Complex Y chromosome structure in East Tyrol (and more IE thoughts)

Cultural diversity can disappear in a few generations, but genetic diversity -barring major genocides or disasters- usually persists.

The East Tyrol region in Austria has been Germanic-speaking since the Middle Ages, but historical evidence documents the presence of Romance, Germanic, and Slavic groups in its territory. How can we untangle the origin of the different groups when they are all jumbled up together now, and all Germanic-speaking? Previous research has shown that patrilineal groups can be isolated on the basis of surnames, but in the case of East Tyrol, the wide adoption of surnames happened after the region had become linguistically Germanic.

The authors of the new paper exploited the structure of local toponyms, in particular pasture names. The figure on the left shows the concentrations of Slavic (panel A), Romance (panel B), and Germanic (panel C) pasture names. While Germanic pasture names are evenly distributed, there is a contrast between those of Slavic and Romance origin. From the paper:

From the 853 analyzed pasture names in East Tyrol 71% were derived from Germanic (Bavarian) etymons, 17% from Slavic etymons, and 12% from Romance etymons. While pasture names with Germanic etymons were evenly distributed in high density within the whole study area the names with Slavic etymons were spatially focused in the east and north of East Tyrol (Fig. 2). Geographically, these are the lower Drau, Isel, Kals, Virgen and the Defereggen valleys (Fig. 1). No names with Slavic etymons were found in the southwestern Puster valley (Fig. 2). The pasture names with Romance etymons focus mainly in the southern part of East Tyrol (Gail, Puster, and Villgraten valley, Fig. 2). The slight northeastward trend observed in the distribution of Romance etymons is solely caused by the Kals valley, a medieval Romance linguistic enclave, which was separated from the Romance main territory in the 10th century [36]. On the basis of these results, East Tyrol was divided into two regions of former Romance (Puster, Gail, and Villgraten valley; region A) and Slavic (Isel, lower Drau, Defereggen, Virgen, and Kals valley; region B) main settlement (Fig. 2).

The authors dissected the occurrence of different haplogroups in the two contrasting regions (A: Romance, and B: Slavic) in some great detail:

Splitting the East Tyrolean population sample into regions A and B resulted in a partitioning of haplogroups E-M78, R-M17, R-M412/S167*, and R-S116*. E-M78, R-M17 and R-S116* Y chromosomes were exclusively found in region B whereas samples assigned to R-M412/S167*, R-U106/S21, and R-U152/S28 reached higher frequencies in region A (Fig. 3, Table S7). When attributing the samples to the fathers' and grandfathers' places of birth/residence, as reported by the participants, practically identical patterns were obtained for most of the haplogroups (Fig. 3). 

Y chromosomes belonging to haplogroups G-P15, I-M253, and J-M304 showed much lower regionalization in their frequencies (Fig. 3) at all three generation levels.

The non-localization of the G-P15, I-M253, and J-M304 seems reasonable as these may represent what is common in these populations (and one could indeed speculate -on the basis of current ancient DNA knowledge- that they correspond to Neolithic, Paleolithic, and Bronze Age processes respectively)

Two of the most interesting findings are:

Haplogroup R-M412/S167* was found at low frequencies in the combined East Tyrolean sample. However, the R-M412/S167* chromosomes were sorted by the subdivision of the study area and reached in region A levels of ~14% whereas their frequency in region B was well below the 5% threshold. At the probands and fathers level of analysis region A featured approximately fourfold higher frequencies of these chromosomes than region B. This ratio changed to about nine when placing the samples at the grandfathers' places of birth/residence. These contrasts remained statistically significant after correcting for multiple comparisons [22] at the fathers and grandfathers analysis level.

and:

The western border of the geographic expansion of haplogroup R-M17 Y chromosomes is to be found in Central Europe and largely follows the political border separating present-day Poland (57%) and Germany (East: ~30%, South: ~14%, West: ~10%) [42]. Frequencies of about 15% and 10% were also found for Austria [18] and North-East Italy [48], respectively. In South Italy and in West Europe R-M17 chromosomes are not present at informative frequencies. 

