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Monday, December 1, 2014

Latest speculation about R1b


The events outlined on the map below purportedly took place during the Neolithic. Perhaps, but we simply don't have enough ancient DNA yet to be sure of much as far as the spread of R1b into Europe is concerned. Nevertheless, I'm just wondering, what's the chance that R1b was established in Western and/or Central Europe before the very late Neolithic, which is what this paper is suggesting, as far as I can tell, but has not shown up in any of the relevant ancient samples to date?

Citation...

Hovhannisyan et al.: Different waves and directions of Neolithic migrations in the Armenian Highland. Investigative Genetics 2014 5:15. doi:10.1186/s13323-014-0015-6

See also...

The story of R1b: it's complicated

154 comments:

Grey said...

I like that they show an overland and a sea route - just cos.

As to the question - I guess it would depend on whether they were talking about the coppery end of the neolithic spectrum or the flinty end.

Tone said...

I've always wondered if R1B was hiding out in the far Northwest of Europe as part of a boreal, HG group sharing an ANE cline with R1A groups in the East The I2 Loschbour HGs (related to middle-eastern J) were south of these R Hunter Gatherers who were descended from mammoth hunters and related to Amerindian Q.

Maybe R1B first exploded out of the Northwest with the Megalith builders. But I know nothing. Just speculating. . .

Chad Rohlfsen said...

I think they point to Varna, Cotafeni, onto Baden, remedello, and Iberia beaker. Earlier than that doesn't make sense. Right now, Remedello prior to 2400 and Iberia 2200, doesn't add up.

Chad Rohlfsen said...

More of that stelae, dagger stuff.

Chad Rohlfsen said...

No R1b in 24 southern French samples, circa 3000bce, nullifies anything prior to that imop.

Chad Rohlfsen said...

https://www.academia.edu/2452426/Metal_in_South-Eastern_and_Central_Europe_between_4500_and_2900_BCE

https://www.academia.edu/1112588/The_Relation_between_Copper_and_Flint_Daggers_in_Chalcolithic_Italy

Krefter said...

As of far we know that the 2600BC R1b dudes from east Germany were of mostly Yamna decent and were very similar to modern central-north Europeans.

Was R1b(L11) somewhere else in west Europe at that time, we don't know. But it's pretty important to get autosomal info from people who had it nearly 5,000 years ago.

Chad Rohlfsen said...

If they want to put R1b in West Asia, it had better be coming from the Caucasus or Western Iran. It can't be any closer to the Anatolian farmers. That is the only way that they can tie it to people like CO1 and Oetzi, with 4-7% ANE. As an R lineage, it should be tied to a decent amount of ANE. The huge jump in WHG in heavy R1b areas kind of kills the West Asia hypothesis though. Unless the south Caucasus was WHG/ANE, with little EEF spill-over prior to 5000BCE, and prior to migrating to the Balkans. That seems kind of unlikely though.

bellbeakerblogger said...

Another map that cuts off North Africa, beautiful.

Anyway, the only way to make R1b clinal is to make its progress Early Neolithic. Of course then we'd have to disregard the archaeology and the pile of genetic studies from Neolithic Europe.

Anyhow, cheers

http://bellbeakerblogger.blogspot.com/2014/11/apparent-cline-vs-real-cline.html

Krefter said...

"If they want to put R1b in West Asia, it had better be coming from the Caucasus or Western Iran."

We can't say R1b had to of been here or there, as a fact. Because there are autosomally radically different people carrying R1b today.

We should all agree that the history of R1b in all of Eurasia, not just west Europe, is a mystery. There is no conclusive answer as to whether R1b-L11 was spread by a non-Yamna-type people in west Europe or was spread by Yamna-type people. Obviously though Bell Beaker has something to do with it.

spagetiMeatball said...

When do most of the R1b carried by European men today in coalesce? I've seen eupedia mention sometime 5,000 years ago? Doesn't that suggest the late neolithic?

Alberto said...

How many Mesolithic (or earlier) Y DNA samples do we have from Western Europe?

Two?

Loschbour (I2a) and La-Brana-1 (C6). And Loschbour is borderline "Western" Europe.

Too many theories based on two little samples (and besides, not a single R1b sample from pre or early Neolithic Near East or Eastern Europe).

The guessing game is fun, but it's just that.

Davidski said...

Yeah, from memory, based on full Y-chromosome sequences, most of the main R1b subclades aren't older than the Bronze Age, or even the Iron Age.

In any case, it's a pity we didn't get any male genomes from Gamba et al. dating to the early Bronze Age, like BR1, which belonged to the Mako Culture. Mako is apparently a type of the Vucedol Culture, which has been described as basically a Yamnaya-influenced Bell Beaker Culture. If this is true, then BR1 might be a good example of one of the populations that moved into Western Europe from the Carpathian Basin, potentially taking with it that Northeastern European-like admixture that BR1 had as well as R1b.

Krefter said...

There are many more than 2 Y DNA samples from ancient western Europe. There are well over 30.

Richard Rocca said...

Given that the two Iron Age R1b Britons are the best proxy for Bell Beaker to date, and they each have ~0% East Mediterranean ancestry, the notion that R1b would have arrived from anywhere near the Levant is flawed IMO. As others have stated, we have a couple of dozen Western European Y-DNA from the Late Neolithic and none of them are R1b+. In fact, that Otzi and the Late Baden sample are extremely close to other Neolithic samples makes me think that a sort of Blitzkrieg of sorts led R1b to Western Europe, and the near identical modal for all L11+ lineages seems to support that.

Alberto said...

@Krefter

It depends on what you mean by "ancient" and what you mean by "Western Europe".

I was clear about meaning Paleolithic and Mesolithic (i.e, before Neolithic migrations), and by Western Europe I mean areas where nowadays R1b is clearly dominant (Iberian peninsula, British islands and France, mostly).

With these requirements, to my knowledge we have 2 samples. If you know about others, would you point me to them?

Davidski said...

A Mesolithic presence of R1b in Iberia doesn't make any sense, unless we're talking about an R1b subclade that is now extinct. Even the early Neolithic is a massive stretch.

I was actually thinking not long ago that R1b might have moved into Europe from the southern Balkans/western Anatolia during the middle Neolithic with the Secondary Products Revolution. But it's obvious now that the initial stages of the SPR were facilitated in Europe by people like NE1, NE7 and Oetzi, so that looks like a dead end.

I'm now betting that BR1 would've carried R1b if it was a male sample, and I suspect that Central European Beakers, like those from Kromsdorf, will be almost indistinguishable from BR1 in terms of genome-wide genetic structure.

But even if that's true, it's now clear that at some point there was also an influx into Western Europe of people similar to IR1. They may have also brought R1b with them.

Helgenes50 said...

The West Asian has certainly brought by the second, this component is not present in the bronze samples.

Chad Rohlfsen said...

I wouldn't be surprised to see a couple r1bs in late Baden, around CO1's time.

Chad Rohlfsen said...

I don't think Vucedol expansion has to do with R1b. R1b is almost nothing in that part of Europe. It like 8% of the ydna in Croatia.

ZeGrammarNazi said...

Chad, the link is that Mako is an extension of Vucedol, and the BR1 sample from the Mako culture plots between to the modern day French and Orcadians.

There is certainly a connection between Mako and modern day Western Europeans, and part of that connection could be due to Y-DNA R1b, but we just don't have enough data right now to make a clear association between the two.

The archaeological relationship between Yamnaya, Bell Beaker, Vucdeol, Mako is there, we just need more ancient DNA from these places to understand that relationship better.

Chad Rohlfsen said...

I'm saying that r1b didn't come from vucedol, not that it didn't mix with them, in the Carpathian Basin or Austria. Bell Beaker is seen as part Vucedol.

ZeGrammarNazi said...

Where do you believe the majority of R1b in Europe came from or expanded from, Chad?

Chad Rohlfsen said...

Probably the Carpathian basin via Kemi Oba and Pontic Kurgan related groups, then to Austria and Germany, where the Chalcolithic population boom happened.

Chad Rohlfsen said...

I think that the Beaker samples will be almost identical to the Iron Age Celts. IR1 had low IBD sharing with the Brits. I think that 45/40/15, is what we will see for EEF, WHG, and ANE.

ZeGrammarNazi said...

That isn't too far from what is being proposed with a link to the Vucedol/Mako cultures.

Vucedol is thought to be an Indo-European culture by many. Gimbutas modeled Bell Beaker as a mixture of Vucedol and Yamnaya cultures, and being that Kemi Oba predates Vucedol by ~700 years, it is not impossible that the Yamnaya influence found in Late Neolithic/Early Bronze Age samples from Europe is due to migrants from Kemi Oba moving into Vucedol territory.

Gimbutas models BB as Yamnaya + Vucedol, Vucedol gave rise to Mako, and the one Mako individual we have DNA from looks like a modern day Western European. I believe these connections are more than coincidences, although figuring out where exactly R1b was in all this madness is not really possible with the data we have.

It could very well be that R1b males were part of the Kemi Oba culture and that they mixed with Vucedol or related groups as they moved further West into Europe. I certainly believe Western Europeans can trace most of their autosomal roots to Early Bronze Age cultures such as Mako, but sex-specific markers are a different story.

ZeGrammarNazi said...

Chad, if you look at the plot David posted of the handwritten and drawn notes from the ASHG 2014 meeting, it appears you could model the LNE/EBA samples as a group that was ~60 MNE and 40~ WHG that then received ~50% admixture from Yamnaya. Then again, it would probably be possible to model LNE/EBA as ~55% MNE and ~45% EHG too.

Krefter said...

Alberto there are over 10 Mesolithic Y DNA samples from Europe, and 1 Upper Palaeolithic.

Krefter said...

Davidski, you think people like IRI went all over Europe(even Baltic region) during the Iron age? Couldn't he had just been from an eastern immigrant population like the Huns, who made little genetic impact? If BR1 represents the people who brought R1b-L11 to west Europe, then who brought the extra ANE?

la señora bibiloni said...

As far as I recall, the oldest R1b found so far is associated with Bell Beaker in Germany around 2600 BC. Until some ancient R1b is found in a Yamna context, I wouldn't put my money on Gimbutas' IE expansion to the west theory. Waiting for Reich's paper...

Davidski said...

IR1 isn't a Hun. Most of his ancestors were probably Cimmerians from the Kuban Steppe. It seems that a lot of them moved west during the Iron Age, setting up various Thraco-Cimmerian cultures in the Carpathian Basin and Balkans.

