Neolithic patrilineal signals, repopulation, and the Armenian Plateau

[A. Tamar Chabadi]

Neolithic patrilineal signals indicate that the Armenian plateau was repopulated by agriculturalists

Kristian J Herrera, Robert K Lowery, Laura Hadden, Silvia Calderon, Carolina Chiou, Levon Yepiskoposyan, Maria Regueiro, Peter A Underhill and Rene J Herrera

Abstract
Armenia, situated between the Black and Caspian Seas, lies at the junction of Turkey, Iran, Georgia, Azerbaijan and former Mesopotamia. This geographic position made it a potential contact zone between Eastern and Western civilizations. In this investigation, we assess Y-chromosomal diversity in four geographically distinct populations that represent the extent of historical Armenia. We find a striking prominence of haplogroups previously implicated with the Agricultural Revolution in the Near East, including the J2a-M410-, R1b1b1*-L23-, G2a-P15- and J1-M267-derived lineages. Given that the Last Glacial Maximum event in the Armenian plateau occured a few millennia before the Neolithic era, we envision a scenario in which its repopulation was achieved mainly by the arrival of farmers from the Fertile Crescent temporally coincident with the initial inception of farming in Greece. However, we detect very restricted genetic affinities with Europe that suggest any later cultural diffusions from Armenia to Europe were not associated with substantial amounts of paternal gene flow, despite the presence of closely related Indo-European languages in both Armenia and Southeast Europe.

European Journal of Human Genetics , (16 November 2011) | doi:10.1038/ejhg.2011.192

 

[A. Tamar Chabadi]

Armenian Y-chromosomes revisited (Herrera et al. 2011)

 

[sparkey]

The case of haplogroup J is particularly problematic. If we had to guess, by looking at present-day distribution, which lineage tracks population movements from the Near East to Europe, there is simply no better candidate: every map of this haplogroup, and especially of its J2a sublineage shows an unambiguous pattern of radiation, with a core area consisting of Southern Italy, Greece, Anatolia, West Asia, Mesopotamia and the northern parts of the Levant. All these regions are crucial to the story of the Neolithic, so the absence of J in Neolithic Europe is perplexing.

I don't find the issue to be that we can't come up with an explanation... the issue is more that there are many possible explanations, and we don't know which one, or which combination, is correct. It's probably a combination of: J has expanded over time at the expense of G in the Near East; sample bias in Neolithic European samples (very Western so far); founder effects and small populations magnifying G early and J later; etc...

Finally, the occurrence of some I2, E-V13, and, perhaps, J2b in Armenia may point to Balkan contacts. But, when did these contacts occur? Are they traceable to the migration of Phrygians to Anatolia, according to the Herodotean account of Armenian origins, or can they be attributed to later contacts with Greeks or other Europeans?

Why the assumption that E-V13 and J2b come from the Balkans? And Armenian I2 isn't I2a1-Din but I2a2-Cont3 and I2c-B (probably from different sources... I2a2-Cont3 probably spread there earlier given its age).

 

[A. Tamar Chabadi]

I just lost everything I typed...wow...this timed log out thing is annoying.

I will come back and type it later.

 

[sparkey]

I just lost everything I typed...wow...this timed log out thing is annoying.

I will come back and type it later.

If you log back in on the login screen that comes up after you are timed out, it's supposed to restore what you've written. Contact Maciamo or write in the site feedback forum if that isn't the case for you.

 

[A. Tamar Chabadi]

First of all, I would like to say I am sorry for complaining, I won't anymore...just have to get accustomed to the non-Facebook rules...hahahaha...I am accustomed to posting links and attachments at will. I apologize to you, Maciamo, and the many other participants in the Eupedia Forum.

Also, I would like to thank Sparkey for shortening my post down to a link...greatly appreciate it.

