it's looking more and more as if R1b was one of the Y haplotypes in the IE homeland, which is not what I was expecting. And I guess one might conclude that the "Karelian" and "Armenian" components adopted a common language and culture without a lot of genetic mixing, given the lack of R1b in the Baltic (which was the main reason I was skeptical about R1b in the IE homeland).

Click to expand...

Well, I think the phylogeny indicates that R1b expanded in an east to west direction. The question is when and with whom. I have a problem with the whole R1b out of Iberia thing because the most frequent R1b clade in Iberia is very "young". Of course, if there was a migration into Central Europe by an upstream clade and then a back migration by a "younger" downstream clade that might explain it. Still, the question would remain as to how and when and with whom they got to Iberia in the first place, given the upstream clades in West Asia, the Caucasus etc., as well as Central Asia.

One well known theory has the Italic-Celtic forms of R1b moving like this:
http://s1000.photobucket.com/user/txumi_ledauno/media/StelaePeople_zps24790e03.jpg.html

It may turn out to be correct, but there's no archaeological evidence for it except for a few stelae, which could just as easily have spread by coast hugging seafarers from the eastern Mediterranean, and after all, absent ancient Dna, that's supposed to be the underpinning for theories, that and linguistic analysis, which doesn't particularly favor this model over any other east to west migration route.

As I've said before, the only thing that gives me pause is that Wolfgang Haak, who worked with Guido Brandt on the recap of ancient Dna paper which posited an R1b out of Iberia movement is also a co-author of the upcoming Corded Ware paper with Lazaridis, so you would think he would be privy to the yDna results the Reich lab has for Yamnaya, Samara people.
http://www.eupedia.com/forum/threads/30646-Human-Paleogenetics-of-Europe-Brandt-and-Haak

What they should do is see if there are useable human bones from the intrusive Beaker finds in Iberia which we've discussed before, or any early Beaker remains in Iberia for that matter and test them. That's really the only way we'll know.

As for their comments about the "Indo-European" languages, as Mallory pointed out I think it all depends upon the "Anatolian" question, as in the "Anatolian" branch of Indo-European. Regardless of whether or not people from Anatolia crossed into the Steppes from the south Caucasus, the question is whether "Anatolian" was the first language to break off from PIE speakers on the Steppes, and made a move through the Balkans into Anatolia (a scenario for which David Anthony, at least, provides no archaeological evidence), or whether perhaps it was the first PIE language and stayed put when all the others moved to the Steppes (Grigoriev provides a great deal of archaeological proof for the Gramkelidze Ivanov out of the Near East model, as Mallory points out).
http://www.jolr.ru/files/(112)jlr2013-9(145-154).pdf

In terms of the whole Karelian/Armenian thing, I don't think we know enough to speculate, but speculating is fun, so... while some people might say the fact that Yamnaya can be modeled as 50% ancient Karelian and 50% Armenian doesn't mean there was an actual admixture, that's splitting rather too many hairs for me. Some sort of admixture between pretty widely divergent groups does seem to have taken place in Yamnaya, unless when the actual paper comes out they disavow these comments. (We have seen this sort of reversal rather recently.) At the same time, I don't know that it says much about the "ancient Karelian like" peoples that might have remained further north and influenced more northern European peoples.

There's also the comment about Corded Ware people being 75% Yamnaya, or at least that this is the model which is the "best fit" for the Corded Ware people. If Yamnaya was 50% ancient Karelian, then Corded Ware would be 35% ancient Karelian just from their Yamnaya input. It remains to be seen if the prior people were unremarkable late Neolithic/Copper Age rather EEF people, or whether they had absorbed some additional H/G of an eastern variety before the arrival of the Yamnaya people. If the former is the case, it's still unclear how the people of that area today got their additional WHG.

One thing I can guarantee, people are already probably busy spinning like tops in order to square their prior theories with this new data, when in fact it contradicts them.

Angela said:

I've only quickly skimmed it, but I'm not sure if I buy all of their conclusions. If I understand them correctly, while they acknowledge that the "G" y Dna lineages might have been pushed to forested mountain refugia, they also believe that their presence in these mountains can be explained by the fact that yDna "G" originated in the Caucasus during the LGM and deliberately chose and kept the forested mountains because as a specific type of hunter gatherer they were accustomed to that terrain. I'm not sure I buy that. What then about all the yDna "G" men who became farmers and brought farming into Europe?

