Two observations on the ancestry of Armenians

I was thinking a bit on how to interpret the findings of the new Haber et al. preprint, and especially the idea that "29% of the Armenian ancestry may originate from an ancestral population best represented by Neolithic Europeans." I looked at the globe13 proportions, and strangely enough, I had estimated that the three Armenian samples (Armenian_D, Armenians, and Armenians_15_Y) have 28-29% of the Mediterranean component that is modal in Sardinians. This seems like a curious coincidence which has raised my confidence that Haber et al. is picking something real.

Looking back at my inferences of Armenian ancestry, it seems (according to globe13) to come completely from West_Asian, Mediterranean, and Southwest_Asian. The Mediterranean component seems real enough as it seems to match Sardinians/early European farmers well. I am not so sure about the Southwest Asian component which is modal in Yemen Jews and may represent population-specific drift in relatively recent Arabians. The West_Asian component is bimodal in Caucasus and Gedrosia, so it can't be the result of a very drifted population in either region (unless there is spooky action at a distance). 

Another curious finding is the lack of North_European in a latitudinal "column" of populations from the Yemen, through the Levant to the South Caucasus (Georgians and Armenians). It seems that North_European is the only one of the four major Caucasoid components that Armenians lack to any important degree. There is a rather abrupt change between the South Caucasus (~1%) and the North Caucasus (15-20%). It seems that the Greater Caucasus did act like a barrier to gene flow. The K=4 analysis of the same dataset that produced K=13 (globe13) also shows the same barrier: all three Armenian samples and Georgians have ~0% of "Amerindian" (which is surely correlated to "Ancient North Eurasian" ancestry and via it with North_European), but North Caucasians and Europeans have 4-10%.  It's clear that this influence did not cross the Greater Caucasus, as Armenians and Georgians lack it. 

Bronze Age mixing of multiple populations => Armenians (?)

As far as I can tell, the hypothesis of "several mixtures" comes from looking at many pairs of populations and seeing that different types of pairs seem like they mixed to make Armenians. Possibility (1) is that Armenians have multiple mixtures, and possibility (2) is that none of the sources work very well.

Hellenthal et al. did not find mixture in Armenians, but they worked with a different methodology and smaller sample size. Either, the N=173 sample size enabled detection of this admixture, or differences in methodology account for differences in conclusions. If true, the admixture dates in this paper would be some of the earliest discovered by looking at modern populations (without the help of ancient DNA).

The TreeMix analysis (Figure 4) is inconclusive about admixture from a population best represented by Neolithic Europeans. There is no plot of residuals in this figure, so this model with one migration event may not be adequate. Prior knowledge suggests that it isn't, as Pakistani and European populations have no admixture in Figure 4.

It's great that the authors will share their data!
ftp://ngs.sanger.ac.uk/scratch/project/team19/Armenian
As of this writing, the data is not "live"; it might appear when the paper is published.

bioRxiv doi: http://dx.doi.org/10.1101/015396

Genetic evidence for an origin of the Armenians from Bronze Age mixing of multiple populations

Marc Haber et al.

The Armenians are a culturally isolated population who historically inhabited a region in the Near East bounded by the Mediterranean and Black seas and the Caucasus, but remain underrepresented in genetic studies and have a complex history including a major geographic displacement during World War One. Here, we analyse genome-wide variation in 173 Armenians and compare them to 78 other worldwide populations. We find that Armenians form a distinctive cluster linking the Near East, Europe, and the Caucasus. We show that Armenian diversity can be explained by several mixtures of Eurasian populations that occurred between ~3,000 and ~2,000 BCE, a period characterized by major population migrations after the domestication of the horse, appearance of chariots, and the rise of advanced civilizations in the Near East. However, genetic signals of population mixture cease after ~1,200 BCE when Bronze Age civilizations in the Eastern Mediterranean world suddenly and violently collapsed. Armenians have since remained isolated and genetic structure within the population developed ~500 years ago when Armenia was divided between the Ottomans and the Safavid Empire in Iran. Finally, we show that Armenians have higher genetic affinity to Neolithic Europeans than other present-day Near Easterners, and that 29% of the Armenian ancestry may originate from an ancestral population best represented by Neolithic Europeans.

Link

DNA & the Origins of Peoples: The Armenians

An excellent lecture from last summer:

Hovann Simonian and Peter Hrechdakian delivered the 18th Annual Vardanants Day Armenian lecture on their work using social media to expand the nonprofit Armenian DNA Project, a Facebook group of more than 1,000 members around the world involved in researching Armenian family history through genetic testing.

The place of the Armenian language in the Indo-European family

A very interesting talk at the Library of Congress making a good case for a Greek-Phrygian-Armenian clade within the Indo-European family.
 


Of course the testimony of Herodotus is -at least for me- the most convincing argument for an ultimately Balkan origin of the Phrygo-Armenians.

Modern Armenians are quite distinct from modern populations of the Balkans but who knows how both they and the populations of the Balkans have changed since the beginning of the Iron Age when the Phrygians established themselves in Asia Minor? The recent study by Hellenthal et al. did not find any good evidence for recent admixture in Armenians but this might be due to (i) Armenians being unmixed descendants of Proto-Armenians, (ii) Armenians being near-unmixed descendants of pre-Armenians, or (iii) the method not having enough power to detect admixture.

My guess is that the relative remoteness of the Armenian highlands coupled with the heterodox position of the Armenian church hindered substantial gene flow into the Armenian population over at least the last 1,500 years.

The place of Armenian in the Indo-European language family

Journal of Language Relationship • Вопросы языкового родства • 10 (2013) • Pp. 85—137

The place of Armenian in the Indo-European language family: the relationship with Greek and Indo-Iranian*

Hrach Martirosyan

The main purpose of this paper is to present lexical correspondences that unite Armenian with Greek and/or Indo-Iranian. They include shared innovations on the one hand, and isolated lexemes on the other. These two lexical corpora — lexical innovations on an inherited basis and isolated words — can be placed within the same temporal and spatial framework. After the Indo-European dispersal Proto-Armenian would have continued to come into contact with genetically related Indo-European dialects. Simultaneously, it would certainly also have been in contact with neighbouring non-Indo-European languages. A word can be of a substrate origin if it is characterized by: (1) limited geographical distribution; (2) unusual phonology and word formation; (3) characteristic semantics. The material presented here, albeit not exhaustive, allows to preliminarily conclude that Armenian, Greek, (Phrygian) and Indo-Iranian were dialectally close to each other. Within this hypothetical dialect group, Proto-Armenian was situated between Proto-Greek (to the west) and Proto-Indo-Iranian (to the east). The Indo-Iranians then moved eastwards, while the Proto-Armenians and Proto-Greeks remained in a common geographical region for a long period and developed numerous shared innovations. At a later stage, together or independently, they borrowed a large number of words from the Mediterranean / Pontic substrate language(s), mostly cultural and agricultural words, as well as animal and plant designations. On the other hand, Armenian shows a considerable number of lexical correspondences with European branches of the Indo-European language family, a large portion of which too should be explained in terms of substrate rather than Indo-European heritage.