Now, for what I said earlier...I agree with you, Sparkey, although, I do not think Haplogroup G was ever very widespread at high frequency...I think it was only prevalent in certain places, most likely it was heaviest in the place where it is currently found in high concentration. Perhaps the Y-Hap G's migrated out to the west first, followed by the Y-Hap J2's...perhaps the J2's stayed close to the coasts which would explain the non-existence, so far, of J2's found in neolithic samples further inland. The G's appear to have traveled further inland. The distribution, to the west, of the Cardial Culture seems to match the pattern of J2's across the Med. Like you said, Sparkey, there are many explanations and not just one thing that explains everything...sometimes, things just aren't cookie-cutter simple. This appears to be a complex migration pattern...more of a diffusive “oozing forth” (think...lava flow) from the Near East rather than surges of any kind...a slow and steady progression or “spreading out/ diffusion” that moved across Neolithic-era Europe. We can’t think that the genes spread from the Near East, spread from there individually...they spread together. I think we tend to think they spread spread individually...like the G’s went first, then the J’s, then the R1b’s, etc, etc...that is not how it worked.

As for Y-Haps E-V13 and J2b...I think they are older in the Balkans...perhaps some E-V13 spread through the Levantine corridor...but there is scanty evidence of this left genetically. There are also competing ideas about the spread of E-V13...Cruciani et al. 2004, Battaglia et al. 2007, and Bird et al. 2007...Bird et al. 2007 is not necessarily in disagreement with either Cruciani or Battaglia. Bird’s conclusion can be included as secondary diffusion of E-V13 out of the Balkans during the Roman Period....I think anyway. J2b is often associated with E-V13 as a Neolithic signature...it appears to be associated with Neolithic Greece also. There appears to be no consensus on where it may have come from. It’s association with Neolithic Greece appears to be why it is considered from the Balkans.


I think we agree about Y-Hap I in this region...

 

[sparkey]

Very good post, I agree almost entirely but would like to explore some nuances...

We can’t think that the genes spread from the Near East, spread from there individually...they spread together. I think we tend to think they spread spread individually...like the G’s went first, then the J’s, then the R1b’s, etc, etc...that is not how it worked.

Yes, but certain populations would have been dominated by certain haplogroups, and some that are now common in Europe are obviously later arrivals than others. I think, for example, that we can nearly rule out significant R1b L11+ as a Neolithic arrival to Europe. However, not even the R1b L11+ wave into Western Europe would have been 100% R1b L11+... in fact, that population probably included J2 and G2, meaning that we can't even call all J2 and G2 in Europe "Neolithic." It's an interesting complication that warrants drilling down to more refined subclades.

As for Y-Haps E-V13 and J2b...I think they are older in the Balkans...perhaps some E-V13 spread through the Levantine corridor...but there is not scanty evidence of this left genetically. There are also competing ideas about the spread of E-V13...Cruciani et al. 2004, Battaglia et al. 2007, and Bird et al. 2007...Bird et al. 2007 is not necessarily in disagreement with either Cruciani or Battaglia. Bird’s conclusion can be included as secondary diffusion of E-V13 out of the Balkans during the Roman Period....I think anyway. J2b is often associated with E-V13 as a Neolithic signature...it appears to be associated with Neolithic Greece also. There appears to be no consensus on where it may have come from. It’s association with Neolithic Greece appears to be why it is considered from the Balkans.

E1b-V13 is still a bit of a mystery, although finding it (finally) in a Neolithic sample cleared some mist about it. Is it true that it's older in the Balkans than in the Levant? I know that the Levant has E1b M78+ significantly but I haven't looked closely at E1b-V13 diversity analyses. I have generally assumed that it originated in the Near East and expanded on Europe, but with some doubt.

I really doubt that J2b is older in the Balkans, though... isn't it quite diverse in the Caucasus? And it stretches to India, indicating an affinity to R1a.

 

[A. Tamar Chabadi]

Well, as for R1b-L11+, I think many subclades of R1b underwent rapid expansion...I don't know if R1b is entirely neolithic in Europe...maybe some subclades are and maybe some are older. I am a strong believer in the Kurgan theory or some version of it. I am not entirely sure that Y-Hap R1a is the only signature of the the spread of the Indo-European languages. I mean, again, it is much more complicated than that...R1a existed before what we now call PIE (Proto-Indo-European). I believe, at least, some clades of R1b are also involved. But maybe the majority of the older R1b's may have spoken something akin to Basque. I have also read the paper by Busby et al. 2010 "Increased Resolution Within Y-Chromosome Haplogroup R1b M269 Sheds Light On The Neolithic Transition In Europe"...the paper states, "Rather than originating with the farmers from the East, we suggest that the sub-structure of R1b-M269 visible in Europe today, and thus the great majority of European paternal ancestry, is the result of the interaction between the Neolithic wave of expansion and populations of early Europeans already present in the path of the wave."

