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Monday, August 24, 2015

Pre- and Post-Kurgan Europe


The Principal Component Analysis (PCA) below is based on four sets of D-statistics. The second image shows what they are and how they affect the components. The datasheet is available here. If you don't know what EHG, SHG and WHG stand for, see here.

Note that the post-Kurgan Europeans are shifted east, towards the Bronze Age steppe groups (most of which are in fact classified as Kurgan cultures), relative to the pre-Kurgan Europeans. Coincidence? Certainly not. Interestingly, the West Asians show a similar shift to the east, although it's not yet clear who caused it and when.


In this analysis I used samples from the Allentoft et al., Haak et al. and Lazaridis et al. datasets, all of which are publicly available. The latter two are found at the Reich Lab site here.

Update 12/09/2015: Matt posted these graphs in the comments. The first graph shows Yamnaya-related ancestry proportions for a series of points along the Yamnaya-Middle Neolithic continuum, which can be used to estimate Yamnaya-related admixture in samples that cluster near these points.

See also...

Smarter than the average bear

248 comments:

1 – 200 of 248   Newer›   Newest»
Simon_W said...

The change towards ANE in Anatolia means either that the Hatti were not descended from the early Anatolian farmers and that they instead moved in later from the east - or that the Anatolian IEs did. At least physical anthropology seems to favour the second hypothesis, as the arrival of the Hittites was marked with a change in cranial metrics, whereas the previous inhabitants of central Anatolia were similar to Danubian farmers. (Further to the east there were also other factors like the Hurrians with their possible Caucasian connections and the Indo-Aryan Mitanni.)

Davidski said...

Needless to say, I can't wait to stick many more ancient samples on this plot, like Barcin, Kumtepe and the Vucedol R1b guy. The Vucedol sample is said to be getting genotyped at Harvard.

DMXX said...

Regarding West Asia, I anticipate the future pre-IE samples (excluding those from western Anatolia) already were more eastern-shifted than the pre-IE Europeans. South-Central Asians probably also resided in a similar part of the west-east cline as their modern representatives.

The overall plotting here is quite reminiscent of what an ANE or EHG (x-axis) vs. "pure" WHG (y-axis) graph would look like.

Aram Palyan said...

Simon if there is some rational in connecting Abkhaz-Adygean to Hattic then of course Hatti people should have high ANE.
But then again the same question. How to discern who is non-IE Hattic who is Hittite.
In Steppe theory Hittites should have some extra EHG.

Matt said...

Yeah, more or less. PCA on two variables is always basically like a rotation so the axis of correlation between A and B is the X (or Y) axis, and the remaining "lack" of correlation is the remaining axis, and SHG and EHG are pretty close enough to linear combinations of WHG / ANE (there's some difference, but not a lot - an interesting one is that when you plot the EHG vs MA1 stat alone, the Scots Argyll has a little more MA1 affinity than would be expected for its EHG affinity compared to other NW Europeans, showing a "Gedrosia" thing again, but that's not a very large deviation in absolute terms).

Since you mention pure measures, the D stat measures here are sort "pure" measures. Depends on what you mean by pure, as BedouinB is the "floor" for both stats here, and is not neutrally related to EHG+WHG, but I think more related to WHG.

So if the stat D(Ju_hoan_North,WHG)(BedouinB,X) is lower than D(Ju_hoan_North,EHG)(BedouinB,X), e.g. like for Georgian, that does not necessarily mean X shares less absolute drift with WHG because BedouinB is a floor closer to the ceiling for WHG.

If you were running a stat like D(Ju_hoan_North,WHG)(Dinka,X) / D(Ju_hoan_North,EHG)(Dinka,X) that might provide a more pure measure, since Dinka should be symmetrically related to EHG and WHG, as a number, although the relative positions when plotted would not change.

It would be interesting to see where the Sweden MN sample sits on this plot as well.

Maju said...

Those are nice interesting graphs, although sorta deja-vu admittedly. I do miss Gokhem, which is the only Atlantic Chalcolithic reference so far and an important one in my understanding.

"West Asians show a similar shift to the east"

That's not really clear, I must say. On one side the ancient references (except Ma1) are all European and can hardly be applied at all to West Asia. We barely know from there that some West Turkish early farmers appear to be somewhat similar to EEFs but we don't yet know the fine detail nor have other ancient West Asian references that allow us to issue judgment.

On the other side the bulk of West Asians (and Spaniards by the other side) fall roughly around the Spain_MN - Sicily line, which goes just over the Iranian sample, what is strongly suggestive of a Mediterranean specific axis of admixture that is not related to the Kurgan one and should have other origins (be them Phoenician, Roman or older, like the proposed Chalcolithic civilizational colonizations hypotheses that still lurk around). If we shift the axis to overlap with the Balcanic cluster (rather than Sicilians), then all West Asians fall to the left of it (only marked exception are Lezgins, who are actually Europeans by geography and in any case exceptionally deviated relative to West Asians). Obviously this second axis should have some Kurgan influence (~35% CW-like vs 65% Spain_MN-like in Greeks, Albanians and Bulgarians), so the truth is somewhere in between.

What is clear is that (Lazaridis dixit), Sicilians cannot be modeled as {WHG+EEF+ANE} and show instead a strong distinctive West Asian deviation. So at the very least everything that falls on or left of the Spain_MN-Sicily axis in West Asia (i.e. Iranians, Armenians, Adana Turks, Druze, etc.) cannot be modeled with Kurgan admixture in mind. What falls right of it might or not have Kurgan admixture but the issue still needs clarification. Obviously the North Caucasians (who are not technically West Asians and are a bit over-represented) do appear to have more Kurgan-like admixture but it is not wholly clear what caused it (if it is Kurgan or pre-Kurgan Northern Euro-Asian influences/substrate in general.

It is notable that the EHG and Ma1 notational axes that you so kindly annotated are barely differentiated. However I'd dare to say that the same that EHG is relatively stronger than Ma1 in East and North Europe, Ma1 seems stronger than EHG in Central Asia and the Caucasus (and for whatever is worth, in Anatolia), what seems to make good sense if we don't attempt to interpret these data in an Eurocentric way.

Maju said...

@DMXX: "The overall plotting here is quite reminiscent of what an ANE or EHG (x-axis) vs. "pure" WHG (y-axis) graph would look like".

That's only the second (horizontal) dimension. Notice that the plot is artificially compressed vertically to enhance that dimension but that in fact the first (vertical) dimension is always more important.

If you measure them, you notice that 0.016 in the D1 axis is only 60 pixels, while 0.012 in the D2 axis is 107 pixels. If anything the distortion should favor the D1 axis, which always weights more by definition (unsure how much more though). So there is a very strong distortion that aggrandizes the WHG-EHG axis and minimizes the West Asia - Europe one. IMO this is a conceptual error, a bias, that can cause much confusion potentially.

Krefter said...

@Maju,
"On one side the ancient references (except Ma1) are all European and can hardly be applied at all to West Asia."

There could have been closely related people in West Asia.

Maju said...

@Krefter: Could, might, maybe...

Excepted EEF for a particular West Turkish case (whose details are not yet published and could well be significantly different from this preliminary assessment) we don't know anything like that. Wild speculation and the very similar wishful-thinking doesn't help.

Personally I think that modern Iranians seem pretty much baseline for "Highland West Asia" but, until we get further info, this is also a speculation (although at least based on some data such as the Sicily-Spain_MN line I mentioned before or the Raghavan measures of ANE, which also converge towards reading Iranians as "Zagros baseline").

Davidski said...

Maju,

I can tell you that Starcevo_EN isn't much different from West and Central Anatolian Neolithic farmers.

So that should give you an idea of the scale of the genetic shift in much of Anatolia.

Davidski said...

By the way chaps, I've added Sweden_MN to the plots. This includes all of the Swedish MN samples currently available, but is mostly made up of Gokhem2.

Aram Palyan said...

Well if the Anatolian farmers come out similar to European Neolithic farmers then the Basque language connection to Armenian (trough Urartean), Georgian ( and other West Asian isolates ) will get some basis.

Mike Thomas said...

Aram

It will only get some basis if the tide of linguistic opinion swings back to that direction, one (afaik) which has been rejected.

That farmers from the South Caucasus, Anatolia and europe were genetically similar doesn't mean they spoke the same languages. You're making the same cardinal errors that everyone else does with the entire EHG/ R1= Pre-PIE nonsense

Aram Palyan said...

Mike

I agree with You and this is because I used the word some basis.

I do believe that before the apparition of complex social structures and some forms of states the languages were not stable and had tendencies to easily drift from each other. So I don't believe in linguistic continuities over vast areas in Palaeolithic and even Early Neolithic.

But nevertheless some cognates are real. I consulted a linguist who has researched this story. He thinks it is perhaps related to some hypothetical Mediterranean substratum.

Mike Thomas said...

Ok then. It would be interesting to see new takes on such connections

a said...

Thanks for the nice graphs.Perhaps in future you could add the Samara H.G.[pre-Kurgan Steppe sample;since it was found in the same region/location[1.5km+/-]as some Yamnaya samples.
Adding age would also be of interest.
For example age difference=
Yamnaya-SamaraH.G.-2600+/-
Yamnaya-Sintashta-1000+/-difference
Yamnaya-Khvalynsk culture-[0-2600+/-]taking in account possible stratified burials within region.
Yamnaya-Corded Ware-500+/-
Untested but future potential samples from groups like the I.E. tribes like Oscans/Osci- buried beneath 3-23 meters of volcano fallout.[L51+ or Z2103+?]>79.A.D.

"Evidence

Oscan is known from inscriptions dating as far back as the 5th century BCE. The most important Oscan inscriptions are the Tabula Bantina, the Oscan Tablet or Tabula Osca[5] and the Cippus Abellanus."

Simon_W said...

@ Aram

Not necessarily! If the hypothetical proto-language behind Hattic and Northwest Caucasian was spoken by ENF-like people without ANE and the ANE in the Caucasus is from an older hunter-gatherer substrate population, then Hatti people needn't have had ANE. One hint that this may be true: yDNA haplogroup G2a was associated with early farmers who had no ANE, but it's also very common in Northwest Caucasians.

But yep, according to the steppe theory Hittites should have some extra EHG. But in my opinion this may be fairly low, like in the Bronze Age Armenians. If the ancestors of the Anatolian IEs came from just north of the Caucasus, then they may have been more like North Caucasians than like Yamnaya people or modern Europeans. It's clear, if the Anatolian IEs came from the Balkans, from an early pre-Yamnaya steppe expansion, as envisaged by David Anthony, then they must have been a thin elite who hardly influenced the DNA of Anatolia. I think that's the most common notion of what the Hittites were like: barely IE, just linguistically assimilated, and with a strong substrate influence in their language. But the more I think about it, the more it makes sense to me that it may have been quite different: That the Anatolian IEs reshaped the genetics of Anatolia quite considerably.

Of couse this applies only to Anatolia proper, in other parts of Asia there was definitely old, non-IE ANE.

Maju said...

"By the way chaps, I've added Sweden_MN to the plots".

Thank you, David. That helps because the Sweden_MN - Corded Ware line leaves only the two Baltic samples "above" it, allowing to perceive better the process of admixture for most cases in terms of simple Neolithic-IE. It's also almost parallel to the Dim2 axis, what seems to qualify this axis in such terms of (late) Neo-European vs. Eurasian Steppe (IE).

Chad Rohlfsen said...

Gokhem samples aren't Atlantic Chalcolithic. They're from an island in the Baltic Sea, and northern Funnel Beaker has roots in Northern Germany and Poland. It has nothing to do with the Atlantic. Also, as David said, EEF is basically identical to everything that stretches near the South Caucasus region. There was a drastic shift in West Asia, probably linked to Afro-Asiatic expansions. This continued beyond the Iron Age, as we see Armenians go from North Caucasus like, to more Balkan like, then to their current make-up, after the Iron Age.

Matt said...

It's tough to know how representative any of Baalberge, Remedello, Spain_MN and Sweden_MN, each were of the general run of European Middle Neolithic / Chalcolithic populations.

Collectively, I think viewed through this prism you could make a case for Northern and West Europe to have been mostly around the average of where Spain_MN, Sweden_MN and Baalberge MN were (properly I don't know if this would really be an Atlantic phenomenon as such), prior to IE. Then they are affected by IE and in some regions pan-Mediterranean gene flow via the Roman Empire (and probably to a lesser extent later empires). Not a very strong case though, as so much is hard to know for sure of the ancient demography with these samples (representativeness).

I think the resemblance to the ENF, WHG, ANE PCA is really striking in any case. The symbol and colour scheme really makes the MN European cline clear as well. I added in the Hindu-Kush region samples that David included in the datasheet he posted up in the last comment thread as well http://i.imgur.com/qom5d1L.png, as they have an interesting position to compare. However this kind of PCA on these four variables can't distinguish between ENA related shift and Basal Eurasian related shift, as they both have effect the variables under test in an equal way (not that Basal Eurasian and ENA admixture reduce affinity to WHG and EHG to the same degree, but they do not affect either WHG or EHG in different proportions, being both equally related to both).

Mike Thomas said...

Very nice Matt. But I'm not sure why mesolithic samples keep being excluded

Maju said...

I am so uncomfortable with the distortion of the graph in favor of dimension 2 and against dimension 1, that I had to re-distort it into a minimally compensated one, what implies a 4x expansion of the vertical axis (at least, but I don't know the datum of each component's exact weight). On top of it I added the best triangle of admixture I can fathom, which has vertexes at Sweden_MN, Starcevo_EN and Yamna (if I use CW, Balcans get excluded and per Lazaridis analysis they should not): → LINK.

I also estimated the 50% admixture cuts, resulting in three annotated "thirds": Atlantic (Basques and, less markedly, other West-North Europeans), Mediterranean (most Iberians, Italians and Balcanic peoples) and NE (Sámi and Kargopol). Estonians and Lithuanians (as well as Corded Ware) fall above the triangle limits but that makes some sense because that area was never affected by the EEF replacement and instead seems to have extra East European and Siberian "HG" blood.

The most important issue I'd like to underline with this is anyhow that the internal "Neolithic" European variation (mostly defined by greater or lesser WHG admixture) is at least as important as the variation between these and Yamna, and should not be ignored nor trivialized because it is quite apparently a very important element in the formation of the European peoples.

Mike Thomas said...

I think Albanians are an interesting vignette for southeastern europe. If Remedello can be roughly representative of Copper Age southern Europe (although I know we have CO1), and if Albanians can be considered to be representative of pre-Slavic Balkans (although they too absorbed considerable slavic admixture), then the case in southern -Central and southeastern Europe appears to be one of pure Eastern shift, rather than northeastern. In which case, we still need to define the exact population which caused the eastern shift in SEE.

Alberto said...

Yes, usually the graphs exaggerate the east-west shift compared to the north-south. Probably because Europeans don't have that much difference in ANE (most of them fall in the 10-15% range, which is a pretty small difference to make a visual impact).

BTW, here is a spreadsheet with IBS sharing with Loschbour, MA1 and Samara_HG, based on figures that David provided long ago. Results are normalized in %, taking Masai as the baseline (0%) and whichever population has the highest sharing as 100%. Unfortunately only modern populations are included (220 from around the globe. Maybe I could remove most and limit it to West Eurasians if that's useful for anyone).

https://docs.google.com/spreadsheets/d/1za0ptfjF_TgrZGdtw_zaYWTQVsw3c-qCK88xwaariaQ/edit?usp=sharing

Alberto said...

@Mike

Yes, that's an interesting question. It's been speculated that the eastern shift in SE Europe might be due to 2 (or 3?) different populations: First Yamnaya, shifting them north east, then from West Asia shifting them South East. And maybe finally Slavs shifting them north.

But the thing is that we don't have much evidence so far of big Yamnaya impact in SE Europe. Up until now it just didn't show up in Copper Age samples. And if the population that shifted them east was like modern West Asians (say Druze), then they should be much more southern (unless that's how they were before the Slavic migrations). But the third possibility would be if SE Europe was shifted east by a "teal" population (that would cluster around Lezgins/Tajiks).

I hope that the Bulgarian and Greek samples that are said to be coming will clarify this.

Matt said...

Mike: Very nice Matt. But I'm not sure why mesolithic samples keep being excluded

My thinking is, IIUC, if you want to include the Mesolithic samples, then you need to be able to first leave some of them aside to do so.

To explain, if you went D(Ju_hoan_North,WHG)(BedouinB,Loschbour), and WHG included Loschbour, La Brana and KO1 all together, then you would get a weird result, inflating the similarity of Loschbour to "WHG" as the measure would be confounded with the sample's similarity to itself, which would be perfect.

You could substitute for this by sort of measuring say D(Ju_hoan_North,La Brana)(Bedouin,Loschbour) and using that as a substitute for D(Ju_hoan_North,WHG)(BedouinB,WHG)... but that's arguably an apples to oranges comparison if everyone else is really tested as D(Ju_hoan_North,WHG)(Bedouin,X), so I could understand why David might not include that.

Mike Thomas said...

I agree Alberto, conditioned by two major facts.

1) the slavic impact was massive , and felt all the way to southern greece. So the pre-Roman balkans must have been more southern in certain areas.

2) the LBA Montenegro indeed looks kinda "Spanish" ( if I recall David correctly).

3) some mediterranean admixture mihjt have confounded more souhern mixture, howeeve, eg soldiers from the Greek East, Armenian settlers, etc.

So there was definitely some Yamnaya impact, but this was after ~3000 BC; as per the palaeogenomics and radiocarbon datings.

Clouding the issue more, however, is the recent emphasis of Balkan scholars for what Id term the 'ante-Steppe" (not anti-Steppe !) path: Caucasus to Balkans via Anatolia. (Ivanova calls it the "royal road".) so this might have been an additional pathway of ANE-like expansion to the balkans; perhaps from a "Teal sans Yamnaya " group.

In fact, I hope its not too blasphemous to suggest that even some Z2103 arrived that way.

Mike Thomas said...

Ok thanks Matt.
I just thought it completes the overall picture. without considering the technical problems .

Krefter said...

Samples from Hungary or peninsula Italy might not be representative of people who lived in the Balkans in the same era. Balkans might have been a differnt world than the rest of Europe(EEF/WHG world) and had constant gene flow back and forth with the Near East.

Mike Thomas said...

Yes of course. Possibly it's own microcosm. I guess we'd need a diachronic, region-wide sample set.

Matt said...

Mike, yeah, ideally you're right and it would.

Alberto, with the IBS here's some quick correlations between the types of stats that overlap between the two sets:

http://i.imgur.com/v5RqO0w.png

The correlations are pretty tight between the relevant D-stat and the relevant IBS stat. Tightness of IBS to D-stat is WHG>EHG>MA1. The pairs appear to deviate most for MA1 over the Hindu-Kush / SCA populations where the IBS measure is a good deal higher than would be expected based on the D-stat measure and D-stat to IBS correlation (for whatever reason).