Link (pdf)

Prince William's Indian matrilineage?

William has a hint of Indian in his DNA, find British researchers

Researchers have sourced William’s Indian ancestry to Eliza Kewark, his great-great-great-great-great grandmother, who was assumed to be Armenian, but now has been revealed as an Indian by genetic research.

...

“Through genealogy we traced two living direct descendants of Eliza and by reading the sequence of their mtDNA, we showed not only that they matched, but also that it belongs to a haplogroup called R30b, thus determining Eliza Kewark’s haplogroup,” the research team revealed.

The haplogroup, which is a group of related ancestral lineages, in this case was revealed to be rare and found only in South Asia. Other related branches of R30a and R30* are also entirely South Asian.

“This confirms therefore that the mtDNA of Eliza Kewark of Surat was of Indian heritage. R30b is rare even in India, where roughly 0.3 per cent of people carry this lineage,” the researchers revealed.

Chaubey et al. (2008) is an article that touches upon the subject of R30b.

Personally, I wouldn't be so quick in discounting the traditional genealogical story. A lineage that occurs at a frequency of 0.3% will almost certainly be missed in any small sample if it occurs at similar trace frequencies in other populations.

ESHG 2013 abstracts

A couple of weeks before the ESHG 2013 conference, here are a few abstracts that caught my eye. Feel free to point to more interesting presentations in the comments section.

• J16.05 - Checking the hypothesis of a Balkan origin of the Armenians
• P17.5 - Y chromosome haplogroup analysis to estimate genetic origin of Balts
• J16.27 - Armenian Highland as a transition corridor for the spread of Neolithic agriculturists
• J16.14 - In Search of the Origin of Haplogroup J1-P58
• J16.03 - Y-chromosome haplogroup analysis in the Besermyan ethnic group
• P17.4 - Mitochondrial DNA Analysis of the Southeast European Genetic Variation Reveals a New, Local Subbranching in Hg X2
• P16.085 - Population diversity and history of the Indian subcontinent: Uncovering the deeper mosaic of sub-structuring and the intricate network of dispersals
• J16.43 - Ancient mtDNA diversity in Bulgaria
• P16.072 - Mitochondrial DNA diversity in medieval and modern Romanian population
• P16.012 - Frequency analysis of the CCR5-Delta32 allele in medieval and modern Romanian population
• J16.78 - The gene pool of Argyn in the context of generic structure of Kazakhs according to data on SNP-Y-Chromosome markers.

Armenian origin of Hamshenis

Hum Biol. 2012 Aug;84(4):405-22

Paternal lineage analysis supports an armenian rather than a central asian genetic origin of the hamshenis.

Authors: Margaryan A, Harutyunyan A, Khachatryan Z, Khudoyan A, Yepiskoposyan L

Abstract

The Hamshenis are an isolated geographic group of Armenians with a strong ethnic identity who, until the early decades of the twentieth century, inhabited the Pontus area on the southern coast of the Black Sea. Scholars hold alternative views on their origin, proposing Eastern Armenia, Western Armenia, and Central Asia, respectively, as their most likely homeland. To ascertain whether genetic data from the nonrecombining portion of the Y chromosome are supportive of any of these suggestions, we screened 82 Armenian males of Hamsheni descent for 12 biallelic and 6 microsatellite Y-chromosomal markers. These data were compared with the corresponding data set from the representative populations of the three candidate regions. Genetic difference between the Hamshenis and other groups is significant and backs up the hypothesis of the Armenian origin of the Hamshenis, indicating central historical Armenia as a homeland of the ancestral population. This inference is further strengthened by the results of admixture analysis, which does not support the Central-Asian hypothesis of the Hamshenis' origin. Genetic diversity values and patterns of genetic distances suggest a high degree of genetic isolation of the Hamshenis consistent with their retention of a distinct and ancient dialect of the Armenian language.

Link

Armenian Y-STR haplotype data

The same data was studied by Herrera et al. (2011) where it was shown that haplogroup R2 was one of the distinguishing features of the Sasun community.

I decided to try the batch version of the Haplogroup Predictor on this data, and I include my results in this spreadsheet. This is useful as test data because real haplogroup assignments and Y-STR data are known for the same individuals, so they can be cross-checked against the predicted haplogroup.

Haplogroup prediction was made on the basis of the highest posterior probability and with equal priors. There were a few errors, some of which are understandable (for example, the 23-marker version does not include haplogroup R2, so the R2 samples were assigned to various other haplogroups).

With a few other discrepancies aside (e.g., some R1b1a2 assigned to L), the overall performance seems robust, and one can probably use this tool for published Y-STR data, with the caveat that some predictions for the less frequent haplogroups -for which there were probably fewer training samples- may be off the mark.

Legal Medicine doi:10.1016/j.legalmed.2012.10.003

Sub-population structure evident in forensic Y-STR profiles from Armenian geographical groups 

Robert K. Lowery et al


Over the course of its long history, Armenia has acted as both a source of numerous indigenous cultures and as a recipient of foreign invasions. As a result of this complex history among populations, the gene pool of the Armenian population may contain traces of historically well-documented ancient migrations. Furthermore, the regions within the historical boundaries of Armenia possess unique demographic histories, having hosted both autochthonous and specific exogenous genetic influences. In the present study, we analyze the Armenian population sub-structure utilizing 17 Y-chromosome short tandem repeat (Y-STR) loci of 412 Armenians from four geographically and anthropologically well-defined groups (Ararat Valley, Gardman, Lake Van and Sasun). To place the genetic composition of Armenia in a regional and historic context, we have compared the Y-STR profiles from these four Armenian collections to 18 current-day Eurasian populations and two ancient DNA collections. Our results illustrate regional trends in Armenian paternal lineages and locale-specific patterns of affinities with neighboring regions. Additionally, we observe a phylogenetic relationship between the Northern Caucasus and the group from Sasun, which offers an explanation for the genetic divergence of this group from other three Armenian collections. These findings highlight the importance of analyzing both general populations as well as geographically defined sub-populations when utilizing Y-STRs for forensic analyses and population genetics studies.


Link

The comings and goings of Near Eastern and European domestic pigs (Ottoni et al. 2012)

This is an excellent paper whose findings re: pig domestication seem to parallel many of my own observations regarding the flow of human populations. It is open access, so you can read it for yourselves, but the following figure illustrates the situation admirably:

The left-right arrangement of the columns corresponds to a west-east longitude across West Asia. It can be easily seen that some of the early domestic samples (yellow, bottom row) are concentrated in the west (Y1 haplotype), while others (blue, Arm1T) in the east.

