Monday, June 22, 2015

Improving Our Understanding of R1b in Western Europe

Maju analyzes some new data on predominantly Western European clades of Y-DNA R1b and fits that into a working hypothesis based on the larger picture of R1b distributions in West Eurasia.

In a nutshell, it appears that R1b-S116 expands into three different daughter clades in three directions from somewhere in the vicinity of central France, one to Normandy and the British Isles, one to Central Europe and the Alps, and one to the Southeast (especially Iberia).

R1b-S116, in turn, could have origins in Iberia via Southern Italy, or in Central Europe, or both in parellel routes to central France.  These way stations could in turn come from R1b-M269 in the Balkans or Highlands West Asia (e.g. including Iran).  The answer to this question (when and where this happened) is one of the hottest questions in European archaeogenetics, because it is in furtherance of the goal of determining the archaeological culture(s) which were involved in this massive genetic transmission of the dominant Y-DNA clade today in Western Europe to its current location.  To answer this question is to unveil the face of the patrilineal ancestors of most Western European men.

Maju figures that from where ever the launching point might be that the most basal R1b would have its ultimate origins in West Asia.  This seems plausible enough, with R2 centered in South Asia and Western Iran, particularly in the Indus River Valley which is mostly Pakistan today, and R1 found in Iran and much of Europe and Central Asia, as well as appearing in South Asia in close correlation with hypothetical Indo-Aryan invaders (particularly R1a).

But, the fact that R1a and R1b were both present and bifurcated on the Russian steppe in the Mesolithic era, along with ancient DNA that seems to point to the Altai is the deeper place of origin for the entire Y-DNA R clade, suggests that West Asia may have received R2 from the steppe, during the Paleolithic era, rather than the other way around, with R1a and R1b arriving later in the Copper Age or early Bronze Age by different routes.  It is hard to distinguish between the scenarios given the available data.

Analysis

Recent ancient Y-DNA studies put examples of Y-DNA R1b and R1a on the European Steppe by the early Bronze Age or even a few centuries earlier.

In particular, European hunter-gatherer populations in Russian have ancestral Y-DNA R1 to the herders and farmers who subsequently live there, suggesting that the spread of R1 to the Russian steppe was

Corded Ware people, as expected, are predominantly R1a and have origins to the North of the Yamnaya people.  The story of R1a in Europe, to quote Eurogenes, can be summed up as follows:
- Mesolithic Hunter-Gatherer from Karelia: R1a (xM198)

- Late Neolithic Corded Ware pastoralist from Germany: R1a (M198, M417, xZ282)

- Late Bronze Age Urnfielder from Germany: R1a (M198, M417, Z282, Z280).
The Corded Ware people and their successors were almost surely Indo-Europeans.

The story of R1b is complicated, however, and the question of how R1b got to Western Europe remains more cryptic.

For example, Tarim Basin mummies are R1a, but the Yamnaya people, the hunter-gatherers from the area where the Yamnaya culture emerged (the Samara Valley), and the Afansievo people are R1b. This means that the Yamnaya and Afansievo people are unlikely direct ancestors for the Tarim Basin mummies, despite the fact that the Afansievo people arrived nearby before the Tarim Basin culture arose.  It suggests a more Northerly route of arrival for the Tarim Basin people, or a later one, as the Indo-Iranians are predominantly R1a.

All ancient Bell Beaker Y-DNA to date are R1b, but the sample to date isn't very representative and is limited largely to Germany, while missing core areas where the Bell Beaker culture was found,

The modern distribution of R1b in Europe, however, is a decent fit to the range of the Bell Beaker culture (with declining frequencies in places where it was more marginal).

Importantly, as well, the Basque are very high in R1b, suggesting strongly that R1b was brought to Western Europe by people who were not Indo-European linguistically and that Indo-European languages only arrived in Western Europe with the Celts or realistically, with their predecessor Urnfield people whose language was probably the source of both the Italic and Celtic languages.  In other words, Indo-European languages arrived in Western Europe only around the time of Bronze Age collapse.  (A similar argument for an origin of Germanic languages in Northwestern Europe around the time of Bronze Age collapse is also consistent with the evidence.)