Iron Age people similar to IR1 certainly had an impact on the modern European gene pool. You can see that in the video clips I posted recently.

So it's a mistake to assume that the present levels of ANE in Europe, or even Western Europe, are overwhelmingly from a single event. There were different waves of population movements to the west, mostly from the Carpathian Basin, but also via the North European Plain, where Corded Ware and Bell Beakers mixed. Also, don't forget isolation-by-distance, which can smooth out ancestral components over time if there's regular mixing across a wide area, even if it's very gradual.

All we can say with any confidence at the moment is that the ANE present across most of Europe today came from various Indo-European groups that pushed west during the Bronze and Iron Ages.

Chad Rohlfsen said...

BR1 is rather late (1k years after Kemi Oba, 600 years after German Beakers, 300 years after Beaker folk are in Britain), plus it's at the exact time that we see Aegean and Balkan influence into the Carpathian Basin. I think we will see that most ANE in Western Europe can be traced to the Bell Beaker expansions.

Chad Rohlfsen said...

R1b L23 is very interesting. It is usually dated to 3500BCE, around the time that Cotafeni started, but its highest diversity is found in Pashtuns, ASFAIK.

Chad Rohlfsen said...

From Armenian genetic projects...

In addition, to determine whether genetic differences among these four Armenian populations are the result of differential affinities to populations of known historical influence in Armenia, we utilize 27 biogeographically targeted reference populations for phylogenetic and admixture analyses. From these examinations, we find that while close genetic affiliations exist between the two easternmost Armenian groups analyzed, Ararat Valley and Gardman, the remaining two populations display substantial distinctions. In particular, Sasun is distinguished by evidence for genetic contributions from Turkey, while a stronger Balkan component is detected in Lake Van, potentially suggestive of remnant genetic influences from ancient Greek and Phrygian populations in this region."

The more isolated southern and eastern regions have high frequencies of a microsatellite defined cluster within haplogroup 1 that is centred on a modal haplotype one step removed from the Atlantic Modal Haplotype, the centre of a cluster found at high frequencies in England, Friesland and Atlantic populations

In short, the heavy R1b areas show Balkan influence... I think, from both Anatolian and later Armenian and Greek speakers.

Chad Rohlfsen said...

Another interesting bit of info there... maybe someone with knowledge on G, can weigh in.

"[...] First, we calculated haplogroup diversity using data in Supplementary Table S1 for the 52 instances when total population sample size exceeded 50 individuals and Z5 hg G chromosomes were observed. Then we applied a 10% overall hg G frequency threshold and the additional specification that both haplogroup G1 and G2 lineages also be present. In the ten remaining populations, haplogroup diversity spanned [...] to highs of 0.88 in Azeris (Iran) and 0.89 in eastern Anatolia and 0.90 in Armenia. We estimate that the geographic origin of hg G plausibly locates somewhere nearby eastern Anatolia, Armenia or western Iran. [...]"

If that's true, R1b can't be there before the Chalcolithic.

Davidski said...

Mako is a late derivative of Vucedol.

So BR1 had ancestors who were in the Carpathian Basin much earlier than the formation of Mako, and they probably caused the shift in genetic structure in the region from CO1-like to BR1-like.

Then came the minor influence from the south, creating people like BR2.

Alberto said...

@Krefter: "Alberto there are over 10 Mesolithic Y DNA samples from Europe, and 1 Upper Palaeolithic. "

I stressed the word "Western", and further explained I refer to the territories where nowadays R1b is highly prevalent, mostly British Islands, France and Iberia. Which are the other 8 Mesolithic Y-DNA samples from this are you know about?

The point is to know if R1b was in Western Europe before the Neolithic or if it arrived during or after the Neolithic. We have 2 samples that I know of from before the Neolithic, and none are R1b. But that's just not enough to jump to conclusions and make theories. We need more data, and till then all is just a guessing game.

ZeGrammarNazi said...

Alberto, we see known European hunter-gatherer y-dna lineages (I2, C6) appear in Neolithic sites from Southwestern France (Treilles) and Northern France (Dolmen of La Pierre Fritte) and Hungary, but we haven't seen any R1b yet.

As you said, we still need more samples before anything is certain, but an early or mid-Neolithic presence of R1b in Western Europe would be a surprise for many at this point; myself included.



Grey said...

Tone

"I've always wondered if R1B was hiding out in the far Northwest of Europe"

I'd like that to be true as it's so neat: R1 split in two by the ice followed by a repopulation after the LGM but... no dna, at least so far and at the same time there is Beaker dna.

As to where else, dunno but given that the R1b expansion in western Europe seems so explosive - I assume because they had some way of exploiting an under-used niche - then there might not have been that many of them at the start of the process which might be a reverse clue i.e. a clue to who they weren't.

Grey said...

I did read up on this once and am about to check again so not just laziness but is there evidence of stuff coming *out* of the fertile crescent before farming or only from the regions *around* it?

(i.e. the coast and surrounding highlands)

Chad Rohlfsen said...

Alberto,
No way does R1b enter Europe before the Chalcolithic. Not a single sample prior to the Kromsdorf Beakers, has R1b.

We have many Mesolithic samples and tons of Neolithic samples, no R1b. It can't be in Western Europe, or the Near East.

Loschbour, LaBrana, Koros1, Kostenki, Motala hunters, Pitted Ware hunters, no R1b.

Just go here...
http://www.ancestraljourneys.org/adnaintro.shtml

Krefter said...

"Loschbour, LaBrana, Koros1, Kostenki, Motala hunters, Pitted Ware hunters, no R1b.

Just go here...
http://www.ancestraljourneys.org/adnaintro.shtml"

I'm almost done breaking down every Upper Palaeolithic-bronze age mtDNA sample on that site. I'll have google maps, spreadsheets, and documents to present the data in a much more accurate, informative, and convenient way than ancestral journeys.

There are alot of miss calls from the original studies, which are posted on ancestral journeys.

Now that I have a predicting-base for most west Eurasian mtDNA hgs, I can break down 20-30 samples in ~1 hour.

By the time the Reich-Laz paper comes out we'll have a very detailed reference for other ancient(and some modern) west Eurasian mtDNA-Y DNA. It'll be easy to know how the ancient people they sample relate to modern and ancient west Eurasians in terms of mtDNA-Y DNA.

I'm tired of having to look back at ancestraljourneys and Eupedia whenever I want to compare modern-ancient mtDNA-Y DNA. If a bunch of people post detailed info from studies it'll help everyone.

Chad Rohlfsen said...

David,
Didn't that Wilde study on the Pontic Kurgan group from Bulgaria, Romania, and Ukraine, basically have them about identical to modern Ukrainians, in terms of aDNA?

Krefter said...

The ancient steppe mtDNA samples are of low coverage, so if they're compared to any modern Euro pop at that coverage they'll seem similar. mtDNA relatedness tests, can't account for drift and low coverage(two lineages separated by 40,000 years, can be perfect HV1 matches). There's no easy and accurate way to compare two pops mtDNA if they're of low coverage(pretty much all ancient mtDNA is).

But anyways, they did have around 50% HG mtDNA and 50% near eastern mtDNA. Which is consistent with them fitting as 50% EHG and 50% Armenian. Modern north Euros might have only 15-20% HG mtDNA, but over 50% HG blood because EEF's HG blood was mostly expressed via Y DNA, which was mostly replaced. Also, IE immigrants blood may be mostly expressed via Y DNA, and so HG mtDNA only rose from 5-10% to 15-20%.

andrew said...

There is R1b in Georgia and Armenia, but given the percentages and distributions and what we know about the history, there is a good case to be made that R1b was received in the region sometime after G was predominant there first.

One place where R1b does appear at the right time and in about the right place in Minoan Crete. Western European Bell Beaker R1b and R1b in Crete may have a common origin, perhaps from Anatolia, and more deeply from Iran where the most basal R1b is found via the Southern Caucasus.

Krefter said...

It looks like all Europeans, except Basque, have post-Neolithic Middle eastern ancestry. So, copper-iron age east Europeans can't be the only source of ANE in Europe.

http://www.eupedia.com/forum/threads/30677-Shriner-et-al-(2014)-Genome-wide-genotype-and-sequence-based-reconstruction-of-the-1?p=445424#post445424

Chad Rohlfsen said...

Andrew, m269 is in Europe and Asia. M343 is most common is SE Kazakhstan.

Davidski said...

There's no evidence in that Shriner et al. study that all Europeans except Basques have post-Neolithic admixture from the Middle East.

It's based on modern DNA, so it can't tell us about direction or source of gene flow. In other words, it can't say whether Europeans have post-Neolithic Middle Eastern ancestry, or whether Europeans and Middle Easterners both have the same kind of post-Neolithic ancestry from a third source, like stuff from the steppe.

Davidski said...

Holy shit, that Shriner study is really bad.

I can't believe it got published in 2014 (almost 2015).

Krefter said...

"It's based on modern DNA, so it can't tell us about direction or source of gene flow. In other words, it can't say whether Europeans have post-Neolithic Middle Eastern ancestry, or whether Europeans and Middle Easterners both have the same kind of post-Neolithic ancestry from a third source, like stuff from the steppe."

Okay, yes I did over react and I'm jumping to quick to conclusions.

We can at least say this paper suggests post-Neolithic Middle eastern ancestry in Europe. GEdmatch admixtures say pretty much the same thing to.

Davidski said...

No, this study is not informative about direction or timeframe of gene flow.

Here, read this, it explains why...

http://biorxiv.org/content/early/2014/03/21/003517

Alberto said...

"Loschbour, LaBrana, Koros1, Kostenki, Motala hunters, Pitted Ware hunters, no R1b."

Again, only the first 2 of those are pre-Neolithic samples from WESTERN EUROPE (as in FRANCE, IBERIA or BRITISH ISLANDS).

TWO samples. 2 samples. Two single individuals in the whole western part of Europe (and I'm even including Loschbour as Western, even if it's close to the border with Central-North Europe where haplogroup I is more common).

Couldn't we just wait to have more data before making so many theories? Oh well, guessing is fun, I admit that. So let this game go on, as long as we all know it's just guessing and any theory could be proved wrong any day when more data arrives.

Chad Rohlfsen said...

Albert, look at all of the ydna from the Neolithic. Stop being rediculous.

Richard Rocca said...