In this study, the proportion of Y chromosomes carrying the derived M17 allele was 14.1%, a value that nearly perfectly matched those reported for West Austria (North Tyrol, 15.4%) and South Germany (Munich; 14.3%) [18], [42]. However, haplogroup R-M17 was completely absent in the East Tyrolean sub-sample from region A, but made up to 16% in region B. This result remained practically unchanged when assigning the probands to their respective fathers' or grandfathers' places of birth/residence (Fig. 3).

The new study reinforces my belief that R-M17 was not originally present in the Italo-Celtic branch of Indo-European. Together with the paucity of the same lineage in Albanians (~5%), Armenians (less than 5%), and its quite uneven distribution in Greeks, it is becoming increasingly clear that R-M17 may represent a late entrant that affected minimally southern and western Europe.

The fountain of its spread was probably a trans-Caspian (?) Central Asian staging point that followed a counter-clockwise route into Europe that spawned the northern (Germanic and Balto-Slavic) groups of Europe and the Indo-Iranians, who remained longer in their BMAC homeland, finally breaking down during the 2nd millennium BC. This would also harmonize with the increasing evidence for complementary R-M17 distributions in Europe and Asia, associated with the Z93 marker. 


It might appear that Z93+ chromosomes may track the later expansion of the Indo-Iranian world. I have observed before that R-M17 seems distributed in a long arc north and east of the Caspian, and it is perhaps in different points along this arc that the dominant European (NW) and Asian (SE) types emerged out of the early Neolithic population.

Combining this insight with an analysis of Y chromosome variation within the Graeco-Armeno-Aryan group, it appears that Graeco-Armenian is characterized predominantly by J2+R1b related lineages, while Indo-Iranian by J2+R1a related lineages. The evidence for Tocharian would involve J2+R1b related lineages.  Overall, it would appear that the earliest J2 core of PIE affected two different groups of populations living on complementary sides of the Caspian:

• The trans-Caspian R-M17 population followed an early (3rd, or late 4th millennium BC?) north-west trajectory into Europe (associated with northern European groups such as Balto-Slavic and Germanic) as well as a later expansion (2nd millennium BC? associated with climatic deterioration in BMAC) that brought Iranian speakers to the steppe, as well as to Iran, and Indo-Aryans to South Asia.
• The cis-Caspian, trans-Caucasian R-M269 population followed an early (late 4th millennium, early 3rd millennium?) expansion into Europe, probably together with J2 in the Balkans (Graeco-Phrygian, perhaps Thracian), and arriving in the form of Bell Beakers in Western Europe (Italo-Celtic), as well as a later (2nd-1st millennium BC?) expansion to the east (Tocharians)

This long excursus was necessary as a preamble to an explanation on what happened in Europe itself, which brings us back to the topic of the current paper:

• The lack of structure between regions A and B with respect to haplogroup J, together with the great difference in levels of this haplogroup between Italy and the Celtic world,  suggests that Italian J-related lineages  may have been inflated in proto-historical and historical times. There are candidates a-plenty: Greeks, Etruscans, Trojans to name but three. Excess of J in Italy, relative to the Celtic world, clearly relates to the abundant traditions of eastern origins for the historical groups of Italy.
• It would appear that during proto-history, most of Europe was dominated by three sets of IE people (R-M269 in the west, who had transmitted Proto-Celto-Italic; R-M17 in the northeast of Proto-Balto-Slavic speech, and Proto-Germanic in-between, participating in both worlds, and --appropriately-- often linked with either Italo-Celtic or Balto-Slavic linguistically)
• There were other (now-extinct) groups as well: the Illyrians vs. Thracians in the Balkans with complementary Y chromosome distributions, the former including an extra chunk of aboriginal legacy (haplogroup I), no doubt due to the much more difficult terrain of the western Balkans. These are contrasted with our final group, the Greeks who straddled three worlds (the Paleo-Mediterranean world of the first farmers, the Thraco-Phrygian world linked to the Indo-Iranians at a deeper level, and the Anatolian world)

The boundaries between these various groups were a little blurred in the course of history. But, apparently, they were still a little clearer during the Middle Ages, and probably much clearer before the Völkerwanderung of the Germans, and the expansion of the Slavs.