Taken to its logical conclusion, if the Yamnaya people are the forebears of the Bell Beaker and other Western European Y-DNA R1b peoples, and while the Corded Ware people are the forebears of the European R1a people, the Yamnaya people may not have been Indo-Europeans after all.  Instead, they could very well have been linguistically Vasconic (i.e. linguistically in the same family as the Basque language).

As Dienekes notes, the data tend to show that "the later steppe cultures of the Sintashta and Andronovo (putative Indo-Iranians according to some), were not a continuation of the Yamnaya-Afanasievo people," as had been widely assumed before the ancient DNA data was available. Dienekes also notes genetic continuity between modern Armenians and those of the early Bronze Age.

As Maju notes, it isn't clear if R1b took a continental route, Mediterranean route, or both, to Western Europe from an origin in Southeast Europe and/or West Asia.

Maju demurs on the timing of R1b's arrival, but suggests that Atlantic Megalithic as well as Bell Beaker were likely involved, rather than a later arrival with Bell Beaker people alone.

13 comments:

Maju said...

I don't know yet what role played the Western Neo-Chalcolithic in R1b spread. We will need much more aDNA from Western Europe in order to understand that ("molecular clocks" don't work). So I'm just considering all possible scenarios.

Anyhow, Bell Beaker is not any independent "culture", much less "people": Bell Beaker is a phenomenon, a subcultural element, maybe even just prestige items of an epoch. While it's possible that there is some BB distinctiveness in Central Europe (where maybe it served as cultural liason for some sort of social reaction to the excesses of Corded Ware Indoeuropeanization) in Western Europe that is not generally the case: instead what we see is just irregular appearance of beakers and related elements like V-perforated buttons within previous cultural contexts, often but not always Megalithic. Bell Beaker style burials (individual in pits with very specific grave goods) almost do not exist West of the Rhine, when they happen, as with the Amesbury archer, they may be related to foreigners (which the Amesbury archer was) or some very specific subcultural group.

Instead Megalithism is a much more clearly important cultural phenomenon, and may be related to demographic changes because it is often associated with the first Neolithic in many Atlantic regions (British Islands, Denmark-Scania, Galicia even), or it is associated with important cultural (and possibly demographic) changes such as the expansion of Michelsberg culture in Germany and North France (where it's known as Seine-Oise-Marne culture). So I would say that what is found in Central European individuals of so-called "Bell Beaker culture" may reflect these previous changes, which pre-date Corded Ware, rather than being a signature of BB itself.

Maju said...

As for R1a, I wouldn't even associate it nowadays with Indo-Aryan expansion. It's obvious since Underhill 2014 that R1a must have spread from Iran to both Europe and South-Central Asia and then re-expanded in these two regions independently. It's possible that much of the European R1a was re-expanded by Corded Ware but the association of Asian R1a to Kurgan cultures remain impossible to accept - it must be mostly previous to them (Neolithic most likely). I don't issue any opinion on R2 because it is not yet properly researched but judging on R1 upstream diversity (which is centered around Pakistan) and other P1 sublineages, IMO it probably arose in North India.

I have not the slightest idea how can you claim that R originates in Altai, which only harbors highly derived R1a-Z93 sublineages. Altai was probably dominated by Q1, maybe with some R* of the Mal'ta type too (but must have been minor because it is today missing and has left no legacy anywhere), until the Neolithic. Q1 is still an important Altaian lineage. All the evidence for R (and related P1) origins point to South Asia (R1 in Pakistan or nearby areas of India, R2 in North India surely, P1* in Bengal, Q probably in Iran...).