We have seen in Central Europe that farmers (haplogroup G) moved in and hunter-gatherers (haplogroup I) eventually took up farming as well. There is no reason why the same would not have happend in Western Europe. You would expect that if R1b were present in some remote corner of Western Europe, it would have manifested itself in several Western European Neolithic and Copper Age samples, and to date it hasn't. For that reason, the low number of Mesolithic samples is not as problematic when we already have post-Mesolithic samples. Now, one can make the argument that R1b hunter-gatherers were part of some closed society, and didn't migrate out of Western Europe until the late Copper Age, but members of a closed society would not have learned how to plow a field, smelt Copper, tame a wild horse, etc. Then we have the east to west splits that are obvious in the phylogenetic tree of R1b. No, there is really no likely scenario where R1b is held up in some magical hunter-gatherer wonderland in Western Europe only to expand at exactly the same time as R1a but from a completely opposite direction, regardless of how few Mesolithic samples there are to date.

Alberto said...

He Chad, nice to meet you. You look like a nice guy.

Could you tell me what's ridiculous about saying and repeating to deaf ears that there are only two samples of pre-neolithic Y-DNA from Western Europe? It's only a fact that someone was having a hard time admitting.

What do you mean by "look at all the Neolithic DNA"? Do you mean that it mostly proves that R1b was not brought to Western Europe by Neolithic Farmers? I can basically agree with that, but who's discussing that?

Right now the two most likely speculations about why R1b is so prevalent in Europe are either that it was there since the Paleolithic (but we have no data to support this), or that it came after the Neolithic from the east with IE speakers (but we neither have any data to support this). The only data that we have at this point is precisely the one you point out: that Neolithic farmers didn't carry R1b, so it's unlikely that they brought it to Western Europe.

Now go back to sleep, and when you wake up next time try to be polite and to make some sense in what you say. It always helps.

Chad Rohlfsen said...

The data speaks for its self. We have what, 40 or so neolithic ydna samples. None of it is R1b. M343 is in Central Asia. M269 is most common NW of the Black Sea. L23 spreads from Mongolia to Spain. It didn't come from the west.

Grey said...

"It didn't come from the west."

I think it's possible that a relatively small group came from the east and had a population explosion in the west leading to an expansion from the west even though they originally came from the east.

(basically "hare" celtic (coastal copper hunters) and "tortoise" celtic (overland full tribes) with the hare celtic being p and tortoise q and the q eventually catching up.)

But that's just an idea.

Chad Rohlfsen said...

I don't think it's a coincidence that L23 is heavier in Northern Anatolia, Southern Italy, France, and Spain. Those are all areas of Greek colonization. Plus, Northern Anatolia has the Armenian related Hemshin. Armenia and Anatolia look like they're as much of a destination as Western Europe.

Chad Rohlfsen said...

I've seen over at Anthrogenica, someone post that all m-269 today, descends from a man whom lived around 2500BCE. Has anyone else seen this?

Chad Rohlfsen said...

Were those three Ukrainians that tested as P-25+, m-269-, from Varzari, in 2006, ever revisited?

Tone said...

Stelae People?

https://s-media-cache-ak0.pinimg.com/originals/2d/f0/f0/2df0f095523a8f15a29abaac515b3ec4.jpg

Davidski said...

My hunch is that those Stelae don't represent Yamnaya-derived migrations from the steppe to Iberia, but rather maritime links between trading posts along the Med and Black seas.

I just can't imagine fairly large groups of people pushing west overland from Ukraine to Portugal. A gradual development of trading links, followed by a Greek-like maritime colonization makes more sense. And if we're talking about Greeks, then why not posit that the ancestors of the Bell Beakers came from around the Aegean somewhere, rather than Ukraine?

But I could be wrong.

Chad Rohlfsen said...

David,
It might be both, but probably over land.

Chris Davies said...

I'm not massively clued up on Y DNA haplogroups or R1b. However has the possibility of a migration from North Africa into Iberia &/or S.France, &/or N.Italy via Sardinia/Corsica been considered as one alternative?

Chad Rohlfsen said...

Chris,
Not a chance.

Davidski said...

It has been considered, because early Bell Beakers show influences from North Africa.

http://bellbeakerblogger.blogspot.com.au/p/genetics-projects.html

But there's no direct evidence for it yet.

Chad Rohlfsen said...

How would R1b be in North Africa, prior to being in Europe? The only way is through the Near East, and their is no Neolithic R1b. It's not an option. If anything, that influence came in the form of y-DNA E.

Ebizur said...

I think one reason some people have considered a North African route of entry of R1b1a2-M269 into Europe is the prevalence of R1b1c-V88 in some parts of Africa (especially in the vicinity of Lake Chad).

However, it is now known that R1b1a2-M269 is more closely related to R1b1a1-M73, a typical Central Asian & Southern Siberian (Turko-centric?) clade, than either of those clades is related to R1b1c-V88. (Furthermore, the rare clade R1b1b-M335 has been found among Turks and Chinese-speaking Muslims, who are proximate to Turkic peoples. R1b-M343(xM335, M73, M269) also has been found in populations of the same general area.) We must trace R1b1a2-M269 back to its MRCA with R1b1a1-M73 before conjecturing about the location of the MRCA of R1b1a-P297/L320 and R1b1c-V88.

Chad Rohlfsen said...

m-269 is supposedly dated to 4000BCE, m-73 to 5000BCE. The thing though, is apparently the 67 marker test has all modern m-269* dated to only 2500BCE. So, it is basically as old as P-312. L23 is from another m-269 line that is extinct, as far as we know it (at least it isn't found yet). If that is the case, then all R1b in Europe is less than 5500 years old. AFAIK, there is no migration from the West Asia to Europe at that time, so I think that would make all m-269* from Europe, with IE migrations, of some kind. If those P-25's in Ukraine are legit, that would be huge! It looks like a massive boom occurred above the Black Sea first, with L23's descendants reaching as far as Mongolia and the Pashtuns. I can't think of anything else moving this haplogroup as far as it is, without IE migrations. To my knowledge, there is no movement from West Asia that would coincide with the ages of m-269* and L-23.

Chad Rohlfsen said...

If that data is correct, it looks like we have m-343 in Central Asia, P-25 in Ukraine, and m-269*/L23, arising in Europe.

Grey said...

"How would R1b be in North Africa, prior to being in Europe?"

If R1b is somehow connected to a trading / early metal working group they could have been all over the place but only in very small numbers and only expanded to large size under unusual circumstances.

If so you might find traces anywhere there was soft metal.

Grey said...

https://s-media-cache-ak0.pinimg.com/originals/2d/f0/f0/2df0f095523a8f15a29abaac515b3ec4.jpg

Judging solely by that graphic there appears to be a gap in the stelae distribution round about where all those advanced copper working Balkan cultures would have been before they disappeared.

Ebizur said...

Perhaps many of you already know that a certain person has argued on the Anthrogenica forum for a European origin of Y-DNA haplogroup N on the basis of an extremely rare subclade that seems to have been found in a couple modern Serbo-Croats and in an Iron Age (9th or 10th century BCE) individual from Hungary. (He has never attempted to explain which Y-DNA lineage may have accompanied the direct matrilineal ancestors of modern Serbo-Croats in D4j, F1b, etc. on their hypothetical migration from East Asia -- perhaps he thinks that these are the descendants of Amazons or that they have originated in Europe alongside Y-DNA haplogroup N. Who knows?) I admire his freethinking, but it has made me wonder about the strength of evidence that has been presented for or against other hypotheses of haplogroup origins.

Please consider the following data from Zhong et al. (2011):

(Non-Muslim) Han Chinese total
44/1729 = 2.54% Q1a1a1-M120
1/1729 = 0.06% Q1a1b-M25
4/1729 = 0.23% Q1a2-M346
49/1729 = 2.83% Q total

16/1729 = 0.93% R1a1a-M17
2/1729 = 0.12% R1b1a2-M269
3/1729 = 0.17% R1b1a1-M73
1/1729 = 0.06% R1b-M343(xM335, M73, M269)
4/1729 = 0.23% R2a-M124
26/1729 = 1.50% R total

75/1729 = 4.34% P1-M45 total

The People's Republic of China is comparable to the subcontinent of Europe in land area, although the PRC contains nearly twice the population of Europe. The Han Chinese account for more than 90% of the total population of the PRC. Anyway, the population of "ethnic" Han Chinese in China is much greater than the population of "ethnic" Europeans in Europe.

The percentage of all Europeans who belong to R-M17 or R-M269 is surely many times greater than the percentage of all Han Chinese who belong to R-M17 or R-M269. However, the situation regarding other subclades of R (or Q for that matter) is not so clear-cut. It is even more perplexing when one considers the fact that there are many more Han Chinese in the PRC than there are Europeans in Europe.

I am not arguing that Y-DNA haplogroup R has originated in China; I am just wondering how strong the evidence to reject a hypothesis that haplogroup R has originated in China might be.

Davidski said...

Well, R is by far the most common Y-hg in Europe, and yet Europeans don't carry any East Asian genome-wide ancestry, except those affected by Turkic and Uralic migrations. So that pretty much kills any idea that R moved into Europe from an area of at least moderate East Asian ancestry.

So the idea that R is East Asian isn't parsimonious. In fact, I would say that N isn't East Asian either, but rather North Asian or Central Asian. The idea that N came from southeast Asia looks ridiculous to me.

Shaikorth said...

Basques are heavily R, differ significantly from other Western Europeans in that they can be modelled without ANE, show only weak negative "Sardinian, ANE" f3 signals and unlike other Western Europeans don't show any "Sardinian, ENA" f3 signals. But this can't really be interpreted to mean their R ancestors did not have at least ANE at some point.

Besides dilution of ancestral components it is possible that some of them cancel each other out for certain methods like ADMIXTURE. The Bedouins for instance were successfully fitted by Everest @ Anthrogenica (with a Z-score deemed acceptable in Lazaridis et al.) as two-way mixture of Chechens (~25% ANE) and about 30% Yoruba in ALDER. In most tests, including ALDER itself when just measuring SSA in Bedouins and not fitting them as 2-pop mixes, these Bedouins show SSA but much less than in the Chechen-Yoruba fit and no ANE. For Europe the component to look at is mostly Middle Eastern/Basal Eurasian, or maybe its African affinities.

There should be more full genome sequencing in Asia plus obviously more ancient DNA to resolve MNOPS origins.

Davidski said...

Basques are a small and unusual isolate. I'm talking about a whole continent here, where, at least for the most part, R arrived after the Neolithic.

Ebizur said...