Geneticists are executing a remarkable pincer movement, zeroing in on the period of European ethnogenesis from both the remote past and the present: through a study of ancient DNA from the dawn of history, they are beginning to understand how Europe was peopled, layer after layer of settlement; and through the study of surnames and toponyms they are drilling ever deeper into the pre-genealogical past. Together with much anticipated technological progress related to full genome sequencing and ancient DNA extraction, it will not be long before the history of Europe will be laid bare in remarkable detail.

PLoS ONE 7(7): e41885. doi:10.1371/journal.pone.0041885

Pasture Names with Romance and Slavic Roots Facilitate Dissection of Y Chromosome Variation in an Exclusively German-Speaking Alpine Region

Harald Niederstatter et al.

The small alpine district of East Tyrol (Austria) has an exceptional demographic history. It was contemporaneously inhabited by members of the Romance, the Slavic and the Germanic language groups for centuries. Since the Late Middle Ages, however, the population of the principally agrarian-oriented area is solely Germanic speaking. Historic facts about East Tyrol's colonization are rare, but spatial density-distribution analysis based on the etymology of place-names has facilitated accurate spatial mapping of the various language groups' former settlement regions. To test for present-day Y chromosome population substructure, molecular genetic data were compared to the information attained by the linguistic analysis of pasture names. The linguistic data were used for subdividing East Tyrol into two regions of former Romance (A) and Slavic (B) settlement. Samples from 270 East Tyrolean men were genotyped for 17 Y-chromosomal microsatellites (Y-STRs) and 27 single nucleotide polymorphisms (Y-SNPs). Analysis of the probands' surnames revealed no evidence for spatial genetic structuring. Also, spatial autocorrelation analysis did not indicate significant correlation between genetic (Y-STR haplotypes) and geographic distance. Haplogroup R-M17 chromosomes, however, were absent in region A, but constituted one of the most frequent haplogroups in region B. The R-M343 (R1b) clade showed a marked and complementary frequency distribution pattern in these two regions. To further test East Tyrol's modern Y-chromosomal landscape for geographic patterning attributable to the early history of settlement in this alpine area, principal coordinates analysis was performed. The Y-STR haplotypes from region A clearly clustered with those of Romance reference populations and the samples from region B matched best with Germanic speaking reference populations. The combined use of onomastic and molecular genetic data revealed and mapped the marked structuring of the distribution of Y chromosomes in an alpine region that has been culturally homogeneous for centuries.

Link

Frachetti on the multiregional emergence of mobile pastoralism

I had previously posted on horses not being important for the emergence of steppe pastoralism, in which Frachetti and Benecke documented how the early (4,500ky) Begash culture of Kazakhstan had developed full-blown pastoralism without apparently relying on horses.

This is in contradistinction to both the Botai culture where there is abundant evidence for horse use, apparently for food, as well as the Eneolithic cultures of the European steppe where horse bones are much more prevalent than in early Begash.

The simple model of the emergence of pastoral nomadism proposes the spread of this mode of subsistence from the European (Pontic-Caspian) steppe, together with horses and horse-drawn vehicles. Popularized by David Anthony in recent years, this model views horses and wheels as the great enablers of pastoralism, and views the emergence of pastoral cultures across the steppe as the result of movements of mobile pastoralists -atop their horses, and with their herds- across the Eurasian steppe.

Frachetti is a critic of this model, and proposes instead the importance of the (hitherto neglected) Inner Asian Mountain Corridor as important in facilitating prehistoric contacts between east and west. In a new paper in Current Anthropology he elaborates on his proposed "multiregional" model of the emergence of mobile pastoralism.