Neolithic European samples possessed the Y1 haplotype, but lacked the Arm1T one. So, the authors conclude that:

The ancient Anatolian data presented here reveal that both wild and possibly domestic Neolithic pigs (identified using traditional metrics) possessed Y1 haplotypes ... The presence of these lineages corroborates the supposition that the earliest domestic pigs in Europe originated from populations originally domesticated in the Near East, conclusively linking the Neolithization of Europe with Neolithic cultures of western Anatolia (Larson et al. 2007a; Haak et al. 2010).

I have repeatedly highlighted the "puzzle" of the early European Neolithic: the signature Y-haplogroup G2a was unaccompanied by other common Near Eastern lineages, and the modal "West Asian" ancestral component in present-day West Asian populations seems to have been absent in early Neolithic samples, which were dominated by a "Sardinian-like" population. I have argued that this meant that the European Neolithic was drawn from a limited founder source that was more "Mediterranean/Southern" autosomally than "West Asian", at least in terms of the components identified by the Dodecad Project.

In Europe itself, the early Near Eastern domestic pigs were replaced by European ones:

Ancient DNA extracted from early Neolithic domestic pigs in Europe resolved this paradox by demonstrating that early domestic pigs in the Balkans and central Europe shared haplotypes with modern Near Eastern wild boar (Larson et al. 2007a). The absence of Near Eastern haplotypes in pre-Neolithic European wild boar suggested that early domestic pigs in Europe must have been introduced from the Near East by the mid 6th millennium BC before spreading to the Paris basin by the early 4th millennium BC (Larson et al. 2007a). 

By 3,900 BC, however, virtually all domestic pigs in Europe possessed haplotypes  from an indigenous European domestication process (Larson et al. 2007a) only found in European wild boar. This genetic turnover may have resulted from the accumulated introgression of local female wild boar into imported domestic stocks, or from an indigenous European domestication process (Larson et al. 2007a).

We have seen that early Neolithic domestic pigs came from Western Anatolia, but apparently these did not last, but were replaced in Europe by pigs carrying mtDNA of European wild boar. An additional possibility is that the European wild boar were better adapted to local conditions in Europe, so the stock of European farmers gradually became "local" due to artificial/natural selection favoring the local "European" type. It might also be that in accordance with Bergmann's rule, European-descended pigs were simply bigger, and thus more economically productive.

In any case, the interesting thing is that pigs carrying the "European" haplotype went the other way, crossing from Europe to Asia. The beginning of this process seems to have occurred in the Middle Bronze Age:

The temporal and geographic distribution of genetic haplotypes presented in our study demonstrates that the first AMS dated pig with European ancestry (haplotype A) appeared almost 1,000 years earlier than the Armenian samples in a Late Bronze Age context (~1,600-1,440 BC) at Lidar Höyük (fig. 1). An even earlier Middle Bronze Age specimen from the same site also possessed a European signature, but a direct
AMS date for this specimen could not be obtained.

I have written how increased mobility and long-range networks associated with the new metallurgical class facilitated commerce during the Bronze Age. The authors suggest the possibility of Minoan-Mycenaean/Hittite involvement during the Bronze Age, which are certainly plausible conduits for European pigs to have crossed the Aegean at this time. But, as you can see from the figure, the "European" pigs are still outliers during the Middle and Bronze Ages, but become common in the Iron Age sample from Lidar Höyük, and eventually replacing local types throughout Anatolia and Armenia, but, apparently, not Iran:

The frequency of pigs with European ancestry increased rapidly from the 12th century BC, and by the 5th century AD domestic pigs exhibiting a Near Eastern genetic signature had all but disappeared across Anatolia and the southern Caucasus. Though we did not detect European signatures in the ancient Iranian samples (fig. 1), the eastward spread of European lineages may have continued into Iran later than the Iron Age since European lineages have been found in wild caught modern Iranian samples (Larson et al. 2007a).

Of course a 12th century BC increase in European domestic pigs is entirely consistent -chronologically- with the Phrygian/Armenian settlement in Anatolia, and this association is further reinforced by the lack of European signatures in pigs from Iran where Phrygo-Armenians did not settle. The increase in European pigs could later be mediated by the Greek colonization, and the increase in trade during antiquity, just as trade would later introduce East Asian pig DNA into Europe.

The beautiful temporal transect presented in the Figure may also prove useful for students of ancient human DNA. I'd love to see how humans living close to sites #14-16, dominated by Arm1T haplotypes throughout history might differ from those of Neolithic West Anatolia, and whether the "mixed" Iron Age sample from Lidar Höyük shows evidence of the arrival of European-like human populations to accompany the European pigs.

Mol Biol Evol (2012) doi: 10.1093/molbev/mss261

Pig domestication and human-mediated dispersal in western Eurasia revealed through ancient DNA and geometric morphometrics

Claudio Ottoni et al.

Zooarcheological evidence suggests that pigs were domesticated in Southwest Asia ∼8,500 BC. They then spread across the Middle and Near East and westward into Europe alongside early agriculturalists. European pigs were either domesticated independently or appeared so as a result of admixture between introduced pigs and European wild boar. These pigs not only replaced those with Near Eastern signatures in Europe, they subsequently also replaced indigenous domestic pigs in the Near East. The specific details of these processes, however, remain unknown. To address questions related to early pig domestication, dispersal, and turnover in the Near East, we analyzed ancient mitochondrial DNA and dental geometric morphometric variation in 393 ancient pig specimens representing 48 archeological sites (from the Pre-Pottery Neolithic to the Medieval period) from Armenia, Cyprus, Georgia, Iran, Syria and Turkey. Our results firstly reveal the genetic signature of early domestic pigs in Eastern Turkey. We also demonstrate that these early pigs differed genetically from those in western Anatolia that were introduced to Europe during the Neolithic expansion. In addition, we present a significantly more refined chronology for the introduction of European domestic pigs into Asia Minor that took place during the Bronze Age, nearly 1,000 years earlier than previously detected. By the 5th century AD, European signatures completely replaced the endemic lineages possibly coinciding with the demographic and societal changes during the Anatolian Bronze and Iron Ages.

Link

rolloff and ALDER analysis of Turks

I carried out rolloff analysis of the Behar et al. (2010) sample of Turks together with the sample of Uzbeks from the same, and the Yunusbayev et al. (2011) sample of Armenians. A --geno 0.03 flag was applied for merging and SNPs available in the Rutgers recombination map for Illumina chips were used.

The exponential decay can be seen below:

The signal of admixture seems pretty clear and extends up to several cM. Of course, as always, this does not mean that exactly these two populations mixed to form the Turks sample, but it does mean that they are reasonable standins.