I agree with you about there not being 100% saturation in the migratory populations. There had to be a mix...exactly how much of each Y-Hap may always a mystery.

I think that most Y-Hap J2 carriers were non-Indo-European...many, if not all, of the languages where J2 is found in high frequency are or were, at one point, non-Indo-European. I think the same for Y-Hap E-V13.

I have read about Y-STR ages for Y-Hap E-V13...they say that this group arose in the Near East, but they must have been almost entirely bred of of the population or most migrated to the Balkans. Much of what is in the Near East, that is E-V13, can be traced to the Hellenistic era, Roman Era, Byzantine Era, and to a time, before the Hellenistic Era, when there was trade between the Aegean ("Minoan", Mycenaean, and Philistine) and the Near East by boat. The percentages of E-V13 in Armenia are small so it must not have been a large number of people. I think if E-V13 were older in Armenia, I think there would be a higher percentage than exists there right now. The frequency of the few lineages of E1b1b1a-V68 that are there are very low (3.9%). The percentage of E-V13 in Armenian plateau populations is .5% (.004842619%) in this paper.

The Cypriots have greater than 10% Y-Hap E-V13...this can be explained by very long time Aegean contact. Even the "Minoans" had a colony on Cyprus...Amathus. This is where the "EteoCypriot" (better to call it Amathusian) dialect of "Minoan" existed...

There is also Y-Hap E-V13 among the Druze...it was originally thought to be of the "Delta" cluster based on Y-STRs. The Druze are native to where they are...the genes they carry are very similar to those around them (Levantine). The >10% of E-V13 in their population MAY BE of the same origin as the rest of the E-V13 in the Near East (mostly the Balkans).

As far as I know, E-V13 is less diverse in Anatolia and the rest of the Near East than the Balkans. I think the dispersal of E-V13 is more complex than just one catch-all explanation as we said before, but studies point to the Balkans as the source of it because of the much higher diversity there.

Outside of J2b-M12, there is another Y-Hap J2 lineage that is associated with Chalcolithic/ Bronze Age Greece...it is J2a4h1a1c-M318, which is also found in the Armenian Plateau at 4.9%.

As for J2b being more diverse in the Caucasus...in Balanovsky et al. 2011 "Parallel Evolution of Genes and Languages in the Caucasus Region"...J2b-M12 appears in 1 person out of 142 Circassians in the 14 Caucasian populations sampled (the overall sample = 1525)

In Yunusbayev et al. 2011 "[SIZE=-1][SIZE=-1]The Caucasus as an Asymmetric Semipermeable Barrier to Ancient Human Migrations"[/SIZE][/SIZE]...their samples of 24 populations across the Caucasus (overall sample size =1952), J2b appears in 2 Avars (n=42), 2 Kumyks (n=73), 1 Nogay (n=163), 3 Adyghe (n=154), 1 Karbadin (n=140), 1 Abkhazian (n=120), and 3 Armenians (n=57) for a total of 13/ 1952 or about .7% overall (.006659836%).

According to the Family Tree DNA M102+ Project...J2b has a frequency peak in the Balkans and two variance peaks, one in the Balkans and one in Central Italy. Their data is based on Pericic et al. 2005 "High-Resolution Phylogenetic Analysis of Southeastern Europe Traces Major Episodes of Paternal Gene Flow Among Slavic Populations." Also according to them..."The absence of J2b-M12 in regions T1 and T8 (Cinnioglu et al., 2004), i.e. those next to the land bridge from Anatolia to Greece, suggests that the first farmers of Greece and the Balkans are less likely to have come overland."

The problem with the India thing is that R1a in India is very old, perhaps Paleolithic and Mesolithic...J2b is Neolithic at the oldest. Some of the deeper (older) Y-Hap clades are found in India, para-groups C*, F*, K*, P*, R*, R1*, and R1a*.