You can use the regression equations from the correlations above (slope + intercept) to normalise the IBS to the same scale as the D-stat and then they PCA all together nicely as follows (this is 99% of variance together, so the other PCA dimensions are not important to these measures):

http://i.imgur.com/OzLWla4.png

Krefter said...

@Matt,

Awesome visuals. If I gave you mtDNA data in the future could you make PCAs and other visuals?

Davidski said...

Based on that talk on the Kumtepe genome given recently, it doesn't sound like Late Neolithic Western Anatolians were much different from Copper Age Southern and Central Europeans.

Krefter said...

@Davidski,

Are you getting inside information?

Davidski said...

I wouldn't call it inside information. It was a public talk and a number of people were there.

Davidski said...

Indeed, in regards to the Balkans being something unique during the Bronze Age, I also don't think so. This is where a Bronze Age Bulgarian clusters on the usual type of West Eurasian PCA (just west of Tuscans)...

https://lh3.googleusercontent.com/asUl7MkRUPGCiHyQUKUl0f_UWRcEQzxVOi8XiwnjiVo=w970-h631-no

Source:

http://docslide.us/science/decoding-ancient-bulgarian-dna-with-semiconductor-based-sequencing.html

So Bronze Age Bulgarians weren't all that different from Bronze Age Hungarians, just more southern.

That sample does clearly have some ANE, because it's shifted east of Sardinians, but it obviously doesn't look very Balkan. That's probably because it lacks the West Asian proper admixture that arrived in southeastern Europe during the Iron Age and early Medieval period.

For comparison, a similar type of PCA featuring a variety of other ancient samples:

https://lh3.googleusercontent.com/VwWhxXrYo8Ent1scgWhVhhEsEfTFJbD589V0Wb8kM2U=w879-h631-no

Gaspar said...

@mike Thomas

You need to study your geography, Remedello culture sits in northern Italy ( Brescia )not far from the admixture marker called Bergamo. It is no-where near southern Europe , its classified as central Europe.
Remedello is closer to Bavaria than Albania.

Check you latitude and longitude maps

cheers

Mike Thomas said...

@ Gaspar

Did you even go to school ?

Because I always learned that Italy is geographically and culturally part of "Southern Europe" ?

https://en.m.wikipedia.org/wiki/Southern_Europe

Certainly, no source or definition ever includes it (even *northern* Italy) in Central Europe

Sorry if that offends your chauvinism.

Fanty said...

Hm Yeah,
Italy as a whole is in "Southern Europe" no matter what definition.

Even if one uses the definition that the Alps are the mountain range that divides Europe into Northern and Southern.

France would, by this definition be a hybrid. Northern France beeing in Northern Euope and Southern France beeing in Southern Europe.
That definition would also include the whole Balkan peninsular into "Southern Europe".

Mike Thomas said...

Definitions vary, and are based on economy/ political and cultural factors.; not just geography.

Eg Croatia and even other former Yugoslav countries are sometimes considered "Central European" because they are not really in eastern Europe, not are they mediterranean.

The case of Italy, however, is more cut and dry.

Alberto said...

Thanks Matt. Indeed the correlation is pretty tight between D-stats and IBS, probably because HGs behave in a very stable way in D-stats. They seem to be quite neutral to ENA ans BEA. D-stats only become problematic with populations that have one of those compared with other that has the other.

@David

Thanks, that sample seems to be P192-1. I think that's a real Thracian sample, from the Iron Age (800-500 BC). It does seem to have a low amount of ANE (8-10%?), but the shift is clearly different from the Hungarian Bronze Age ones. The Hungarians seem to have Motala-like admixture, while this Thracian is shifted towards the Caucasus.

I hope they're resequencing the other Thracian samples too. They were too noisy to really know how they looked like.

Mike Thomas said...

@ Davidksi

Thanks for that link. But I can't access it with my iPhone (im currently travelling around europe) . Is it a new study/ different samples from that of the Basamante lab?

@ Alberto

Motala -like admixture in Hungary BA? I know we've discussed this before, but what does that mean to you ? I doubt actual Scandinavian foragers :)

Matt said...

Krefter: Awesome visuals. If I gave you mtDNA data in the future could you make PCAs and other visuals?

Hah, they only looked OK because I was freeloading off the colour scheme in Davidski's datasheet. I'm usually too lazy to make them look OK (although it doesn't take that long). The basic graphics are really easy to produce though, the Past 3 software Davidski has linked before is pretty easy to use - http://folk.uio.no/ohammer/past/.

It's a little 5 meg program, just stick in the tables of mtdna frequencies like its a spreadsheet and then run some analysis functions and you're ready to go. Check out some of the datasheets Davidski has stuck up for examples.

Here's an example where I took the data from the Supplement 3 - Table 12 http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4039305/ from the "Ancient DNA reveals key stages in the formation of Central European mitochondrial genetic diversity" Brandt 2013 paper - http://i.imgur.com/1rINiOL.png. These took a matter of minutes to make and running the PCA or clusters on these tables takes seconds. Another example using frequencies from the website Eupedia - http://i.imgur.com/N5Mbj0Y.png.

Gaspar said...

@mike Thomas

You should refrain from using modern national borders with ancient cultures. You do realise the people living in the bronze age of Italy are not the same as the people of today, I hope you realise this.

Anyway, the Altheim, Mondsee and Pfyn cultures of Austria are identical in goods, daggers etc to the Lagozza and Remedello cultures of Italy...........is Austria also southern Europe?

kind regards

Alberto said...

@Mike

By Motala-like I just meant that they are shifted very north, but not very east. They have a huge amount of WHG ancestry and a low amount of ANE (like 60% WHG, 10% ANE). It's a mystery at this point where they came from, but the pattern looks like EEF + Motala (whether that's the actual mixture or not is a different matter, but we don't know for now).

Mike Thomas said...

Yes I see. Perhaps it's not "motala like" at all, but simply high WHG-containing EEF (indeed like much of MN Europe) with minor Yamnaya admixture; from the kurgans that appeared East of the Tisza?

Mike Thomas said...

@ Gaspar

Your comments are pointless

Southern Europe is southern Europe, chronology is irrelevant.

Anyhow, Remedello was even more ""southern"" back than compared to modern Northern Italy, and not northeastern shifted like its contemporaries in Central Europe (Germany) .

It's no surprise we'd see analogies either side of the alps, but that doesn't mean northwen italy was 'central european' in the copper age, anymore than the presence of Mycenean- like objects in Scandinavia making Sweden to be Aegean.

Maju said...

@Alberto: "Yes, usually the graphs exaggerate the east-west shift compared to the north-south. Probably because Europeans don't have that much difference in ANE (most of them fall in the 10-15% range, which is a pretty small difference to make a visual impact)".

Well, that's precisely my contention: that the Kurgan (or ANE or Yamna) admixture is being hyped at the expense of an even more crucial element in the European demic genesis, which is the variation among Neo-Europeans (populations of Neolithic affinity, which includes pretty much all modern SW Europe as well). Once we correct that distortion, we can see that the Stuttgart-Gokhem differential is brutal: at least as large as the Gokhem-Yamna one, probably quite larger in fact.

This is no trivial matter at all, and triviliazing it by means of distortion of the graphs is not helpful at all.

Maju said...

@Mike: what do you make of Hungary_BA being so extremely non-Yamna, when they "should" in your paradigm of obligated demic replacement? Or will you tell me that Vucedol (and hence, in your paradigm, all Balcanic Kurgan-derived cultures) were not actually Kurgan/IE either? If so, where do ancient Greeks come from?

Davidski said...

I rotated the graphs/plots to fit geography, since the focus here is on the Kurgan expansion into Europe from the east, rather than the Neolithic expansion from the south.

The source of the eastern genetic shift across Europe associated with the Kurgan expansion, which, by the way, was massive and doesn't need to be exaggerated, was not geographically close to Western or Central Europe, and this is what the plots are designed to highlight.

Mike Thomas said...

But call it what you will, I was merely using relative terms for sake of comparisons

Matt said...

Maju, I agree with your contention that PC1 really should be the long axis, even though that doesn't fit as nicely on most screens.

For the relative size of each, it's true that in this datasheet, PC1 is 86.5% of variance vs PC2 12% of variance.

On the other hand, bear in mind, this analogous genotype based PCA that Davidski has posted before, which has a similar form to these PCA. https://drive.google.com/file/d/0B9o3EYTdM8lQcVNqUDBMZUt3WkE/view?pli=1 - PC1, which spans in that version all the way from BedouinB to Loschbour (so a longer PC1 than here, which only goes from Druze to Lithuanians-Yamnaya) is only twice as large as PC2 (0.99% vs 0.45%). So with this variability I think it is not worth arguing about which shift is more important, we can only say that seems like both are of some importance to distinguish present day Europe from Early Neolithic.

Maju said...

@David: I'm not blaming you. At least thanks to the figures you included in the graph's axes, I could easily spot the distortion. It's a problem that seems generalized: one aspect (the Kurgan admixture) is made very much central (and it is of course important) and the other aspect is being just not addressed and even dissimulated. It's not just this graph or analysis: it's widespread.

I just perceive that, assuming that the Yamna-Gokhem and the Starcevo-Gokhem axes of admixture are equal, the vast majority of modern Europeans (i.e excluding Eastern exceptions) are distributed through most of the Starcevo-Gokhem axis and only through a small fraction of the Yamna-Gokhem or Yamna-Starcevo ones, and always tending to the Neolithic edge. This is very different from the "Yamna (or Corded) replaced them all" perception that has been stated sometimes and, even more importantly, from the "all Neolithic Europeans were nearly the same" one. The reality seems rather that (1) Kurgans had "only" a 20-40% influence, depending on which population, and (2) the variation in the Atlantic-Mediterranean axis is much more important Europe-wide, implying a major shift in the Chalcolithic or even Late Neolithic, surely associated to Megalithism, prior to that Kurgan/IE impact.

Maju said...

@Matt: "For the relative size of each, it's true that in this datasheet, PC1 is 86.5% of variance vs PC2 12% of variance".

Wow! That's much more than I could expect. After knowing that even my version of the graph is still heavily distorted in favor of the Yamna exaggeration (as it would roughly reflect a 50-50 or 55-45 apportion) The reality truly does not fit most screens vertically, as it's 16x taller than the original graph and 4x taller than my version.

I'd like too see the properly proportioned plot for the record anyhow. It's the kind of image that speaks volumes.

Maju said...

I correct: 32x taller than the original graph and 8x than my version (approximately).

Davidski said...

Maju,

The ratio of variance carried by component 1 vs component 2 has no relevance to the amount of steppe admixture in Europe.

Even if this ratio was 99% vs 1%, it still wouldn't change the fact that there's around 50% of steppe admixture across much of Europe that only arrived there after the Middle Neolithic.

What you're actually focusing on is the fact that there's a lot more difference between WHG and EEF, than between EEF and steppe ancestry. But no one ever said that there wasn't.

Maju said...

Well, nevermind, I tried at home and it becomes almost a line between Iraqi Jews and Lithuanians. Europe-wise seems to be almost all structured along this single axis of EEF-HG admixture with only some of the ancient Kurgan samples deviating somewhat to the right. Impressive to watch anyhow because it makes all this Kurgan issue seem almost like "splitting hairs" and maybe explains why characterizing the Kurgan admixture is a bit complicated sometimes.

Davidski said...

We're not splitting hairs.

A new population moved into much of Europe after the Neolithic and caused a major population turnover.

The fact that this population wasn't very different to what was already there doesn't lessen the impact of the migration.

I think what you seem to be missing in this discussion is that when you squeeze the plot horizontally the Bronze Age steppe groups are also affected by this, so that their positions relative to everyone else don't change, thus the impact of their migrations doesn't change.

Maju said...

@David: "Even if this ratio was 99% vs 1%, it still wouldn't change the fact that there's around 50% of steppe admixture across much of Europe that only arrived there after the Middle Neolithic".

Rather like 40%-20% (depending which population) but I concede in the essentials, yes.

What caused my astonishment was that even more distorted graphs were used in Haak 2015, what led some to argue that there was almost no EN/MN change (i.e. very low change attributable to the Atlantic-rooted secondary migration) and actually, when we properly apportion the graph, we can see that it is many times as important, in absolute terms, as the Kurgan-Gokhem differential (40% of very little, 12%, is even less very little: 5%) and therefore something very very important that we cannot just ignore.

My point is that the "correctly apportioned" graph shows a huge variance in Neolithic Europeans (and their descendants, i.e. us) that is not being discussed. And it should.

Prior to Kurgan impact but also afterwards, this variance explains the Basque-Andalusian variation and underlines it as very large (for European ranges), and that same variation (in very similar amounts in spite of the larger geographic distance) is what makes apart Czechs or English from Bulgarians or Greeks for example. We are talking after all of >80% of the variance including West Asia and maybe 60 or 70% when only considering Europe.

You may consider it a less interesting issue but I don't see why it would be.

Davidski said...

What you're talking about is the WHG rebound that happened from the Early to Middle Neolithic.

This was discussed in Haak et al., but we'll need more samples from across time and space to discuss it in more depth.

However, the post-Middle Neolithic population movements from the east had a bigger impact on European Y-chromosomes, linguistics and culture, that's why they'll always take center stage in this context.

Mike Thomas said...

@ maju

The 10-20% of ‘ANE’ in BA Hungary accords exactly as I anticipated. Before I explain further, you really need to grasp a couple of basic, preliminary facts (that you have thus far failed ☺ )

1) There was no Gimbutas # 1 “Kurgan” wave, which I believe she dated to c. 4500 BC. She ascribed – incorrectly – the end of Neolithic Balkan cultures like Vinca to this ‘1st wave’. Rather, the changes are reflective of internal changes within the Balkans, and broader late Neolithic Eurasia, as well as certain social tensions. Some of the ‘internal changes’ were a shift toward more dispersed settlements, more pastoralism, more mobility, the wheel, etc. These were widespread changes that weren’t introduced by people from the steppe, but had their roots in the Late Neolithic socieites in which they occur, and diffused so widely and rapidly that any search for specific point of origin impossible. We also know that these changes weren't caused by steppe invasions because (1) there is no evidence for Yamnaya in Balkans and central Europe before 3000 (based on hundreds of radiocarbon dated kurgans) and (2) the chronology of changes occur from west to east (ie the abandonement of tells, etc, occurs in northern Serbia (4500 BC), then Bulgaria (4200)) (3) The late copper age Hungarian has no ANE/ Yamnaya admixture. (4) Such apparent “Kurganization” is even seen in Italy – but here no ANE/ Yamnaya admixture even as late as late M3. And your idea of an ethereal, extra-corporeal, non-tangible Kurgan migration is obviously the most stupid thing you've ever said.

2) Kurgans west of the Black Sea steppe only appear after c. 3000 BC (to re-iterate) . We are talking about Hungary, northern Bulgaria, etc. So there is clearly a hiatus of several hundred years.

3) Kurgans in Hungary occupy a very specific part – the steppe region east of the Tisza. The rest of the Carpathian basin is occupied by a variety of cultures which show continuity with the preceding Copper Age. So, indeed, the 10-20% ANE admixture in BA Hungary makes perfect sense to me.

This is the genetic / demographic situation. I will for now abstain from the “IE question”, because I think it was a far more complex issue than arrival of ANE or Yamnaya, and unlike you, and indeed most people, I like to treat different modalities of evidence separately, and not engage in circularity. But as for where did Greeks come from,- my honest answer is im not sure. But I can imagine that movements from the Caucasus region also reached the southern Balkans at some point in the Bronze Age, before 1600 BC. Whether this was the steppe route or the Anatolia route, I really don’t know.

Mike Thomas said...

@ Maju

Almost forgot about northern Europe.
Having lesser density , movements from the east had greater impact, naturally. But I'm still not sure about corded ware being "derived from Yamnaya". At best, it's a "kurgan inspired culture", but I suspect it has its own, separate genetic roots; from the forest-steppe Carpatho-Ukraine, and not the Caspian.

As for R1b-M269 and Western Europe, I'm leaning toward Carpathian groups spreading it up the Danube. I'm not sure of exact chronology, because no one is .

Davidski said...

Carpatho-Ukraine is too close to the Carpathian Basin for Corded Ware to have come from there.

There's no reason why there should be such a massive divide in genome-wide structure, mtDNA and Y-chromosomes between western Ukraine and Hungary well into the Bronze Age. Saying the Carpathians were a barrier isn't a very persuasive argument.

Corded Ware obviously came from somewhere around the Don.

Mike Thomas said...

Well it's certainly possible. But they sampled all the way to the Don (Allentoft), didn't they? All R1b apart a later Mezhovskaya sample.

Matt said...

Maju: Wow! That's much more than I could expect.

An IBS based PCA based on those three IBS variables (MA1, Samara, Loschbour) Alberto posted up that maps pretty well to these, actually has exactly the same pattern for the overlapping populations, with the first dimension dominating around 86% variance, and the second 13%. So it seems like there is a strong case that with these different Mesolithic HGs (WHG, EHG, SHG), the relatedness to them (either as measured by the D stat or IBS) is very strong explained as fitting on a single dimension, which is 6x larger than another dimension discriminating between different kinds of HG ancestry.

However, from Davidski's other genotype PCA, based on distances of modern populations, from one another and the ancients, these are not so dominated by a single dimension, with the first dimension being only 2x as large as the second one. These plots here is not based on the distances of modern samples from one another, only each of them *to* the Mesolithic samples. So I would be wary of saying these are more correct than those.

Mike Thomas said...

Btw by Carpatho-Ukraine", I meant somehwere from the Carpathians to anywhere in Ukraine (and perhaps more north in Belarus and ME Russia), not the western most Oblasts.

Dmytro said...

"they sampled all the way to the Don (Allentoft), didn't they? All R1b apart a later Mezhovskaya sample." (Mike Thomas)

Has there been Y-DNA sampling of "in between" cultures like Ukraine Corded Ware, Dnipro-Donetsk, Usatovo, Globular Amphora, Western Yamna, Late Trypilia (for starters)? I thought there was still a huge gap between the Don and Central Europe remaining untested.

Mike Thomas said...

I meant all the way backwards from the east. So no, nothing between Vistula and Dnieper yet, apart from mtDNA studies of the Black Sea steppe and some CT samples

a said...


Blogger Gaspar said...

" @mike Thomas

"You need to study your geography, Remedello culture.....

cheers"

By memory I think Po valley/& region stick out as having R1b-Z2103? and or red hair? and or other unique mutations? :)
Perhaps a connection with Afanasevo culture.?

Alberto said...

Probably another issue with European plots is the overrepresentation of some "small, extreme" populations. Like Sardinians, Basques and East Baltic, but maybe also Cyprus, Malta... When looking at the whole European population these are quite a minority, yet they take a lot more space and attention than, say, Germans.

Spaniards, as a bit outsiders too, are also quite overrepresented due to regional sampling.