The jackknife gives an admixture time estimate of 27.622 +/- 5.348 generations or 800 +/- 160 years, which of course makes perfect historical sense as it is a date between the first arrival of the Seljuks in Anatolia and the final consolidation of power by the Ottomans. Note also that this probably applies principally to this particular sample (which I believe is from Cappadoccia) and there were perhaps different admixture dynamics elsewhere.

I had started this analysis before the announcement of ALDER, but since it is very fast, I decided to give it a go as well. Below is the raw output:




                    *** Admixture test summary ***

Weighted LD curves are fit starting at 1.45 cM

Pre-test: Does Turks have a 1-ref weighted LD curve with Armenians_Y?
   1-ref decay z-score:    0.09
   1-ref amp_exp z-score: -0.01
                                  NO: curve is not significant

Pre-test: Does Turks have a 1-ref weighted LD curve with Uzbeks?
   1-ref decay z-score:    6.56
   1-ref amp_exp z-score:  5.02
                                  YES: curve is significant

Does Turks have a 2-ref weighted LD curve with Armenians_Y and Uzbeks?
   2-ref decay z-score:    5.61
   2-ref amp_exp z-score:  5.58
                                  YES: curve is significant

Do 2-ref and 1-ref curves have consistent decay rates?
   1-ref Armenians_Y - 2-ref z-score:                  0.01   ( 13%)
   1-ref Uzbeks - 2-ref z-score:                       0.69   ( 11%)
   1-ref Uzbeks - 1-ref Armenians_Y z-score:          -0.00   ( -1%)
                                  YES: decay rates are consistent

Test FAILS (z=5.58, p=2.4e-08) for Turks with {Armenians_Y, Uzbeks} weights

DATA: failure 2.4e-08 Turks Armenians_Y Uzbeks 5.58 -0.01 5.02 13% 23.92 +/- 4.26 0.00002930 +/- 0.00000525 27.18 +/- 302.36 -0.00000082 +/- 0.00013129 26.84 +/- 4.09 0.00002316 +/- 0.00000461

DATA: test status p-value test pop ref A ref B 2-ref z-score 1-ref z-score A 1-ref z-score B max decay diff % 2-ref decay 2-ref amp_exp 1-ref decay A 1-ref amp_exp A 1-ref decay B 1-ref amp_exp B

The age estimate appears to be very similar, and most curves appear to be significant, except the one with Armenians_Y. This makes good sense. From Loh et al. (2012):

Also, if a reference A' shares some of the same admixture history as C or is simply very closely related to C, the pre-test will typically identify long-range correlated LD and deem A' an unsuitable reference to use for testing admixture.

In our case, A'=Armenians and C=Turks. We can be fairly sure that Armenians lack the same admixture history as Turks (because they were not affected by Central Asian Turkic invasions), but we can try a 1-ref analysis of Armenians with Uzbeks to substantiate it. The admixture lower bound estimate is a huge interval 7.6 +/- 88.2 and the jackknife is unable to estimate the admixture time. Thus, more plausibly, the second explanation applies, and because Armenians_Y are very closely related to Turks, they are deemed as an inappropriate reference to test admixture.

Finally, the lower bound of the admixture fraction for Turks with an Uzbek reference is estimated as:

Mixture fraction % lower bound (assuming admixture): 29.8 +/- 4.0

This is a very interesting number. We can be fairly sure that Central Asian Turkic people who invaded Anatolia carried with them an East Eurasian component, but in what proportion to their West Eurasian one? The East Eurasian element in Turks has been rather consistently estimated at ~5-7% with various methods, so perhaps this formed the minority element in the Turkic people who arrived in Anatolia. 

On the other hand, this case is rather muddled by the occurrence of by-directional gene flow: Uzbeks may have West Eurasian ancestry of ultimate West Asian origin, just as Turks have Central Asian ancestry. And, indeed, when we estimate the admixture fraction of Uzbeks with the Turks as a reference, we obtain:

Mixture fraction % lower bound (assuming admixture): 46.7 +/- 2.4

The age estimate for this is ~16 +/- 2 generations = 460 +/- 60 years. Very similar time estimates appear when Armenians are used as a West Eurasian reference. So, this might indicate that the Uzbek population was formed by admixture after the Anatolian Turks were so formed.

I see no easy way to solve the problem of estimating admixture proportions when both extant populations have been both donors and recipients of gene flow, but in any case, these numbers are something to think about.

Analysis of Turks with a variety of Turkic and East Asian populations

I subsequently formed a new dataset by merging the sample of Turks with a variety of Turkic and East Asian populations (same procedure for SNP choice).

For the calendar year calculation, I arbitrarily set the birthdate of the modern sampled individuals at 1980; I have no idea on the age profile of the individuals comprising the Behar et al. sample of Turks. I have also used a mindis=0.5cM which facilitated the convenient automated extraction of the dates from the ALDER output and also gave a level playing field for all the reference populations. The age picked by ALDER using its own adaptive threshold did not usually differ from the reported one by more than a few generations.

The results indicate two things:

• The % of admixture depends on the choice of population, with highest amount using Uzbeks  as a reference, and lowest using the far Asian populations from China. This indicates our uncertainty regarding the East/West Eurasian-ness of the people who settled in Anatolia.
• Admixture times, on the other hand appear to be fairly constant and appear to frame an important watershed moment of Anatolian history, the Battle of Manzikert which paved the way for the eventual Turkification of the peninsula. The Turkmen sample appears as an outlier in this respect, which might indicate that limited migration of Turkmen tribes may have occurred at a later date.

rolloff analysis of South Indian Brahmins as Armenian+Chamar

The first analysis of this population showed that there were negative f3(Brahmin; X, Y) signals when X were a variety of West European, Balkan, and West Asian population, and Y either the Chamar or North Kannadi. In the first analysis I used Orcadians and North Kannadi. I have now carried out a new rolloff analysis on 470,559 SNPs, using Armenians_Y and Chamar_M as the reference populations.

The exponential fit can be seen below.

The admixture date is 142.814 +/- 15.010 generations, or 4,140 +/- 440 years, which seems to correspond quite well with commonly accepted dates for the formation of Indo-Iranian.

I have previously observed that:

These patterns can be well-explained, I believe, if we accept that Indo-Iranians are partially descended not only from the early Proto-Indo-Europeans of the Near East, but also from a second element that had conceivable "South Asian" affiliations. The most likely candidate for the "second element" is the population of the Bactria Margiana Archaeological Complex (BMAC). The rise and demise of the BMAC fits well with the relative shallowness of the Indo-Iranian language family and its 2nd millennium BC breakup, and has been assigned an Indo-Iranian identity on other grounds by its excavator. As climate change led to the decline and abandonment of BMAC sites, its population must have spread outward: to the Iranian plateau, the steppe, and into South Asia, reinforcing the linguistic differentiation that must have already began over the extensive territory of the complex.