 

[sparkey]

Well, as for R1b-L11+, I think many subclades of R1b underwent rapid expansion...I don't know if R1b is entirely neolithic in Europe...maybe some subclades are and maybe some are older. I am a strong believer in the Kurgan theory or some version of it. I am not entirely sure that Y-Hap R1a is the only signature of the the spread of the Indo-European languages. I mean, again, it is much more complicated than that...R1a existed before what we now call PIE (Proto-Indo-European). I believe, at least, some clades of R1b are also involved.

Right, I'm arguing that only R1b L11- is likely to have been part of the Neolithic haplogroup array, and almost certainly as a minority. I find it to be a mystery when R1b L11+ dominant populations starting speaking Indo-European, or if that's even looking at the question right (suppose, for example, that the "original IE people" had R1b as a minority clade, and a population bottleneck caused it to become dominant among a certain population expanding on Europe). We do get some hints at an upper bound by observing that Assyrians (non-IE population obviously) have a lot of L23+ L11-.

But maybe the majority of the older R1b's may have spoken something akin to Basque.

Do you place the origin of the Basque language with R1b peoples? I'm not so sure. I consider it a possibility but I find its origin with an I2a1a-dominant population that eventually had much of its Y-lines replaced with R1b a slightly more viable theory. If my guess is right, then "older R1b's" are unlikely to have spoken anything like Basque.

I have also read the paper by Busby et al. 2010 "Increased Resolution Within Y-Chromosome Haplogroup R1b M269 Sheds Light On The Neolithic Transition In Europe"...the paper states, "Rather than originating with the farmers from the East, we suggest that the sub-structure of R1b-M269 visible in Europe today, and thus the great majority of European paternal ancestry, is the result of the interaction between the Neolithic wave of expansion and populations of early Europeans already present in the path of the wave."

Busby has had poor speculation based on the evidence IMHO, with the Busby 2011 paper then claiming that R1b STR analyses are meaningless and almost implying that we should assume Paleolithic origin of it (that's certainly how the media took that paper). If anything, Busby's work as a whole has been pretty good in showing that R1b fits the European Neolithic picture poorly.

I think that most Y-Hap J2 carriers were non-Indo-European...many, if not all, of the languages where J2 is found in high frequency are or were, at one point, non-Indo-European. I think the same for Y-Hap E-V13.

I agree.

According to the Family Tree DNA M102+ Project...J2b has a frequency peak in the Balkans and two variance peaks, one in the Balkans and one in Central Italy. Their data is based on Pericic et al. 2005 "High-Resolution Phylogenetic Analysis of Southeastern Europe Traces Major Episodes of Paternal Gene Flow Among Slavic Populations." Also according to them..."The absence of J2b-M12 in regions T1 and T8 (Cinnioglu et al., 2004), i.e. those next to the land bridge from Anatolia to Greece, suggests that the first farmers of Greece and the Balkans are less likely to have come overland."

Pericic et al isn't very useful here if they didn't extend their tests deeper in Asia than Anatolia. We can also see that they get an incorrect variance analysis of I2a-Din based on sampling issues, so I'm not going to read much into their conclusions about J2b either. Is there a better analysis that includes the rest of the Middle East and India?

 

[A. Tamar Chabadi]

Well, Y-Hap R1b(xL11/P310)...is Anatolian by most considerations...the parent group to those in Ht15 by the old, and now defunct, RFLP method.


I agree that it must have assuredly belonged to the Neolithic array...perhaps much more than you think...ht35 is the dominant R1b group in the Balkans according to most studies. There is ht15, but at much smaller frequencies than ht35. R1b-L11+ is thought to have originated in one of several places Central Europe, the Lower Danube basin, the Middle Rhine, or Paris basin.

According to the "R1b1a2 (P312- U106-) DNA Project (aka ht35 Project)"...R1b-L11+ (U106-, P312-) is still haplotype 35...the Armenians are rife with R1b-L23+.

From the paper originally cited in this post, “Shortly after the arrival of early farmers in Armenia and Anatolia (8 kya), agriculture spread to Greece and the Balkans, before rapidly expanding across Europe. Furthermore, the classification of Armenian as an old Indo-European language with similarities to the ancestral Proto-Indo-European languages has led to the supposition that agriculturalists migrating from Armenia into Europe were responsible for the establishment of Indo-European languages in the continent. However, despite the close linguistic relationship between Armenians and the Indo-European speaking populations of Europe, we see little genetic support for this claim. The derived M412 allele, which is found in nearly all haplogroup R1b1b1*-L23 chromosomes in Europe, is absent in the sampled Armenians, which also exhibit a scarcity of haplotype sharing with Europeans, suggesting a limited role for Armenians in the introduction of R1b into Europe.