I have not read the paper's 38 pages carefully yet (including CA comments and response), but this is clearly a seminal work on the studied topic that will be referenced for years to come. I will post any specific comments in updates to this post. For the moment, I will limit myself to a couple of observations:

• Recent work in Y-chromosome phylogeny has established a fairly disjoint division within haplogroup R1a1a; in particular the R-Z93 subhaplogroup seems to abound in the "Asian steppe", as well as Asia in general, while being generally absent in Europe; the R1a1a and Subclades Y-DNA Project is keeping track of new developments in this field.
• My own research on autosomal DNA, suggests the confluence of two "streams" of ancestry onto the steppe: a west-to-east stream emanating from eastern Europe, and associated with the Atlantic_Baltic (K7b) or North_European (K12b) ancestral component; as well as a West_Asian (K7b) or Caucasus/Gedrosia (K12b) component emanating from the highland regions south of the Caspian and south of the steppe (the traditional Silk Road territory).

These lines of evidence certainly appear to be consistent with a "multiregional" model of early mobile pastoralism. In particular they testify to the non-uniformity of ancestry of steppe groups and are inconsistent with their derivation from a single source. The picture is further complicated by the historical movements of nomads across the steppe (including Scytho-Sarmatian type people as well as Turkic-Mongolian ones). Charting the emergence of steppe populations will require a great deal of sleuthing in the genomes of modern steppe inhabitants, as well as a great deal of work on ancient DNA.

In any case, the new paper by Michael Frachetti will provide important new insight to all those who seek to understand "what actually happened" in Eurasian prehistory, and it is a very welcome addition to the ongoing debate.

Current Anthropology Vol. 53, No. 1, February 2012

Multiregional Emergence of Mobile Pastoralism and Nonuniform Institutional Complexity across Eurasia

Michael D. Frachetti


Abstract

In this article I present a new archaeological synthesis concerning the earliest formation of mobile pastoralist economies across central Eurasia. I argue that Eurasian steppe pastoralism developed along distinct local trajectories in the western, central, and (south)eastern steppe, sparking the development of regional networks of interaction in the late fourth and third millennia BC. The “Inner Asian Mountain Corridor” exemplifies the relationship between such incipient regional networks and the process of economic change in the eastern steppe territory. The diverse regional innovations, technologies, and ideologies evident across Eurasia in the mid-third millennium BC are cast as the building blocks of a unique political economy shaped by “nonuniform” institutional alignments among steppe populations throughout the second millennium BC. This theoretical model illustrates how regional channels of interaction between distinct societies positioned Eurasian mobile pastoralists as key players in wide-scale institutional developments among traditionally conceived “core” civilizations while also enabling them to remain strategically independent and small-scale in terms of their own sociopolitical organization. The development of nonuniform institutional complexity among Eurasian pastoralists demonstrates a unique political and economic structure applicable to societies whose variable political and territorial scales are inconsistent with commonly understood evolutionary or corporate sociopolitical typologies such as chiefdoms, states, or empires.

Link

Y-chromosomes of Marsh Arabs

What do the Marsh Arabs have to do with ancient Sumer? Nothing that can be determined on the basis of this data. There are plenty of ancient Sumerian skulls, so how about we study them directly?

As far as I can see, the only link between Marsh Arabs and Sumerians presented in this paper comes from dating Y-STR variation of their major J1-Page08 group using the evolutionary mutation rate, with a divergence time of 4.5 +/- 2.6 ky. Even if that mutation rate was correct (it is not) and the assumptions on which the confidence interval are based were exhaustive (they are not), we still have +/- 2.6 ky leeway to deal with, which spans not only the Sumerians but plenty more besides.

Not to mention that the evolutionary mutation rate is wrongly applied to every case under the sun, and that Y-STR based age estimation in general has been conclusively shown to be a rather futile exercise.

Nonetheless, the paper does have value in demonstrating the paucity of J2 and R1 in the Marsh Arabs compared to the more cosmopolitan general Iraqi population:

Different from the Iraqi control sample, the Marsh Arab gene pool displays a very scarce input from the northern Middle East (Hgs J2-M172 and derivatives, G-M201 and E-M123), virtually lacks western Eurasian (Hgs R1-M17, R1-M412 and R1-L23) and sub-Saharan African (Hg E-M2) contributions.