They also seem to give a nod to what I think might be the out of the Armenian highlands theory of the Indo-European languages, but I couldn't find either of the papers they cited.
"Archaeologists suggest that the first Indo-Europeans expanded to the Southern Caucasus from the southwest in the 3rd millennium BC (Melikishvili, 1959; Melaart, 1970). Currently, relatively high proportion of the haplogroup R1b is found in Indo-European speaking Armenians, but also in Georgians from the areas south of the Lesser Caucasus (this study) and in some Daghestanian ethnic groups (Yunusbayev et al. 2012).


I did find this interesting in light of some of the recent discussions on this Board:

"Expandingtribes (most likely dominated by haplogroups J1 and J2) that had survived the Ice Age south of the Caucasus probably started settling in the Caucasus both before emerging early agricultural settlements in theFertile Crescent about 9.5 KY ago (Allaby et al. 2008) and after that."

It also provides a nice summary of the connection, or disconnection, between y lineages, language and religion in the region. I think it might serve as a cautionary tale for those who tend to see Indo-European and certain y lineages as indissolubly linked.

Click to expand...

The G-P15 among Caucasians may not have spread farming to Europe. These could be very old isolated branches. I don't believe the G-P15 among Ossetians have the same downstream mutations as those found in Europe, or at least there is a separation point. It is likely that the major expansions of G-P15 were spread with farmers, but independently to different areas. For instance some branches pushed into the Caucasus mountains, where as other branches pushed into Europe. I find a linear situation of Middle-East -> Caucasus -> Europe as highly unlikely.

It's possible later farmers like J2 or J1 pushed the earlier G-P15 farmers into the mountains. Perhaps a group like I2c is the oldest in the Caucasus.

Angela said:

Well, I think the phylogeny indicates that R1b expanded in an east to west direction. The question is when and with whom. I have a problem with the whole R1b out of Iberia thing because the most frequent R1b clade in Iberia is very "young". Of course, if there was a migration into Central Europe by an upstream clade and then a back migration by a "younger" downstream clade that might explain it. Still, the question would remain as to how and when and with whom they got to Iberia in the first place, given the upstream clades in West Asia, the Caucasus etc., as well as Central Asia.

One well known theory has the Italic-Celtic forms of R1b moving like this:
http://s1000.photobucket.com/user/txumi_ledauno/media/StelaePeople_zps24790e03.jpg.html

It may turn out to be correct, but there's no archaeological evidence for it except for a few stelae, which could just as easily have spread by coast hugging seafarers from the eastern Mediterranean, and after all, absent ancient Dna, that's supposed to be the underpinning for theories, that and linguistic analysis, which doesn't particularly favor this model over any other east to west migration route.

As I've said before, the only thing that gives me pause is that Wolfgang Haak, who worked with Guido Brandt on the recap of ancient Dna paper which posited an R1b out of Iberia movement is also a co-author of the upcoming Corded Ware paper with Lazaridis, so you would think he would be privy to the yDna results the Reich lab has for Yamnaya, Samara people.
http://www.eupedia.com/forum/threads/30646-Human-Paleogenetics-of-Europe-Brandt-and-Haak

What they should do is see if there are useable human bones from the intrusive Beaker finds in Iberia which we've discussed before, or any early Beaker remains in Iberia for that matter and test them. That's really the only way we'll know.

As for their comments about the "Indo-European" languages, as Mallory pointed out I think it all depends upon the "Anatolian" question, as in the "Anatolian" branch of Indo-European. Regardless of whether or not people from Anatolia crossed into the Steppes from the south Caucasus, the question is whether "Anatolian" was the first language to break off from PIE speakers on the Steppes, and made a move through the Balkans into Anatolia (a scenario for which David Anthony, at least, provides no archaeological evidence), or whether perhaps it was the first PIE language and stayed put when all the others moved to the Steppes (Grigoriev provides a great deal of archaeological proof for the Gramkelidze Ivanov out of the Near East model, as Mallory points out).
http://www.jolr.ru/files/(112)jlr2013-9(145-154).pdf

In terms of the whole Karelian/Armenian thing, I don't think we know enough to speculate, but speculating is fun, so... while some people might say the fact that Yamnaya can be modeled as 50% ancient Karelian and 50% Armenian doesn't mean there was an actual admixture, that's splitting rather too many hairs for me. Some sort of admixture between pretty widely divergent groups does seem to have taken place in Yamnaya, unless when the actual paper comes out they disavow these comments. (We have seen this sort of reversal rather recently.) At the same time, I don't know that it says much about the "ancient Karelian like" peoples that might have remained further north and influenced more northern European peoples.

