search this blog

Saturday, March 28, 2015

Population genetics of Copper and Bronze Age inhabitants of the Eastern European steppe


I'm hoping like hell that the samples from this thesis eventually get the same treatment as those from Haak et al. 2015.

Summary: This dissertation presents the first genetic study of prehistoric populations in the Pontic-Caspian steppe from the Upper Thracian Plain to the Volga. Hypervariable region I (HVR I) and 30 short sections of the coding region containing 32 clade- determining polymorphisms on the mitochondrial DNA, as well as 20 putatively naturally selected autosomal SNPs and a sex-determining locus were analysed using a combination of multiplex PCR and 454 sequencing. Data analysis was performed on the HVR I of 65 of the 180 Eneolithic and Bronze Age samples. (Partial) genotypes were generated from 61 individuals. Published ancient DNA data from Central and Eastern Europe and Central Asia, as well as modern DNA sequences were consulted for comparison.

The genetic data support the inference that early Neolithic farmers from Southeast Europe were involved in establishing pastoralism in the steppes by demic diffusion. The consistently low values of the FST-statistic (the range includes zero) between the Yamnaya Culture of the steppe and a succession of Neolithic cultures in Central Europe indicate continuous or recurrent contacts between the two regions. Between the Yamnaya Culture and its successor, the Catacomb Culture, the incidence of haplogroup U4, which is at high frequency in hunter-gatherer populations of Neolithic Scandinavia and Mesolithic Northwest Russia, rises from approximately 5 % to above 30 %. It is possible that immigrants from Eastern Baltic hunter-gatherer refugia were involved in the genesis of the Catacomb Culture.

The low FST values between the prehistoric steppe populations and the modern populations of Central and Eastern Europe indicate genetic continuity. This is supported by the nuclear genotype frequencies. According to current knowledge the modern European gene pool can be explained by three roots: indigenous Mesolithic hunter-gatherers, early farmers from the Near East, and an ancient North Eurasian component with an Upper Palaeolithic origin. Maybe the third ancestry component was introduced into the late Neolithic European genome by the North Pontic population.

Source: Wilde, Sandra, Populationsgenetik kupfer- und bronzezeitlicher Bevölkerungen der osteuropäischen Steppe, 2014, Dissertation

102 comments:

Nirjhar007 said...

'' Published ancient DNA data from Central and Eastern Europe and Central Asia, as well as modern DNA sequences were consulted for comparison.''
Which aDNA from Central Asia David?

Nirjhar007 said...

Another Interesting Thing Is that ''it is been suggested'' that Catacomb Cultures Burial ritual had roots in South-Western Turkmenistan from the early 4th millennium (Parkhai cemetery).

Colin Welling said...

The consistently low values of the FST-statistic (the range includes zero) between the Yamnaya Culture of the steppe and a succession of Neolithic cultures in Central Europe indicate continuous or recurrent contacts between the two regions.

Uhhhhh, what does that mean? In one line they say how similar the steppe was to neolithic central europe but then they also say the steppe introduced ANE to central europe in the late neolithic.

Is there any word on ydna? I was expecting it in Wilde's first paper, but we know how that turned out.

Davidski said...

What she's saying is that the Near Eastern mtDNA in the steppe groups probably comes from the Neolithic farmers just west of the steppe, not directly from the Near East. This contradicts Haak et al., but the Yamnaya samples here are from much further west than the Samara.

Mike Thomas said...

It is basically saying what I always maintianed ; late Cucuteni-Tripolye groups settled the Pontic steppe to a large degree, as well as late Neolithic Polish groups, on which basis the Yamnaya culture emerged; with further input from the Caucasus. The type of pastoralism in the western steppe derives from late Neolithic eastern and southeastern European groups, which different to central eurasia and eastern eurasia in turn.

I further maintain that the corded ware culture is also indigenous to the north European plain and not steppe derived.

This was always obvious to anyone who's examined the issue closely ; and was merely a matter of time genetic evidence collated. I hope soon we might get fuller data from CT and other late Neolithic and residual forager groups from Poland and the baltic, including Y DNA groups.

Davidski said...

Yes, and then the Indo-Europeans expanded from the steppe, taking ANE and R1a/R1b to Europe, and R1a/R1b to Asia.

Nirjhar007 said...

'' and R1a/R1b to Asia.''
Another Prophecy David?.

Krefter said...

Wilde. 2014 and Haak 2015's data suggest the Pontic Steppe had very differnt mtDNA than Neolithic Central Europeans. How could this new data from the Pontic Steppe completely contradict the old data?