Once you weigh the populations accordingly, the bulk of Europeans would appear in a north to south cline, with quite small variance in the east-west axis. Spaniards would be a kind of a bridge between north and south, but shifted slightly west from the main cline. And Russians, who are a big group, obviously, and also a bit outsider because of the Asian influences, would fall a bit to the east of the northern group.

So yes, the main difference in most Europeans is marked by the WHG/ENF ratio. All those populations in the main north-south cline are about 12-16% ANE, but the WHG ancestry goes from some 30% to 50%.

But anyway I think we all know this. The numbers are there in any admixture run. And it doesn't change anything, really. Regarding the influence of the Eastern populations that brought ANE to Europe, if they originally carried about 1/3rd ANE themselves, we'd have to multiply by 3 the figures of ANE in modern populations to get the approximate influence they had. And it's pretty big. Taking the 12-16% range of ANE, that would put the impact at around 36-48%.

CroMagnon said...

@ Alberto

"Once you weigh the populations accordingly, the bulk of Europeans would appear in a north to south cline, with quite small variance in the east-west axis. Spaniards would be a kind of a bridge between north and south, but shifted slightly west from the main cline. "

So Theyd essentially mirror the pattern found in earlier european autosomal studies, like Lao etc

Davidski said...

These plots aren't European plots, they're West Eurasian plots, and they mirror West Eurasian plots. See here...

http://eurogenes.blogspot.com.au/2015/07/the-ancient-dna-case-against-anatolian.html

There's no point running European only plots when looking at West Eurasian variation.

Alberto said...

Ok, but that doesn't change the point. If you see this:

https://drive.google.com/file/d/0B2ZfdVZaNXDxT3A3by1hYVl5dEE/view?usp=sharing

The only 2 populations that are big and stay out of that north to south cline are Spaniards (by a bit) and Russians (not by much either, and not due to ANE but to East Eurasian). All the rest are in that cline, except the small, extreme populations that I mentioned above: Sardinians, Basques, East Balts (and these last ones again not by much, and partly also because of East Asian admixture).

Which mostly means that the ANE variation in Europe is pretty small among the large majority of populations (Irish, Brits, Germans, Swedes, Danes, Austrians, Czechs, Hungarians, Romanians, Croatians, Serbs, Bulgarians, Greeks... all have the same amount of it. Frenchmen and Italians a tiny bit less, but still in the cline), and the main difference is the WHG/ENF ratio.

Your last K12 also shows it quite clearly. Almost all Europeans are about 28-33% Afanasievo. If that cluster is about 50% ANE, that's some 14-16% ANE:

https://docs.google.com/spreadsheets/d/1ajolEB_2NXAnxtGJXSbwKSF1CvDvFiGy5otpq5f-C1g/edit#gid=1554639156

Don't we all agree about this?

Davidski said...

Even if I left out some of the modern European groups I'd still be running the ancient populations, and the latter are the real outliers in many cases. So it wouldn't make any difference.

Alberto said...

David, I'm not complaining about anything. I'm just explaining it because of the question that Maju raised about the graphs being distorted.

As I said, this does not change anything, nor it's anything new (the numbers have been out there all the time).

And I'm fine with including Sardinians, Basques, Finns,... but I'm just stating that they have a big influence in the look of the plots when in reality they are a pretty small percentage of the European population. Just something to keep in mind.

Davidski said...

You're making the same argument I was making several years ago, before the ancient samples came on the scene.

I used to think that Sardinians were just a local isolate and superfluous to understanding ancient population movements in Europe. But I was wrong, because West Eurasian dimension 2 is really defined by EN/MN Europeans vs Yamnaya/Afanasievo. So it makes no difference whether the Sardinians are dropped or not.

But the useful thing about Sardinians, and also the reason why they can't be left out of these sorts of analyses, is this:

"The island of Sardinia was indo-Europeanized relatively late by the Romans. Intermediate pre-Roman, but Indo-European linguistic strata are lacking. For this reason, pre-Roman words in Sardinian derive from pre-Indo-European times (Hubschmid 1953: 16)." Page 118

http://www.wuj.pl/page,art,artid,1107.html

Actually, that whole paper is really interesting.

Krefter said...

La Brana-2 had C1 just like La Brana-1. La Brana-1 was negative for a bunch of mutations found in modern C1a2. This makes sense because we have very few examples of C1a2 today, and after 8,000 years modern C1a2s has many ancestors La Brana-1/2 don't.

https://genetiker.wordpress.com/2015/08/27/more-y-snp-calls-for-la-brana-2/

Geneticker has refined the haplogroup label of many Hunter gatherers. I2a1a2a has been found in Motala, PWC, and Hungary(K01+NE7). pre-I2a1b in Motala and Loschbour. I2a1b1a in PWC. I2c2 in Motala. I1 in Baltic Island by Sweden.

Alberto said...

David, you must be misunderstanding me. I'm not saying that Sardinians or Basques are not important for European history, or superfluous.

I was only trying to explain why a graph where the east-west variation is amplified is visually more appealing than one where it isn't. You surely remember this graph:

https://drive.google.com/file/d/0B2ZfdVZaNXDxSmI2S1NNWU4zS2c/view?usp=sharing

So personally I prefer your plots, and I don't want you to change them. But answering Maju's question, yes, the bigger variance in Europeans is WHG-ENF. That's a fact that we all know.

Aram Palyan said...
This comment has been removed by the author.
Aram Palyan said...

Mike

In the link provided by Davidski there is a mention about Hispano-Caucasian substratum.

""The oldest linguistic substratum in southern Europe is the Euro-African substratum. It has been proved to have existed in Upper Paleolithic (Hubschmid 1953: 101). It is
followed by the Hispano-Caucasian substratum, which may have borrowed several
loans from the Euro-African substratum. ""

http://www.wuj.pl/UserFiles/File/Studia%20Linguistica%20129/SLing-129_8.pdf

Matt said...

Alberto: But answering Maju's question, yes, the bigger variance in Europeans is WHG-ENF. That's a fact that we all know.

Yeah, it's correct to say that, from these stats, overwhelmingly there is a single dimension of HG related variation in Europe.

(http://i.imgur.com/BkTUlva.png - for modern Europeans only PC2 which loads differently on WHG-EHG is tiny (4%)compared to PC1 (94%) where they both load similarly and positively).

I might not call that WHG-ENF, necessarily, as that seems like it would imply that the populations who have more HG affinity have a WHG:EHG relatedness ratio inclining to WHG, which by these stats not really.

Agree with Davidski's point that there isn't much use to a European only plot of these stats as it just tells you less than a West Eurasian plot without adding any more information though, even that's not what you're saying, exactly. At least looking at European only samples on genotype PCA reveals trends in genetic drifts that are unrelated to the WHG, EHG, etc. splits (usually better with linked than unlinked PCA) but with these stats you wouldn't get any more information than the whole WE PCA already shows.

Aram Palyan said...

Simon

That theory sounds reasonable. It explains how the highest levels of G2 are associated with highest levels of 'teal' in West Caucasus. It is possible that the high level of ANE was already present in Maykop.

This is an excerpt from Rootsi et al. 2012

""Concerning the presence of hg G in the Caucasus, one of its distinguishing features is lower haplogroup diversity in numerous populations (Supplementary Table S1) compared with Anatolia and Armenia, implying that hg G is intrusive in the Caucasus rather than autochthonous.""

http://www.nature.com/ejhg/journal/v20/n12/abs/ejhg201286a.html

But of course the variance should be taken with grain of salt, because population movements can increase the variance like in the case of Tatars.
This is the variance table. For example Adygheans have very low variance.

http://www.nature.com/ejhg/journal/v20/n12/extref/ejhg201286x4.xls

Alberto said...

@Matt

Yes, I agree, and I prefer a West Eurasian plot with Caucasus, Sardinians, Palestinians, etc... I actually don't disagree with anything that David said.

Regarding WHG-ENF, I was referring to the components more than to populations. It's a bit tricky to know how much WHG came from actual WHGs, and how much from EHG (or even from EEFs or "teal" populations).

The normalized IBS sharing I posted above also has to be understood with its caveats. For example, Estonians have the highest IBS sharing with both Loschbour and Samara_HG, so the value was 100% for both. But that doesn't mean they have the same affinity to both. A stat like D(Mbuti, Estonian)(Loschbour, Samara_HG) has a significantly negative value. Using admixture it's quite tricky to measure too. A cluster with pure WHG and another with EHG also makes Europeans cluster with WHG by far and large. If ANE came exclusively from EHG it would be easier to measure, but since it doesn't, we don't really know how much EHG admixture there really is in Europe, though test comparing directly to WHG show that not much (maybe wrongly? I don't know).

Matt said...

Alberto: A stat like D(Mbuti, Estonian)(Loschbour, Samara_HG) has a significantly negative value

This is likely, but will it vary significantly for Europeans? That's complicated to answer though as well as more admixture with Basal Eurasian would also make that stat tend to 0, even more imbalanced WHG:EHG ratio in the HG part of the ancestry (as BE would be equally related to both).

I think real EHG admixture in Europe is probably significant, but that seems like you can answer it only well through combining our knowledge of the stats with the temporal sequence of pre-Yamnaya steppe-> Yamnaya->Corded Ware as a process of EHG admixture, rather than strictly through statistics alone, due to the closeness (if not identity) of ANE+WHG with EHG.

Alberto said...

@Matt

Here are some of those stats:

https://docs.google.com/spreadsheets/d/1FP-oL5qInnMACDdR_0rF1ooqGiDLXPPlt1FdOmGRIn4/edit?usp=sharing

I think we still don't know exactly why this shift towards WHG happened, but the stats are clear:

Loschbour EHG Yamnaya Chimp -0.0354 -6.545 341312
Loschbour EHG Corded_Ware_LN Chimp -0.0189 -3.32 341342
Loschbour EHG Estonian Chimp 0.0159 3.121 341763
Loschbour EHG Lithuanian Chimp 0.0189 3.687 341763
Loschbour EHG Scottish Chimp 0.0202 3.771 341763

Also as you point out, more Basal Eurasian reduces the difference, with Cypriots having the closer to 0 among European populations:

Loschbour EHG Cypriot Chimp 0.0056 1.097 341763

But yes, I agree this is tricky to measure reliably. Probably we'd need many ancient samples to understand the process.

Maju said...

@Mike: "And your idea of an ethereal, extra-corporeal, non-tangible Kurgan migration is obviously the most stupid thing you've ever said".

Don't Indians and Nigerians speak English? Cameroonians and Senegalese French? That's what I'm talking about. It's quite "extra-corporeal", you have to admit. And it's just the first phase of acculturation with occupations that lasted less than a century in many cases.

A more clear case (because it's already consolidated) is the influence of Latin, which is not linked to any obvious migration from Italy (or anywhere else).

Obviously some people did move, but not enough to be strikingly apparent. Same with Turks, etc. So Corded Ware's rather massive impact seems to be exceptional rather than the norm. It's a piece of evidence that confirms the Kurgan model but not everything Kurgan or Indoeuropeanizing has to imply mass demic replacement, rather the opposite seems to be common, if not "the rule".

"10-20% of ‘ANE’ in BA Hungary"

Not apparent at all in David's graphs here. If Basques have 10% ANE (Lazaridis' scores), that's pretty much like saying "baseline" (Sardinians scored 4-5%, you know, there's nothing lower in Europe). Bringing ANE scores to this is trying to confuse things because there's no "absolute ANE", because the Ma1 sample is way too old to be relevant on its own and because HG pops. other than WHG show a deviation towards it, what adds to confusion. Using Yamna is much better, at least for Europe. Let's stick to Yamna vs "Neolithics": it's much more illustrative.

"Kurgans in Hungary occupy a very specific part – the steppe region east of the Tisza".

That's the early kurgans. But what about Vucedol, centered in North Serbia, which is the relevant culture here?

"We are talking about Hungary, northern Bulgaria, etc. So there is clearly a hiatus of several hundred years"...

You're forgetting Cotofeni. There's no real hiatus. But anyhow I wanted to emphasize Vucedol, which seems to be the cultural context of Hungary_BA and probably originating somewhere else, maybe Corded-related groups.

For me is yet another clear case of conquest by a minority and subsequent assimilation, something like the Slavic expansion of the early Middle Ages in the very same region or Magyars themselves, another success of the "extra-corporeal" invaders, of which we know from history, linguistics, etc. but can't find any genetic signature, maybe a tiny drop or so at most.

"I will for now abstain from the “IE question”, because I think it was a far more complex issue than arrival of ANE or Yamnaya"...

Obviously it was more complex than a linear simplistic genetic approach to the matter. I concur.

...

Maju said...

...

"But as for where did Greeks come from,- my honest answer is im not sure. But I can imagine that movements from the Caucasus region also reached the southern Balkans at some point in the Bronze Age, before 1600 BC".

As far as I know again there is no obvious trail for a second steppe expansion reaching as far as Greece. Even linguistically Greek (or sometimes proposed Greco-Armenian) seems a tad closer to Western IE than to other IE branches, suggesting Central European or at least Sredny-Stog II origins (I personally favor a Vucedol origin, the "megaron" concept is quite telling, although Cotofeni could also do). What we know is that they were still erecting kurgans (tumuli) when they arrived to Greece c. 2000 BCE.

"Whether this was the steppe route or the Anatolia route, I really don’t know".

Neither one seems to be supported by archaeology and in the Anatolian case we have the issue of absolutely no relationship with the Anatolian branch of IE (Hittite, Luwian).

"Almost forgot about northern Europe. Having lesser density , movements from the east had greater impact, naturally".

Agreed that it is a factor, particularly in Poland and nearby areas, where LBK was mostly not too dense.

"But I'm still not sure about corded ware. May best, it's a kurgan "inspired culture", but I suspect it has its own, separate genetic roots; from the forest-steppe Carpatho-Ukraine, and not the Caspian".

Fair enough re. Y-DNA. I would tentatively agree that it is a possibility, judging on R1a evidence. On the other hand autosomal DNA is yelling a very strong steppe (Yamna-like) influence that is mostly or totally new in the region, so... a bit ambiguous. The main characteristic of corded is that it brings more-or-less Eastern genetics to North-Central Europe and in this sense it is the most clear case of a migration with important demic replacement, unlike others.

"As for R1b-M269 and Western Europe, I'm leaning toward Carpathian groups spreading it up the Danube".

There's nothing that can back such speculation. IMO we need to look for the R1b-L11 expansions and origins in Atlantic Europe and hence probably in what David calls "the WHG rebound that happened from the Early to Middle Neolithic". By this I'm not saying that the lineage has to be WHG, it could well be a Neolithic founder effect of course, but it clearly has no relation whatsoever with the Oreintal immigration associated to Corded Ware. All this is particularly true for S116, whose South French origins are very much unmistakable and hence can only be related with pre-IE elements (that area was never a core of IE expansion of any sort, much less one that could affect all Western Europe as S116 does, also many of its carriers were non-IE speakers in proto-historical times, some still are).

Matt said...

@Alberto: Graphing those D(WHG,EHG)(X,Chimp) against the stats in this post - http://i.imgur.com/uFTocOo.png. Weak relationship between "absolute" sharing with MA1 and EHG (as measured against BedouinB), stronger positive relationship with absolute sharing with WHG. This is because in the present day the most HG related populations (Europeans) tend to be WHG related, while the least related (Middle East and West Asia) tend to be a mix of WHG related and EHG related.

http://i.imgur.com/ZkE0vJa.png - this results in D(WHG,EHG)(X,Chimp) pointing weakly north in PCA. Probably would become much more neutral if lots of Yamnaya type samples were included (so overall HG relatedness would become unrelated to the WHG-EHG balance).

Maju said...

@"a" and others who insist on linking Remedello with kurgans. AFAIK there is no such connection: they were more or less directly derived from Impressed-Cardium Pottery culture. An interesting aspect in this development is that in the early Chalcolithic there was a cultural ammalgamation across the Alps (Chassey-La Lagozza) that could well have set the basis for the proto-historical Ligurians, of similar (but shrunk) distribution. Remedello is from a later date but same roots, so I'd argue for a "Ligurian" (and Vasconic) identity and language.

The main (only?) Remedello culture sample is Ötzi, whose genetics are clearly pre-IE.

The IE expansion into Italy only seems to happen later on with the Urnfield culture, which almost certainly brought Italic-speaking conquerors to the area (form Central Europe). Some of these later on (c. 700 BCE) expanded to the central-south peninsula, eventually leading to the Roman Empire. This is a very parallel sequence to what happened in Iberia: early IE arrival (NE) with Urnfields (probably not yet true Celtic, some argue for "Italoid" and include Lusitanian language there) and later consolidation with Hallstatt (early Celts), expanding into Central-West Iberia. The main difference is the (re-)Iberianization of the NE c. 550 BCE, associated to establishment of the Massilian (Phocaean) trade networks, and that Hallstatt had not such an influence in Italy (hence no early Celts there, only later with La Tène).

Maju said...

@Alberto: "Probably another issue with European plots is the overrepresentation of some "small, extreme" populations. Like Sardinians, Basques and East Baltic, but maybe also Cyprus, Malta... When looking at the whole European population these are quite a minority, yet they take a lot more space and attention than, say, Germans".

I am of the opposite opinion. I feel often that the overrepresented ones are Baltic, Finnic speakers and other tiny NE European populations. Or also Icelanders, Orcadians, etc. Which are all as small or even much smaller (and clearly more endogamous in many cases) than the ones you mention. Why Germans should be so important? They (81 mill.) are not many more than Italians (61 mill) or Iberians (57 mill.) The demographic weight of Northern Europe (senso latissimo) was even much smaller historically at the beginning of the Modern Age, with France (smaller than today) alone holding much more people than Greater Germany, which was back then in figures similar to Iberia.

Even by modern figures, excluding the former USSR (Russia and Ukraine have lots of people but are less relevant for this debate), the demographic balance line probably goes through the Alps, more or less, maybe slightly north of them (Switzerland, Austria, Central France). So, even if South Europeans are more or less different, they are still A LOT of people: about half of all Europeans (excluding Eastern Europe).

Also Sardinians and Basques express important polarities of European genetics/demic genesis that should not be minimized or trivialized. For sure they are extreme but that's precisely what polarities are about: the rest are effectively mixtures of these polarities, whose relation with ancestral populations has been by now very much confirmed and is no artifact at all. If we want to use modern populations to describe the European demic/genetic genesis, we can do well using Basques, Sardinians and Kargopol Russians, or maybe adding Lithuanians and Greeks.

Sicilians (and Maltese) are indeed exceptional but that's only them, they still are as many as the Finnish and more than the three Baltic republics together.

Maju said...

@Krefter: it has been recently determined, if my memory is correct, that La Braña 1 and 2 were brothers, so it's only logical they had the same haplogroups.