All the evidence of R1b specifically, which was the only thing I discussed, points to West Asia, with branches heading to Eastern Europe and Central Asia (but also present in West Asia), to Africa (but also to Italy) and to Western Europe, as well as some diversity of all the major subclades and some more left back in West Asia itself. Western European R1b has no relation whatsoever with Eastern European or Central Asian R1b except via West Asia. Please stop mixing things. Mixing these distinct lineages is like mixing I and J, R and Q: it's so shallow that only because it shares the first letter of the nomenclature people think that there must be some greater relation than when they do not!

andrew said...

My basic reasons for doubting that Bell Beaker was a case of predominantly cultural transmission or a thin cultural superstrate, rather than an instance of mass demographic shift via folk migration and expansion of the migrating population, as you suggest in your first comment boils down to the limited ancient DNA evidence we have about the Atlantic Megalithic people prior to the advent of the Eneolithic era and the Bell Beaker phenomena.

Western European clades or for that matter, any clades of Y-DNA R, are entirely absent from the ancient DNA region in the smattering of Western European Megalithic or Western European Paleolithic/Mesolithic ancient DNA samples that we have, prior to Bell Beaker archaeology, despite being predominant by Iron Age at the latest, and probably sooner. Admittedly, our sample size of ancient DNA is only one the order of a dozen or two right now. But, it shouldn't take much of a sample size to find a Y-DNA clade that is now found in more than 75% of Western European men.

There are simply no other good candidates in the right place at the right time to be a source of this Y-DNA, earlier in prehistory, to explain the dramatic shift in Western European population genetics.

This demic impact may have been male dominated and had a much milder impact on the mtDNA mix, although the surge in mtDNA H in most of Western Europe suggests, at least, significant folk migration of whole families once Southern European women are integrated into the new culture at the time of ethnogenesis, but cultural transmission of a culture with little demographic impact wouldn't leave Western European R1b men any significant base to expand from.

andrew said...

Then again, an an R1b-V88 individual in El Trocs, Spain ca. 5100 BCE just confirmed this year arguably shifts the analysis somewhat, although I am still inclined to see this individual as a outlier traveler from the same population that became the linguistically Chadic people of the Sahel (perhaps via Gibralter rather than Southern Europe), and not a direct ancestor of later R1b individuals in Western Europe including Spain.

andrew said...

It looks like my take in the previous comment is pretty widely shared, for reasons similar to those that drive my intuition on the issue.

Maju said...

Interesting the El Trocs data. I was not aware. Here we are again before a typical pre-judged "Neolithic" lineage, such as E1b-V13, G2a or even surely I2a1b. Like the latter, R1b-V88 is common in Sardinia of all places, what underlines its "Neolithic" character. It's very straightforward.

And nope: it has only a remote relation (via West Asia) with Chadic R1b-V88. Again lacking a proper understanding of the geostructure of the lineage causes confusion by shallowness of the kind: "R1b-V88! Chadic!" (nope: Sardinian), "R1b(xM269) in Volga! Same as R1b-L11 in West Europe!" (nope: a different "Asian" clade). Shallow happy-nippiness is what I see rather than sound intuition.

As for the limited knowledge we have of Megalithic aDNA, it's not just prior to Bell Beaker: we also have no knowledge of Bell Beaker aDNA outside of Eastern Germany, so all the extrapolations are futile, more so when Bell Beaker was neither "a culture" nor much less "a people" but a subcultural phenomenon of some complexity. In general we have very limited knowledge of Western European aDNA (barring a few Western Mediterranean early farmers that are of little use) and the little we know about Atlantic aDNA (Basque, North French, Portuguese) shows high frequencies of modern-like mtDNA H (and general pool), so inferring high frequencies of modern-like Y-DNA R1b-S116 is not out of the question at all but rather a logical expectation.

"Admittedly, our sample size of ancient DNA is only one the order of a dozen or two right now".

Our Atlantic Neolithic Y-DNA sample is exactly zero!

"But, it shouldn't take much of a sample size to find a Y-DNA clade that is now found in more than 75% of Western European men".

I hope so. I also hope that someone, some day, bothers doing that research.

BTW, it's not quite true that R1b-L11 is found in 75% of Western European men, the figure is quite lower, excepted some pockets like Basques, Irish and Welsh who are around 90%, or Scots (~75%). It's rather 50-70%, depending on region.