Krefter said...

"Afanasievo, Okunev, Andronovo, Sintashta DNA?"

http://dienekes.blogspot.com/2015/03/afanasievo-okunev-andronovo-sintashta.html

Krefter said...

For the positively selected-SNPs and mtDNA they need to start testing Volga-Pops who autosomally are the most similar to Yamna today.

Ukrainians and Russians are mostly from recent immigrants from Central Europe, with a big chunk of Gok2-like blood.

The East Asian-ancestry in Volga-Pops will mess somethings up, but they might turn out more similar to the ancient Pontic Steppe.

Matt said...

So labels on the phenotype SNP analysis are (name of culture and normal dating, not necessarily the dates they sampled):

ANL - Eneolithic = Mesolithic (pre c.3500 BC?)
JAM - Yamnaya c. 3500 BC – 2000 BC?
KGK - Catacomb Grave Culture c.2800–2200 BC
POL - Poltavka Culture c.2700—2100 BC
KAK - Globular Amphora Culture c. 3400–2800 BC
aDNA - Complete data set of the old sample (a bin of everything published)
mUKR - Modern Ukraine

Points

- Eneolithic lacks both Asian "thick hair" EDAR and blue eyed HERC2, has some derived SLC45A2.
- Catacomb Grave does not shift towards lighter pigmentation. Poltavka Culture does but is still darker than modern Ukraine.
- No LCT A lactase persistent types in the 30 strong Catacomb Culture sample. While I would presume this to be T−13910 which we find in Bell Beaker and in the Basque country at around this time.
Interesting that there is this LCT B though - there is some evidence that an SNP which is almost always found with A−22018 in West Eurasians has independent predictive value in Kazakhs.
- "Between the Yamnaya Culture and its successor, the Catacomb Culture, the incidence of haplogroup U4, which is at high frequency in hunter-gatherer populations of Neolithic Scandinavia and Mesolithic Northwest Russia, rises from approximately 5 % to above 30 %." Infusion of EHG via females?

Krefter said...

Interesting that EDAR pops up again. Mesolithic Motala and now Bronze age Yamna.

Bernard said...

Sandra Wilde published a paper last year: Direct evidence for positive selection of skin, hair, and eye pigmentation in Europeans during the last 5,000 y. See here: http://www.pnas.org/content/111/13/4832.full

Matt said...

Couple more - SLC24A5 derived variant and ABCC11 derived variant

http://i.imgur.com/Ii9M6DV.png

ABC11 is derived variant (frequent in East Asians) linked with dry earwax, which is recessive so a sample would have to have two derived copies to be different (so none of the populations would've really differed on average much on this trait).

However, their test of significance on ABC11 does find significant differentiation from modern Ukrainians on this SNP, albeit at a low level (modern Ukrainians less derived, more wet earwax pheno). Also the alcohol dehydrogenase ADH1B.

Krefter, the individual mtdna haplogroups are in the PDF (http://ubm.opus.hbz-nrw.de/volltexte/2015/3975/pdf/doc.pdf), so perhaps you could look at those if you were interested in how differentiated they seem? (p352 of PDF, page number 336)

There are maps for sample location as well. (p340/num324)

Krefter said...

Thanks alot matt.

M. Myllylä said...

SHG seems to be today highest in Eastern Baltic Sea region, not in Western Scandinavia.

Krefter said...

There's only a handful of IDs in Tabelle 13.12 that weren't in Wilde's previous study. Not much new with mtDNA. The cultural labels have changed, I guess now it's more specfic.

The Globular Amphora samples in this study were Catacomb samples in Wilde's previous study.

The mtDNA is not similar to Neolithic Europeans, beside having WHG-SHG-EHG and Near eastern mtDNA. IMO, this reveals a more simple approach to comparing pop's mtDNA is more reliable than formal statistics.

Davidski said...

Krefter,

Yep, it looks like there's a new paper on the way with ancient DNA from Yamnaya, Sintashta, Andronovo and Afanasevo.

https://drive.google.com/file/d/0B9o3EYTdM8lQQlB1N3hZbFhfUVU/view?usp=sharing

http://translate.google.com.au/translate?hl=en&sl=ru&u=http://polit.ru/article/2015/03/15/kozhintsev/&prev=search

I think this is the same one that I've been hearing about that has ancient DNA from Bronze Age Armenia, and will soon be published in Nature.

Krefter said...