I'm sorry but I don't recall now where I read that (I found an Anthrogenica thread but, as usual, they say I have exceeded my reading quote and that I must register before I can read anymore, what I won't - lol, it's easier to delete the cookies or even reboot the router if it's a matter of IPs but I just get pissed off by that kind of attitude).

Rokus said...

'insist on linking Remedello with kurgans. AFAIK there is no such connection'
Still Mallory (1989) was convinced Remedello was tightly linked to a first wave of Steppe expansions. Actually, Remedello and its kind (eg. Rinaldone) in Italy were an archeological showcase of PIE culture to him. He dismissed assertions of archeologists that Remedello and the kind were local as rubbish, even though he failed to link the horizon with an obvious direct external source. Some evidence seemed to link Remedello to Hungary, though Bell beaker was more to the mark. If IE links existed, it must have been the other way round! History proved Mallory wrong in his Steppe interpretation, though this doesn't undo archeological evidence and the status of Remedello of an archeological PIE showcase. Back on track?

Maju said...

@Matt: "Agree with Davidski's point that there isn't much use to a European only plot of these stats as it just tells you less than a West Eurasian plot without adding any more information though".

Well, the fact is that European-only plots do organize themselves in a cross-shaped way that IMO is informative. One dimension is usually Sardinians vs Russians (or similar) and the other Basques vs East Mediterraneans (or North Caucasians, if included). Obviously the first one indicates EEF vs generic HG and is the most important but the second one indicates an Atlantic (MN/Chalcolithic) vs Eastern dimension that can also be of interest.

On one side of this second dimension stands the Basque or Atlantic or Megalithic pole, which, as discussed previously can't be just ignored (it pretty much dominates the pre-IE non-Eastern European variance), on the other the fact that Maltese or Sicilians, which can't be explained by Lazaridis' triangle, cope the other pole, influencing apparently Italy and the Balcans, is probably revealing, although I'm not sure right now how to interpret it.

Also West Asians clearly fall outside of European variation, nor should be tentatively explained by European polarities or aDNA (as some have tried, causing only confusion IMO), so they are largely irrelevant for the understanding of European origins (there's room for the all West Eurasian plot and analysis but it should not exclude the Europe-only one, which is also important).

When we use the Europe only plot (for example this one by Lazaridis, annotated by me), Basques and Maltese fall pretty much in line with other South Europeans in the dimension one, showing only peculiarities in this intriguing second dimension. Instead of just dismissing it without further ado, I think this Basque-Maltese or sometimes Basque-Caucasian polarity should be researched because it also shapes European diversity.

A tentative explanation is that what happens in the European-only plot is that the polarities are rearranged. Both European polarities are mostly from dimension 1 of the all-WEA plot: one accumulates the EEF-WHG plus the EEF-Yamna main contradictions into a single line and the other is basically an Atlantic (MNE) vs Eastern Mediterranean (Vinca-related???) that is also folded in dimension 1 in the all-WEA plot. This second dimension is extremely intriguing to me and definitely affects all Southern Europeans one way or another: Italians and Balcanic peoples tend to the "Maltese" pole, while Iberians and Gascons tend to the "Basque" pole instead.

From a North European perspective this may be a bit trivial (although French and NW Europeans do tend to the Basque polarity almost as much as Iberians) but from a Southern one it is clearly not. For Basques and related peoples it may be important but for Balcan and Italian peoples it seems even more important to understand this dimension and its causes, which so far are not well understood at all.

Maju said...

@Rokus: ditto. There's nothing specific linking Remedello and related industries to the "Kurgan Europe" of the time.

Alberto said...

@Matt

Thanks, as always, for the graphs.

They do remind me again of the possible importance of these Bronze Age Hungarians in the genesis of modern North Europeans. They seem to be a great match for a more or less 50/50 mix with Corded Ware in modern populations. They are the only samples we have so far that can explain the high levels of WHG (as a component) in modern North Europeans, their higher affinity to WHG vs. EHG, but without dramatically reducing ANE.

The mystery is where is the source of these Hungarians that appear around 2000-1800 BCE looking like a straight mix of EEF and Motala (both of which populations had probably disappeared thousands of years earlier). Other combinations (Yamnaya + MN + pure WHG) are possible, but again, where would pure WHGs come from at that time? Maybe if Vinča or CT show very high levels of WHG (as one Vinča sample suggested), the source could be there.

a said...

@Maju/ Rokus.....

Straight up, I have a conflict of interest. My branch of R1b-Z2103 and subsequent-descending snp's are interwoven with R1a283+[93-] clades [Tomatoes]Eastern branch of R1a,[perhaps a little wider distribution area, in geographical terms.] Yet both of us plot the same. Either, we both descend from common ancestor's within Central/Western/Southern-Europe or Southwest Asia or South Asia[R1a-R1b] {R1b-Z2103 & some R1a}]; or we originate from the area that is already producing 5+/-k-7+/-k samples of R1a &R1b.
For obvious reasons both of you have chosen Europe. I have debated this with Tomatoes several times [1km-1kyears]providing as much leeway as possible. So lets see what the next batch of samples provide us with.
BTW Ive charted my specific snp branch down to 600A.D.+/- more or less, picking up snp matches that lead a trail to Yamnaya.

Alberto said...

@Maju

Me: Probably another issue with European plots is the overrepresentation of some "small, extreme" populations. Like Sardinians, Basques and East Baltic...

You: I am of the opposite opinion. I feel often that the overrepresented ones are Baltic, Finnic speakers and other tiny NE European populations.

Why do I have the feeling that I'm being misunderstood throughout this thread? I don't see that your opinion is opposite of mine. We actually quite agree, if not completely agree.

I'll repeat that of course I consider Sardinians, Basques and Finns important and would never ask for them to be excluded. I'm only putting them into context.

Why Germans should be so important? They (81 mill.) are not many more than Italians (61 mill) or Iberians (57 mill.)

Germans was an example. Same goes for Frenchmen, Brits and many others. If you look at most plots, you'll see something like:

Sardinians (1.7 million): 20 dots
Basques (2.5 million): 20 dots
Finns (5.5 million): 10 dots
Maltese (0.5 million): 6 dots
Germans (80 million): 4 dots
British (60 million): 3 dots
French (57 million): 6 dots
Austrians (10 millions): 0 dots
Dutch (15 millions): 0 dots
Swiss (8 million): 0 dots
Spaniards (45 million): 30 dots
Estonians (1 million): 5 dots
Romanians (20 million): 0 dots
Serbs (7 million): 0 dots
Poles (38 million): 4 dots
Ukrainians (40 millions): 5 dots
etc...

I do understand this bias is to avoid oversaturation in the centre and make the plots more interesting. As long as we know it, no problem. But the point was: Looking at those plots without considering the above, you might think that Europeans are a cline from Sardinians to Finns (going south west to north east). But that's not the reality. The reality is that Europeans are mostly a north to south cline, with some small exceptions that fall out of that cline (Sardinians, Basques, NE Balts), and only 2 big populations that also fall slightly out of that cline: Spaniards and Russians.

I personally don't have a problem with this. I'm fine with the graphs as they are. It was you who complained about them, and I just said that of course you have a point in your complain, but also explained why they are done this way (this was regarding the horizontal distortion, not the sampling bias, but both things are somehow related).

DMXX said...

So long as resolution or scaling issues are recognised, inferences leading off these won't necessarily be subject to error.

Following Alberto's post, for what it's worth, I've always perceived the north-south cline across Europe to be more prominent than west-east on these PCAs. If the qualitative inference does not satisfy, quantitative measures should confirm that much anyway (have no numbers at my disposal, but I'd assume the Orcadian-Russian and Spaniard-Greek GD's will be smaller than Orcadian-Spaniard or Russian-Greek).

CroMagnon said...

Alberto

Id bet off hand that some 75% of the Copper age population in Europe was in Tje Carpathian basin and Lower danube regions . lets see what more data from hungary, Bulgaria,
Romania, western ukraine etc show ...

Rokus said...

@Maju
There was NO "Kurgan Europe" of the time. Yamnaya started about 500 years later than Remedello. Mallory and Adams (1997): "The culture exhibits many similarities, at least in metallurgical types, with cultures north of the Alps and east in the Aegean and Anatolia". If any part of these apparently IE-like cultures reached the Steppes, they must have been severly bottlenecked culturally, linguistically and genetically by the time they arrived. Even then, burials in the flexed position, simple pit graves with weapons in it and claims of horse remains, according to Mallory even claims of suttee practices, were all IE traditions observed in the Steppe and beyond.

Mike Thomas said...

@ Maju

I wholly agree that language spread doesn't have to be demographic , but youre missing the point. For the kurgan/Yamnaya hypothesis to be correct, it has to have been demic. yamnayans were not elites, nor were they any more martial, so your elite conquest or Roman empire theory is non-congruent and irrelevant here.


* Vucedol: is centred on croatia . Not really featured by kurhans . But yes there some Kurgans in northern Serbia . Anyhow Vucedol dates 2800-2200 BC

* Cotofeni : I'm not forgetting about it. As I've explained to you , but I can't grasp- Usatavo & Baden-Cotofeni are considered by ALL relevant sxholars to be derived from local Balkanic groups (the CT and Gulemnitsa, resp). Forgot Gimbutas


* the slavic migration wasn't an elite conquest. It was *massive* with some admixture.

* about Proto-Greeks . There isn't any clear trail from the steppe at all, at any point. Just subtle signs and possibilities. No clear smoking gun. I think here we really need aDNA .

* about R1b .
I was just hypothesising . It's not an area im too expert in. I agree with need for Atlantic adna, but am just going wit the current evidence .

Krefter said...

@Maju,
" IMO we need to look for the R1b-L11 expansions and origins in Atlantic Europe and hence probably in what David calls "the WHG rebound that happened from the Early to Middle Neolithic".

https://docs.google.com/spreadsheets/d/12G2cfjG0wHWarsl5bB99ridFmvUWzqlZfZ6_e_R6oIA/edit#gid=1740513758

Maju this reminds of you're opinion that R1a-Z93 can't be from East Europe solely because of modern Y DNA. Ancient Y DNA is greater evidence than modern. It doesn't matter that we don't have a lot of Y DNA from Neolithic Atlantic Europe and none from Asia. What we have is good enough.

Harvard will find the pre-Beaker R1b guy from Hungary had R1b-L11 and Steppe ancestry. If Bell Beaker isn't the source who is? We have Megalithic and Funnel beaker Y DNA and it isn't R1b. It has to be Steppe.

Grey said...

"But yes there some Kurgans in northern Serbia"

I wonder if they're near the Serbian copper mines.

Grey said...

"I wonder if they're near the Serbian copper mines."

a bit of googling suggests "no" as the answer to that question

Maju said...

@"a": Z2103 is "brother" of M412, hence it is already apart of the main expansion westward in Europe of R1b (which is characterized by R1b-M412 and downstream subclades). As such it belongs to a rather ill-characterized paragroup R1b-L23(xM412) which extends by the Balcans, West Asia and Central-Eastern Europe but is very rare in the West. So what is logical is to think that Z2103 is a branch that coalesced in the East (be it Eastern Europe or West Asia, I at least don't know enough to pinpoint its origins) and that has no particular relationship with the Western subclades of M412 and L11: S116 and U106. They must have got a common origin very long ago but that common origin should be in the Balcans or West Asia.

The most recent common origin I can imagine could be something like Neolithic (either in the Balcans or West Asia) and in that case its expansion to Europe (or non-Balcanic Europe) might be parallel to that of R1a and other lineages like G2a, E1b-V13, J2b, etc. Alternatively the common L23 or M269 origin is older. Hard to decide because the current methods do not offer any certainty re. chronology and aDNA on the matter is still scarce and fragmentary.

Maju said...

@Alberto: maybe you are being misunderstood or maybe I was reading too much on your emphasis on Basques and Sardinians, whom I think are particularly important "fossil" populations (unlike the Baltics or Orcadians or Icelanders). I do agree that for general purposes it is usually best practice to use samples that (loosely) reflect the demographics (historical better than modern probably) and I have argued for the same in other occasions. So it's true that we are not in disagreement in the main argument, rather the opposite.

On the other hand it is well known that North Europeans are relatively similar to each other, unlike Southern ones and unlike some exceptional populations of the Far North, so that may explain why what you argue for is probably not that important after all in this case.

The graph I linked in my last comment yesterday (Europe-only PCA from Lazaridis) has a better representation of NW Europeans, with many French and English among others, not Germans though but also many Russians and various Central Europeans, so you really get a feeling of where Germans and Poles (who are also missing) should be.

But yes, in general terms you are right and your warning is more or less correct.

Maju said...

@Rokus: "There was NO "Kurgan Europe" of the time. Yamnaya started about 500 years later than Remedello".

Good point, although I'd warn again against Yamna-centrism on this matter because Yamna is just a derived branch. Older (West of the Don) are at least Sredny-Stog II and Baalberge but of course they were far away from Italy and hence totally unlikely to be related.

I imagine that the "missing link" in that hypothesis was Baden culture (which Gimbutas proposed as Kurgan-derived) but I don't think they were kurgan in any way, rather continuity of previous Neolithic layers, notably Lengyel, whose sites are continuous in most cases.

More mysterious is a "seed" group, that apparently triggered the formation of Baden culture, known as Bóleraz whose tombs (small stone circles) are intriguingly similar to what we can see in the Pyrenees millennia later (in the Iron Age, what makes any connection most unlikely) but nothing other than the generic state of Funnelbeaker-related upheaval of Central Europe at the time allows to classify them as "kurgan". Rather I'd imagine them being somehow related to Michelsberg and hence having diffuse "Western" roots if anything (speculative anyhow).

In my modeling of European prehistory, Baden is rather the last major "Danubian" (LBK-derived) culture (Foltesti, Cucuteni-derived, in Moldavia lasted for longer but it was much less relevant). The fact that Baden displaced Baalberge in Moravia and may have got a serious presence even as far as Kiev, suggests to me that it worked in opposition to the earliest Kurgan expansion, attenuated it also via cultural influence on Baalberge-derived cultures but eventually yielded to Luboń in the area of West Ukraine and finally succumbed to conquest in its Middle Danube homeland (leading to Vucedol).

Maju said...

@Mike: "For the kurgan/Yamnaya hypothesis to be correct, it has to have been demic. yamnayans were not elites, nor were they any more martial, so your elite conquest or Roman empire theory is non-congruent and irrelevant here".

That's not the usual interpretation. On one side we should de-emphasize Yamna, which is just the Eastern (homeland) branch of the kurgan phenomenon at a time when other branches had already formed. The origins for what affect Europe west of the Don and Dniepr seem to be Samara → Khvalynsk → Sredny-Stog II → various offshoots (notably Baalberge →→ Corded Ware but also others in the East Balcans and some secondary influences from the steppe which are not clearly Yamna-related, like Catacomb culture). It is indeed possible that Yamna was less elitist than its Western relatives originally, I have read some prehistorians who think that way, yes.

But when you look at Baalberge for example, or the Balcan-Tisza kurgans or Vucedol... at least some elitism is apparent: one had not just princely kurgan burials but also a fortified town with higher quality pottery acting as capital, the other had very very rich princely kurgan burials, the latter had the famous "megaron" with smithy and what-not. It's of course subject to interpretation but the elite domination was at least arguably, and very probably IMO, present in all them. In most cases such expressions of elitism were new (only in the Balcans there was an apparent pre-kurgan elitism, first arguably in the urbanism of Vinca-Dimini, later in the "royal" tombs of Varna).

"Vucedol: is centred on croatia".

Rather Northern Serbia (and what they call Eastern Croatia or Slavonia, as well as the Vojvodina, etc.) Vucedol itself is near modern Vukovar, at the Croatia-Serbia border and a disputed area in the last Balcan War. In any case it extended through Hungary, Moravia and much of the former Yugoslavia (→ map), replacing essentially Baden culture and also other groups like Vinca and Impressed-Cardium-derived ones of Bosnia, Dalmatia and Montenegro.

"Anyhow Vucedol dates 2800-2200 BC"...

Some sources say 3000 BCE but whatever. It's somehow related to Corded Ware and the Tisza kurgans but unclear exactly how. Interestingly it does seem to imply some sort of demic replacement because most Vucedol sites are new foundations and previous sites were typically destroyed.

I presume that the Hungary_BA sample is from Vucedol but you tell me if I'm wrong.

...

Maju said...

...

"Cotofeni : I'm not forgetting about it. As I've explained to you , but I can't grasp- Usatavo & Baden-Cotofeni are considered by ALL relevant sxholars to be derived from local Balkanic groups"...

I have still notes from some excellent German author (whose name I don't recall, sadly) on this matter explaining that Cotofeni is characterized by many tumular burials (i.e. kurgans or something very similar). It may retain some substrate elements such as female "idols" but also has relations with early "kurgan" Cernavoda (which is not the same as Usatovo: Cernavoda is from Vallachia, Usatovo from Southern Ukraine or Moldova and possibly some sort of minor kurganized Cucuteni sub-group).

If my interpretation is correct, Cotofeni could well be a mixed culture but with strong kurgan elements, i.e. your typical product of elitist conquest, where the substrate survives but under new management (and probably new language displacing the old ones in the long term).

In any case its tumular burials are one of the most clear cases of kurgan influence in all the Balcans. And that's why I also consider it a candidate for the origin of early Hellenes, who also styled tumular burial (especially early on in the north), unlike Vucedol (who nevertheless correlates with Myceneans in the megaron feature. Maybe the origins of Hellenes are a mix of Vucedol and Cotofeni? It wouldn't be abnormal as conquering elites often form multiethnic alliances (Bulgars and Slavs, Vandals and Alans, Mongols and Tatars...)

"the slavic migration wasn't an elite conquest. It was *massive* with some admixture".

No way. There is no Y-DNA correlation (nor any other that I can see) between the Slavic homeland north of the Carpathians and Southern Slavs. Greek Macedonia has more R1a than any Balcanic Slavs, go figure! Quite obviously the Southern Slavs absorbed a lot of pre-Slavic peoples and were these "new Slavs", if anyone, who made whatever lesser (localized) demic replacements that might have happened. They are not substantially different from Hungarians or Romanians in fact.

Maju said...

@Krefter: other than Treilles (which is a very specific cemetery of relatives and also a very Mediterranean and Cardium-looking population) we have nearly no Y-DNA from Neolithic/Chalcolithic Western Europe (some STR-based speculations, a lineage that can't be discerned at all...). So we can't say yet what was it like. MtDNA, better researched, is very modern, "hyper-modern" even in the case of Portugal. Also there is a good case for Basque demic continuity, with very minor alternations at most, since Neolithic or Chalcolithic at the latest - so why would 90% of Basque Y-DNA be "foreign", while it's only like 50% in the case of Indoeuropeans like French and Spaniards? If anything we would expect the opposite pattern: more R1b among French and Spaniards, less among Basques.

Krefter said...