The same study that I have quoted in my previous comment (Zhong et al. 2011) has found N-M231 in 117/1729 = 6.77% non-Muslim Han Chinese from the PRC. They have found no discernible latitudinal cline (59/876 = 6.74% N-M231 Southern Han, 58/853 = 6.80% N-M231 Northern Han), which in my opinion tends to support a hypothesis that all but an insignificant part of modern Han N-M231 lineages have descended from N-M231 ancestors who have been members of the proto-Han ethnos as opposed to people who have subsequently joined the Han ethnos. (A later addition to the historical Han meta-population ought to exhibit some sort of cline.) There is no salient longitudinal cline either as far as I can tell from eyeballing the study's data table in Figure 2.

Shaikorth said...

I brought up the Basques because while their autosomal DNA differs from other Europeans, never mind non-European R populations, their Y-DNA doesn't differ in a similar way. Autosomals of a modern population can't really be used to gauge the autosomals of the populations that Y-DNA originated in. Neither Basques or Europeans are exactly the hotspots of basal R (or N) diversity.

spagetiMeatball said...

David,

What is east asian then? I mean haplogroup O is east asian for sure, because it is almost entirely limited to southeast asian and east asian populations.

What I mean for example is, does it make sense to speak of a "central asian" haplogroup? Who was living in central asia 10,000 - 15,000 years ago. What about north asia?

Chad Rohlfsen said...

R1b may have started in Central Asia, but it's not East Asian. One source I've found dates the Bashkirs l23 as the oldest. I'm looking for more at the moment.

Alberto said...

"Basques are a small and unusual isolate."

We're talking about a population of about 2.5 millions, who happen to be the single "big enough" population in the world with the highest percentage of R1b, and it is a very high ~85%. And yet their autosomal DNA show them as the most "Western" of all the modern populations. Coincidentally, they're also the only population in Western Europe who speak a non-IE language.

We cannot just dismiss these clues as a "small and unusual isolate". And actually the whole Spanish population (some 42 million people) is pretty "Western" and yet it has higher R1b (around 70%) than they have in France or England (and of course Belgium, Netherlands, Germany, Italy...)

I don't deny the clues that suggest an Eastern origin with IE speakers bringing R1b to Western Europe after the Neolithic. But we cannot deny these other clues that suggest the opposite either.

The reality is that we don't know.

Guessing is fun and I have nothing against it. But let's just all agree that it is just guessing and any theory could be proved wrong any day when more data comes in.

(Yes, I did notice the tile of the post containing the word "speculation". Just wanted to remind it here in the comments again).

Richard Rocca said...

Sorry to disappoint...

Irish = 90% R1b+ (Busby 476 samples)
Basques = 83% R1b+ (Martinez-Cruz 581 samples)

Ireland Population = 6.4 Million
Basque Country Population = 2.5 Million

So the Irish have the highest percentage of R1b on the planet in a population more than double that of the Basques in an area many times larger than Basque Country combined.

Chad Rohlfsen said...

Basque has Celtic influence. It's more a case of endogamy after the first push. If we want to go that direction, I can find plenty of groups with as much r1a as a lot of Eastern Europeans, yet speak non-ie. One or two isolates is nothing. Bashkirs have a ton of r1b. Should we make r1b Turkic? Of course not....don't get hung up on one or two groups, as stated, I can do the same with R1a.

ZeGrammarNazi said...

The Ouldeme are 96% R1b and they are autosomally Sub-Saharan African. I guess that means R1b comes from Africa, right...?

Modern day frequencies and autosomal figures do not trump the SNP trail; which doesn't point to an Iberian or African origin for R1b lineages.

Chad Rohlfsen said...

I wouldn't even go into V-88 and V-69, those branched off a long time ago, in a place that we have no idea, but took off around 3-5kya. It wasn't in the Neolithic groups, so it may be from a further South P-25 group. It might be the one South of the Caspian, expanding into the Levant and Africa in the metal ages.

The big thing, is that we have modern m-269* spread from Western Europe, West Asia, Mongolia, and to Central Asia. PF7558,7562, and 7563 descendants are apparently only dated to 2500BCE. So this one guy's kids branched around at that time. Which people migrated to those places at that time? I can only think of one group. L23 is branched into a Western and Eastern group. The Eastern group follows R1a around, to the East, the other one looks to have expanded in the Balkans, leading to L-51. Armenians, with a lot of L23, are the ones that are Balkan shifted. All other Armenian groups plotted closer to people they were surrounded by. Basically, everything we have, with Eurasian L23, happened in the last 5500 years.

Show me a migration into those areas , that is not IE related, at that exact timeframe. There is no migration out of West Asia into those locations. Only steppe groups spread at that time, to those locations.

Lathdrinor said...

@Ebizur studies on ancient DNA in East Asia have shown that N, thus far, has primarily been found in the area immediately to the north of the Chinese agricultural heartland. It was discovered at great quantities in the vicinity of eastern Mongolia/western Manchuria, where it experienced diachronic change, being eventually largely displaced by O3 and C3 during and after the Bronze Age. It was also discovered at a Neolithic Yangshao site in Inner Mongolia.

While I agree that N entered China before/during the proto-Han period, as indicated by finds ~3,000 BP showing N in China proper, and indeed lived side-by-side O3 even prior to that, there are no finds, to my knowledge, of N in ancient southern China and southeast Asia. Thus, even though modern SNP diversity leads us to the belief that N came from southeast Asia via southern China, ancient DNA requires us to take up a 'wait-and-see' policy towards the problem. To this end, the lack of ancient DNA data from southern East Asian regions remains a great obstacle to understanding how it all came together, one that I hope is going to be addressed in the future.

Alberto said...

Ok, I took figures from Eupedia where it sums up different sources and Ireland is rated at 81% R1b with over 1000 samples. Anyway it's not important if Basques are 1st or 2nd in the world.

So there seems to be some consensus here that R1b spread from the Yamna Culture? Ok, maybe that's right. I can't wait to see some ancient DNA samples from that culture and see if they where R1b, R1a or something else.

Does someone here expect to see R1b in the samples from the Samara region that were recently analysed (but not yet published)? If not, why not? R1b was isolated to Ukraine and not spread to the rest of the steppe?

Davidski said...

Steppe groups were patriarchal and patrilineal, so they were often dominated by single Y-haplogroups, even more so than most modern populations. For example...

http://eurogenes.blogspot.com.au/2009/05/ancient-siberians-carrying-r1a1-had_24.html

So even if R1b was present at the western edge of the steppe, in a group like, say, the Ezero, it might have been lacking totally among the Yamnaya groups of the Samara Valley.

http://books.google.com/books?id=tzU3RIV2BWIC&pg=PA188&lpg=PA188&dq=ezero+culture&source=bl&ots=wWk305abaL&sig=i1O8-ewUhKDXJwKeKWVD1A5LpSY&hl=en&sa=X&ei=7lp-VPPDLIi-ggSX7oAg&ved=0CDkQ6AEwAw#v=onepage&q=ezero%20culture&f=false

Ebizur said...

More data from that same study (Zhong et al. 2011):

Uyghur/Xinjiang total
2/187 = 1.1% Q1a1a1-M120
3/187 = 1.6% Q1a1b-M25
4/187 = 2.1% Q1a2-M346
2/187 = 1.1% Q1b1-M378
11/187 = 5.9% Q total

55/187 = 29.4% R1a1a-M17
6/187 = 3.2% R1b-M343(xM335, M73, M269)
3/187 = 1.6% R1b1a1-M73
3/187 = 1.6% R1b1a2-M269
6/187 = 3.2% R2a-M124
73/187 = 39.0% R total

84/187 = 44.9% P1-M45 total

9/187 = 4.8% N-M231

26/187 = 13.9% O-M175

35/187 = 18.7% NO-M214 total

Hui (Chinese-speaking Muslim) total/Ningxia (n=62) + Yunnan (n=10)
2/72 = 2.8% Q1a1a1-M120
1/72 = 1.4% Q1a2-M346
3/72 = 4.2% Q total

1/72 = 1.4% R1-M173(xR1a1a-M17, R1b-M343)
3/72 = 4.2% R1a1a-M17
1/72 = 1.4% R1b1a2-M269
1/72 = 1.4% R1b1b-M335
2/72 = 2.8% R2a-M124
8/72 = 11.1% R total

11/72 = 15.3% P1-M45 total

7/72 = 9.7% N-M231

25/72 = 34.7% O-M175

32/72 = 44.4% NO-M214 total

Note that N-M231 appears to be more common among Han Chinese throughout the PRC than it is among Turkic-speaking Muslims in Xinjiang. These Turkic-speaking Muslims (so-called "Uyghurs") have been sampled from four locations in northern Xinjiang (i.e. the former Dzungaria), where many toponyms are of Mongolic origin; the Turkic- and Tungusic-speaking inhabitants of this region, like the Chinese-speaking ones, mostly have invaded the area since the Oirat (western Mongol) extermination campaign ordered by the Manchu government of the "Great Limpid Empire."

An origin of Han Chinese N-M231 directly from the west appears implausible. It could have come from the west via Siberia followed by Mongolia and/or Manchuria, but I do not think lgmayka's argument regarding N-Y6503 is a compelling reason to prefer a hypothesis of a European origin over a hypothesis of an East Asian origin of N-M231.

Chad Rohlfsen said...

David, that's possible. The thing is that Bashkirs in that valley, look to have the oldest l-23. R1b looks like it may have been very small and blew up around 5500 years ago, possibly in Cotafeni, with the l51 branch. It may have been the Ezero that blew up first. Bell beaker not having west Asian mtdna, but corded does, may point to that.

Chad Rohlfsen said...

Ezero may be too old for L23 though. It's hard to say. We could have basal L23 in the Samara area... which turns into Eastern L23, those that go West turn into the Western branch, which entered West Asia. The ones that stayed in the Balkans become L51, and swamp out Western Europe during the Copper Age.

Chad Rohlfsen said...

That Ezero connection with Troy would be a neat story for how L23 entered Anatolia. You may be spot on. I was thinking Ezero was older. Ezero and Cotafeni are about the same age.

ZeGrammarNazi said...

How about R-M269(xS127) hanging out on the West Coast of the Black Sea around 3000 BC in the form Ezero and related cultures that introduced arsenical copper to area(look at Fig. 2 in Busby's 2011 R1b paper), then moving through Trans-Danubian complex (Vucedol, Baden, etc.) sparking the Remedello II phase in the Alps, and also becoming part of the Eastern Beakers?

ZeGrammarNazi said...

Chad, I am going to send you a PM over at Anthrogenica with some interesting info on the topic we are currently discussing.

Chad Rohlfsen said...