Rather than "Sumerian", it seems that the Marsh Arabs have rather preserved a more pristine Semitic patrilineal gene pool compared to the cosmopolitan Iraqi samples that have absorbed pre-Arab and pre-Semitic population elements.

BMC Evolutionary Biology 2011, 11:288doi:10.1186/1471-2148-11-288

In search of the genetic footprints of Sumerians: a survey of Y-chromosome and mtDNA variation in the Marsh Arabs of Iraq.

Nadia Al-Zahery et al.

Abstract (provisional)

Background
For millennia, the southern part of the Mesopotamia has been a wetland region generated by the Tigris and Euphrates rivers before flowing into the Gulf. This area has been occupied by human communities since ancient times and the present-day inhabitants, the Marsh Arabs, are considered the population with the strongest link to ancient Sumerians. Popular tradition, however, considers the Marsh Arabs as a foreign group, of unknown origin, which arrived in the marshlands when the rearing of water buffalo was introduced to the region.

Results
To shed some light on the paternal and maternal origin of this population, Y chromosome and mitochondrial DNA (mtDNA) variation was surveyed in 143 Marsh Arabs and in a large sample of Iraqi controls. Analyses of the haplogroups and sub-haplogroups observed in the Marsh Arabs revealed a prevalent autochthonous Middle Eastern component for both male and female gene pools, with weak South-West Asian and African contributions, more evident in mtDNA. A higher male than female homogeneity is characteristic of the Marsh Arab gene pool, likely due to a strong male genetic drift determined by socio-cultural factors (patrilocality, polygamy, unequal male and female migration rates).

Conclusions
Evidence of genetic stratification ascribable to the Sumerian development was provided by the Y-chromosome data where the J1-Page08 branch reveals a local expansion, almost contemporary with the Sumerian City State period that characterized Southern Mesopotamia. On the other hand, a more ancient background shared with to Northern Mesopotamia is revealed by the less represented Y-chromosome lineage J1-M267*. Overall our results indicate that the introduction of water buffalo breeding and rice farming, most likely from the Indian sub-continent, only marginally affected the gene pool of autochthonous people of the region. Furthermore, a prevalent Middle Eastern ancestry of the modern population of the marshes of southern Iraq implies that if the Marsh Arabs are descendants of the ancient Sumerians, also the Sumerians were most likely autochthonous and not of Indian or South Asian ancestry.

Link

R1b founder effect in Central and Western Europe

Post will be updated after I read the paper. (Last Update: Aug. 29)

UPDATE I:

From the paper:

The ages of various haplogroups in populations were estimated using the
methodology described by Zhivotovsky et al,30 modified according to Sengupta
et al,10 using the evolutionary effective mutation rate of 6.9 x 10^-4 per 25 years.
The accuracy and appropriateness of this mutation rate has been independently
confirmed in several deep-rooted pedigrees of the Hutterites.

Of course readers of the blog are aware that I disagree with the use of the evolutionary rate. My comments on the Hutterites paper will be posted separately after I read it. I will simply say that there are numerous cases where the use of the genealogical rate makes better sense of the evidence than use of the "evolutionary" rate. Off the top of my head, the genealogical rate harmonizes with the Genghis Khan cluster, the expansion of Na-Dene speakers into the Americas, the expansion of Balto-Slavic, the Bronze Age spread of Semitic speakers, in accordance with the linguistic evidence, the expansion of Bantu in Angola, more recent British surnames, the formation of Arabian kingdoms, Greek colonization of Sicily, and the Bronze Age origin of Indo-Aryans and Finno-Ugrians (and I skipped a few).

UPDATE II (Aug 26):

Here is the phylogeny of R-M207 from the paper. For reference, the R-M207 page from ISOGG.


UPDATE III (Aug 26):

Going through the material in this paper in a systematic manner is not easy, so I will probably do a potpourri of updates covering various topics of interest.