@Maju,

"The most recent common origin I can imagine could be something like Neolithic (either in the Balcans or West Asia) "

I can understand the regional-diversity of R1b-L23 and its western subclades, it's very simple. No one knows more than the other in this regard. And I can understand that the main thing that determines when you think R1b-M412 arrived and expanded in West Europe is based on how old you think each regional-specific subclade is(Not archaeology, etc.). Archaeology is used to find a population who spread R1b-M412 in the time period you think it expanded in.

So, it seems your opinion that R1b-M412 expanded within West Europe before ANE arrived is based on how old you think it is. If it wasn't for Ancient DNA I'd agree with you.

I'm open to a pre-3000 BC arrival and local expansion, and I'd be pretty exited if a Neolithic R1b-L11 was found. Outside of Ancient DNA it makes sense.

The pre-Beaker guy from Hungary who had R1b, will probably be L51 or L11. Earliest known ancestor of Western R1b. His autosomal profile will give clues to the origin of Western R1b, that's probably why Harvard is getting his genome(Davidski said so). If he has Steppe-ancestry it supports a Steppe-origin if he's EEF/WHG it proves a local origin.


" MtDNA, better researched, is very modern, "hyper-modern" even in the case of Portugal."

How do you define hyper-modern?

Shaikorth said...

There are a couple of studies incoming about the Slavic migrations soon. If they don't close the case, ancient DNA from pre-Slavic expansion Macedonia and Northern Greece will.

R1b being an Indo-European marker and its high frequency in Basques is not really a contradiction even with Basques having less I-E autosomal influence, because isolate populations are more affected by drift and founder effects. Anyone taking a look at the Basques in the Human Origins set, or other sets including them, can see that their levels of within-population IBS sharing and homozygosity are higher than pretty much any other European population's there, Sardinians aside. Founder effects can easily cause major changes in uniparental distribution of such populations.

Mike Thomas said...

@ Maju

1) "That's not the usual interpretation. On one side we should de-emphasize Yamna, which is just the Eastern (homeland) branch of the kurgan phenomenon at a time when other branches had already formed. The origins for what affect Europe west of the Don and Dniepr seem to be Samara → Khvalynsk → Sredny-Stog II → various offshoots (notably Baalberge →→ Corded Ware but also others in the East Balcans and some secondary influences from the steppe which are not clearly Yamna-related, like Catacomb culture). It is indeed possible that Yamna was less elitist than its Western relatives originally, I have read some prehistorians who think that way, yes. ""

--> Well I've explained to you your Samara ->Khvalynsk -> Yamnaya sequence is simplistic and best, entirely wrong at worst. Wholly outdated, but you cannot see / or do not wish to accept it. And what you think is the "wider Kurgan phenomenon" is merely the secondary products revolution which has no specific cultural-genetic clink.

2) "But when you look at Baalberge for example, or the Balcan-Tisza kurgans or Vucedol... at least some elitism is apparent: one had not just princely kurgan burials but also a fortified town with higher quality pottery acting as capital, the other had very very rich princely kurgan burials, the latter had the famous "megaron" with smithy and what-not. It's of course subject to interpretation but the elite domination was at least arguably, and very probably IMO, present in all them. In most cases such expressions of elitism were new (only in the Balcans there was an apparent pre-kurgan elitism, first arguably in the urbanism of Vinca-Dimini, later in the "royal" tombs of Varna). "

Baalberg is not Kurgan. The general view (e.g. Mallory) is that it has nothing to do with the Pontic steppe, and the similarities are generic/ coincidental. The recent genetic studies (in case you've missed the last 2 years of discourse) confirm this .

The other thing that you're just not getting is that you can't just lump all 'kurgan cultures' together. The Yamnaya ones specifically are the poorest of all, the least elite, and the sparsest populated. The richest kurgans of M4 appear in Majkop and Usatavo; which are not part of your Samara-Khvalnysk-yamnaya pseudo-sequence, or indeed Gimbuta's overall (wrong) thesis.


3) "Some sources say 3000 BCE but whatever. It's somehow related to Corded Ware and the Tisza kurgans but unclear exactly how. Interestingly it does seem to imply some sort of demic replacement because most Vucedol sites are new foundations and previous sites were typically destroyed. "

Well that's your speculation. Vucedol is a copper age culture with probably local origins. Volker Heyd sees it as an *oppositional* block to new steppe intruders. Whatever the case, Ill rest my judgement until we see its aDNA profile. "

Mike Thomas said...

@ Maju

4) "I have still notes from some excellent German author (whose name I don't recall, sadly) on this matter explaining that Cotofeni is characterized by many tumular burials (i.e. kurgans or something very similar). It may retain some substrate elements such as female "idols" but also has relations with early "kurgan" Cernavoda (which is not the same as Usatovo: Cernavoda is from Vallachia, Usatovo from Southern Ukraine or Moldova and possibly some sort of minor kurganized Cucuteni sub-group). If my interpretation is correct, Cotofeni could well be a mixed culture but with strong kurgan elements, i.e. your typical product of elitist conquest, where the substrate survives but under new management (and probably new language displacing the old ones in the long term). "

--> Incorrect. Usatavo is derived from Cucuteni. Its earlier than Yamnaya, thus cannot derive from it. Nor does it derive from Khvalynsk, which was long gone, and the steppe had a kurganless hiatus for several centuries. Cernavoda is derived from local Bulgarian group. Kurgans are generic burial categories, like cremation. There is no need for genetic links between any/ all groups using similar tumular styles. All it does is represent the social position of the deceased. It does;t mean that everyone & anyone buried in a kurgan is from Samara or Khvanlynsk. This is a very crude and backward understanding of archaeology.

But I see you will only be convinced when aDNA from Usatavo and Cernavoda comes back Balcanic. If it does not, I'll happily take back my words.


Mike Thomas said...

5) ""In any case its tumular burials are one of the most clear cases of kurgan influence in all the Balcans. And that's why I also consider it a candidate for the origin of early Hellenes, who also styled tumular burial (especially early on in the north), unlike Vucedol (who nevertheless correlates with Myceneans in the megaron feature. Maybe the origins of Hellenes are a mix of Vucedol and Cotofeni? ""

-> As I said im not sure. But I am sure its not as simple as 'fusions' of 'cultures'. Such interpretation is about 4 decades too old. Modern Anlophone archaeologists don;t simplistically equate material cultures with "tribes" or "peoples" like you, or the sources you rely on, do.

Mike Thomas said...

@ Maju


"the slavic migration wasn't an elite conquest. It was *massive* with some admixture. ..
No way. There is no Y-DNA correlation (nor any other that I can see) between the Slavic homeland north of the Carpathians and Southern Slavs. Greek Macedonia has more R1a than any Balcanic Slavs, go figure! Quite obviously the Southern Slavs absorbed a lot of pre-Slavic peoples and were these "new Slavs", if anyone, who made whatever lesser (localized) demic replacements that might have happened. They are not substantially different from Hungarians or Romanians in fact."

--> South Slavs absorbed *Some* pre-Balkan groups, of course. But what your clueless of is the massive demographic crash in the Balkans c. 600 AD, and the subsequent mass replacement by Slavs. You're probably unaware that there have been several extensive land surveys in various Greek regions - and all show massive depletion of rural populace in the 7th century, followed by appearance of a very Slavic looking material culture in the 8th. Granted, some areas like Corinth, Athens, Thessalonki and several Peloponessian towns continued

The non -concordance of genetics is only apparent if one doens;t understnad the genetic and historic material. The large amounts of R1a in ""Greek" Macedonia" is expected, because Greek macedonia was one of the earliest and densest regions of Slavic settlement (several tribes around Thessaloniki: Dragubitai, Rynhini, Strymonoi, Belgezets, Sagudati), and most modern Greek Macedonians are Hellenised/ Byzantinized Slavs (go figure that !). When you combine all the R1a and I2a in the Balkans, you'll easily see something like 90% replacement in Herzegovina, and as high as 30- 50% in Greece.

You can't just indiscriminately apply models from Visigothic Spain (which was indeed a case of elite dominance) to other regions of Europe which you apparently know little about.

That Slavs aren't; too different from Romanians doesn't mean anything for your argument. Romanians are basically Latin speaking Carpathian-Slav

Kristiina said...

I checked yfull’s R1b tree and noticed that R-Z2110 (S12460) is only found in Europe and it is formed 6200 ybp and its TMRCA is 6200 ybp. It is not very wide-spread in Europe today, but it could be an early arrival to Europe unrelated to IE languages. Earlier branches of this line are also found in China and India.

Also R-Z2111 (Z2118/PF7589) is formed already 5800 ybp and its TMRCA is 5100 ybp. It is restricted to Europe and looks like unrelated to IE.

However, the huge majority of European R1b is derived from R1b-L51 which is formed 6200 ybp with TMRCA 5800 ybp. It is interesting that the earliest split separates Sardinian Z2115 from the rest of the Europeans. Therefore, European R1b could precede Yamna and be contemporaneous with Sredny Stog. There are far too few samples from the Mediterranean area to be sure of the northern Corded Ware origin of R1b-L51.

I am inclined to think that R1a and R1b did not spread together and R1b-L51 may be originally non-IE. Understandably, many people here are IE-centered and think that all Bronze Age innovations are the same as IE, hence all innovative Bronze Age cultures in Europe must be linked with IE speaking R1a or R1b. However, there were many non-IE languages in Caucasus, Europe and also in the steppe and R1b was also found in them.

To my astonishment, I also noticed that in yfull the earliest R1b branch is found in Peru. It is even older than V88. If the earliest branch-off means the origin of a y-line, as is sometimes claimed, the astonishing conclusion is that R1b spread from America. I do not really support that but it should confirm the North Eurasian origin of R1b.

Mike Thomas said...

Kristiina

I share your observations. Most people are either ignorant of this, or too R1b-IE'centric, thus just sweep these inconvenient truths under the rug, or make hogwash explanations like R1b is actually IE, and anyone who is R1b +ve but isn;t IE just switched their language- like all the Basque, Caucasian, Arabic, Jewish Rb +ve groups, and Im sure we'll soon find out - ancient Etruscans. Special pleading par excellence.

But disagree with your focus on Sredni Stog. Its insignificant to European history .

Mike Thomas said...

Oh I forgot Turkic !

Kristiina said...

Mike, I am not a specialist in archaeology. I chose Sredni Stog just to find a culture that preceded Yamna close to the area from which R1b may have entered Europe.

Go and see R1b haplotree in http://genome.cshlp.org/content/suppl/2015/02/18/gr.186684.114.DC1/Supplemental_Figures.pdf

David very recently stated that it's highly unlikely that any Proto-Indo-Europeans ever stepped foot on Sardinia. Now, when I look at the above mentioned haplotree I see that Sardinians abound in rare early variants of R1b. They have R1b-V88, R1b-M269(xL23), R1b-L23 (xM412) and R1b-L51(x R-L151). The huge majority of R1b in Europe is under that R-L151.

Mike Thomas said...

Yes. So obviously R1b in Europe E is due to recent founder effect. The questions which remain are how it got introduced into the BB networks and how / when it became IE

Maju said...

@Krefter: it's not about "how old" because we just don't know with any certainty, so disregard. It's about the geostructure of S116 and U106, which is not at all pointing to an eastern origin but the opposite.

"So, it seems your opinion that R1b-M412 expanded within West Europe before ANE arrived"...

I would think so, yes.

"... is based on how old you think it is".

Nope. It's based on the fact that S116 MUST have expanded from somewhere near South France and (probably) U106 from somewhere near the Netherlands. There's nothing pointing East other than migration arrows from these most likely origins. These two haplogroups expanded from the West almost without any doubt, regardless that upstream R1b did in general expand from the East at some point.

This point is quite apparently a pre-Yamna date, as Yamna has Z2103 and M412 should not be expected to be more recent. So the common ancestor of Z2103 and M412, i.e. R1b-L23, is necessarily older than Yamna. And judging by Yamna's very local, static, origins and R1b-L23 structure, several millennia older at least. I'm here reading the aDNA evidence you insist on very straightforwardly but with a totally opposite conclusion than yours. Now tell me: is my logic good or not? I believe it is very sound: Yamna's dates are a "minimum ante quem" and just a bit of common sense applied practically forces. You can read it as being Neolithic or pre-Neolithic but never as being originated in Yamna or in general the Kurgan homeland.

To do that, you need to force-feed some highly unlikely and very questionable assumptions: (1) that Kurgans expanded largely demically as single long-range vector, without any intermediate co-option of other populations (what contradicts the different haplogroups found in Corded Ware, Yamna, etc.), (2) that all evidence of M412* in East Europe has been erased by statistically impossible random events, (3) that Kurgan peoples did not only speak Indoeuropean but seemingly also Vasconic (and maybe even Afroasiatic also, judging for what we see in Cameroon), (4) that all the mtDNA evidence suggesting continuity in the Basque is absolutely irrelevant and must be totally ignored, and (5) probably other absurdities I can't even fathom.

Maju said...

@Saikorth: R1b is not a valid category for this discussion. Western European type R1b, i.e. under M412, is the valid category. This haplogroup is low to very rare in the area of early Indoeuropean expansion, including the Corded Ware area. Even in Germany it clearly begins decreasing relative to more westerly areas: North Germany 36%, Brittany: 80% (Eupedia figures); South Germany and East/North France: 47-56%, West France: 75-80%. It grows in frequency to the West, so it does not make sense it is weak where IEs were for millennia and so strong where IE arrival was as late as the late Iron Age, just a few centuries before the Romans, or where they never arrived at all: Basques 85-90%. R1b (Western subclade) is clearly a pre-IE marker and, if Neolithic, a Megalithic or similar kind of marker. If you have that lineage, and roots West of the Rhine or North Sea or south of the Alps, it is almost certain that 3000 or 3500 years ago (at most) your patrilineal ancestor did not speak a IE language, and, regardless of geography, 99.99% sure that your patrilineal ancestor 5000 years ago did not speak IE. Actually it is quite certain that he spoke some sort of Vasconic.

Aram Palyan said...

Kristiina

That Peruvian is R1b1c*, so he has an African origin. I don't know how yfulltree categorize his data but I think most probably he is a post-Columbian settler. Nothing surprising.

Maju said...

@Mike: "I've explained to you your Samara ->Khvalynsk -> Yamnaya sequence is simplistic and best, entirely wrong at worst".

You have not explained anything. You have merely CLAIMED and somehow assumed that I would take your word for it. Sorry but nope.

"Baalberg is not Kurgan".

Baalberge has burials in tumuli that resemble kurgans. These burials became more popular or common as time passed. Baalberge is also a direct ancestor of Corded Ware (and not Yamna). Why is it "not kurgan"? Just because of aDNA? Sorry but all the aDNA on Earth cannot characterize a CULTURE: culture is not genetic, even if it may (or not) be related to genetic or demic flows.

"... you can't just lump all 'kurgan cultures' together".

In this I agree. You have to study their origins and trajectories as much as possible in the particular details. But that does not mean that the Kurgan macro-culture did not exist and should not be considered as a ultimately unified whole with its complexity and diversity.

"The richest kurgans of M4 appear in Majkop and Usatavo; which are not part of your Samara-Khvalnysk-yamnaya pseudo-sequence"...

Read again what I said. I said Samara → Khvalynsk → offshoots, one of which is Yamna, another is Maykop, yet another is Afanasevo, and another Sredny-Stog II, which is plausibly at the roots of all or most European IE. I am all the time de-emphasizing Yamna if you'd care to pay attention to what I say: Yamna is just the Eastern branch (in Europe, Afanasevo is of course more easterly) and other than a late expansion into parts of Ukraine and its likely (or at least plausible) role at the origins of Sintashta-Andronovo much later, it does not seem very active.

I do agree that where richer kurgans appear it is generally associated to a greater emphasis in hierarchy and probably even powerful and showy monarchy (surely the case of Maykop). So what? Obviously not all kurgan cultures would be equal and it seems somewhat logical that the original homeland would retain more equality, while conquered areas could be more prone to hierachization. But that's a bit too nuanced anyhow and I don't know if we can afford to waste time discussing such complicated details, many of which would have been produced by locally "chaotic" evolution which does not follow any too obvious patterns. Why this or that kurgan was erected and why of its details? Maybe they were the caprice of certain but not other princes, much like the Egyptian pyramids. What matters are the general cultural patterns and the apparent connections and disconnections between cultures and layers.

"Vucedol is a copper age culture with probably local origins".

No way. And it's not "my speculation", I bothered looking online for confirmation because now and then I do get some doubts, you know. Just check Wikipedia if you care.

Shaikorth said...

Maju, certain R1b clades are very high frequency in small western populations, and it is something one would expect considering their lower diversity makes them more suspectible to founder effects.

Y-Full dates the R-M269 MRCA to 6400 BP. All its subclades including M412 have theirs later. M269 has old subclades such as R-PF7562 with a distribution in Belarus and Northeast Turkey, which may help in showing R1b's movements. Right under U106 there's a Polish-Ukrainian subclade R-Y15627, so there's no reason to assume origin near the Netherlands is a necessity. Lets wait for more Y-DNA from Neolithic Iberia and South France to see if there really was any R-L151 (MRCA 4900bp, which makes S116 and U106 less than 5000 years old) there.

Mike Thomas said...

Maju

I've already provided references (Rassamakin and Mazura about the Yamnaya sequences- who are *the* premier archaeologists on Yamnaya; and Mallory that Baalberg is not kurgan) but you choose to ignore then because you either lack faculties to understand, have a meagre /pathetic reference dataset or you're too biased in your dogma. I suspect it's all 3.

But there's no point in arguing with someone who's not even wrong .

Aram Palyan said...

The recent study on Albanians showed that they have significant number of R1b-L51 xP311

Gheg Albanians:
E-V13: 38%
J2b: 25%
R1b-L51 xP311: 12%
R1b-M269 xL51: 4.2%
I2a-xM26,M223: 3.3%
R1a-M17: 2.5%
I1-M253: 3.3%

Tosk Albanians:
E-V13: 29%
J2b: 12%
R1b-L51 xP311: 8%
R1b-M269 xL51: 6%
I2a-xM26,M223: 11.5%
I2a-M223: 5%
R1a-M17: 6%
I1-M253: 3.8%

http://www.nature.com/ejhg/journal/vaop/ncurrent/full/ejhg2015138a.html

Aram Palyan said...

Mike

From Your list of modern nations the only hard case is the Basques. Assyrians also but they are have E1b1+J1 majority at last.

The rest is easy to explain. Some of them don't even have a relative majority of R1b.

But let's wait Hittite Y DNA.

a said...

@ Maju said...

@"a":
" Z2103 is "brother" of M412, ..... "
R1b-Z2103 & R1b-H.G. clustered on the Steppe in Samara.
R1b-Z2103 was already 1000+/- years old; by the time Sintashta came into the region.
If you like Rassamakin's work on Skelya culture, the same cluster using the R1b H.G.pre Kurgan[5600B.C.+/-] sample instead of the Kurgan R1b-Z2103 sample predates Skelya culture by about 1000+/- years more or less.