Z,
Very interesting stuff. I still don't think the link goes back to Iran, but the Northern Steppes, where the tanged daggers started. They were received from Maykop between 4000-3000BCE.

There's a decent chance that anything m-269 in SE Europe and West Asia is only 4500yo, and too young for this, but part of the migration into Anatolia.

Chad Rohlfsen said...

Same origin, via Iran, into Maykop, but probably not involving R1b genes. L23 is supposedly older in the Bashkir, than Europe, or West Asia.

Chad Rohlfsen said...

Plus, we have the WHG problem. Where R1b is heavier, so is WHG. We have over a 20% increase in WHG from Baden to mako. It's not from corded, so it had to be from R1b. Therefore a route through west Asia is not likely.

ZeGrammarNazi said...

Then how about this: The R1b, steppes, Ezero, Mako and arsenical copper working link may be attributed to the people described here:

http://www.ufg.uni-kiel.de/dateien/dateien_studium/Archiv/201011_furholt_hinz_lesenswert/03_sitzung/gimbutas1979.pdf

Page 120, Section II.

Even has a connection with Jean's Stelae theory.

ZeGrammarNazi said...

R-M269 may represent the Second Wave of Kurgans described by Gimbutas and R1a clearly represents the Third Wave of steppe peoples represented by Yamnaya, which led to Corded Ware.

To further this speculation, it may be that R1b-U106 represents a Global Amphora lineage that was absorbed by Corded Ware, represented by R1a guys as the Third Wave moved into town wrecking everything.

Alberto said...

Davidski, that would make sense if those samples belonged to the same clan.

But those samples from the Samara region span a period of 6000 years. Do you know how long is that? From the time of Jesus till the present it's "only" 2000 years. From the time the big pyramids were built till the present it's "only" 4500 years. 6000 years is a HUGE amount of time.

If someone proposes that R1b came from the Eurasian Steppe Proto-Indo-Europeans, who are known for having great mobility and being warriors by nature (fighting for dominating important resources, like the Volga river in this case), he should expect those samples from Samara to be around 50% R1b - 50% R1a (at least in the older samples from 5000-9000 y.a). That's what the logic would say if one is objectively looking at the facts and not just trying to push some preconceived theory.

If R1b is not present in those Samara samples, but later it is found in cultures from the Western coast of the Black Sea (Cernavoda, Ezero,...) It could still suggest a post-Neolithic eastern origin, but the possibility of coming from the southern coast of the Black Sea (Anatolia) instead of the northern coast (Ukraine) might really make more sense.

Davidski said...

There's absolutely no reason why we should expect the Samara Valley Yamnaya samples to be 50/50 R1a/R1b. Like I said, because of the patriarchal nature of the Indo-Europeans we should expect the opposite; different clans living next to each other with vastly different frequencies of closely related but different Y-hg lineages.

And I really don't see how we can squeeze in a major post-Neolithic migration into Europe from Anatolia, considering that Europe became less Near Eastern after the Neolithic, unless we're talking about a few regions only.

Alberto said...

"Like I said, because of the patriarchal nature of the Indo-Europeans"

But you still ignore the fact that those samples span a period of 6000 years! If R1b was in the steppes, it must have been in the Volga river at some time during those 6000 years. There actually is no reason why to expect the samples to be R1a at all, instead of 100% R1b.

I say 50-50 just as a way of saying that if both groups where there (in the steppes, sharing the same language, religion, burial, practices, culture,...), during those 6000 years both must have passed and stayed in that region. They were moving people, and 6000 years is a long, long time.

I don't propose an origin through Anatolia. I just say that IF there are no R1b samples in Samara, and IF they are later found in Cernavoda, Ezero and related cultures (which is what you seem to be suggesting), then it would suggest a more likely passage through Anatolia than from Ukraine, for obvious reasons. But none of these two things have happened yet, so let's not get ahead of ourselves.

Chad Rohlfsen said...

Mako and Bell Beaker have too much WHG for West Asia. Armenians suggest the movement was in the other direction. That Ezero to Troy link from David is pretty fascinating, and fits the timeframe. It might be a bit of a fantasy, but L23's age and its dominance in West Asian R1b would fit. Eastern L23 is not found in West Asia, that is another killer of that route.

Grey said...

@spaghetti

"What is east asian then?"

*Now* I'd say it was the end result of their version of farmer expansion from whichever sub group was the epicenter for that.

Before then it may have been different - I'd guess the local variety of ASE derived HGs.

(If ASE is a real thing.)

Grey said...

Alberto

"So there seems to be some consensus here that R1b spread from the Yamna Culture?"

If you look at maps of the distribution of LBK and Atlantic Megalith then there is a big gap along the western and northern edges of Europe which I assume means those regions weren't suitable for neolithic crop-centric farming at that time.

So if a small population arrived in those regions who had a method of exploiting that niche (e.g. cattle and LP) then they might expand very dramatically.

(The pastoralist habit of maintaining paternal lineages is possibly important here.)

If that is the case then it seems at least possible that R1b *might* have originally been quite small in which case it could conceivably have come from a lot of places.

So personally I'm open to suggestions although currently leaning to the steppe.

Nothing would particularly surprise me about R1b though.

Grey said...

"If R1b was in the steppes, it must have been in the Volga river at some time during those 6000 years. There actually is no reason why to expect the samples to be R1a at all, instead of 100% R1b."

Assuming it was the steppe area for the sake of argument then if there was a split into R1b and R1a then it might it have occurred at a natural geographic / climate border.

So say R1b were marsh HGs around the Black Sea (I assume it was very marshy when the sea level was lower) while R1a were in the interior

(or alternatively R1a were the same but around the Caspian).

And under the influence of farmers to their south the R1b develop a mixed farming / pastoralist model on the edge of the steppe while the R1a develop a full pastoralist model because farming isn't viable at all in their region.

Then assume full pastoralist > half pastoralist > full farmer in terms of hassling people and you have the mechanism for a conveyor belt across Europe from the steppe to the west.

Chad Rohlfsen said...

https://www.familytreedna.com/public/r1b/default.aspx?section=ycolorized

Chad Rohlfsen said...

https://www.academia.edu/9666892/Y-Chromosomal_Haplogroup_R1b_Diversity_in_Near_East_is_Structured_by_Recent_Historical_Events

Interesting, but I'd say the majority came 2-3k before this. There is l-11, in Armenians, and I think that if saw a p-312, in an Assyrian, L51, in a Lebanese. It's on that R1b ftdna link that I posted.

Davidski said...

Nah, this paper is a bunch of horseshit. This guy produces stuff that's even worse than we normally see from academia.

Chad Rohlfsen said...

Most of it is wrong, but there are typically European subclades there.

Chad Rohlfsen said...

Going by 10 strs, Pashtuns have the highest L23 diversity. We can probably thank Greeks, Turks, and Mongols for that. Second is actually Poland. Next is Turkey, but that probably includes anatolians, phyrgians, Greeks, celts, Romans, and Armenian diaspora communities. Third and fourth is Romania and Bulgaria.

Chad Rohlfsen said...

Fourth and fifth, excuse me.

Chad Rohlfsen said...

DYS19—DYS388—DYS389I—DYS389II—DYS390—DYS391—DYS392—DYS393—DYS439—DYS461

Modal(L23+)----L23+----956-----14—12—13—16—24—11—13—13—12—11

Modal(L23xL51)----L23----212-----14—12—13—16—24—11—13—12—12—10

European Modal(L23+)----L23+----812-----14—12—13—16—24—11—13—13—12—11

Western European Modal(L23+)----L23+----632-----14—12—13—16—24—11—13—13—12—11

Western European Modal(L23xL51)----L23----29----14—12—13—16—24—11—13—12—12—10

Caucasus Modal(L23+)----L23+----33----14—12—13—16—24—11—13—12—12—10

Caucasus Modal(L23xL51)----L23----33----14—12—13—16—24—11—13—12—12—10

Turkey Modal(L23+)----L23+----69----14—12—13—16—24—11—13—12—12—10

Turkey Modal(L23xL51)----L23----33----14—12—13—16—24—11—13—12—12—10

Everyone has the same modal ancestor. Nothing deeper in Turkey. Ezero, with some in Cotafeni?

Chad Rohlfsen said...

Myers has Romania as the most diverse, at 8 STRs, and Pashtuns the most diverse at 10, with Romania in second.

Davidski said...

Population genetics based on 8-10 STRs is a lost cause.

Chad Rohlfsen said...

Not great, but it's all that is out there. The Bulgarian project from 2013, had L-23* as oldest in Bulgarians. Older than that in the Caucasus or Near East. The Caucasus and Iraq was older than Turkey, but not by much. Kind of strange, possibly two entry points. There was no testing done on Ukrainians though. Something else that's interesting is that Iraqi Kurds are more R1b than R1a. 33% more, to be exact. They also have elevated I y-dna. 16.8%, the same as R1b. Most of it is I2, I believe. That makes a Balkan route look even better, IMHOP.

Simon_W said...

@ Tone, Alberto

Neither STR diversity nor the presence of old R1b paragroups supports an R1b origin in northwestern or western Europe, to the contrary.

Also R1b wasn't present in the Scandinavian HGs known so far, they all belonged to I2. And what is often forgotten, we've got 2 y-chromosomes from the megalithic SOM culture, and these were I2.

@ David

There is no way that Vucedol could be a Bell Beaker culture. What has been suggested was some Vucedol influence in the eastern Bell Beakers in southern Germany and nearby. But this doesn't make Vucedol a Bell Beaker culture. Bell Beaker spread to the Carpathian Basin later, from the west.
Furthermore Vucedol was rather a „kurganised“ culture in the widest sense, like Gimbutas used it, which however doesn't always depend on real EHG input as someone in another thread pointed out.

@ Richard Rocca

I don't think so. If you look at early EEFs like Stuttgart or NE1, they have lots of East Med, and also some Red Sea. If you look at later ones, like Gok2 or NE7, their East Med and Red Sea is strongly reduced, and their West Med increased. What has happened? Apparently WHG admixture managed to increase the West Med component at the expense of East Med and Red Sea. And since northwestern Europe probably harboured even more WHG admixture, this may well have completely nullified any East Med that might possibly have been associated with the early R1b folks. But there seems to be some slight South Asian admixture associated with R1b, even on the British Isles, and this couldn't be nullified by WHG admixture. Where did R1b people get this from? Quite possibly from Western Asia.

Simon_W said...