"But, it shouldn't take much of a sample size to find a Y-DNA clade that is now found in more than 75% of Western European men".

We don't know if there was any shift: we just don't have any data to judge. There is ZERO Atlantic Neolithic ancient Y-DNA, I insist.

andrew said...

The timing of the El Trocs remains is about the same as the time of Chadic ethnogenesis.

Maju said...

I wouldn't discard a connection. Chadic are Afroasiatic but there is a complex and ill-understood intermingling between Afroasiatic and Nilo-Saharan and I suspect that Vasconic (which I identify with Thessaly-rooted Neolithic languages) has at least partial Nubian (Nilo-Saharan) roots, also from roughly Sudan.

However I do not see any clear evidence of R1b-V88 being of African, rather than West Asian, origin. The matter still requires much research admittedly.

What I do not accept in any case is any shallow, simplistic and fallacious relation between R1b-V88 (a Mediterranean lineage in Europe, parallel to E1b-V13, G2a, I2a1b or J2b) and R1b-L11.

andrew said...

"Vasconic (which I identify with Thessaly-rooted Neolithic languages)"

Interesting. This is one of the clearest positions I recall seeing you take on the origins of the Vasconic language family.

Maju said...

Strange because I've been at it for some time. Basically if there is Vasconic substrate in Italy and the Balcans (and there is IMO), it cannot be Magdalenian (nor merely Megalithic), so it must be Neolithic.

I also suspect quite strongly that Vasconic is doubly related to:

1. Indoeuropean: there are many old shared Basque-PIE cognates such as "hartz", bear, or "hauts", dust or ash, and they are with PIE not with this or that derived language (that also but that would be Vasconic substrate or adstrate, a totally different kind of interaction in time and space). See also this quadruple mass lexical comparison

2. Nubian languages (Nilo-Saharan): a recent discovery, see this other mass lexical comparison

So possibly Vasconic arose as a hybrid language when a Nubian-like core (Neolithic migrants from Palestine?) met a Indoeuropean-like one (Paleoeuropean peoples from the Balcans or maybe other West Asian farmers?)

Said that... consider and discuss. So far it's just a working hypothesis.

Blasonario Cremonese said...

Maju, your post is interesting: Neolithic migrants that met a Indoeuropean population...

And, if I could ask, what are - according to your view - the Y-Dna inherited by this hybrid population from that hypothetical Neolithic population, and what are the Y-DNA inherited from that Indoeuropean one?

Maju said...

@Blasonario:

1. It would not be Indoeuropean-speakers but something ante-pre-proto-IE, what Roslyn Frank calls Paleoeuropean, whose key shared words are often better preserved in Basque than in modern IE (although fit well with reconstructed PIE). Indoeuropean can only be identified since the steppe Late Neolithic or Chalcolithic, not earlier: its origins beyond that are still to be unraveled.

2. I do not have a well finished theory, because data is missing and we can only await till it is researched. I mean for example Atlantic European and West Asian ancient DNA, certainly Y-DNA (as you ask) but also autosomal DNA (mtDNA too but we already know something of that).

This is critical to answer your question. Right now I could equally lean for an UP or a Neolithic spread of European R1b-M269 (and especially its main sublineages S116 and U106), what I do not accept as valid is the mad conjecture floating around and being waved as quasi-ideological banner by a bunch of Indoeuropeanist fanatics of this lineage originating in Eastern Europe and spread "recently" by Indoeuropean migrants: there's just no evidence and there is no allowance for it with the available data.

The data shows that some R1a is associated with Corded Ware and hence with the NW branch of Indoeuropean expansion and that G2a, E1b-V13, some I2 and now also J2 are associated with the Neolithic expansion in Europe. That's pretty much in agreement with "old school" readings of modern Y-DNA frequencies and geography.

Maju said...

PS: ... "of modern Y-DNA frequencies and geography"... and phylogenetically structured diversity too, of course.