As noted in the other recent paper, and shown in the above Figure from the current one, R-U106 peaks in northern Europe. Its frequency (including the R-U198 sublineage) is 36.8% in the Netherlands, 20.9% in Germany and Austria, 18.2% in Denmark, 18.2% in England, 12.6% in Switzerland, 7.5% in France, 6.1% in Ireland, 5.9% in Poland, 5.6% in north Italy 4.4% in Czech Republic and Slovakia, 3.5% in Hungary, 4.8% in Estonia, 4.3% in south Sweden, 2.5% in Spain and Portugal, 1.3% in eastern Slavs, 0.8% in south Italy, 0.6% in Balkan Slavs, 0.5% in Greeks (i.e. 2 of 193 Cretans, and no mainland Greeks), 0.4% in Turks, 0% in Middle East.

The age of R-U106 is estimated by the authors as 8.7ky BP, which translates to about 2.5ky BP with the germline rate. The existence of R-U106 as a major lineage within the Germanic group is self-evident, as Germanic populations have a higher frequency against all their neighbors (Romance, Irish, Slavs, Finns). Indeed, highest frequencies are attained in the Germanic countries, followed by countries where Germanic speakers are known to have settled in large numbers but to have ultimately been absorbed or fled (such as Ireland, north Italy, and the lands of the Austro-Hungarian empire). South Italy, the Balkans, and West Asia are areas of the world where no Germanic settlement of any importance is attested, and correspondingly R-U106 shrinks to near-zero.

UPDATE IV (Aug 26):

Another informative lineage, as noted in the other recent paper as well is R-U152:


Of interest is the fact that while R-U152 has a clear French-Italian center of weight, the locations exhibiting highest STR variance are Germany and Slovakia, i.e., Central Europe. My guess is that R-U152 originated in Central Europe spreading to the west and south, perhaps with Italo-Celtic speakers or some subset thereof. In its home territory of Central Europe, its frequency decreased by the introduction of the Germanic and Slavic speaking elements which dominate the region.

Irrespective of what the ultimate origin of R-U152 is, it provides us with a good diagnostic marker for population movements out of the French-Italian area. In Italy for example it is noted at 26.6% for the north and 10.5% in the south. It would be extremely interesting to see its occurrence in Balkan Vlachs, as this would confirm/disprove the Italian component in their origin. However, R-U152 occurs in 7.3% of Cretans, suggesting introgression Y-chromosomes of North Italian (Venetian) origin, from the 4-century period of Venetian rule of the island. It also occurs in 4.1% of Greeks, where it might come from any period since the Roman annexation of the Hellenistic states to the Vlachs. However, its presence at only 1.8% of Romanians makes a large Italian contribution to the Romanian population unlikely. Balkan R-U152 chromosomes should be better resolved to determine when they arrived from the northwest.

The paucity of R-U152 in Turks (0.6%) make tales of wandering Galatians less likely to be true. There is no doubt that Galatians settled in Anatolia, but they were probably so few in numbers that they did not permanently alter the population. Knowledgeable readers should chime in about the Lebanese Christian R1b which was posited as a signature of the Crusades a couple of years ago, and its position in the phylogeny.

UPDATE V (Aug 26):

The most commong R1b subgroup in Europe is R-M269 and the most common subgroup is R-L23 which encompasses the vast majority of European R-M269 chromosomes. It is interesting to see where R-M269(xL23) is concentrated. In Europe I see cases in Germany, Switzerland, Slovenia, Poland, Hungary, Russia, the Ukraine. It is most prominent, however, in the Balkans, where every population except Croatia mainland (N=108) possesses it. In the Caucasus it does not exist except in the northeast. In Turkey and Iran there is some, albeit it is not clear in which regions.

UPDATE VI (Aug 27):

The authors write with respect to haplogroup R-V88:

With the exception of rareincidences of R1b-V88 in Corsica, Sardinia13 and Southern France(Supplementary Table S4), there is nearly mutually exclusive patterning of V88 across trans-Saharan Africa vs the prominence of P297-related varieties widespread across the Caucasus, Circum-Uralic regions, Anatolia and Europe. The detection of V88 in Iran, Palestine and especially the Dead Sea, Jordan (Supplementary Table S4) provides an insight into the back to Africa migration route.