Kristiina said...

Aram Palyan, then it is not a proof of pre-Columbian R1b in America. In any case, according to yfull tree, the oldest branches of R1b are in Africa and Saudi-Arabia. Apart fron that Peruvian sample, there is also R-Y7771* from Nigeria and R1b1c3 (V69) from Saudi Arabia, and R-L389* from Puerto Rico. I am wondering if also that HG00640 Puerto Rico R1b is originally from Africa.

Mike Thomas said...

@ Aram

This is what I'm seeing

R1b P25

(1) V88 - northern Africa, Sicily, Neolithic southern Europe.

(2) M335 - Anatolia

(3) P297

(3a) M73 - ancient Samara HG (pre-M73), modern Turkic groups in central Eurasia, Mongol groups

(3b) M269

(3bi) PF7558 - Sardinians, Arabs, Jews.

(3bii) L23

(3biia) Z2103 - various Balkans, Caucasian speakers, Anatolians, Armenians, Assyrians

(3biib) L51 - Europeans (almost exclusively), incl Basques.

Spin that into a nice story please.

Dospaises said...

@Kristiina

All of the samples at YFull that have and id that start with HG or NA are from 1000 Genomes or Hapmap which are from people that were living at the time the the country and ethnicity indicated by the 3 digit code. None of them are ancient samples so if one of the samples from the Americas has a haplogroup or subclade that seems to be post-Columbian there is little doubt it is.

These FAQs from 1000 Genomes might be helpful.

http://www.1000genomes.org/faq/which-populations-are-part-your-study
http://www.1000genomes.org/faq/can-i-get-phenotype-gender-and-family-relationship-information-samples
http://www.1000genomes.org/faq/does-1000-genomes-project-use-hapmap-data

Fanty said...

"The Yamnaya ones specifically are the poorest of all, the least elite, and the sparsest populated. The richest kurgans of M4 appear in Majkop and Usatavo;"

The richest and most dense populated regions of Indoeuropeans are in the USA as far as I know.
That doesnt actually mean that the Indioeuropeans spread of the USA.

Fanty said...

Hm. Just sprungs to my mind that the richest Germanic peoples are also not the ones "home" but the once that established an elite goverment somewhere while "at home" where they originate they are a primitive lumb of barbarians.

The richest Mongol Khans are also the ones who are almost Chinese already and not the ones from the Mongol steppe.

The rule: The richst and densest populated place is the origin of a people or culture seems wrong. Usualy its exactly the oposite.

Grey said...

"Spin that into a nice story please."

Family groups of metal workers spreading out in all directions from a central origin creating a branch like pattern: so V88 south, L51 west etc.

Remaining a minority caste in most places they settled , a significant minority when they settled somewhere which also had a source of copper and a majority only along the Atlantic coast where numbers expanded dramatically (probably cos acid soil + LP imo).

Maybe pre-IE or maybe ex-IE as a minority settled among a majority culture.

.

If this map of copper working expansion is at least semi close to accurate then the choice of origin might be one of the three dark areas - or maybe somewhere situated in between those three - like Sredny.

http://cache.eupedia.com/images/content/Copper_Age_Europe.png

Maju said...

@Mike: Mallory claims that Remedello is kurgan... so, with all due respect, let me ignore his opinion. Re. Yamna, nothing relevant.

Maju said...

@Mike: "(1) V88 - northern Africa, Sicily, Neolithic southern Europe".

You can't talk of V88 ignoring Central Africa and West Asia. The lineage seems to originate either in West Asia or the Balcan-Italian arch (see: here and here). This is pretty much the same we see re. other R1b taken as a whole.

Aram is correct emphasizing the importance of the Balcans (and reasonably Albanians, who can well be considered, at least the coastal ones, a "cuasi-fossil" of early Cardium peoples prior to Westward expansion) on these matters. Even R1b-M269 could well have expanded from the Balcans (see here and here), although, as it happens with R1b-V88, the issue is not fully clear cut and a West Asian ultimate origin is at least as plausible, if not more.

Maju said...

@Grey: not "metal workers" (what a strange obsession, that comes up once and again) but much more likely to have been just part of the wider early farmer pool, causing different genetic founder effects here and there. Of course, pre-Neolithic expansions can't yet be fully discarded either because we really don't have any good chronology but in any case the Yamna and Beaker findings are "terminus ante quem" and back in those days specialist metallurgy was still wearing diapers.

Alberto said...

@Maju

When you say that you follow quite strictly Gimbutas hypothesis, what do you mean exactly? Because to me there are 2 important matters there that I still don't know what's your take on them, so I would like to ask your opinion.

First, the whole concept of the "Old Europe": egalitarian, pacifist, matriarchal, worshipping mother earth... that was destroyed and replaced by Steppe people who were war-like, elitist, patriarchal, sky worshippers,...

I know that you are much better informed than me about prehistorical cultures to know that this is simply not correct, and rather naive at that (and I obviously don't want to take any merit from Gimbutas pioneering work and her insights into this matter, but some things must be revised). You've mentioned above the Varna culture as an obvious example of stratified society that predates IE migrations. But there are many others too, some that have been considered Indo-European based on some of those traits and now aDNA makes it more doubtful (Baden, Baalberge, Remedello). You'll excuse me for quoting Wikipedia, but it's easier this way. About Remedello:

"The Copper Age graves contained a single body in a crouching or supine position with the head facing north-west.

The male set was represented by arrows, stone daggers and polished stone axes, among the tombs few are those with axes and daggers or ornaments made of copper."


But to mention one that is without a doubt pre-IE, we could just refer to the Iberian Copper Age culture of Los Millares:

"Analysis of occupation material and grave goods from the Los Millares cemetery of 70 tholos tombs with port-hole slabs has led archaeologists to suggest that the people who lived at Los Millares were part of a stratified, unequal society which was often at war with its neighbours."

Unequal, stratified, at war with neighbours, in fortified settlements, building tholos tombs (a much more sophisticated form of burial mound than Kurgans),...

So basically this means that the changes that were happening in the 4th millennium and that intensified in the next one, are independent of the IE migrations since they precede them, and besides they are natural changes related to any society that reaches certain level, and not something exclusive from any people or culture.

The second question is about elite conquer vs. migration. You've mentioned examples of language or cultural shift without genetic impact, and of course there are many of them. However, the eastern genes that were characteristic of IE people have reached all European populations equally. IOW, the genetic impact has been the same across Europe (slightly decreased in SW Europe, especially Sardinia). So I do agree that some of the language spread was due to elite conquering (Latin is the obvious and best attested example, but it's also a much later phenomenon), but for most cases we don't need an elite conquering to explain the language shift simply because sooner or later the genetic impact did happen. Italians and Greeks have the same amount of eastern genes than people from the Corded Ware area. Maybe the migrations that caused the genetic shift didn't cause a language change (since some small elite had already caused it before, and the mass migrants adopted the already IE local language). All is possible. But in general it's just not necessary (and unless it can be proved, as in the case of Latin, it should be the less obvious choice, I would think).

a said...

Mal’ta South - Central Siberia F999914 M U 24,000 years
and Stuttgart F999916 using K7 & Kurds list from Anthrogenica.



MA1 Stuttgart Saudi Mbuti -0.0587 -11.262 70474
MA1 Stuttgart RISE_baArm Mbuti -0.0273 -3.085 46181
MA1 Stuttgart Lithuanian Mbuti -0.0199 -3.773 70474
MA1 Stuttgart Estonian Mbuti -0.0142 -2.804 70474
MA1 Stuttgart RISE_baSin Mbuti -0.0076 -1.036 64594
MA1 Stuttgart Corded_Ware_LN Mbuti -0.0009 -0.145 70307
MA1 Stuttgart RISE_baAndrov Mbuti 0.0077 1.115 70291
MA1 Stuttgart Kalash Mbuti 0.0096 1.798 70474
MA1 Stuttgart RISE_baYam Mbuti 0.0212 3.283 68788
MA1 Stuttgart RISE_baAfan Mbuti 0.027 3.826 69299
MA1 Stuttgart Yamnaya Mbuti 0.0273 4.602 70317




Kit Number: F999916 Elapsed Time: 16.26 seconds
Population
ANE -
ASE -
WHG-UHG 43.28%
East_Eurasian 0.21%
West_African -
East_African -
ENF 56.51%



Kit Number: F999914 Elapsed Time: 14.36 seconds
Population
ANE 49.08%
ASE 11.22%
WHG-UHG 33.58%
East_Eurasian 1.54%
West_African -
East_African 4.58%
ENF -



Mike Thomas said...

Fanty
That's a silly example. Because Majkop, Yamnaya etc were all do temporary. uSA came 40000 years after europeans existed

Davidski said...

Mikop was wealthy because it was exporting copper to Mesopotamia. But what does this have to do with the direction of cultural influences?

The dating of Maikop kurgans has always been very iffy. Repin Kurgans way up north on the steppe show the same inflated C14 dates, so there's no way of knowing which Kurgans are older and how the idea spread, nor if language change came with it.

On the steppe Kurgans appear to be just another expression of the Indo-European sun and moon cult. The reason they're round is to make them look like the sun. Some of the Kurgan burials also contain two high ranking people, so they're probably copying an early version of the Divine Twins.

a said...


Blogger Fanty said...

"The Yamnaya ones specifically are the poorest of all, the least elite, and the sparsest populated. The richest kurgans of M4 appear in Majkop and Usatavo;"

The richest and most dense populated regions of Indoeuropeans are in the USA as far as I know.
That doesnt actually mean that the Indioeuropeans spread of the USA."

Great example. I suppose you could also use ydna Q/Siberia and North America as an example since they were the main population before Europeans started to settle .
Wealth in those days- measured in livestock and or even grains[not gold bars], used in special ceremonies.

Mike Thomas said...

Dave

Its more than wealth or the act of building kurgans themselves. Its important to understand what was happening, rather than having a blind understanding of the kurgan theory of 20 years ago, pace Maju

First of all, the dating of Majkop is now incontrovertable, early M4, and thus far earlier than Yamnaya. But its not about the wealth, but also the technology they must have possessed - metallurgy (?better weapons). This, afterall, is why they acquired such wealth. They were the intermediary from the south to the Eurasian steppe and Carpathian basin.

Its also demography. The Caucasian foothills region was more densely settled than the Yamnaya steppe, or the latter Repin predecessors - which were units of individual households rather than entire communities in Majkop.

So *if* we look to a stimulus for an eastward shift emanating from somewhere, then the stimulus must have been Majkop, and whatever lay behind that in turn. Because, quite frankly, Repin-khvalynsk-Yamnaya were nothing. All the more given that Anthony;s reconstruction if false. They didn;t ride horses, had no chariots, didn;t invent the wheel (the only area were there is solid evidence for horse domestication was the site at Botai - which is in the peripheral far east and can;t be falsely generalize for all of the steppe).
The early M4 dating of Majkop also coincides immediately after the final demise of the Old Balkan metallurgical centres . The "Yamnaya expansion", IMO, is only an apparent one. What it actually is is various communities settling the steppe - and steppe-like areas of the northern Balkans during the humid phase (3000-2500 BC), all adapting / sharing similar cultural features.

The kurgan itself is irelavant - another thing Maju can't understand. Like cremation - it can spread without demic movements, like any religious idea. Whether you believe in a pan-indo-European religion or not, what kurgans symbolized was the chiefly nature of its interred. There are kurgans/ mounds in the Balkan area before Yamnaya - those at Usatavo and Cernavoda - which archaeologists are derived from the final phases of the Old Balkan cultures - and Im sure aDNA will prove to be correct. Interestingly, Usatavo kurgans are richer than yamnaya also - possibly becuase they were directly connected to Majkop, and the relative exclusion of Yamnaya in between. Whether actively avoided, or simply Yamnaya was incapable of mastering such metallurgy (which requires a threshold of development) - im not sure.

But that is how it went.

And we shouldnt forget that the Majkop - Caucasus expansion went to the far east as well - Urals, etc, which accords with the aDNA evidence also.

Davidski said...

Language change in ancient times happened either via massive migrations or conquest.

Maikop lived and died in the North Caucasus. They didn't migrate anywhere and they didn't conquer the steppe, Europe or Central Asia.

You can't just claim that Indo-European languages spread with metallurgical skills and/or cultural inspiration from Maikop. There's no evidence of this.

Massive migrations from the steppe are simply a better alternative to explain the Indo-European expansion.

The only way around this is to argue that these migrations didn't happen, which is what a lot of people were doing before this year, because they knew it was a crucial point. But now we're past that. You need something really special to argue against the Kurgan/steppe theory.

Mike Thomas said...

I'm not arguing against massive populaton changes in the period in question., afterall , how do you know Majkop people didn't migrate ?

Have we really proven that Yamnaya (specifically) did Migrate ? What's the ultimate source for Western European L23??

And are we being so so loose as to what we call "Yamnaya" that basically anything East of the Dnieper is being admitted as "Yamnaya" or "Kurgan" ? Is proto-Corded ware Yamnaya ?(the earliest cord ornamentation occurs in Tripolje)

I think we can do better than that. And my main aim is to construct tighter historical narratives based on a better reading of archaeology. There is only so much aDNA itself can tell us (ie point us in general directions).

Let's wait South asian, greek, Ukrainian and bulgarian aDNA . It might all indeed fall into place.

But I already know the kurgan hypothesis is wrong. The "badasses" who invaded india, if indeed from east Central Europe, themselves invalidate the kurgan hypothesis, which stipulates Yamnaya-catacomb-andronovo-BMAC - central asia . It didn't stipulate anything about some convoluted west then East then South scenario, did it ? In fact, Catacomb might come out quite different to Yamnaya .

Kristiina said...

@Grey "Family groups of metal workers spreading out in all directions from a central origin creating a branch like pattern: so V88 south, L51 west etc."

I agree with Maju and Mike.

The Bronze Age Maykop culture is dated ca. 3700 BC—3000 BC and we do not know their yDNA.

R1b-V88: formed 16700 ybp with TMRCA 7300. This could fit with introduction of pastoralism to Africa.

R1b-P297: formed 15800 ybp, TMRCA 13300 ybp. This can only fit with an Epipalaeolithic culture.

R1b-M73: formed 13300 ybp, TMRCA 7300 ybp. This could in fact be linked with introduction of pastoralism to the Eastern steppe

R1b-M269: formed 13300 ybp, TMRCA 6400 ybp. This lineage may have been involved in introduction of agriculture to the Mediterranean basin.

L51: formed 6200 ybp, TMRCA 5800 ybp. This lineage may have been involved in introduction of agriculture to the Western Mediterranean.

Mike Thomas said...

@ Maju

*V88- yes my mistake. I meant central and west Africa , not north

* "Aram is correct emphasizing the importance of the Balcans (and reasonably Albanians, who can well be considered, at least the coastal ones, a "cuasi-fossil" of early Cardium peoples prior to Westward expansion.."""

False. There is nothing all too special about Albanians, certainly nothing like sardinians which we all know are ~~ neolithic europeans. Albanians have absorbed slavic admixture , not to mention whateve went on during the bronze and iron ages. And Aram is incorrect, their xP311 is only apparent- from lack of SNP complete testing

* in making the false statement that Remedello is kurganoid , Mallory was merely following your feminist/ communist idol- Gimbutas. It seems you're happy to pick and choose what's wrong and right from Her works. Sounds truly like the self-contradictory pseudoscience that has come to characterise your doctrinal views

Mike Thomas said...

Kristiina

I have no strong views at present about introduction of M269 specifically in Europe. All I know is aDNA testing is incomplete . Could be copper age could be somewhat earlier

As clever as those YFull guys are, there are error margins to consider. And the TMRCA could represent when it first began to expand, not when it first reached Central Europe

Plus the advent of "ANE" needs to be accounted for.

Kristiina said...

@Mike

That's why I said "may have been involved". I agree that much more ancient yDNA is necessary. However, accumulation of early R1b in Neolithic Sardinians and prevalence of R1b in non-IE Basques is problematic for the northern Corded Ware origin and in general for the IE origin of R1b.

Kristiina said...

Correction: I meant Sardinians who are today genetically most Neolithic of all Europeans.

Aram Palyan said...

Mike, Maju

I didn't make any claims about Albanians. I just showed that they have L51, probably not from West European branch. That is all.

Mike

Concerning Catacomb culture. I checked their mtDNA during developed Catacomb culture. Their main difference from preceding periods is that they are dominated by U4 and U5. If my understanding is correct we should find there a lot off R1a.

For the 'nice story' I will answer little bit later.

Maju said...

@Alberto: "When you say that you follow quite strictly Gimbutas hypothesis, what do you mean exactly?"

I never said that. Gimbutas is a pioneer of the Kurgan model and she described it very well, however she also has some (probably less important) questionable claims: on one side a bit exaggerated manichean claims about the cultural nature of "Old Europe" and the Kurgans, which may have a kernel of truth but is anyhow excessively loaded, on the other side her qualification of Baden culture as kurgan does not fit the data, at least not the modern one. And there are probably other things. So I do not follow Gimbutas strictly at all. In fact I have only read a little of her work and most of my knowledge and opinions come from more modern materials.

I do agree with her in tracing the Kurgan culture's roots to Khvalynsk and Samara. Nothing new stands against that assessment. I also agree with qualifying Baalberge as kurgan and as the ultimate root of Corded Ware (although via its Cuyavian offshoot, not directly from East Germany). But that I agree with her in such issues does not mean that I follow her in any strict way, even the best ones get it wrong sometimes and we must correct what must be corrected.

"First, the whole concept of the "Old Europe": egalitarian, pacifist, matriarchal, worshipping mother earth... that was destroyed and replaced by Steppe people who were war-like, elitist, patriarchal, sky worshippers"...

I don't believe it was quite that way. "Old Europe" is a complex reality and it was not matriarchal in any case. Matriarchy is a myth, just like the amazons. What is real in an anthropological sense are more egalitarian societies (but almost invariably still dominated by the men) and matrilocal (aka uxorilocal) kinship systems in which the husband goes to live to the wife's community and inheritance is largely passed via the maternal line (from uncle to nephew in the case of males). A good historical example are the Iroquois. I don't even think that there is any clear evidence of matrilocality for Old Europe anyhow.

What there is evidence for, and quite a bit, is for what I call a gender-dualistic cuasi-monotheism, similar to Basque mythology or ancient religion or what Hesiod describes as the roots of the Greek mythology, or what can still be found residually in some Hindu sects (Sakhtism, some variants of Shaivism that claim Shiva, along Sakhti, as only God). This religion hence probably had a concept of the Divine as divine couple whose mating kept the world alive (their son was Odei, the storm god, in Basque legends). In all this religion the importance of the Mother Goddess, Gaia for the Greeks (gai(-a) meaning in Basque matter or substance, but also capacity, potential), was central, and in some way it has persisted even in Christianity, with the very dominant devotion to Mary, which overshadows all other aspects in popular religiosity in many many societies. The core value of this religion would be, beyond that mythology of perpetual re-creation of the World or fertility cult, social responsibility, something the priestesses (sorginak = witches) watched for, punishing the cheaters in the name of the Goddess, or so the legends seem to imply.