There's also the comment about Corded Ware people being 75% Yamnaya, or at least that this is the model which is the "best fit" for the Corded Ware people. If Yamnaya was 50% ancient Karelian, then Corded Ware would be 35% ancient Karelian just from their Yamnaya input. It remains to be seen if the prior people were unremarkable late Neolithic/Copper Age rather EEF people, or whether they had absorbed some additional H/G of an eastern variety before the arrival of the Yamnaya people. If the former is the case, it's still unclear how the people of that area today got their additional WHG.

One thing I can guarantee, people are already probably busy spinning like tops in order to square their prior theories with this new data, when in fact it contradicts them.

Click to expand...

I still believe that different subclades of R1b arrived in western Europe by different routes and I think we should consider the possibility that L21 and DF27 were in distributed along the Mediterranean and Atlantic prior to the Bronze Age, with only U152 being part of the "Armenian" portion of the IE population and U106 entering Europe via the Balkans. I'm not concerned about the close connection between L21, D27 and U152 if we're thinking in terms of a dispersal from Anatolia. And I would interpret a comment about Corded Ware being 75% IE as meaning more "Karelian" and less "Armenian". But I'll have to think more about Mallory's paper.

Edit: or the 75% IE refers to Corded Ware having absorbed 25% Neolithic European after arriving in Neolithic Europe - that seems more likely, now that I think of it.

Aaron1981: It's possible later farmers like J2 or J1 pushed the earlier G-P15 farmers into the mountains.

Click to expand...

That makes sense to me too.

As to the "G" lineages, there just hasn't been enough work done on them, in my opinion. In the Boattini et al paper for Italy they found five separate clusters of "G" in Italy, all with different dates. Any vast generalizations about "G" in general are going to be off, in my opinion.

See: Sarno, Boattini et al:
http://www.plosone.org/article/info:doi/10.1371/journal.pone.0065441

Figure 2 shows the five different "G" clusters and their geographic distribution. The STR's are provided in the supplement.

There's also a discussion here about their second updated paper. Their conclusions as to the date of arrival of the different y lineages seems rather prescient given recent ancient dna results.
http://www.eupedia.com/forum/threads/29842-southern-Italian-paper-2014?highlight=U152+Italy

Aberdeen said:

I still believe that different subclades of R1b arrived in western Europe by different routes and I think we should consider the possibility that L21 and DF27 were in distributed along the Mediterranean and Atlantic prior to the Bronze Age, with only U152 being part of the "Armenian" portion of the IE population and U106 entering Europe via the Balkans. I'm not concerned about the close connection between L21, D27 and U152 if we're thinking in terms of a dispersal from Anatolia. And I would interpret a comment about Corded Ware being 75% IE as meaning more "Karelian" and less "Armenian". But I'll have to think more about Mallory's paper.

Click to expand...

Well, you would think that if Corded Ware was 75% ancient Karelian they would have said so, but until we get the paper, I'm leaving all the options open. We have to also consider the fact that Haak and Brandt said that there was increasing absorption of hunter gatherer women in the more northern regions less hospitable to farming, and then a movement of these people back into central Europe. Perhaps this is the source of the additional WHG. That would, of course, have nothing to do with the Indo-Europeans.

If people keep changing the definition of Indo-European to fit whatever group of people they think are their ancestors, the term will wind up meaning nothing in particular. If the tweets from the conference are correct, and substantiated, it seems that these researchers have concluded that Indo-Europeans=Yamnaya (which isn't to be wondered at given that David Anthony is a consulting author, no matter what the reality might be), and if David Anthony is correct that means the people on the Pontic Caspian steppe from 4,000-3000 BC. (which he specifically stated), and the "ethnic mix" there was half Near Eastern like and half ancient Karelian like hunter/gatherer. I mean, that area hasn't changed all that much, if you think about it. Aren't southern Russians/Ukrainians about 47% EEF? Plus, that's after centuries of Russia planting colonies of more northern Slavs in the region. The "ethnic mix" of the Corded Ware people and the Central European Beakers in the period we're discussing is another matter. They were the result of further admixtures, and then you have further changes after that which produces the modern clusters. I think there's a tendency to back project modern people far into the past. Modern populations "jelled" pretty recently, in my opinion, especially in northern and central Europe.