Haplogroup R-V88 has been the subject of a recent study and was associated with the migration of Chadic speakers in Africa. It is difficult to say whether or not the authors' results really provide any insight into an alleged movement of this haplogroup from Asia to Africa, as it occurs in only a single Palestinian, and a single Iranian. Neither is the higher frequency (13.7%) observed in the Amman and Dead Sea area of Jordan really evidence of its antiquity there.

Neither the aforementioned paper nor the current one presents any evidence (e.g., Y-STR variance) for any great antiquity of the Asian R-V88 with respect to the African one. Indeed, with the exception of the aforementioned Jordanian sample, R-V88 is rare in Asia, while it is widespread in African Berbers. I see no clear reason at present to think that it migrated to Africa from Asia, and not to think of it as a relic of an older, widely dispersed R1b population leading to R-V88 in Africa itself.

UPDATE VII (Aug 28):

The paper repeats the standard claims about the origin of R1b and its main sublineage R-M269 in Asia, but presents no new information that would support this claim. With the state of the evidence, I see no real reason to prefer a West Asian to a Southeastern European origin for this haplogroup.

I don't give much credence to small differences in Y-STR variance, due to the large confidence intervals associated with such estimates, and it is interesting that the authors do not present an argument from Y-STR variation about the origin of R1b, preferring to make broad statements about Mesolithic-Neolithic movements into Europe.

A study of supplementary table S2 which gives coalescent times reveals that there is no clear pattern of greater Asian diversity within haplogroup R1b or its subclades. And, while Central-Western Europe does appear to be an outgrowth of R1b rather than a place of origin (with the dominance of derived R-M412 lineages) there is nothing in the paper that would make one prefer West Asia to Southeastern Europe as a place of origin.

Personally I think the issue cannot be settled yet, but there are reasons to prefer the latter option. An Asian origin of R1b has a major parsimony hurdle: it would require a seemingly directed drang nach westen for R1b, into Europe, and into North Africa, with a paucity of R1b in the opposite direction (among Arabians and to the south and in South Asia) and a scattering of very young R-M73 and R-M269 to the east of Europe.

UPDATE VIII (Aug 29):

R-S116 shows maximum Y-STR diversity in France and Germany but maximum frequency in Iberia and the British Isles. In the latter region it is represented mainly by R-M529 with the R-M222 subclade being particularly prominent in Ireland but also North England. It would be interesting to see data for Scotland, and I do not doubt that R-M222 would be prominent there as well. R-S116 also shows signs of being a Celtic, or Celtiberian-related lineage.

European Journal of Human Genetics doi: 10.1038/ejhg.2010.146

A major Y-chromosome haplogroup R1b Holocene era founder effect in Central and Western Europe

Natalie M Myres et al.

The phylogenetic relationships of numerous branches within the core Y-chromosome haplogroup R-M207 support a West Asian origin of haplogroup R1b, its initial differentiation there followed by a rapid spread of one of its sub-clades carrying the M269 mutation to Europe. Here, we present phylogeographically resolved data for 2043 M269-derived Y-chromosomes from 118 West Asian and European populations assessed for the M412 SNP that largely separates the majority of Central and West European R1b lineages from those observed in Eastern Europe, the Circum-Uralic region, the Near East, the Caucasus and Pakistan. Within the M412 dichotomy, the major S116 sub-clade shows a frequency peak in the upper Danube basin and Paris area with declining frequency toward Italy, Iberia, Southern France and British Isles. Although this frequency pattern closely approximates the spread of the Linearbandkeramik (LBK), Neolithic culture, an advent leading to a number of pre-historic cultural developments during the past ≤10 thousand years, more complex pre-Neolithic scenarios remain possible for the L23(xM412) components in Southeast Europe and elsewhere.

Link