There clearly seems to be a founder effect at the beginning of R1b in western Europe. L23* is still an eastern marker, found chiefly in the Balkans, Western Asia, the Caucasus and the Urals. The westernmost pocket of it seems to be in the Valais in the Swiss Alps. Descended from L23* is L150*, about which little is known. It does have a Near Eastern daughter clade however. And its other daughter clade is M412* which is already clearly western European. It's most common in Southeastern France, France in general and eastern Ireland. From this L11* is descended. So maybe R1b dudes from Western Asia sailed to the Western Mediterranean by boat and landed in Southern France perhaps. Certainly not with the early Neolithic dispersals, but quite possibly later, Copper Age. And in France they got incorporated into the expanding Bell Beaker culture. If that's true, they may have carried the K15 West Asian component to Western Europe as well.

The increased WHG in Mako may have been brought by R1a people.
Eastern L23 isn't found in West Asia? And basal L23 is found where?

Simon_W said...

Also see my comment in the other thread: I now have read the paper linked by David, by G. Kulcsar and V. Szeverenyi, it's exciting! I've drawn four main conclusions from it, and an additional fifth insight:

1. The source for the EHG admixture in the Mako culture must be the Yamnaya east of the Tisza - there really is no other explanation, and it seems possible, if this was from the early western Yamnaya periphery where the Caucasus admixture may still have been minimal.

2. Although these Yamnaya people had an important effect on the Mako gene pool, culturally they were not really the dominant source for the Mako culture. Vucedol seems to have had the bigger formative influence. Which means that we cannot conclude that the Yamnaya people imposed their language on the Carpathian Basin. Mere biological importance via genes isn't as good an indicator for linguistic dominance as cultural dominance is. And Vucedol was quite a dominant culture. It was considered a "Kurganised culture" (i.e. patriarchalic, warlike etc) by Gimbutas and they had fortified hilltops etc, really hard to overthrow. 

3. Yet Vucedol probably wasn't EHG admixed! The early Vucedol in the south and the west of the Carpathian Basin was contemporaneous with the late Baden in the north of the Carpathian Basin. The latter were typical EEFs, and I really don't believe that people to the west of them were more EHG admixed than them. 

4. The idea that R1b in German Bell Beakers is in any way Yamnaya-derived can be thrown into the rubbish bin. Obviously there was never any Yamnaya in Germany. The idea has been put forward that Vucedol influence may have carried Yamnaya admixture westwards along the Danube. But as I have just argued above, Vucedol probably wasn't Yamnaya admixed. They were culturally „Kurganized“ (in the widest possible sense), but not Yamnaya derived.
And in case someone thinks the late Vucedol, which was contemporaneous with the early Mako, might have taken R1b to the German Bell Beakers - this doesn't work out either, because late Vucedol was a Bronze Age culture, while German Bell Beakers still were on a Copper Age level. Forget it.

5. It's possible that the original R1b carriers in western Europe didn't have EHG admixture, but picked it up from the GAC/Corded substrate in Germany. After all, we now know that the dominant haplogroup of mondern Scandinavians, I1, was from EEF people. Hence, probably neither IE language nor strong HG ancestry was associated with the haplogroup that is now dominant in northern Germanics. With R1b it may have been similar. Or partly so.

Simon_W said...

Hovhannisyan et al.'s suggestion that it all has to be explained with the earliest Neolithic wave is quite silly. And contradicted by tons of yDNA of the actual earliest Cardium and Cardium derived Mediterranean farmers. G was predominant among them and they had neither J2 nor R1b, at least not in any significant numbers. The author's data plotting the STR diversity on maps is more interesting, but important populations are missing, especially Iberia and between central and southeastern Europe. The heatmaps of the Fst are quite interesting, though I'm missing the Ukraine there.

Chad Rohlfsen said...

MJost has been doing lots of work on the higher STRs and SNPs. The highest variance for L-23, is found in Romania. Second is Bulgaria. It can't be older than 4300BCE, and is likely younger. He dates L-11 to 3500BCE max, probably younger, and M-269 to 5000BCE. Who was in the West Coast of the Black Sea at that time? Cernavoda, the ancestor of Cotafeni and Ezero. Cotafeni and Ezero end up in the Carpathian Basin. Ezero also ends up in Troy by 2800BCE. That is probably how L23, L51, and L11 were introduced to Anatolia. R1b springs from an ANE group in Central Asia. It can't be in West Asia, as Neolithic groups had no ANE. Yamnaya won't have enough WHG vs ANE do explain BR1. I'm pretty sure we won't see Yamnaya having at least twice as much WHG as ANE. Pontic Kurgans are my best guess. That allows two modes of entry into West Asia, for L-23. There are no migrations from West Asia at that time to plant L-23 there, and then spread it to Mongolia and Spain. That is illogical.

Chad Rohlfsen said...

From MJost :

In my L23* combined variance number, Romania had considerable more 2.8 than western Bulgaria. Hungary had 1.86 in the eastern half and 2.0 in the western half. I didn't see any Ukraine data but I would assume its at a smaller percentage yet since Poland and Russia are in the variance range of 1.0 and 1.2 respectively.

Chad Rohlfsen said...

The final blow to the West Asian hypothesis is the fact that only Corded Ware showed West Asian mtDNA. Bell Beaker had none. They only had Neolithic, Pontic Steppe, and EHG. It was in that paper from a while back.

Chad Rohlfsen said...

Bell Beaker mtDNA
http://img196.imageshack.us/img196/9937/g9sn.png

Notice Bulanova too...

Corded Ware mtDNA
http://3.bp.blogspot.com/-L4xDPSyXvDs/Ulb1KodX6KI/AAAAAAAAJLs/ETRnvsd-tmw/s1600/UC.jpg

Bell Beaker did not get its ANE from Corded Ware. They don't share much in the way of haplogroups. If anything, Bell Beaker lost its ANE from mixing with European farmers more than Corded did.

Chad Rohlfsen said...

Excuse my mistake. Too far West for Bulanova, but you get the idea.

Chad Rohlfsen said...

Sorry, that map was of Unetice. I will post the Corded Ware.

Chad Rohlfsen said...

Corded Ware
http://dienekes.blogspot.com/2013/10/ancient-central-european-mtdna-across.html

in here

Simon_W said...

Well, Cotofeni may well have had some ANE/EHG admixture. I've read that Kurgan-type crania were present in Cernavoda, among others, since that population was very diverse. And Ezero was related with Cernavoda III, too. And then the Bronze Age in Ezero was earlier than in the Carpathian Basin, meaning that there may have been some southeastern influence at the beginning of the Bronze Age Mako culture. So the change from CO1 to BR1 might indeed possibly be ascribed to Cernavoda III-Cotofeni-Ezero related admixture. But I'm not sure, it depends on how strongly EHG admixed these latter cultures really were. With Yamnaya at least we'd still have a more or less pure thing, geographically closer too.

But either way, this doesn't quite explain the R1b in Kromsdorf. The German Bell Beakers were a Copper Age culture, Ezero was Bronze Age. In my opinion a migration that was formative on the Bell Beaker gene pool must have been on a similar cultural level, otherwise the German Bell Beakers would have become a Bronze Age culture too. Cotofeni was a Copper Age culture in the Carpathian Basin, but to the west of it (hence closer to Germany) was Baden, which was autosomally like CO1. Maybe they had R1b, but they were like EEFs.

What was the L23* variance in West Asia like? Any data for this?

I'm not an expert for these late Copper Age cultures of southeastern Europe, but Baden-Kostolac-Cotofeni-Cernavoda-Ezero are generally seen as forming a complex of related cultures. And the importance of the Kurgan/steppe element is controversial, Baden for instance also shows relationships with Anatolia. I guess in that case the others of this complex will show that too. So you can't say there are no migrations from West Asia at this time. If L23* was from West Asia, it might have got to Mongolia via the Silk Road, perhaps with the Tocharians. This doesn't have to have been at the time when L23* came into being (split time =/ migration time). Indeed the Tarim mummies were all R1a. But we don't know if they spoke Tocharian or something else. And according to the Eupedia map of R1b-L23 (the map includes L51, L11 and Z2103) southwestern Iberia also has somewhat elevated frequencies of these old clades. The variance will probably be much lower than in southeastern Europe, as the Iberian ones are farther away from the origin. But, the frequency is similar to parts of Bulgaria. And incidentally the Bell Beakers originated in southwestern Iberia.

You say R1b couldn't have sprung from West Asia as Neolithic groups had no ANE. But how about this idea: R1b migrated from central Asia through Iran and arrived at a relatively late date in eastern Anatolia, long after the establishment of EEFs in Europe.

And no, it's not true that Bell Beakers had no West Asian mtDNA. The map you posted shows the overall distance between populations, and because of the high share of haplogroup H in Bell Beakers, which is rare in West Asia, the distance to West Asian populations is large. But Bell Beakers did have T1 which is the most important of the late Neolithic/EBA haplogroups linked to West Asia. They also had I1.

There are quite some commonalities between German Bell Beaker and Corded Ware mtDNA. There is even some U2 in Bell Beakers. In the recent paper by Anna Szécsényi-Nagy, Guido Brandt, Victoria Keerl et al. BBC and CWC plot together both in the PCA and in the MDS.

Simon_W said...

I've thought again about what I wrote yesterday, and it's really not as simple as it may seem.

First of all, the K15 South Asian component is partly an ANE component and thus not in every case from West Asia. For instance, it's 1.71% in Kargopol Russians and even 3.86% in Mari. And to me it looks very likely that the relatively strong South Asian component in Hinxton4 (1.9%) is ANE related, and not from West Asia. Moreover at present it's stronger in northwestern Europe (especially in Orcadians and Danes) than in the southwest. And thus it's clearly related with the elevated Gedrosia and Indo-Iranian components in northwestern Europe.

There are three interesting questions that may look connected to each other at first sight, but which we'd better treat seperately: 1. Where did the R1b originate that exploded in western Europe? 2. How and when did the strong ANE admixture in northwestern Europe arrive? 3. How did Italo-Celtic languages spread?
We don't know for sure that the answer to all three questions is the same.

Two things are quite clear to me at the moment: R1b in Kromsdorf Bell Beakers wasn't derived from Yamnaya. And the German Bell Beakers didn't have particularly high levels of ANE, they had considerably less than the Corded people.

So, to me it seems likely that later, Bronze Age invasions from the east must have increased the ANE in central Europe again. Because in the end this must have spilled over to northwestern Europe and the Bell Beakers probably wouldn't suffice, because surely there was no complete population replacement in northwestern Europe. There is the theory by Grigoriev about Seima-Turbino influence in the Middle Bronze Age Tumulus culture, and there is the spread of chariot technology from Siberia/central Asia at about the same time. I think the Bronze Age Dane M4, with his surprisingly strong shift towards Finns in the PCA may be an example for one of these later, ANE-rich invaders.