On the other side the Indoeuropean religion has nothing to do with that: it's clearly polytheistic, has no creation myths of relevance, and does not hold even an ethical or moral code worth that name. Its core beliefs seem to be destiny and glory, honor being found in war above any other thing. It seems a militaristic kind of religion, at least largely. And of course the culture we find among IEs is invariably strongly patriarchal and not egalitarian. The individual, the individual adult male, powerful, aristocratic, warrior, is the center of everything. In a few variants (Hindu, Druidic) priests also have a major role but these priests or their concept at least was almost certainly borrowed from the pre-IE substrate (IVC in India, British Megalithism in Europe).

...

Maju said...

...

"Analysis of occupation material and grave goods from the Los Millares cemetery of 70 tholos tombs with port-hole slabs has led archaeologists to suggest that the people who lived at Los Millares were part of a stratified, unequal society which was often at war with its neighbours."

Absolutely. Civilization and in general the changes of Chalcolithic generated the same sort of dynamic everywhere or at least in many other places. Particularly in civilized sets like Los Millares, etc., the division of labor and subsequent social stratification is something that just does happen without any need of any invasion. And definitely their descendants, the historical Iberians, were also stratified. These changes however were probably much weaker in less civilized areas like Cantabria, etc. but we cannot say they did not exist at all. We also see stratification in a non-IE population like ancient Etruscans or, say, ancient Egyptians. However we also see that women had a much better status in all those societies. So if we can talk of patriarchy at all, that patriarchy was very "soft", while the patriarchy of Indoeuropeans was almost invariably very hardcore. One of the easiest ways to contrast the IE socio-cultural paradigm and a non-IE control is analyzing the Latin-Etruscan relations, which were intense but not without mutual distrust. The Romans for example would consider Etruscans "immoral" because they treated women liberally and even (woe!) dined with them.

But then of course we also have non-IE extremist patriarchy in at least some Semites. If one dares to actually read the Bible (OT especially) he/she will find a very extremist treatment of women (and children) as chattel. And there are surely other examples. But in any case, in Europe at least, probably in India too, IEs were a major factor in the extension of these patriarchal cultural paradigms. And it took millennia and revolutions to change them. I just recently heard someone in a documentary about ancient Egypt saying: what is most striking is that Egyptian women had rights, such as managing their own money, that most women in the USA only obtained in the mid 20th century. Said that, Egypt also had elements of patriarchy, as it's apparent in the rare cases a woman became a pharaoh - but in general the society was more gender-egalitarian than, say, Greece or Rome or Vedic India, and even much more recent IE examples.

"However, the eastern genes that were characteristic of IE people have reached all European populations equally".

I don't have that very clear: different analysis produce different results. You're probably judging based on Haak's ADMIXTURE graphs but Alentoft's results were much more reasonable, IMO, with zero Kurgan influence among Basques for example and a much more subtle and clinal distribution of this component in general.

The problem with autosomal DNA is that it can only be processed with statistical methods and these are not rocket science at all. So it's good to contrast different approaches and fall in doubt when these contradict each other.

Maju said...

@Mike: That Albanians (or other Balcanic peoples) have of course absorbed admixture (from Indoeuropeans mostly) does not mean they can't retain many pre-admixture elements, notably in uniparental lineages of either line. It's very apparent for example that Albanians (and Greeks) retain in large amounts the important Neolithic lineage E1b-V13, so why not others? There are some indications re. R1b-M269 that could make it original from the Balcans, although West Asia is also a possible origin. Most flows between West Asia and Europe went through the Balcans in any case, and very clearly the Neolithic one, so... there is very good reason to pay attention to that often neglected region of Europe, which we can well say that is "the craddle of Europe" or at the very least one of the craddles, and a major one.

Maju said...

@Aram: "I just showed that they have L51, probably not from West European branch".

Of course. But it may well be relevant for the (older) M269 stage of R1b expansion in Europe and nearby areas of West Asia, which I have documented that has some sort of major relation with the Balcans, be it as origin or as corridor. The frequencies in Albanians are relevant because in most other Balcanic populations R1b is quite scarce.

Alberto said...

@Maju

Thanks for clarifying that. I was quite sure your thoughts went in this line, but some of your latest comments made me doubt a bit about it.

About the "eastern" genes (presumably IE, but maybe not all of them), I do look at many different analysis, so I'm quite confident that they are about the same in most Europeans (descending only in SW Europe: Spanish > Basque > Sardinian). What about the Allentoft paper are you referring to exactly? The admixture graph? For what I see there, it's again that "teal" component, that is higher in Greeks than in Russians, Norwegians or Estonians. Italians are missing, but they are about as Greeks.

And regarding the patriarchal culture, well, I rather think that Europe in general has always been quite an exception in being relatively equal (meaning: at least not horribly unequal). Between presumed PIE and, let's say, Semites, I don't see much difference. Only when IE entered Europe this cultural trait relaxed to a very large extent (from Celts to Vikings, the role of women is hugely different to most extra-European cultures, be them IE or Semitic). But this is a wholly different debate in any case, not too relevant to this thread.

Alberto said...

@Maju

I forgot, but I also wanted to ask your opinion about something else: the origin of the steppe cultures.

I think that specially important is the transition from gathering food to producing it, or in more basic terms, from hunting to having domestic animals. If we place this transition somewhere after the Samara_HG related cultures, and around the origin of Khvalynsk, how do you think this transition happened?

- A local development (hunters learning how to domesticate and exploit goats)?
- A cultural borrowing from their southern neighbours, but without demic movements?
- Migrations from the South (and east?) Caspian area to the north bringing domesticated animals with them?
- A different option?

I personally go for the third option: Iranians (lato senso) migrating to the steppe and bringing these developments with them, a model proposed since long ago (G. Matyushin, 1986) and supported now by ancient DNA (still waiting for those unpublished Khvalynsk samples, but Yamnaya/Afanasievo are already clear about it). Do you have any opinion about it?

Aram Palyan said...

Maju
The link provided by Kristiina is interesting.
http://genome.cshlp.org/content/suppl/2015/02/18/gr.186684.114.DC1/Supplemental_Figures.pdf

You can see there the most part of Slavic I2 is a recent expansion. Two times more recent than R1b/R1a/I1 expansion. This supports the theory that Slavic I2a has a young expansion.


Kristiina

The Kyrgyz R1a is even more younger. Thus supporting the idea that the Kyrgyz R1a-Z93 was a recent founder effect. That is why despite having the highest level of R1a-Z93 in Central Asia they do have a large East Asian component and speak Turkic language.

I don't know what branch is the R1b-M73 there. But most probably it is the branch where Mongols and Bashkirs split. This split is 2 times younger than the split between Armenian and Assyrian R1b-Z2103. If it is confirmed then we can assume that the modern distribution of R1b-M73 doesn't reflect it's ancient distribution.

BTW Kristiina what is the title of this study?

Matt said...

Btw, on the D stats from this post, noticed there is some marginal signal of Sweden_MN having sharing with SHG that is greater than would be expected from their WHG and MA1 sharing:

http://i.imgur.com/7mkIcyd.png - comparing variables directly
http://imgur.com/179hmNf,DMXmAHr#0 / http://imgur.com/179hmNf,DMXmAHr#1 / http://i.imgur.com/hG8F8y2.png- PCA / graph of just the EN and MN samples together

So them being admixed with SHG locally for some of their ancestry (in addition to other WHG admixture from elsewhere) might make more sense than we had thought.

Very small divergences though, so poss just within the boundary of error.

Aram Palyan said...

Mike

I also think that the chances are high that Etruscans will have many R1b-L51. Possibly like Basques with lower ANE.
But, this will be much stronger hit to the Vasconic theory of R1b-L51 than to the IE.
The problem is that Etruscan has no any relation to the Basque language as far as I know. And assuming that a same group of L51 created two completely unrelated languages in so short time is far less probable than to assume that at last one (or even two) L51 groups had a linguistic shift.

Kristiina said...

Aram, the paper is available here with name “A recent bottleneck of Y chromosome diversity coincides with a global change in culture”: http://evolbio.ut.ee/
Only supplementary material is free.

Often we have seen that a high frequency coincides with a young age, and often modern distribution does not reflect the ancient distribution, and more so farther back in history we go.

Many Mongolic groups are probably quite recent, e.g. Kazaks should have formed with Genghis Khan Empire.

Aram, do you think that Etruscan and IE languages belong to the same family? If Vasconic, IE and Etruscan are all unrelated, your logic does not work. However, it may be turn out that Basque is R1b dominant language and Etruscan is I2 dominant language.

Wikipedia says that Etruscan substantives had five cases, and a singular and a plural. Nouns merge the nominative and accusative; pronouns do not generally merge these. Gender appears in personal names (masculine and feminine) and in pronouns (animate and inanimate); otherwise, it is not marked.

Both Basque and IE languages have cases, singular and plural, but proto-IE had also dual, like Slovenian today. Basque has no dual. PIE had eight or nine cases and Basque even more. Accusative/nominative distinction is important in proto-IE and cases do not merge while Basque is an ergative language and accusative/nominative distinction does not exist. Proto-IE has three genders and all nouns have a gender, while Basque does not make any gender distinctions.

Maju said...

You're not wholly correct about Basque gender Kristiina: Basque shows animate/inanimate "gender" distinction in many declension cases (for all purposes plants are inanimate), Basque also shows personal gender in "hika", an old style form of 2nd person singular, still in use in some parts of the countryside, which conjugates the verbs differently depending on the interlocutor's gender. So if I'd use it here with the expression "you have written [DO]", I'd say "[DO] idazten dun" when addressing you but "[DO] idazten duk" when addressing a man.

Basque has not dual... nor do most surviving Indo-European languages. I'm not going to claim that Vasconic had dual but we cannot say for sure considering that we only have one survivor, while in IE we have a lot more to choose from and compare with, what may help shedding light on obscure topics like this one. If all the IE that survived would be English, for example, we could not ascertain the nature of early IE so precisely. The same but even more dramatic problem happens with Tyrsenian, because only a limited array of texts in two closely related languages (Etruscan and Lemnian) are known. It seems enough to guarantee IMO that Etruscan is not related to Basque in any striking way but deeper in time who knows.

As for associating language families with particular Y-DNA branches that seems to make no sense in southern latitudes. It's true that it seems to work very well in Northern Eurasia at least for Uralics, but the Boreal specialization is an extreme case; otherwise the populations involved seem to only rarely have reached the low numbers needed to cuasi-fixate their diversity in one particular lineage. If we consider, as we should, the (pre-Vinca) Neolithic Europeans as ethno-linguistically related, then we see a number of lineages showing up (at least G2a, E1b-V13, I2a, J2, etc.). They appear to have been diverse in this aspect, almost as much as modern Europeans. Locally a particular lineage or group of lineages may have reached quasi-fixation via founder effects (may be the case of some R1b subclades or possibly R1a too) but overall they look very diverse within West Eurasian variability.

rozenfag said...

@Kristina

Kazakh people speak Turkic language. Their case is actually unique - we actually know when they "formed", in 15 century. In short, at that time, after collapse of the Golden Horde, there were a number of successor states. One of them was Khanate of Abu'l-Khayr Khan. His descendents eventually became Khans of Bukhara. However, during his lifetime, two other Chingizids, Janybek Khan and Kerey Khan, rebelled, declaring independence and forming their own state - Kazakh khanate. Presumably Kazakh here means "free, independent". Therefore, we know exactly when the group of people gathered together and called themselves Kazakhs - it's the middle of 15th century.

Kristiina said...

Thank you Rozenfag, it is very recent indeed!

Thank you Maju for your clarification. It means that gender in Etruscan and Basque is more similar than I presented above compared to IE protolanguage.

Maju, generally, I agree with you that a smaller population number should favour fixation of one or two y lines, but I do not really agree with you that all Uralic groups have a fixation with yDNA N. On the other hand, yDNA N reaches high frequencies in some IE, Turkic and Paleo-Siberian groups.

Maju said...

@Kristiina: the IE and Turkic peoples you mention are almost certainly recent assimilated peoples from other backgrounds. As for Paleo-Siberians not sure which ones you have in mind (Kets?) but in any case the N1 correlation with Uralic peoples is extremely strong, while the main Paleo-Siberian lineage seems to have been rather Q1 - but as the N1 migration began probably soon after the LGM, there's been plenty of time for the buildup of some complexity. But the exception does not make the rule.

I don't mean that I know every detail about the genesis of Uralic peoples but I can discern the main tendency of it being tightly and patrilocally associated to N1, with two more specific founder effects which are N1b and N1c. Similarly we can identify proto-Amerinds or Paleo-Siberians with Q1 very much. But these "extreme" correlations with a single lineage are very rare or non-existent further south, at least after Neolithic. And not just in Europe, also in Africa and elsewhere. We typically see three or five or more lineages together in the big picture and very often at least two at the local level (aDNA). This still allows for some sort of founder effects in specific areas but we just can't associate the whole ethno-linguistic family to any single lineage, only particular branches at most (would be R1b-S116 or L11 in the Atlantic region the most notable, assuming it is indeed Neolithic).

Onur said...

Thank you Rozenfag, it is very recent indeed!

Kristiina, it is the formation of the Kazakh tribal confederation that is recent, but the formation of most (if not all) of the tribes that comprise the Kazakh tribal confederation are much earlier than the formation of the Kazakh tribal confederation and surely earlier than the Genghisid times.

the IE and Turkic peoples you mention are almost certainly recent assimilated peoples from other backgrounds.

Luis, based on the current distributions and some ancient DNA evidence, I am almost totally sure that Y-DNA haplogroup N was one of the main constituents of the Y-DNA haplogroup pool of Proto-Turkics. But I think there was already some local variation in its distribution levels in Proto-Turkics.

Maju said...

@Onur: proto-Altaic rather than specifically proto-Turkic. Yes, why not? But there is not such an outstanding specificity of N (often not N1) and rather other lineages like O3 and C2 dominate instead. So it's a blurrier situation in which various populations, with various "paleo" founder effects converged through time into a more or less unified macro-ethnicity (hard to discern the particulars of that process of convergence anyhow).

Notably Taiga Altaics show distinctive genetics, at least in the mtDNA pool. See this and scroll down for the update. This strongly suggests that they were assimilated and that they were not initially Altaic but something else.

Mike Thomas said...

Thanks Aram
I'm certainly under no pretense that despite being genetically very similar all farming groups spoke the same language. Quite the opposite. And I certainly don't support the Vasconic theory etc. Similarly later R1 bearing pastoralist groups all didn't speak the same language. That's been my point all along

Kristiina said...

Onur, I do not disagree with you at all. I am also quite sure of what you say, but the question is which clades and from which period. N1c-Z1936 looks very much Turkic and its split from the North-Eurasian branch dates c. to 2862 BC. However, expansion of N1c and N1b in northern Siberia may be relatively recent as I am only aware of one N from the Bronze Age West Mongolia (may be xN1c, N1b), one from the Iron Age N(xN1c, N1b) from Afontova Gora and two N1c from the Hunnu period from Egyin Gol, but it is true that we do not yet have any ancient yDNA from the truly northern parts.

Maju, I agree with you that Basques have a fixation with R1b (c. 85%), Eastern Finns have a fixation with N1c (71%) and Udmurts with N1c and N1b (85%), but on the other hand I have recently seen a graph showing that they are all high in homozygosity, Eastern Finns more than Udmurts and Basques more than Eastern Finns. Maybe it is the isolation that has slowly eliminated other y-lines in small communities without much foreign input.

Kristiina said...

Onur, for the sake of completeness, I forgot to mention that Shorian branch is wholly Turkic and must have been somewhere in Central Asia from a very early period, as well as Yakut branch. Interestingly, according to this Karmin et al paper, Yakut - Lebanese split dates to c. 2694 BC.

It looks like N1b was incorporated into the proto-Turkic gene pool at c. 1540 BC.

All estimates are from Karmin et al.

I do not know where these proto-Turks were in the Iron Age / Bronze Age and if Allentoft paper includes groups that were proto-Turkic speakers. Do you have any idea?

Grey said...

@Davidski

"Language change in ancient times happened either via massive migrations or conquest.

(snip)

You can't just claim that Indo-European languages spread with metallurgical skills..."

According to Wiki (which may be wrong)

https://en.wikipedia.org/wiki/Anatolian_languages#Hittite

"The records show a gradual rise to power of the Anatolian language speakers over the native Hattians, until at last the kingship became an Anatolian privilege. From then on, little is heard of the Hattians, but the Hittites kept the name. The records include rituals, medical writings, letters, laws and other public documents, making possible an in-depth knowledge of many aspects of the civilization."

That doesn't sound like direct conquest to me - also keeping the old religion sounds more like an internal takeover by a minority group. (Perhaps also why they turned out to be fragile.)

For that kind of takeover to happen it seems to me the minority group would have to have a significant advantage of some kind: mercenaries maybe (which would require horses imo), horse traders? artisans/traders of some other kind but whatever it was would have to be valuable enough for them to gradually become the elite.

Stuff like that makes me wonder if *some* of the earliest IE expansion wasn't by tribal groups at all but something else.

This might explain things like some of the IE languages having a lot of input from local languages, early kurgans behind enemy lines etc.

Mike Thomas said...

Kristiina

"
I do not know where these proto-Turks were in the Iron Age / Bronze?"

Wouldn't they be near mongolia and environs, if the earliest attested Turkic speakers (Xiongnu) came from around there ?

Aram Palyan said...

Grey

I don't think using a word 'enemy' is wise. The ancient people interactions were not based on the linguistic nationalism approach that arise in the last centuries with secular states.

In fact it is Nesites (Hittites ) who must be thanked for the preservation of Hattic texts. They also borrowed many religious concepts from natives. But the inverse is also true. For example the leading god of Hurrians Teushpa is a typical IE thunder god. And his story of fight with his father Kumarbi is basically the same story as the Greek Zeus fighting Kronos. So non-IE people were also borrowing ideas from IE people.

Of course there is also the inverse theory were it is the Greeks and Hittites who borrow the thunder god. But the fact is that Hurrians were using Indo-Aryan terms for many things related to horse.

So there was a deep cultural interactions between this various groups.

Onur said...

I do not know where these proto-Turks were in the Iron Age / Bronze Age and if Allentoft paper includes groups that were proto-Turkic speakers. Do you have any idea?

Not sure exactly which ones of the Allentoft et al. samples were Turkic-speaking. The strongest candidate is RISE504, the Iron Age Kytmanovo sample, which is dated to as recent as the 8th-9th centuries AD and has Kazakh-Kyrgyz-like autosomal results.

Maju said...