Looks like Steppe groups had more 374F(rs16891982 CG or GG) than Neolithic Central-West Europeans. Bronze age Germany-Hungary has the most of all of them.

Here's the frequency of 374F in Pre-Historic Euros.

Mesolithic.
WHG: N=3: 100% CC(3)
SHG N=9: 22.2% CC(2), 22.2% CG(2) 55.6% GG(5).
EHG: N=2: 50% CG(1), 50% GG(1).

Neolithic:
Central-North N=20: 55% CC(11), 25% CG(5), 20% GG(4).
Spain N=6: 66.7% CC(4), 33.3% CG(2)

Bronze age.
Steppe: N=28, 39.28% CC(11), 35.7% CG(10), 25% GG(7)
Germany+Hungary N=18: 11.1% CC(2), 39% CG(7), 50% GG(9).

I'm pretty sure Bronze-Iron age Siberians had close to 100% GG.

Krefter said...

Thanks for the info. Steppe migrations towards Asia and Europe from the same time period will finally be proven.

The samples from Keyser 2009 traced their ancestry to the Steppe not EHG, east Asian, etc., etc. people running around northern Siberia who miraculously became just like Yamna mtDNA wise.

Brothers R1a-Z93 in Asian Steppe-like people and R1a-Z283 in European Steppe-like people from 1000BC, I wonder where their father-clade came from?

Chad Rohlfsen said...

Anyone speak Russian?

http://dienekes.blogspot.com/2015/03/afanasievo-okunev-andronovo-sintashta.html?m=1

Krefter said...

@Chad,

Davidski gave a link to an English-translation of that article.

http://translate.google.com.au/translate?hl=en&sl=ru&u=http://polit.ru/article/2015/03/15/kozhintsev/&prev=search

Here's a picture he showed in the same post.

https://drive.google.com/file/d/0B9o3EYTdM8lQQlB1N3hZbFhfUVU/view?usp=sharing

Mike Thomas said...

@ Krefter

"Steppe migrations towards Asia and Europe from the same time period will finally be proven. "

Krefter I very much think it was more complex than this. and the evidence is mounting , everything points to what I'm saying

"EDAR present in Motala and BA steppe"..

Clearly then EDAR was selected out in Europe, which has less to do with extinction of SHG type peoples


@ M Myllyla

You're correct. Modern Baltics are basically SHG ! Something I pointed out few days ago

@ Davidski

"PIE expanded from steppe, taking R1a/R1b to europe ... Central Asia "

Possibly; Dave. But every new study merely points to the more complex scenarios I've been intimating at (which dovetails with any more -than-superficial analysis of other lines of evidence (archaeology, linguistics)).

Even if you want to narrow down to one Y haplgroup - R1a- the evidence is still not there for a "steppe origin". And what's more important is the mechanisms and range of its expansions of M417.

I trust more studies will soon reveal this.

Chad Rohlfsen said...

Thanks, I was just hoping for a more precise translation. We're home now too. I'm done driving for a while!

Tobus said...

@Krefter: I'm pretty sure Bronze-Iron age Siberians had close to 100% GG.

What makes you think that? Modern Siberians have very low frequencies (Chukchi highest on ALFRED with 32%, most are under 20%)

Here are the overall frequencies of the G allele from the stats you quoted

Mesolithic.
WHG: 0%
SHG: 78%
EHG: 75%

Neolithic:
Central-North: 22%
Spain: 17%

Bronze age:
Steppe: 43%
Germany+Hungary: 70%

It looks like an introgression from SHG/EHG into Europe diffusing to Steppe to me - in which case Siberians would more likely be 100% CC than GG... no? Have I missed something here?


Davidski said...

Bronze Age Siberians...

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

Tobus said...

Thanks for that David - they include SLC45A2 in the study (rs16891982) but unfortunately only give results as part of a 3-SNP "Eu" or "Asian" test, so no way to really tell for sure what the frequency is - you don't know if they published the raw SNP result (or genomes) anywhere do you? I couldn't see it in the Supp. info.

Extrapolating from the data, assuming each "Eu" result was homozygous derived we'd have 88% "GG", assuming heterozygous would be 44%, so the real frequency is probably somewhere in between... much higher than I would have expected. Makes me increase the idea of "EHG territory" I have in my head.

Davidski said...

The Andronovo and Scytho-Siberian nomads are almost all R1a-M417 with Yamnaya-like mtDNA, so they're early Bronze Age migrants from Eastern Europe to south Siberia.

Krefter said...