But now the question arises: What have they got to do with the spread of R1b? Was the R1b in Kromsdorf a dead end? After all, only M269 has been verified for them. Did the ANE-rich Bronze Age invaders carry R1b too? Or did R1b explode earlier, in Bell Beaker times, with less ANE? On the one hand there is the fact that the Welsh have nearly zero R1a. So their ANE/Gedrosia/Indo-Iranian almost has to have arrived with R1b people. On the other hand, not all R1b-rich populations in western Europe have lots of ANE. The Basques and probably the Catalans too, both carrying more than 80% R1b, don't have a lot of ANE. Furthermore R1b in southwestern Europe looks positively correlated with non-IE languages. It's strongest in southwestern France and northern Spain around the Basque country, where Aquitanian and Basque were spoken. It's equally strong in Catalonia, where Iberian was spoken. Areas where IE Lusitanian and Celtiberian were spoken have considerably less R1b. And in Corsica R1b seems to be stronger in the interior, where according to Seneca the younger also an Iberian language was spoken - IE Ligurian was confined to the coast. This seems to suggest a pre-IE spread of R1b without large amounts of ANE. But how do we have to explain the above observation on the Welsh? Maybe it's comparable to the explosion of EEF derived I1 in Scandinavians, i.e. maybe it has little to do with the autosomal make-up.

Simon_W said...

But where did the Kromsdorf R1b come from at all? I think that's quite an open question at the moment. It might have come from southeastern Europe or from Iberia, since according to the Eupedia map of R1b-L23 which includes L51, L11 and Z2103, southwestern Iberia also has somewhat elevated frequencies of these old clades. Ultimately it originated in the east, but not in Yamnaya, rather West Asia.

Now finally, what about Italo-Celtic, with what was its spread associated? The quite extensive presence in southwestern Europe in antiquity of non-IE languages makes me doubt an association with Bell Beaker in general. (According to Pseudo-Skylax and other writers Iberian was present up to the Rhone, mixed with Ligurians.) Some eastern Bell Beaker people might have been IE speaking. This might have arrived from southeastern Europe or with the Corded people or with both. Although an Italo-Celtic origin in the Corded Ware would make the entry of the early q-Italic branch into Italy a bit difficult. I cannot completely rule out an arrival with the middle Bronze Age eastern invaders either.

Chad Rohlfsen said...

I'll post the variances when I get home. Iberia wasn't close. That bit in the south is the site of Greek colonization. The same as Southern Italy. It was Bulgaria, Romania, Georgia, Armenians, and I think next was Pashtuns. If you follow German beakers, bronze follows within two hundred years almost everywhere. They gave bronze technology to Corded Ware. The connection to Baden and Troy is all after Ezero, nearly 1000 years after the first incursion at Cernavoda. They had eastern burials with ochre, and elite males. I know of no migration from Anatolia in the L23 phase.

ZeGrammarNazi said...

Simon, the mtDNA from the Kromsdorf site doesn't look very Western European, so it is hard to imagine the people found there had their origins in the Iberian peninsula.

The mtDNA looks more like something you would find in the East, so it is hard to use this site as a benchmark for Beaker peoples. I believe most beaker sites analyzed from Western Europe are more than 75% mtDNA H.

Kromsdorf had the following:

U2e
W5a
I1a1
K1
U5a1
T1a

U2e and U5a1 have both been found in a steppe context and I believe T1a has too, but I'm not as certain of that one.

Anyways, the 2 Kromsdorf R1b finds raise more questions than answers, imo.

ZeGrammarNazi said...

Simon, Ezero is listed as existing from 3300-2700 BC, and the Kromsdorf Beaker site is from 2600-2500 BC.

I do not see anything anachronistic about a possible genetic relationship between the two.

Davidski said...

Another question worth asking is why La Tene and Hallstatt remains show signals of copious millet consumption?

http://www.researchgate.net/publication/223618070_Diet_and_social_status_during_the_La_Tne_period_in_Bohemia_Carbon_and_nitrogen_stable_isotope_analysis_of_bone_collagen_from_Kutn_Hora-Karlov_and_Radovesice

Millet is unheard of in Western Europe, so if Celts came from the west, then they changed their diet in a very important way when they arrived in Central Europe. This is possible, but hard to believe.

In fact, millet is thought to have arrived in Eastern Europe during the late Neolithic from China via Central Asia and the north Caucasus. It then made its way to Central Europe and parts of southern Europe during the Bronze Age, probably along with the habit of drinking animal milk.

The Middle Bronze Age north Italian site of Olmo di Nogara, for example, shows evidence of both milk drinking and millet consumption.

http://www.academia.edu/1214628/Stable_Isotope_evidence_for_the_consumption_of_millet_and_other_plants_in_Bronze_Age_Italy

http://eurogenes.blogspot.com.au/2014/12/milk-consumption-in-late.html

These Bronze Age Italians probably replaced the Oetzi-like population of the region. What types of Y-haplogroups did they carry?

Chad Rohlfsen said...

H1 and H3 are in Germany, with the Salzmunde and Baalberg cultures, 800 years prior to Bell Beaker. Balkan Yamnaya is loaded with H.

Chad Rohlfsen said...

https://drive.google.com/file/d/0By9Y3jb2fORNR0xpSENNWmpZSHc/view

This one does not have a lot of markers, but appears pretty consistent up to 10. He is going to do one of at least 20 when he receives info from Walsh on how the new names match up to the old ones.

Chad Rohlfsen said...

BR1 is likely admixed. The LNE/EBA boom took place in I think, Bohemia and Moravia. So they could've been more of the original invader.

Anyway, they said Beaker was less ANE than Czechs, not Basque, not French, and not Tuscans. Czechs are 16%. If these Beakers create individuals at 15%ANE, then that is enough to pretty much cover the Welsh, Irish, English, French, and anyone West of there. Ireland had the highest population boom in the LNE/EBA, where the population increased four-fold. Don't be surprised if Beakers plot with the Irish, or by the Hinxton Celts. Maybe just a hair closer to BR1. Bell Beaker plotted much closer to Corded Ware than they did to MNE cultures. I think Corded is going to be around Norwegians/Swedes.

Chad Rohlfsen said...

That Cimmerian IR1 and Hallstatt millet could tie together. Could've been a fad that kept on, or it proved an easier crop. It's pretty damn tough and has a short growing season. You could possibly crank out an extra harvest with it.

Chad Rohlfsen said...

It certainly could be used as cattle feed too. More feed= more cattle= more power= more ladies= more kids.

Chad Rohlfsen said...

Numbers posted by MJost

My old STR data for M269 (280 67marker HTs.) was 5,821.5 SD+- 1,358.0. Thus at the max SD = 7, 179.5 ybp (or 5180bc).

Since I have been recalibrating using my SNPs back to DF13 and adding back all ancestral SNPs known, M269 was spawned around 7233ybp (or 5233bc).

Using my estimated 129 years per mutation dating SNP blocks above DF13 at 3750 ybp (my suggested DF13 age) working backwards similar to Wei Wei, Qasim Ayub, [...], and Chris Tyler-Smith's results.

R1b1a2 - M269 - (903 yrs in this block of unordered seven SNPs to L23) = Max 7233 (5233bc)

block of unordered SNPs - max ages
L23 6330 ybp (4300bc)
L51 5943 ybp (3943bc)
L11 5427 ybp (3427bc)
P312 4524 (2524bc)
L21 4266 (2266bc)

So, none of these lines can seemingly be older than this. This should put L23 around Bulgaria or the Western Steppes. There is nothing obvious from West Asia, after Dudesti, around a 1000 years prior the max age of L23. West Asian L23 should therefore all be from Europe.

Simon_W said...

Chad, sorry for the late reply, I had to read a little.

Greek colonization in Iberia was negligible. To my knowledge there were only three Greek colonies in Spain and none in Portugal. There was Emporion in the very northeast, close to the French border. Then there was Hemeroskopeion in the east, south of Valencia. And finally Mainake in the south near Malaga. However, according to that Eupedia map I was referring to, elevated levels (5 – 10%) of the older R1b clades cover the entire southwestern third of Iberia, i.e. almost all of Portugal and the south and southwest of Spain. I don't know if that map is correct, but I assume it was made with the necessary care. This pattern in any case cannot be explained with Greeks.

There is one big problem with the idea that L23 originated in the western Pontic: Bashkirs have lots of L23, but at the same time also M73, which is an old sister branch of M269 which is virtually absent from Europe. M73 and L23 must have reached the Bashkirs together, at the same time. Because the distribution of L23 and of R1b in general is so patchy in eastern Europe and Siberia, it would be a stretch of mind to say L23 came from the western Pontic and joined the M73 in the Bashkirs that was already there, while avoiding all neighbouring populations without M73. We've seen similar things in India, where R1a has a rather high STR diversity, but nonetheless we no longer assume that R1a originated in India.

German Bell Beakers didn't give Bronze technology to the Corded Ware, they were a Copper Age culture. The bronze technology spread later, from the Carpathian Basin, giving rise to Unetice, which was related with Corded people, and the North Alpine Bronze age, which was related with Bell Beaker. Also note that e.g. in Britain the Bronze Age began later and so the Bell Beaker people there lived on a Copper Age level for about 600 years, instead of 300 in the case of central Europe. Apparently, not the Bell Beaker people were the cause for the early start of the Bronze Age in central Europe, but the close proximity of Bronze Age cultures in the Carpathian Basin.

The burials of the eastern Bell Beakers look more like a reflection of Corded Ware burials than any burials on the steppe. Typical for the Corded people was a sex differentiated burial rite: men were oriented west-east, women east-west, both facing towards the south. Bell Beaker burials were more varied, there were some in collective graves, some with cremation, but in the eastern groups a common pattern was: men were oriented north-south, women south-north, both facing towards the east. I would be surprised if this wasn't developped after contacts with Corded people. On the other hand, in the North Pontic steppe there was no sex differentiated orientation.

Simon_W said...

As for Anatolian influence in the Baden culture, Alexander Häusler mentioned details of the cart construction which in his view has parallels in Anatolia, and he mentioned more, but I don't remember what. Well, now that we have NE7 and CO1, we see that the difference between them is tiny. CO1 does have slightly more of the East Med component, but it isn't a big difference, and she had no West Asian and no Gedrosia either. So I don't think there was R1b in Baden.