Quite possibly, as the West Asian element began mixing with the Sardinian-like population in Greece, another branch of the Indo-Europeans made its appearance east of the Caspian, in the territory of the BMAC, admixing with South Asian-like populations. Thus, it might seem that the Graeco-Aryan clade of Indo-European broke down during the Bronze Age, with one branch heading off to the Balkans, and another to the east. 

This scenario would also explain how the likely J2-bearing population associated with the earliest Proto-Indo-Europeans may have acquired the contrasting pattern I have previously described: the western (cis-Caspian) population would have admixed with R1b-bearers who occupy the "small arc" west and south of the Caspian, while the eastern (trans-Caspian) populations would have admixed with R1a-bearers who occupy the "large arc" in the flatlands north and east of the Caspian. It would also explain how the "western" branch (Graeco-Armenian) would have picked up Sardinian-like "Atlantic_Med" admixture, which is absent in the "eastern" Indo-Iranian branch.

At the same time, this scenario would explain the lack of "North European" admixture in the "western" branch (since this was shielded by the Caucasus and Black Sea from the northern Europeoids who may have lived north of these barriers), and explain it in the "eastern" branch (since the BMAC agriculturalists were in contact with presumably northern Europeoid groups inhabiting the steppelands, unhindered by any major physical barriers). (The relative absence of this admixture in the Graeco-Armenian branch may be advanced on the strength of its absence in Armenians, the evidence of a Sardinian-like Iron Age individual from Bulgaria, and the historical-era timing of admixture for the Greek population.)

It would be interesting to carry out similar experiments on Iranian groups, to see if they, too, present a similar pattern of admixture.

Armenians as Phrygian colonists, or, rolloff analysis of Armenians as a mixture of Sardinians+Balochi

I analyze the Yunusbayev et al. Armenians_Y sample in a similar manner as the South Indian Brahmins. The 30 lowest f3 statistics are:

Sardinian Velamas_M Armenians_15_Y -0.00349 0.000264 -13.23 239451
Sardinian Piramalai_Kallars_M Armenians_15_Y -0.003213 0.00028 -11.484 239389
Sardinian GIH30 Armenians_15_Y -0.002983 0.00023 -12.986 241310
Balochi Sardinian Armenians_15_Y -0.002837 0.000193 -14.681 241698
Sardinian Sindhi Armenians_15_Y -0.002794 0.000203 -13.757 241928
Sardinian Muslim_M Armenians_15_Y -0.002761 0.000295 -9.351 238639
Indian_D Sardinian Armenians_15_Y -0.002743 0.000224 -12.226 241916
Sardinian Kanjars_M Armenians_15_Y -0.002727 0.000281 -9.722 239240
Iyer_D Sardinian Armenians_15_Y -0.002718 0.000263 -10.322 238943
Brahui Sardinian Armenians_15_Y -0.002715 0.000196 -13.882 241885
Sardinian Dusadh_M Armenians_15_Y -0.002666 0.000281 -9.502 238994
Sardinian INS30 Armenians_15_Y -0.00265 0.000237 -11.162 240965
Iyengar_D Sardinian Armenians_15_Y -0.002624 0.000297 -8.847 238564
Sardinian Dharkars_M Armenians_15_Y -0.002501 0.000267 -9.359 239505
Sardinian North_Kannadi Armenians_15_Y -0.002463 0.000288 -8.545 239278
Sardinian Chamar_M Armenians_15_Y -0.002445 0.000275 -8.904 240102
Sardinian Kshatriya_M Armenians_15_Y -0.002372 0.000267 -8.897 239047
Pathan Sardinian Armenians_15_Y -0.00224 0.000199 -11.264 241759
Sardinian Brahmins_from_Uttar_Pradesh_M Armenians_15_Y -0.002189 0.00025 -8.774 239395
Jatt_D Sardinian Armenians_15_Y -0.001806 0.000273 -6.608 238465
Cypriots Kanjars_M Armenians_15_Y -0.001699 0.00026 -6.547 238237
Cypriots Velamas_M Armenians_15_Y -0.001642 0.000275 -5.965 238392
Cypriots Muslim_M Armenians_15_Y -0.001618 0.000279 -5.798 237762
Cypriots Dusadh_M Armenians_15_Y -0.001611 0.000279 -5.779 238096
GIH30 Cypriots Armenians_15_Y -0.001608 0.000223 -7.216 239819
Iyer_D Cypriots Armenians_15_Y -0.001562 0.000251 -6.217 238012
Sindhi Cypriots Armenians_15_Y -0.001544 0.000209 -7.383 240276
Cypriots North_Kannadi Armenians_15_Y -0.001464 0.000276 -5.298 238273
Cypriots Kshatriya_M Armenians_15_Y -0.001438 0.00026 -5.534 238076

I carried out rolloff analysis using the Balochi and Sardinians as references, for a total of 510,844 SNPs. Note that the Burusho were not used in this experiment, because they were culled due to more than 5% East Eurasian admixture, as per the followed procedure. 

The Balochi are very similar to the Burusho otherwise, and this also gives me the opportunity to see a Sardinian+Balochi population pair to complement a previous analysis of French as Sardinian+Burusho, which presented an f3 signal of quite similar intensity. The exponential fit is seen below.

The jackknife gives an age estimate of 113.194 +/- 14.674 generations, or 3,280 +/- 430 years, assuming a generation length of 29 years.

I had somewhat expected the Armenians to show a more recent signal of admixture than the French, as they lived much closer to the boundary of Europe and Asia, and may have had more opportunity to admix between Sardinian-like populations of Europe and "West Asian"-like populations of Asia.

But, the inferred date also raises another possibility. Herodotus says of the Armenians who were part of the army of King Xerxes:

the Armenians were equipped like Phrygians, being Phrygian colonists" (7.73)

Now, the Phrygians became masters of Central Anatolia during the tumultuous events near the end of the Bronze Age (12th century BC), following the collapse of the Hittite Empire. And, their ancestral homeland was in Thrace. And, there is fairly good evidence that Armenian is the closest language related to Greek within the Indo-European language family. And, we have some tantalising evidence that even during the Iron Age, the population of Thrace was Sardinian-like. And, the Armenians do contrast with their Caucasian neighbors in possessing ~10% of the Sardinian-like Atlantic_Med component that South and Northeast Caucasians lack.

All of the above combine to make a pretty compelling story. Could it be that Armenians preserve a legacy of admixture between a linguistically Indo-European speaking, genetically Sardinian-like population, which arrived in Asia Minor from the Balkans at the end of the Bronze Age, finally settling in the Armenian Highlands, and mixing with the local people they encountered?