Tobus,

My guess is all the "Eu"s had GC or GG, because pretty much no one in Asia has GC or GG except Middle easterns(still rare).

That means almost all of them had GC or GG. The "Asian"s can be maybe be explained by East Asian admixture.

Here you can see their frequency of the derived allele is comparable to Bronze age/Late Neolithic Germans and Hungarians.

https://drive.google.com/open?id=1PVpN5zC3vW-FC_IUzFMaf6JI2XezGooJgsoIRagKyy8&authuser=0

Samples dating to 2,500BC-1,000BC have a significantly higher frequency of 374F than earlier(and some from the same era) samples and are comparable to modern Europeans in other markers to.

Mike Thomas said...

@ David

"with Yamnaya-like mtDNA, so they're early Bronze Age migrants from Eastern Europe to south Siberia"

But we now know this "yamnaya-like " mtDNA is actually Central and southeastern european and West Asian . ....


Davidski said...

The Yamnaya-like mtDNA structure is specific to Eastern Europe.

The fact that it was formed from the mixing of indigenous Eastern European hunter-gatherers and Near Eastern-derived farmers during the Neolithic is irrelevant to the question of whether the steppe hypothesis is valid or not.

Mike, you can't keep using the Neolithic transition as an argument against the steppe hypothesis. I'd say that anyone but complete novices in this area can see through this argument. What's the point?

Mike Thomas said...

Davidski

There was no neolithic transmission on the steppe .

Anyone but a complete novice knows this :)

Once you come to terms with this fact; then everything will fall into place for an overall understanding of what's happening

Nirjhar007 said...

Well all i can say that any person who believes in Logic and Reasoning will say that We should look for Central Asian aDNA in large numbers and also should be aware that Steppe Hypothesis has the advantage not because its practical but its very popular and well sold one it will take some time to people in numbers to understand this.

Nirjhar007 said...

David,
''The Andronovo and Scytho-Siberian nomads are almost all R1a-M417 with Yamnaya-like mtDNA,''
That's irrelevant because they are close to the European area so similarity is obvious but not defining, we need aDNA from south to establish a agreeable genetic origin of those people...

Mike Thomas said...

Sorry
That's should have said "no neolithic transition on the steppe.

Ie : what we're seeing with this mtDNA evidence is not some earlier, remote arrival of farmers , but the very catalyst behind the Yamnaya phenomenon.

It's all clear, David , I don't know why you're befuddled.

Davidski said...

Maternal lineages from Neolithic farmers entered the gene pool of western steppe populations well before 3300 BC, and the Neolithic transition did move right up the the Dnieper on the steppe.

These two events might well be related for at least a part of the Yamnaya horizon. In fact, they look like the same process to me.

You can call it what you like, if you don't want to call it the Neolithic transition, but whatever you call it, it still won't have any relevance to the validity of the steppe hypothesis.

Tobus said...

@David/Krefter:

Yep, makes sense... and frequency has since been ameliorated by Persian/Turkic/Mongol/Han etc. expansions.

Mike Thomas said...

Davidski

Simply: the Yamnaya pastoralists dervive from certain Eastern European late Neolithic groups. So it seems a bit arbitrary that you're assigning yamnaya as "point zero " when the trail clearly begins elsewhere and earlier.

And I suspect that R1a expanded from the forest steppe; not the steppe , at least initially

Nirjhar007 said...

Mike, On Pastoralism of Yamna it seems they have learnt it from the west Siberian tribes, their eastern neighbors, and, if so, it was of Central-Asiatic, Iranian derivation.

capra internetensis said...

@Nirjhar

Do you have any actual evidence for that? I mean besides speculation by Matyushin from 30 years ago.

Mike Thomas said...

No Nirjhar, Yamnaya pastoralism isn;t an offshot from Iran, and the archaeo-zoological evidence is clear.


Goats and sheep dominated assemblages in Central Asia, as they did in the far eastern steppe (possibly having spread from Central Asia via the Tien Shan mts)

On the other hand, cattle and pigs were dominant in the western (Black Sea) steppe, as they had been in Late Neolithic central - eastern Europe

The central Eurasian steppe had its own assemblage peculiarity - masses of horse bones, which were killed for food.

The independent features of steppe economy gradually diffused and became more mixed east & west over the latter third Millenium, into the 2nd.

Skilur said...

@Krefter
After looking at the Fst values Ukrainians seem to be closest (smallest difference) to steppe populations. Or i understood something wrong there? What does negative fst values actually mean?

Nirjhar007 said...