@Kristiina: Basques would be more homozygous than Eastern Finns? I have a hard time believing that. As far as I know Basques show relatively high homozygosity for Southern Europe or in general West Eurasia but nothing of the kind that is found in the high Northern latitudes. Eastern Finns should be more homozygous and probably Western Finns too.

"Maybe it is the isolation that has slowly eliminated other y-lines in small communities without much foreign input".

I've been reading that kind of line since, well, since I first read Razib's blog maybe in 2006 or 2007. It didn't make any sense back then and it does not now. Fixation tends to accumulate in the dominant line, a drip of foreign lines would be simply erased or kept small.

IDK, think Pygmies, who have been absorbing Bantu lines for millennia and yet show a clear distinctive personality. Nobody doubts the Pygmy or Pygmy-like origin of those lineages that, even found also among Bantu, appear to have a Pygmy centrality. So why this strange obsession with R1b? Just because it becomes personal (as we are discussing among mostly people of European origin, many of which have that very lineage themselves, and the discussion has not got the attention of people of African descent).

In any case, sticking to Europe and R1b: forget Basques and think Iberians, who did not speak IE in the eve of written History either, were much more numerous and civilized back then and still have the same kind of Y-DNA structure, having more R1b than Indoeuropeans from the West (Portuguese, etc.) or their likely residuals in the Ebro basin (Aragonese). The whole pattern of R1b-S116 distribution is yelling "not IE!": relatively high among Basques, Iberians, areas of the Atlantic where IE only arrived a few Centuries before Rome (La Tène), noticeably lower in areas where we know IEs were in earlier: Germany, West Iberia, etc. Also the geostructure of the haplogroup R1b-S116 strongly suggests a South French origin.

rozenfag said...

@Onur

Yes, I should have clarified this. Whenever we have creation of a nomadic state (Kazakh khanate, Gokturk Khaganate, etc.), the tribes, that this state unite, are older.

@Mike Thomas

Not all linguists agree that Xiongnu were Turkic speakers. For example Alexander Vovin thinks that they were speaking Yenisean language.

Maju said...

@Grey: What for you does not sound to conquest, it does to me. You may argue against "direct conquest" and indeed it may include a transitional period of ambiguity in which both ethnicities co-existed but the IEs stuck to power positions, most likely as soldiers rather than crafters (what a strange idea!) This would be somewhat similar to what happened in Mesopotamia between the first Semitic expansion into what today is called "the Sunni triangle" (curious how millennia old differences persist through formal ethno-linguistic and religious change) and the Akkadian Empire (which, like Hittites with Hattian, retained for some time the old language, Sumerian in that case).

Aram Palyan said...

Kristiina

Thanks for the link and title.

""Aram, do you think that Etruscan and IE languages belong to the same family? If Vasconic, IE and Etruscan are all unrelated, your logic does not work. However, it may be turn out that Basque is R1b dominant language and Etruscan is I2 dominant language.""

I have no opinion about Etruscans. I use the consensus of linguistic community for my theories. I think that Etruscan, Basque and IE has no relations. Of course I maintain some doubts also.

My logic is that every case should be examined going deep into his own details. All information should be taken into account. The Y DNA, the mtDNA, the autosomal components, the linguistic.

I use also historic analogies. When Kassites invaded Mesopotamia, they didn't impose their language. Despite being rulers. The reason is obvious because there was a much superior culture in Mesopotamia and compared to it they were barbarians. But if the same Kassites invade another region with low density population then a language shift can occur.
Later Kassites themselves started to use Semitic names. The last time were Kassites are mentioned by Greek historians, most probably they were already a Iranian speaking tribe.

Or another example. The case of Hazaras. They have clearly a Turkish/Mongolian genetic input. Quite large. But they speak Persian. Many other nations have much lesser East Asian components but they did have shift to Turkic. How that happened? Because the linguistic shift is not only a matter of genetics but also something related to the people's internal values, political situation and religious believes. But the genetics do also matter of course. My logic is to take as much as possible factors into account.

Shaikorth said...

Maju, the Basques are indeed more homozygous than Norwegians, Finns, British, North Russians, Belarusians and so on. This is obvious if you test HGDP or 1000genomes samples, or the Human Origins set (which is largely taken from the previous ones). So are the Sardinians. Other South Europeans are heterozygous as a general rule, but what applies to them does not apply to these isolates. Perhaps some regional isolates in those North European regions are more homozygous, but broadly they're not.

Kristiina said...

Mike
In ancient Mongolia, there is more R and Q than N, although C and O are most prominent. Look at these figures, p. 92 (http://221.145.178.204/nrichdata/csd/book/file/KBJ05_02_01.pdf)

Also in Altai, there is more Q than R, and no N in this recent analysis: http://dienekes.blogspot.com/2014/06/ancient-dna-from-bronze-age-altai.html

Xiongnu have much more Q than any other haplogroup, so that Yeniseian claim is genetically justified:
Pengyang County: 4x Q1a1-M120
Heigouliang, Xinjiang (Xiongnu kings summer palace): 6 x Q1a*(xQ-M120, xQ-M25, xQ-M3), 4 x Q1b (M378) ja 2 x Q*(xQ1a, xQ1b)
Hengbei site, Shanxi: 9 x Q1a1, 2 x O2a-M95, 1 N, 4 x O3a2-P201, 2 x O3, 4 x O*
Egyin Gol: 2x N1c, Q-M242 ja C-M130
Duurlig Nars: R1a1 ja C3

So, maybe Uralic N1b and N1c will be found in the forest area between the Baltic Sea and Altai and Yakut and Shorian lines in Kazakhstan.

Onur, by the way, I think that Yakuts were decimated by Genghis Khan and other Mongolic troups and that's why a few survivors fled to Sakha.

Onur said...

Onur, by the way, I think that Yakuts were decimated by Genghis Khan and other Mongolic troups and that's why a few survivors fled to Sakha.

That is what both history and genetics seem to point to, but it is up to debate whether Proto-Yakuts (who lived in the Baikal region) were decimated by Mongols, or a minority of Proto-Yakuts participated in the migration to the north with the rest mostly submitting to the Mongol rule and staying in the Baikal region, or Proto-Yakuts were already a tiny group during the Mongolian invasion.

Kristiina said...

Onur, history is fascinating. This is what a DNA paper tells about Yakuts:

The origin of the most frequent Y-chromosomal haplotypes (Ht1 and Ht2) was difficult to establish on the basis of genetic information. Indeed, these two lineages belonging to haplogroup N1c seem to be restricted to Yakut populations, and were probably present since the period they were first located in Central Yakutia. Interestingly, the comparison with archaeological data revealed that the male individuals at the beginning of the 18th century, identified as Clan Chiefs (or tojons) on the basis of their grave goods (weapons, jewelry, silk clothes, richly ornamented saddles and signet rings), belonged to these two haplotypes. Therefore, archaeological data could bring interesting information in tracing back the origin of these enigmatic male lineages. Indeed, the grave goods of the 15th/17th centuries (weapons and horse harnesses) and the construction of coffins with an empty trunk from the 18th century are similar to the burial customs of the Cis-Baïkal area and of the Egyin Gol Necropolis during the 3rd century BC. This suggests that the male ancestors of the Yakuts were probably formed of a small group of horse-riders originating from Northern Mongolia or the Baïkal Lake.

and:

Based on the analyses of the maternal and paternal lineages of ancient Yakuts, we were able to demonstrate that the formation of this population started before the 15th century, with a small group of settlers composed of horse-riders from the Cis-Baïkal region and a small number of women from different South Siberian origins.

(http://www.biomedcentral.com/1471-2148/10/25)

They did not necessarily go to Sakha for fun as it is pretty cold there.

Kristiina said...

Onur, you are right that RISE 504 is close to Kyrgyz. According to Genetiker, Rise504 is yDNA J2a2 and mtDNA C4a1d. I agree that he probably spoke Turkic.

I compared RISE504 with RISE554 from Iron Age Afontova Gora, which according to Genetiker is the same N as IR1, and I see that they are similar. RISE504 is more South Asian and Chinese and RISE554 is more Siberian and Native American. Western Eurasian hunter gatherer ancestry is the same.

RISE554 was found in the same place as RISE553 (yDNA R1a1a1 without further subclade), and they were almost contemporary, c. 1000 BC. Kytmanovo RISE504 is more than thousand years younger. Maybe RISE553 and RISE554 were proto-Turkic speakers, although RISE553 seems to be related to Sintashta as he is the only one to have ENF of these three.

rozenfag said...

@Kristiina

Which program are you using to compare genomes? (I am noob in such questions)

Can you compare RISE504 to Altaians?

Davidski said...

Here you go. Just stick this into a text file and then open with any spreadsheet software.

Sample ID Nganasan Naxi European_HG Afanasievo Atayal Pulliyar Chukchi Papuan Esan Bedouin Karitiana San
RISE_irRus RISE504 0.300618 0.268739 0.057523 0.326976 0.005087 0.00001 0.00001 0.003197 0.00001 0.00001 0.037811 0.00001
Altaian altai280p 0.341442 0.360563 0.083568 0.148127 0.001579 0.012953 0.00992 0.000847 0.00001 0.015229 0.025752 0.00001
Altaian altai431k 0.332276 0.32442 0.082001 0.159317 0.011336 0.007046 0.03171 0.00001 0.00001 0.026807 0.025056 0.00001
Altaian altai283p 0.334053 0.353591 0.096427 0.15367 0.011717 0.007133 0.019133 0.00001 0.00001 0.00388 0.020367 0.00001
Altaian altai380k 0.351025 0.309519 0.087676 0.155509 0.009182 0.009355 0.041912 0.00001 0.00001 0.015917 0.019874 0.00001
Altaian altai363p 0.357725 0.361942 0.062511 0.139038 0.003471 0.018798 0.011515 0.00001 0.00001 0.027685 0.017286 0.00001
Altaian altai409k 0.405306 0.305093 0.066218 0.147322 0.005816 0.007059 0.0281 0.008265 0.00001 0.016022 0.01078 0.00001
Altaian altai419A 0.339048 0.315028 0.082924 0.169663 0.009964 0.005837 0.031234 0.00001 0.00001 0.036706 0.009566 0.00001

Full spreadsheet here...

https://docs.google.com/spreadsheets/d/1ajolEB_2NXAnxtGJXSbwKSF1CvDvFiGy5otpq5f-C1g/edit?usp=sharing

rozenfag said...

@Davidski

Thank you very much!

Simon_W said...

It's striking how much the Chalcolithic and Early Bronze Age central Anatolians from Alishar Hüyük resembled the Danubian farmers: The neurocrania small, longish in shape (dolicho- to mesocephalic) and high vaulted; the face small and narrow, but the nose rather broad in shape. The Early Bronze Age inhabitants from Alishar Hüyük in numbers:

cranial index = 75.4 (low mesocranic)
length-height index = 75.3 (high vaulted)
upper facial index = 54.2 (quite elongated)

In the Middle Bronze Age the crania became different: Large, somewhat broad-shaped (moderately brachycephalic) and low vaulted, with somewhat broader faces and moderate browridges. In numbers:

cranial index = 79.5 (high mesocranic)
length-height index = 69.1 (low vaulted)
upper facial index = 52.5 (mesene)

I think typologically we can say they go somewhat into an "Alpinid" direction (this doesn't imply a connection with the Alps of course). The change is striking and most definitely must have been associated with thorough genetic changes. And I think the Middle Bronze Age may be the time when the ancestors of the Anatolian IEs arrived in Anatolia. Otherwise we would have to speculate that the Hatti people arrived that late.

Aram Palyan said...

Mike

Thanks to invite my attention to the branch R1b-PF7558 - Sardinians, Arabs, Jews.
I looked again at the database and I see that there are many untested Armenians who are probable candidate for that branch. It is certainly something centered to North Near East (most probably Anatolia), later expanded by Jews. And the few Arabs could be related to Turkish expansion.

Aram Palyan said...

BTW Gioiello certainly has found interesting facts. Many basal forms of haplogroups pops up in Italy ( Sardinia included ). Especially in Tuscany.
For example the J2b2 ( http://yfull.com/tree/J-Z1825/ ). Age 16000 years but the basal SNP is in Tuscany. The notable thing of this branch is that it is present in India.
And I notice sometime things like this in other haplogroups also.

Very interesting.
Is it a LGM Franco-Cantabrian refugium effect or something related to Roman Empire, time will show.

Simon_W said...

Now Wikipedia calls the theory of Middle Bronze Age mass migrations in Anatolia "outdated". But I would be cautious about this, since all the old evidence suggestive of violent destructions was hardly all bogus? And then there's the above mentioned cranial evidence. The same article also seems to suggest that Indo-Hittite was native to Anatolia, which is very unlikely in the light of ancient DNA evidence.

https://en.wikipedia.org/wiki/Middle_Bronze_Age_migrations_(Ancient_Near_East)

Check the nice map:

https://en.wikipedia.org/wiki/File:Mass_migration_of_Greece_and_Turkey_in_1900BCE.svg

Maju said...

@Saikorth: I haven't seen that data (waiting for a reference) and I have seen (years ago, don't recall where) the opposite data, sorry.

Just searching around I found this entry where it's apparent that Basque homozygosity is only slightly larger than French one and lower that East Asians in general, so its roughly in the European pattern. Basques are only notable in short ROH homozygosity, which implies ancient bottlenecks (modern inbreeding is defined by long ROH). Sardinians are even less notable and almost average European in all this. Sadly no Far Northerners to compare with (Russians are probably from the center or south).

Incidentally East Asians (all them) are most strikingly homozygous in the medium ROH, so the bulk of their bottleneck is quite apparently more recent than that of Basques (although some southern populations also show an old bottleneck and many show at least some recent inbreeding as well).

For Finns, I found this article, whose data clearly shows that even Southern Finns are more homozygous than Swedes, and of course NE ones are even more. The data is not directly comparable with the previous anyhow but it's clear that there is a quite direct correlation with latitude (Stockholm is quite more to the south than Helsinki) and inland habitat.

I could not find any direct Basque-Finnish (or similar) comparison on this matter.

Onur said...

I compared RISE504 with RISE554 from Iron Age Afontova Gora, which according to Genetiker is the same N as IR1, and I see that they are similar. RISE504 is more South Asian and Chinese and RISE554 is more Siberian and Native American. Western Eurasian hunter gatherer ancestry is the same.

RISE554 was found in the same place as RISE553 (yDNA R1a1a1 without further subclade), and they were almost contemporary, c. 1000 BC. Kytmanovo RISE504 is more than thousand years younger. Maybe RISE553 and RISE554 were proto-Turkic speakers, although RISE553 seems to be related to Sintashta as he is the only one to have ENF of these three.


Well, to make firm suggestions about the location of Proto-Turkic speakers during the Late Bronze or Early Iron ages I would first want to see a broadered range of sample results from the same era.

Maju said...

@Aram: anything pointing to Italy as origin is definitely not "Franco-Cantabrian refuge". Italy was totally out of the Solutrean and Magdalenian cultures (the frozen Alps were effectively impassable in those times) and had instead its own Epi-Gravettian dynamics instead. Lineages stemming from Italy should be thought as Impressed-Cardium legacy most likely (unless a Roman origin can be somehow demonstrated, of course).

Kristiina said...

Maju, here is the link http://www.forumbiodiversity.com/showthread.php/44636-Homozygosity-and-divergence-among-Europeans

Maju said...

@Kristiina: let me be skeptic considering the other data mentioned before. I'd be interested in internal divergence data for Basque and Sardinians, because it seems more consistent with recent endogamy than raw homozygosity, whose figures, as per Lemminkäinen, are almost the same across the board. In internal divergence, Northerners tend to score low (relatively inbred), while southerners score high, all as expected more or less. Instead in his homozygosity scores it's almost the opposite, what suggests that some hidden confusing factor is at play, such as maybe admixture from East Asia or whatever.

Grey said...

Aram

You're clearly right about the literal meaning of "enemy lines". I use it to mean evidence of one culture seemingly popping up beyond their cultural borders without any clear path of conquest.

One common reason for that is mercenaries, another might be traders. Either way it implies the intrusive group had something valuable the receiving group wanted.

Simon_W said...

Regarding the debate about R1b, I think it has become obvious by now that present-day distributions of haplogroups are not a reliable guide to ancient distributions. Who would have expected from present-day distributions that the predominant haplogroups of EN and MN European farmers were G2a and I2a? Who would have expected that Scandinavian hunter-gatherers and their colleagues in Luxembourg were dominated by I2a? And wasn't it surprising that there was C1a in Iberian hunter-gatherers? The picture provided by ancient DNA is one of discontinuity as the rule, not the exception. That has become so self-evident now that I find it disturbing that I even have to write this!

According to Maju the geostructure of R-S116 suggests an origin in Southern France and a non-IE one at that. Yes, R-S116 is rather common in France and all over western Europe. In fact, it's most of all R-U152 that dominates in Southern France (outside Southwestern France), and this marker is even more common in Northwestern Italy and Corsica. Does this suggest a non-IE origin? Not necessarily. In Southern France, Northwestern Italy and Corsica there were the Ligures, an IE population linguistically related to Celts. So how can anyone conclude that R-U152 definitely must be non-IE? After all, non-IEs once used to live almost everywhere except the PIE homeland. So with that reasoning every haplogroup can be declared pre-IE.

I do agree that R-DF27 does look basically non-IE. But it is just one variant of R-S116, it may have branched off from an IE R-S116.

I have to declare here that I'm not a dogmatist. If new aDNA evidence should turn up that makes a pre-IE origin of R-S116 more likely, then I will change my opinion immediately. The same applies to the origin of R-Z93. The thing is just that, based on the currently available evidence, I don't believe this will happen. But anyone is free to favour the theory he or she considers the most likely one, we're not the IS! Nobody is forced to believe anything.

Grey said...

Maju

"What for you does not sound to conquest, it does to me. You may argue against "direct conquest"..."

Quite.

It doesn't sound like a conquering horde to me - especially with all the compromises over language and religion. It sounds like a minority group who managed to take over over time - maybe it started with a conquest of a small region or a migration into terrain that wasn't being used or maybe it started with being invited in for some reason.

.

"This would be somewhat similar to what happened in Mesopotamia between the first Semitic expansion into what today is called "the Sunni triangle""

That's what I was thinking also.

.

"most likely as soldiers"

That would be my first thought if they were known as horsemen already but that seems up for debate.

.

"rather than crafters (what a strange idea!)"

I don't understand why it's strange. How are the massive technological changes that occurred in this era: wheel, metallurgy, domesticated horses etc, going to spread?

I'd say the most likely option is a family of artisans moves from the source region to the nearest trade node in a particular direction and later a younger son is sent to the next node in that direction and a generation later his younger son moves to the next node and so on.

Taking Srebny as the start point for the sake of argument and going mainly by sea then how many nodes to Egypt?

Srebny
->mouth of Danube
->west Anatolia
->Cyprus
->Egypt

four generations? 100 years?

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