As for how the various "young" clades of R1b got to their current locations in Europe, I'm not sure, other than that the older clades had to be in close proximity to each other at one point, and they had to come from the east originally. As to U-152, I don't know why U-152 would be any more likely to be from the "Armenian part" of the steppes (I doubt it was divided up that way. I think they were talking about the Yamnaya as a whole, although we'll soon see), unless you're going by the physical anthropology?

If we go by physical anthropology, the "Iberian beakers" were apparently not "Beaker" like as in like the "Central European Beakers", who indeed have "Brachy" skulls and a flat occiput apparently. However, the "Iberian Beakers" were also no different than their EEF farmer predecessors, which doesn't look very intrusive. Still, the only way to know is by testing the ancient Dna. So far we have "G", E-V13 and "I2a" from the coastal Mediterranean samples, but the number of total samples is still small, so who knows.

Perhaps R1b was in the southern part of the Pontic-Caspian steppe, or perhaps it straddled the Caucasus, or perhaps it moved in a slightly earlier period into the Balkans and then fled the collapse there. However it actually happened, I think it's likely to be involved with the development of the Indo-European "culture", which really, so far as I can tell, was a combination of a lot of different cultural strands coming from different directions. After all, we have a J2a who was part of the Iron Age migrations from the steppe, and I bet a certain clade of G2a will show up as well, so why not R1b?

Angela said:

Well, you would think that if Corded Ware was 75% ancient Karelian they would have said so, but until we get the paper, I'm leaving all the options open. We have to also consider the fact that Haak and Brandt said that there was increasing absorption of hunter gatherer women in the more northern regions less hospitable to farming, and then a movement of these people back into central Europe. Perhaps this is the source of the additional WHG. That would, of course, have nothing to do with the Indo-Europeans.

If people keep changing the definition of Indo-European to fit whatever group of people they think are their ancestors, the term will wind up meaning nothing in particular. If the tweets from the conference are correct, and substantiated, it seems that these researchers have concluded that Indo-Europeans=Yamnaya (which isn't to be wondered at given that David Anthony is a consulting author, no matter what the reality might be), and if David Anthony is correct that means the people on the Pontic Caspian steppe from 4,000-3000 BC. (which he specifically stated), and the "ethnic mix" there was half Near Eastern like and half ancient Karelian like hunter/gatherer. I mean, that area hasn't changed all that much, if you think about it. Aren't southern Russians/Ukrainians about 47% EEF? Plus, that's after centuries of Russia planting colonies of more northern Slavs in the region. The "ethnic mix" of the Corded Ware people and the Central European Beakers in the period we're discussing is another matter. They were the result of further admixtures, and then you have further changes after that which produces the modern clusters. I think there's a tendency to back project modern people far into the past. Modern populations "jelled" pretty recently, in my opinion, especially in northern and central Europe.

As for how the various "young" clades of R1b got to their current locations in Europe, I'm not sure, other than that the older clades had to be in close proximity to each other at one point, and they had to come from the east originally. As to U-152, I don't know why U-152 would be any more likely to be from the "Armenian part" of the steppes (I doubt it was divided up that way. I think they were talking about the Yamnaya as a whole, although we'll soon see), unless you're going by the physical anthropology?

If we go by physical anthropology, the "Iberian beakers" were apparently not "Beaker" like as in like the "Central European Beakers", who indeed have "Brachy" skulls and a flat occiput apparently. However, the "Iberian Beakers" were also no different than their EEF farmer predecessors, which doesn't look very intrusive. Still, the only way to know is by testing the ancient Dna. So far we have "G", E-V13 and "I2a" from the coastal Mediterranean samples, but the number of total samples is still small, so who knows.