The plot thickens. And, this is, certainly, a question that can be answered by ancient DNA research, e.g., by comparing the genomes of historical Phrygians and Armenians with those from Hittite-era, or earlier Anatolians.

Huge study on Y-chromosome variation in Iran (Grugni et al. 2012)

This is the equivalent of a box of candy for anyone interested in Eurasian (pre-)history. I will have digest all the goodies within, and post any of my comments as updates to this post.

UPDATE I: Here is the table of haplogroup frequencies for easy reference:

One of the most interesting finds is the presence of a few IJ-M429* chromosomes  in the sample. Haplogroup IJ encompasses the major European I subclade, and the major West Asian J subclade. The discovery of IJ* chromosomes is consistent with the origin of this haplogroup in West Asia; it is widely believed that haplogroup I represents a pre-Neolithic lineage in Europe, although at present there are no Y chromosome-tested pre-Neolithic remains.

There is also a wide assortment of Q and R in Iran. While some of these may be intrusive (e.g., the 42.6% of Q1a2 in Turkmen, likely a legacy of their Central Asian origins), the overall picture appears consistent with a deep presence of these lineages in Iran. This is especially true for haplogroup R where pretty much every paragroup and derived group is present, excepting those likely to have originated recently elsewhere.

UPDATE II: From the paper:

Although accounting only for 25% of the total variance, the first two components (Figure 3) separate populations according to their geographic and ethnic origin and define five main clusters: East-African, North-African and Near Eastern Arab, European, Near Eastern and South Asian. The 1stPC clearly distinguishes the East African groups (showing a high frequency of haplogroup E) from all the others which distribute longitudinally along the axis with a wide overlapping between European and Arab peoples and between Near Eastern and South Asian groups. The 2ndPC separates the North-African and Near Eastern Arabs (characterized by the highest frequency of haplogroup J1) from Europeans (characterized by haplogroups I, R1a and R1b) and the Near Easterners from the South Asians (due to the distribution of haplogroups G, R2 and L). Iranian groups do not cluster all together, occupying intermediate positions among Arab, Near Eastern and Asian clusters. In this scenario, it is worth of noticing the position of three Iranian groups: (i) Khuzestan Arabs (KHU-Ar) who, despite their Arabic origin, are close to the Iranian samples; (ii) Armenians from Tehran (THE-Ar), whose position, in the upper part of the Iranian distribution, indicates a close affinity with the Near Eastern cluster, while their position near Turkey and Caucasus groups, due to the high frequency R1b-M269 and other European markers (eg: I-M170), is in agreement with their Armenia origin; (iii) Sistan Baluchestan (SB-Ba) that clusters with its neighbouring Pakistan.

UPDATE III: There are lots of little details in the haplogroup distribution that make historical sense. For example, C3 exists in Assyrians from Azarbaijan, and both C*, C3, and O exists in Zoroastrians from Yazd. It is often forgotten that before the spread of Islam, and quite time thereafter, Inner Asia was teeming with Zoroastrians and Nestorian Christians. It seems quite likely that these outliers represent a legacy of these communities.

UPDATE IV: I have a feeling that Razib will take exception with this statement: "Ancient Persian people were firstly characterized by the Zoroastrianism. After the Islamization, Shi'a became the main doctrine of all Iranian people."


UPDATE V: This confirms my observation from the recent studies in Afghanistan, that there is an inverse relationship of J2a and R1a in Iranian-speaking groups, with an excess of the latter among the eastern Iranians, and of the former among the Persians. From the paper:

Among the different J2a haplogroups, J2a-M530 [46] is the most informative as for ancient dispersal events from the Iranian region. This lineage probably originated in Iran where it displays its highest frequency and variance in Yazd and Mazandaran (Figure 2). Taking into account its microsatellite variation and age estimates along its distribution area (Tables S3 and S7), it is likely that its diffusion could have been triggered by the Euroasiatic climatic amelioration after the Last Glacial Maximum and later increased by agriculture spread from Turkey and Caucasus towards southern Europe. The high variance observed in the Italian Peninsula is probably the result of stratifications of subsequent migrations and/or of the presence of sub-lineages not yet identified. Of interest in the M530 network (Figures 2 and S3) is the presence of a lateral branch that is characterized by a DYS391 repeat number equal to 9. Differently from previous observations [46], this branch is not restricted to Anatolian Greek samples being shared with different eastern Mediterranean coastal populations. The M530 diffusion pattern seems to be also shared by the paragroups J2a-M410* and J2a-PAGE55*. In addition, the variance distribution of the rare R1b-M269* Y chromosomes, displaying decreasing values from Iran, Anatolia and the western Black Sea coastal region, is also suggestive of a westward diffusion from the Iranian plateau, although more complex scenarios can be still envisioned because of its non-star like structure.

Of course, the idea that the diffusion of J2a related lineages ties in with early agricultural expansions has been with us for a long time, but it is time to abandon it. First of all, as we have seen, J2a diminishes greatly as we head towards South Asia; it certainly doesn't look like the lineage of the multitude of agricultural settlements that sprang up along the southeastern vector soon after the invention of agriculture. Second, it is lacking so far in all ancient Y chromosome data from Europe down to 5,000 years ago. It seems much more probably that J2 related lineages spread from the highlands of West Asia much later. 


The "age estimates" are the result of using the inappropriate "evolutionary mutation rate", and become even older because of the inclusion of the DYS388 marker that is very stable in many haplogroups but very mutable within haplogroup J. On the left you can see frequency, Y-STR variance, and haplotype network structures for various J-related groups.


It is unfortunate that there is no progress in the phylogeographic assessment of R1a in this paper. There have been substantial discoveries of SNPs within this haplogroup as a result of commercial testing; however there is clearly an ascertainment bias in the newer discoveries, as almost all these SNPs have been detected in Europeans. The new paper confirms the high levels of Y-STR variance in India, Pakistan, and Iran. Together with the cornucopia of related paragroups in Iran, there is little doubt that this haplogroup originated in the general area of Central/South Asia.


Personally, as I have stated before, I would relate this R1a with Neolithic peoples living east of the Caspian, in contrast to the R1b bearers who lived west and south of it. These two populations came under the influence of the Indo-Europeans and spread in different directions. The Indo-Iranians were then initially the mixed descendants of the Indo-Europeans and the R1a old agricultural population, and were formed in the territory of the Bactria-Margiana Archaeological Complex. 


This also explains the contrast between Iranian and Armenian groups: the latter mostly lack the R1a lineage, contrasting with all Iranian groups (even their Kurdish neighbors) who possess it. Conversely, Iranian groups, and especially eastern Iranians and Indo-Ayrans lack the R1b lineage. This is due to the fact that neither R1a nor R1b were originally part of the Indo-European community, but their geographical position was such that they came under the influence of the Indo-Europeans when the latter began their expansion.