According to Grigoriev, archeological influence from West Asia into the Balkan-Carpathian region during the late Copper Age is most clearly seen in metalwork: With the appearance of arsenic bronze, the technique of metal casting, the casting with two-part moulds, stylistic similarities, traces of nickel in the alloy and tanged tools with a stop. But it's also seen in a new breed of sheep, larger and with better wool. Grigoriev also cites Sherratt who regards the appearance of grapes, olives, asses and fortifications constructed using rock as evidence for Anatolian input. But now there is a twist: According to Grigoriev, most of these developments reached the Balkans via the Pontic steppe. So it's not necessarily in disagreement with the Kurgan theory. But at the same time it would necessitate West Asian input in the steppe populations, which is what the upcoming paper seems to have found as well.

Simon_W said...

@ ZeGrammarNazi

Yes, the German Bell Beakers definitely had EHG influence. Imho stemming from the Corded Ware substrate. In some places apparently more than in others, as you rightly point out. But still, if men migrated more than women, the R1b males might be non-local.

Ezero was Bronze Age technology, Kromsdorf Copper Age technology, that's not anachronistic, but imho precludes an important input from Ezero in Kromsdorf.

Simon_W said...

Imho the evidence put forth for the association of R1b with Italo-Celtic, or even with Centum IEs, isn't very compelling, to put it mildly.

See this comparison of maps:
http://jpst.it/v4Fn

As I said, in southwestern Europe, where we are lucky to know the indigenous languages at an early date, R1b is negatively correlated with IE languages. The orange stuff in the east is non-IE Iberian, and the green stuff is Basque/Aquitanian. Iberian is also present in southern France, mixed with IE Ligurians. The most strongly IE area of Iberia is in the west-center, where a pre-Celtic IE population existed, and later the Celts arrived and joined the blend. In the non-IE areas R1b is 80%+, in the most IE areas it goes towards 50%+.

And now see this comparison of maps:
http://jpst.it/v4FE

R1b and I2b have complementary distributions. I2b is stronger, where R1b is weaker. And I2b is stronger in the more easily accessible areas, R1b is stronger in refuge areas, suggesting that I2b arrived after R1b. And note that I2b is not exclusively associated with English input, but is also strong in areas where English admixture was low, like southwestern Scotland and western Ireland. To me this suggests the Celts brought I2b and reduced the pre-Celtic R1b.

Consider also that in the Lichtensteinhöhle on the Harz mountain chain in Germany I2b was by far predominant, R1a and R1b were present as minority admixtures. And this was from the late Bronze Age, in central Europe. So perhaps central Europe in the Bronze Age wasn't as bursting with R1b as some imagine.

Finally I would like to add that I've come across quite convincing linguistic evidence linking Basque with Northeast Caucasian, see this:
http://www.people.fas.harvard.edu/~witzel/mt26s.html

The linguistic evidence alone might perplex you, but consider also that the Northeast Caucasus, especially Dagestan, is a place with quite strong levels of R1b in some groups and some strange autosomal connection with northwestern Europe.

So I'm thinking that R1b went from West Asia to Europe at some time during the later Neolithic. Perhaps to the eastern Balkans first. And from there it probably went to southwestern Europe by sea. Only to spread later with the Bell Beakers. Gedrosia is weak in BR1 and BR2 and absent in CO1, so it probably didn't take the Danubian route. Also it's dubious that R1b reached the eastern Balkans via the northern Pontic, given the low STR diversity in the northern Caucasus. There may have been some R1b in Yamnaya, since even the Bashkirs have it, but it probably wasn't the predominant haplogroup.

Chad Rohlfsen said...

Simon, the British Bronze Age started 200 years after Beaker. Moravian beakers gave corded ware bronze. Bashkir ancestors may have had it first, but it doesn't look to take off until in the Balkans. L23 isn't old enough to come from West Asia.

ZeGrammarNazi said...

Simon, Ezero did not have Bronze in its early stages. It had arsenic-copper alloys, but not tin-copper (proper bronze). Bronze was a later development.

Chad Rohlfsen said...

There is some more interesting info. R1b L51 may actually be the brother to L23 Z2103,Z2105, and is not descended from it. It is now being dated to about the same age. It could be around 3000BCE or just before. The gap between L51 and L11 has grown quite a bit. It also shows a hot pocket in the Carpathian Basin and Austria, while being surrounded on three sides that are more L23 related. This does match up with steppe groups appearing here at this time, and the supposed birthplace of Italo-Celtic. I am looking for variance reports on it. It looks like L23** split into two separate waves. One that went primarily to the SE Balkans and West Asia, the other to Central Europe.

Chad Rohlfsen said...

If that doesn't have a nice ring of Anatolian and Italo-Celtic to it, I don't know what else does. 1st and 3rd PIE breakoffs there. I don't know of any R1a subclade migrations that match up as well. If there are, please point me to them. I am interested in studying them.

Simon_W said...

Again, my apologies for the sluggish reply.

Chad, Bell Beaker in Britain started around 2500 BC. 200 years later would be 2300 BC, that would be earlier than the beginning of the Bronze Age in central Europe (2200 BC). I don't believe this is possible. According to the English wikipedia, „there is no clear consensus on the date for the beginning of the Bronze Age in Great Britain and Ireland. Some sources give a date as late as 2000 BC, while others set 2200 BC as the demarcation between the Neolithic and the Bronze Age.The period from 2500 BC to 2000 BC has been called the "Late Neolithic/Early Bronze Age", in recognition of the difficulty of exactly defining this boundary.“ However further down in the article, it's stated that „at first they (the Beaker folk) made items from copper, but from around 2150 BC smiths had discovered how to make bronze (which is much harder than copper) by mixing copper with a small amount of tin. With this discovery, the Bronze Age began in Great Britain. 
So I would say it took them 350 years to discover that.

Moravian Beakers gave Bronze to Corded people? That is an anachronistic simplification. Strictly speaking, in the central European early Bronze Age neither Corded Ware culture nor Bell Beaker culture existed anylonger. What we have instead are Unetice and the groups of the North Alpine Bronze Age. Generally speaking, the Bronze Age proceeded from southeastern Europe to central Europe. So thus far, we are in agreement. The problem is just that Bell Beaker proper didn't have a strong presence in southeastern Europe and the Carpathian Basin. Bell Beaker wasn't the predominant culture there. Of course Unetice for instance did have Bell Beaker influence, but that was from the local substrate! But where did Unetice have the Bronze technology from? As Wikipedia states: „The Nitra Group, inhabiting southern Slovakia not only precedes chronologically the Únětice culture, but is also strongly culturally related to it.“ Now the Nitra group itself was basically an Epi-Corded group. Their burial rite was identical to the Corded Ware burial rite. And afaik there was never a Bell Beaker group in the area of the later Nitra group. And where did Nitra acquire the Bronze technology? In the Nagyrev group! This indeed seems to root in the Bell Beaker Csepel group. But this was a late easternmost extension of the ultimately western Bell Beaker culture, which in turn must have acquired the Bronze technology from non-Bell Beaker groups further to the southeast.

The question if L51 is a brother to L23 could be easily tested: There would have to be lots of people carrying L51, but no L23.

Simon_W said...

@ ZeGrammarNazi

I think you may be right, and Ezero just had arsenic bronze and not the „real thing“. Ezero ended about 2700 BC. Central European Bell Beaker people (starting no earlier than 2600 BC) still predominantly used more or less copper. Though copper objects with considerable amounts of arsenic have been found in Bell Beaker contexts, these were exceptions and became predominant only after 2350 BC, that is 350 years after the end of Ezero, and anyway clearly after the two Kromsdorf males.

Simon_W said...

correction:

...still predominantly used more or less pure copper...

Simon_W said...

Regarding my older comment on a possible Basque – Northeast Caucasian connection, I've given that idea up again.

As this paper
http://dienekes.blogspot.com/2014/11/paternal-lineages-and-languages-in.html
clearly showed, Northeast Caucasian speaking groups are dominated either by J2 or by J1. The presence of R1b in them is very patchy, and strong only in a few groups immediately bordering the Caspian Sea. On a population level, R1b is most common in IE Armenians (as far as populations in or near the Caucasus are concerned).

And as the paper by Hovhannisyan et al. showed, the STR diversity of R1b-M269 peaks in the area where ancient Hittite, Luwian and Palaic were spoken, and it decreases towards the east, although the frequency increases towards the east. In Lezgins the diversity is particularly low. I think this suggests that R1b-M269 spread from the west eastwards and was originally associated with IEs.

Simon_W said...

The crucial question being what exactly the meaning of high STR diversity is. Obviously you guys don't believe that R1b-M269 originated in southeastern Anatolia. You may well be right. First of all this may be just a hotspot where R1b-M269 exploded at an early date, and secondly, the Hittites, Luwians and Palaians are considered to be the earliest group that split off from the rest of IE, that is they must have had more time to diversify than the others. And they must be regarded as the cusp of the earliest IE wave, and this cusp may have left close to zero trace in the original homeland. Instead it may have completely dislocated from it.

Simon_W said...

I think we have to face the fact that R1b was fairly common in central Germany (otherwise it would have hardly been found in Kromsdorf) centuries before anything like Bronze technology was around. And this R1b may well have expanded to the British Isles with the Bell Beakers.

But I can't ignore the fact that BR2 had as little as 1.14% Gedrosia and 0% MDLP Indo-Iranian. This suggests to me that R1b probably was present there, but didn't have as strong an autosomal impact as in northwestern Europe. So the latter impact may be rather derived from R1b Bell Beakers in Germany, which in turn must have come from somewhere other than the Carpathian Basin.

There were later expansions from the Carpathian Basin. While the central European early Bronze Age cultures were still predominantly of local origin, the Middle Bronze Age Tumulus culture has been regarded as the result of a push from the Danubian area.

Actually, the fact that Bell Beaker proper was so weakly adopted in the Carpathian Basin may have preserved the R1b derived IE language there (probably Italo-Celtic). While those groups which were more strongly affected by Bell Beaker culture, may have adopted a Basque-related language. The very high incidence of R1b in Basques is often interpreted as a testimony to the victorious, penetrative force of Italo-Celtic R1b males, who were only eventually tamed by Basque women. However, it should rather be interpreted as what it is: The fact that some R1b males were assimilated by Basque culture.

If the early R1b expansion in western Europe was associated with Basque-related languages, only to be marginalized to the fringe during the Bronze Age proper, this would explain the patterns of R1b vs. I2b and of R1b vs. IE language in the maps I linked above.