Perhaps R1b was in the southern part of the Pontic-Caspian steppe, or perhaps it straddled the Caucasus, or perhaps it moved in a slightly earlier period into the Balkans and then fled the collapse there. However it actually happened, I think it's likely to be involved with the development of the Indo-European "culture", which really, so far as I can tell, was a combination of a lot of different cultural strands coming from different directions. After all, we have a J2a who was part of the Iron Age migrations from the steppe, and I bet a certain clade of G2a will show up as well, so why not R1b?

Click to expand...

G2a ?
which G2a3a or G2a3b?

Angela said:

Well, you would think that if Corded Ware was 75% ancient Karelian they would have said so, but until we get the paper, I'm leaving all the options open. We have to also consider the fact that Haak and Brandt said that there was increasing absorption of hunter gatherer women in the more northern regions less hospitable to farming, and then a movement of these people back into central Europe. Perhaps this is the source of the additional WHG. That would, of course, have nothing to do with the Indo-Europeans.

If people keep changing the definition of Indo-European to fit whatever group of people they think are their ancestors, the term will wind up meaning nothing in particular. If the tweets from the conference are correct, and substantiated, it seems that these researchers have concluded that Indo-Europeans=Yamnaya (which isn't to be wondered at given that David Anthony is a consulting author, no matter what the reality might be), and if David Anthony is correct that means the people on the Pontic Caspian steppe from 4,000-3000 BC. (which he specifically stated), and the "ethnic mix" there was half Near Eastern like and half ancient Karelian like hunter/gatherer. I mean, that area hasn't changed all that much, if you think about it. Aren't southern Russians/Ukrainians about 47% EEF? Plus, that's after centuries of Russia planting colonies of more northern Slavs in the region. The "ethnic mix" of the Corded Ware people and the Central European Beakers in the period we're discussing is another matter. They were the result of further admixtures, and then you have further changes after that which produces the modern clusters. I think there's a tendency to back project modern people far into the past. Modern populations "jelled" pretty recently, in my opinion, especially in northern and central Europe.

As for how the various "young" clades of R1b got to their current locations in Europe, I'm not sure, other than that the older clades had to be in close proximity to each other at one point, and they had to come from the east originally. As to U-152, I don't know why U-152 would be any more likely to be from the "Armenian part" of the steppes (I doubt it was divided up that way. I think they were talking about the Yamnaya as a whole, although we'll soon see), unless you're going by the physical anthropology?

If we go by physical anthropology, the "Iberian beakers" were apparently not "Beaker" like as in like the "Central European Beakers", who indeed have "Brachy" skulls and a flat occiput apparently. However, the "Iberian Beakers" were also no different than their EEF farmer predecessors, which doesn't look very intrusive. Still, the only way to know is by testing the ancient Dna. So far we have "G", E-V13 and "I2a" from the coastal Mediterranean samples, but the number of total samples is still small, so who knows.

Perhaps R1b was in the southern part of the Pontic-Caspian steppe, or perhaps it straddled the Caucasus, or perhaps it moved in a slightly earlier period into the Balkans and then fled the collapse there. However it actually happened, I think it's likely to be involved with the development of the Indo-European "culture", which really, so far as I can tell, was a combination of a lot of different cultural strands coming from different directions. After all, we have a J2a who was part of the Iron Age migrations from the steppe, and I bet a certain clade of G2a will show up as well, so why not R1b?

Click to expand...

I've certainly never claimed that my ancestors were IE in the genetic sense, and I suspect that the genetic impact is quite minimal in some parts of Europe compared to the linguist and cultural impact although some parts of Europe appear to perhaps experienced a significant genetic impact from IE. The reason I suspect that U152 was either part of IE or quickly absorbed by it is, as I've said before, because of the archeology associated with Hallstatt (and La Tene). And no, we don't have G, E-V13 and I2a from coastal Mediterranean Bell Beaker samples - we don't have any Y DNA from Beaker sites at all, except for the two German R1b samples.

As I've said before, I think physical anthropology can be misleading, since things such as skull shape can apparently be affected by climate and diet. But the archeology and mtDNA evidence suggests that Bell Beaker folk were intrusive.

Unfortunately this article has many shortcomings, most importantly direct linking of language families and meta-haplogroups (e.g. coevolution of Kartvelian language family and G2a, which is absurd), and messed up STR's in support data.

I have had a personal discussion with the authors but they seem to be deaf to the obvious.