Several authors have proposed that the Indo-European language presently spoken by Armenians arose during the Bronze Age, when Indo-European speaking tribes from the Balkans and Greece invaded Anatolia and Transcaucasia, leading to the subsequent spread of their culture and language. In this study, we have detected a number of lineages that are prominent in the Balkans (I2*, I2b*, J2b1 and J2b2) at low levels throughout Ararat Valley, Gardman and Lake Van, the latter of which also contains haplogroups commonly associated with Bronze Age Greece (ie, J2a8-M319 (4.9%), and E1b1b1-M78 and its sublineages (3.9%)). While this may suggest genetic input from early Greek or Phrygian tribes, it is also possible that these low levels of Balkan lineages arrived in Armenia at a later time, such as during one of the many incursions into the area during the reign of the Macedonian, Roman, and Byzantine empires.”

As for Basque being spoken by Y-Hap I2a1a-M26 speakers...I will partially agree especially since there is evidence of an Iberian or Basque substrate in the Sardinian Romance language...Paleo-Sardo is likely to have been more than one language though. Also, the R1b presence is immense in the Basque populations, but is rather low in haplotypic diversity. The variance peaks are in Anatolia and the Caucasus. Which could give some credence to the Vasco-Caucasian theory if Basque was spoken by the at least some groups of the R1b peoples.

As for Busby, again, his paper even made Dienekes give up on Y-STRs...so something in it must have been rather convincing. I found it very convincing. Again, we must not consider all R1b to be Neolithic. Some branches are and others are not...some are older and some even younger than the Neolithic.

As for Pericic et al...the data for Y-Hap J2b is still most useful...no one that I could find had any objections to the Y-Hap J2b observations of the paper. Most other papers focus on J2b1 in India and places where it traveled in Southeast Asia and it’s travel with the Gypsies into Europe. As far as I can find...all J2b in India is J2b1.

Here is a map:



Here is another:

 

[sparkey]

I agree that it must have assuredly belonged to the Neolithic array...perhaps much more than you think...ht35 is the dominant R1b group in the Balkans according to most studies. There is ht15, but at much smaller frequencies than ht35.

ht35 is very interesting, but I haven't really studied it closely. I know it's more distant from R1b-L11+ than even the Assyrian modal haplotype, which shares L150 with L11+. Without additional study on my part, I think that makes it a pretty good candidate for Neolithic entry. But L150+ looks like it originated in Asia and came later.

It's fairly small in comparison to other suspected Neolithic entries in a lot of Europe, though. Perhaps that's genetic drift (in favor of G2a mostly, probably) within the Westward Neolithic migrants. Or perhaps it's just a later entry, or was more dominant in a later wave, or the modern frequencies only really reflect later expansions and contractions... I'm not sure.

As for Basque being spoken by Y-Hap I2a1a-M26 speakers...I will partially agree especially since there is evidence of an Iberian or Basque substrate in the Sardinian Romance language...Paleo-Sardo is likely to have been more than one language though. Also, the R1b presence is immense in the Basque populations, but is rather low in haplotypic diversity. The variance peaks are in Anatolia and the Caucasus. Which could give some credence to the Vasco-Caucasian theory if Basque was spoken by the at least some groups of the R1b peoples.

This is a good summary of the evidence and possibilities for the origin of Basque.

As for Busby, again, his paper even made Dienekes give up on Y-STRs...so something in it must have been rather convincing. I found it very convincing.

Personally, I like Dienekes as an excellent aggregator of information, but I don't always trust his conclusions. Have you read and understood what Mikewww and Ken Nordtvedt (as quoted by me) have to say about Busby on the R1b TMRCA thread? I find them much more convincing than Dienekes here, and I'm still not really sure why you're not with me here. (By the way, let's redirect discussion on TMRCA calculation to that thread so we're not duplicating a discussion between two threads.)