UPDATE VI: I have created my own dendrogram using the Y-haplogroup frequencies and the hclust package of R (default parameters):

From top to bottom, one can identify some clusters:

• Eastern Europe, further broken down into Balkans and Slavic+Hungary
• West Asian/Caucasus
• Iranian Proper
• Arab

These correspond largely to the clusters identified by the authors, with India and the Turkmen sample emerging as the clear outliers. I omitted the Ethiopian samples, since E-M78 was not resolved phylogenetically, causing the Ethiopians to group with the likely E-V13 from the Balkans.

UPDATE VII: I have also run MCLUST over the haplogroup frequency data over the MDS representation of the distance matrix. The maximum number of 10 clusters occurred with 5 MDS dimensions retained. Population assignments in the 10 clusters can be found in the table below:

Iran/Azerbaijan_Gharbi+Tehran_(Assyrian)	1
Iran/Lorestan_(Lur)	1
Iran/Tehran_(Armenian)	1
Iran/Azerbaijan_Gharbi_(Azeri)	2
Iran/Hormozgan_(Bandari+Afro-Iranian)	2
Iran/Hormozgan/Qeshmi	2
Iran/Khorasan_(Persian)	2
Iran/Kurdistan_(Kurd)	2
Iran/Sistan_Baluchestan_(Baluch)	2
Pakistan	2
Iran/Fars+Isfahan_(Persian)	3
Iran/Gilan_(Gilak)	3
Iran/Yazd+Tehran_(Zoroastrian)	3
Turkey/Central	3
Turkey/East	3
Turkey/West_	3
Iran/Golestan_(Turkmen)	4
India	4
Iran/Khuzestan_(Arab)	5
Egypt_(Arab)	5
Iraq/Baghdad	5
Oman	5
Saudi_Arabia	5
Tunisia	5
United_Arab_Emirates	5
Iran/Mazandaran_(Mazandarani)	6
Iran/Yazd_(Persian)	6
Balkarian	6
Georgia	6
Albania	7
Greece	7
Bosnia	8
Croatia	8
Slovenia	8
Czech_Republic	9
Hungary	9
Poland	9
Ukraine	9
Iraq_(Marsh_Arab)	10
Qatar	10
Yemen	10

We can ignore cluster #4 which consists of the two outliers (India + Turkmen). The rest of the clusters seem relatively coherent. Notice, for example, the Arabian cluster #10, Balkan cluster #8, Eastern European cluster #9, Greek-Albanian cluster #7, Mixed Arab cluster #5.

PLoS ONE 7(7): e41252. doi:10.1371/journal.pone.0041252

Ancient Migratory Events in the Middle East: New Clues from the Y-Chromosome Variation of Modern Iranians

Viola Grugni et al.


Knowledge of high resolution Y-chromosome haplogroup diversification within Iran provides important geographic context regarding the spread and compartmentalization of male lineages in the Middle East and southwestern Asia. At present, the Iranian population is characterized by an extraordinary mix of different ethnic groups speaking a variety of Indo-Iranian, Semitic and Turkic languages. Despite these features, only few studies have investigated the multiethnic components of the Iranian gene pool. In this survey 938 Iranian male DNAs belonging to 15 ethnic groups from 14 Iranian provinces were analyzed for 84 Y-chromosome biallelic markers and 10 STRs. The results show an autochthonous but non-homogeneous ancient background mainly composed by J2a sub-clades with different external contributions. The phylogeography of the main haplogroups allowed identifying post-glacial and Neolithic expansions toward western Eurasia but also recent movements towards the Iranian region from western Eurasia (R1b-L23), Central Asia (Q-M25), Asia Minor (J2a-M92) and southern Mesopotamia (J1-Page08). In spite of the presence of important geographic barriers (Zagros and Alborz mountain ranges, and the Dasht-e Kavir and Dash-e Lut deserts) which may have limited gene flow, AMOVA analysis revealed that language, in addition to geography, has played an important role in shaping the nowadays Iranian gene pool. Overall, this study provides a portrait of the Y-chromosomal variation in Iran, useful for depicting a more comprehensive history of the peoples of this area as well as for reconstructing ancient migration routes. In addition, our results evidence the important role of the Iranian plateau as source and recipient of gene flow between culturally and genetically distinct populations.

Link

Craniological and dental signatures of Out-of-Armenia

From the paper on the craniological results:

One can see a clear link between the Armenian highlands samples and the Western Europe samples (the Arcvakar sample - 17 close phenetic links are revealed). The samples from the Georgia (Samtavro /Late Bronze Age - II period) and Iran (Tepe Gissar III), Uzbekistan (Sapallitepe) are identified as the samples with closest affinities samples from Ukraine (Shirochanski) and Poland, Germany (Corded Ware culture) in particular (figure 3). This suggests that some of the European genes do actually stem from this area. So, mediterranean connections from Armenian highlands, Georgia and Central Asia are distinctly fixed in Western Europe and in the Middle-Late Bronze Age. 

... 

If true, it is suggested that the dispersal of the Indo-European languages have been accompanied by migration and some gene flow from the Armenian highlands homeland to the various historical seats of the Indo-European languages. The different rates of genetic drift and external gene flow may have contributed to the morphological differentiation and diversification amongst the different Eurasian populations. Cluster analysis has revealed a craniological series having analogies (on a complex of craniometric, odontologic characters) with representatives of the population of the Armenian highlands, the Caucasus, the Near East and Central Asia. The initial starting area (or one of the intermediate areas), as indicated by the anthropological data, would seem to be the Armenian highlands, and the Caucasus as a whole (Figure 7).

Asian Culture and History Vol. 4, No. 2; July 2012
doi:10.5539/ach.v4n2p48


Bioarchaeological Analysis Mutual Relations of Populations Armenian Highlands and Eurasia Using Craniological and Dental Nonmetric Traits

Anahit Yu. Khudaverdyan1 Institute of Archaeology and Еthnography National Academy of Science, Republic of Armenia

Abstract

Undertaken here is a multidimensional craniometric analysis of more than 254 ethnic groups of the Neolithic and Bronze Ages from the territory of Eurasia. On the basis of the received information, cluster analysis was done and has shown the genetic condensations of ethnoses and vectors of relatives or, conversely, distinctions between them. Craniometric and odontologic investigation of the Bronze Age is interesting and in connection with discussion about the origin of Indo-Europeans and about the place of their ancestral home. Different aspects of the problem of the ancestral home of Indo-Europeans are far from completely resolved and generate lively debate in the pages of scientific publications. New anthropological data allowed identification of alien Mediterranean characteristics influencing various ethnic Eurasian groups and revealed evidence of a migratory stream from the Armenian highlands and the Caucasus. This research provided new evidence of patterns of ethnic contact and intermixture in Western Eurasia

Link

First look at Turkish and Kyrgyz data from Hodoğlugil & Mahley (2012)

The authors of the recent paper on Turkish population structure were kind enough to share their data with me. I will be sure to use this data in future experiments, such as the ChromoPainter and fastIBD analysis of Balkans/West Asia, as well as a ChromoPainter analysis of Altaic speakers, following on the footsteps of my recent analysis of Afroasiatic speakers.

PCA


As a first step, after processing the new data, I carried out a PCA analysis (in smartpca with no outlier removal iterations), combined with various Turkic groups, as well as a few neighbors of Anatolian Turks, combining data from the literature and the Dodecad Project.

The Turkic cline from East to West Eurasia, observed by myself and others in various experiments is again evident.

The blowup of the above, focusing on the West Eurasian portion (top right) is easier to read:

As always, population labels are placed in the average position of each population. So, for example, the Behar et al. Iranians_19 sample is shifted to the left, because of the existence of a few African admixed individuals in this sample. The Iranian_D sample of Project participants seem to lack this admixture.

Also, note that since there is no South Asian reference in this first experiment, Iranians overlap with Turks along the first two dimensions. As we've seen in the Dodecad Project, both Iranians and Anatolian Turks are "eastward-shifted" relative to other West Eurasians, but the former have a strong South Asian- and the latter a Central Asian- tendency.

The new Kyrgyz sample falls between the Kazakh and the Altai along the cline, and is more "eastern" compared to the Uygurs and Uzbeks, and more "western" compared to Altai, Tuva, and Dolgans.

Kayseri and Istanbul Turks overlap with Behar et al. Turks as well as the Turkish_D sample. The Aydin sample appears to be more heterogenous, with a more eastern overall center of weight. More on this below.

ADMIXTURE


I also carried out a K=3 ADMIXTURE analysis of the dataset.

Below are the population portraits for the three new Turkish samples, as well as the Kyrgyz sample:

It is obvious that many Turks have low levels of Asian admixture, lacking in their geographical neighbors, but this is quite variable on an individual basis.

UPDATE (17 Feb):


I have also assessed the new data with the K12b calculator. Below are the normalized median proportions.

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

Armenian Y-chromosomes have been a largely ignored since the publication of the classic Weale et al. (2001) paper a decade ago. The Armenian DNA Project has largely covered the void during the intervening years, but it is nice that the topic is revisited by academics.

Armenia is sandwiched between Anatolia, the Fertile Crescent, the Iranian plateau, the Caucasus, and the Black and Caspian seas, making the study of Armenian Y-chromosomes extremely interesting for the student of Eurasian prehistory.

Gene flow from the surrounding regions may have affected the Armenian population over historical time, but the remoteness of the Armenian highlands, coupled with the national church -- which distinguished Armenians from both the Orthodoxy of the Roman Empire, the Zoroastrianism of the Persians, and, later the Islam of Arabs and Ottomans -- may have prevented it.

My comments on the paper will follow below once I read it.

UPDATE I: The paper spends a lot of time on analysis of Y-STR variance; my opinion of Y-STRs as a tool for inferring past population movements is, to put it mildly, low. When Bahamian Y-STR variance is higher than African one, and E-V13, one of the youngest European Y-haplogroups (in terms of Y-STR variance) turns up in Spain in one of the earliest ancient DNA samples, it goes without saying that the burden of proof is on those who wish to continue to talk about Neolithic or other population movements to make the assumptions of their models clearer. Nonetheless, there is still some utility in Y-STRs, so I reproduce some tree diagrams from the paper (top left), and link to the supplementary info that has a collection of haplotypes that may be useful to genealogists.

From the paper:

However, owing to the contentions associated with the current calibrations of the Y-STR mutation rates,32,34,35,41 as well as the limitations of the assumptions utilized by the methodologies for time estimations, the absolute dates generated in this study should only be taken as rough estimates of upper bounds.

Indeed. We are at the point where Y-STRs are at the end of their utility, but the replacement technology of extensive Y-chromosome sequencing has not quite arrived in an economical way yet.

UPDATE II: I will have some additional thoughts on Y-chromosome distribution in the third update, but, for the time being, the two most important "nuggets" of information are: (i) the unusual haplogroup frequencies in Sasun (high R2 and T), which may be due to a founder effect, but it would be interesting if Armenian historians could find some explanation for their occurrence there, and (ii) the occurrence of R-M269*(xL23) in Ararat Valley. I invite more knowledgeable readers to comment on the issue; the haplotypes are in Table 2 of the supplement.

UPDATE III: The ubuiquity of haplogroup G2a in Neolithic Europe, coupled with the absence of other prominent present-day European haplogroups, has important implications about European discontinuity.

But, it also has implications about West Asian discontinuity. The Neolithic in Europe arrived by all accounts from either of two principal areas: Anatolia or the Levant. Today, in Anatolia and the Levant, we see a set of haplogroups of which haplogroup J is the most important and ubiquitous one. Haplogroup R1b is also quite frequent in Armenia, the east Caucasus, Anatolia, and Iran, but its frequency drops dramatically to the east and south. And, there is a whole assortment of other haplogroups with varying frequency.

Why didn't all these non-G2a haplogroups participate in the early Neolithic colonization of Europe? It could very well be that a very small founder population crossed the Aegean into Europe, one that happened to be G2a-dominated. But, that is ultimately not very satisfying: if there was plenty of J and R1b in West Asia at the time of the Neolithic expansion, why are these haplogroups so conspicuous in their absence -at least so far- from Neolithic Europe?

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.

And, the story has other complications. From the current paper:

The relative expansion times for haplogroup J2-M172 (Table 4) generally correspond with those yielded for R1b-M343, with the exception of Greece and Crete, which, unlike haplogroup R1b-M343, are slightly older than the dates yielded for several of the Near Eastern groups as well as the four Armenian populations.

As mentioned above, I don't give much weight on Y-STR evidence, but observations such as the above certainly add to the feeling of unease that something is not quite right with the default picture of prehistory.

Another observation on the Armenian population, is its very low frequency of haplogroup R1a1. Proponents of the Kurgan model of Indo-European dispersals sometimes associate this haplogroup with the Proto-Indo-European community, and it is strange why -if their ideas are right- Armenia is so lacking in this haplogroup, like its Caucasian neighbors. Why would these hypothetical migrants make such a huge impact in faraway India and barely a dent in nearby Armenia?

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?

The veil of mystery seems to be raised even higher by every new study: we may be less certain of what really happened today than in the days of happy ignorance, ten years ago. Ultimately it is new data, like the ones included in this paper, that will make every piece of evidence fit, and the grand puzzle of the history of Eurasia will be revealed in all its glory.

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

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.

Link