Capra, You can read this book if you like,
https://books.google.co.in/books?id=jwmgRFmKBMIC&q=prehistoric+russia&dq=prehistoric+russia&hl=en&sa=X&ei=RqMXVYfPIYOpuwSOgYLIDg&redir_esc=y
Mike,
Ok tell me What was the Core of Yamna Stock-Breeding? and How important were Goat and Sheep?

Davidski said...

Yeah, Ukrainians show the lowest (most negative) Fst, and thus highest affinity, to Yamnaya (YAM) and Yamnaya West (YAW), and also Corded Ware (CWC).

Mike Thomas said...

Nirj

For the western steppe (from the lower Danube in Romania/Bulgaria to the Urals), the Pontic steppe was dominated by cattle & pig bones. Further east, in the Caspian steppe, goats and sheep dominated.

Further east still, in northern Kazakhstan, the dominant animal remains was horse bone.

Finally, in the Afensievo culture, most animal remains are from wild animals, with cattle, pig and sheep making the rest. The likely 'source' for the Afensievo culture might have been from the Hindu Kush, via the Tien Shan.

So clearly, every region had its own particular economy.

Mike Thomas said...

@ David

"Ukrainians show the lowest (most negative) Fst, and thus highest affinity, to Yamnaya (YAM)"

Dave wouldn't you say, however, that Ukrainians are still quite different to ancient Yamnaya? They are far more a "Baltic"/ EE group.

Davidski said...

It's very unlikely that all of the Yamnaya and Corded Ware sub-populations were identical to each other.

The bulk of modern Baltic and Slavic ancestry comes from the border between Yamnaya and Corded Ware as shown on this map IMO.

https://drive.google.com/file/d/0B9o3EYTdM8lQQlB1N3hZbFhfUVU/view?usp=sharing

But this is where the Haak et al. Yamnaya samples came from.

https://drive.google.com/file/d/0B9o3EYTdM8lQRndsdDBCS2toZTQ/view?usp=sharing

Skilur said...

@Mike Thomas
It is interesting that they also presented Fst results for Yamna populations west (YAW)and yamna populations east of the Dnjepr (YAO). Modern Europeans have a much lower affinity to YAO than to YAW. Ukrainians have for for YAM (-0,0266), YAW (-0,02284) and for YAO( 0, 04566). They have the highest affinity to Yamna of modern Europeans and only Poles have for YAO a higher affinity. I expected that Russians would have the highest affinity but historically traditional Ukrainian core areas where much closer to the steppe located than traditional russian areas so this results make sense. Unfortunately they forgot to compare Finno-Ugrians, Bashkirs and Tatars with ancient steppe populations. They would have very likely an even higher affinity to this groups

Matt said...

Krefter: I'm pretty sure Bronze-Iron age Siberians had close to 100% GG.

I think that's pretty possible - based on Mathieson 2015's data points, it looks like the derived allele in SLC45A2 (rs16891982) should have swept to fixation in Europe between around 3000 YBP to 2000 YBP, with the precise time depending on how much weight you put on each data point (also, whether you disaggregate the Neolithic Spaniards and Germans, but then each has a very low sample size). 3000 YBP to 2000 YBP is the Bronze-Iron Age transition in Europe (except Greece?).

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

The Siberian late Bronze Age / Iron Age in the Keyser paper gives dates of 3400 YBP to 2800 YBP for LBA and 3000 YBP to 1900 YBP for Iron Age, so it sounds like the same selective story as in Europe, even though they don't break down the exact frequency of rs16891982 itself (with selection on eye pigment maybe matching closer to North Central Europe).

Alberto said...

"The genetic data support the inference that early Neolithic farmers from Southeast Europe were involved in establishing pastoralism in the steppes by demic diffusion."

I just skimmed through the paper (I can't read German), but does this mean that cultures like Varna and Cernavoda show affinity to those of Western Yamna and Maykop (or Maykop/Yamna lower Don area) cultures?

If so, and related to the Aegean civilizations conference, could this mean that this 3rd population (ANE or whatever) entered the Balkans around 4000 BC (if not earlier) from the South Caucasus/North Anatolia? And after they colonised the steppe from both sides of the Black Sea?

Or were these Balkan people just EEF and unrelated to Caucasus populations?

Tesmos said...

Davidski,

What about the bulk of mdoern Germanic ancestry?

Davidski said...

Well, in theory Germanics should be a mix of Corded Ware and the Yamnaya bands that fed into the Bell Beakers via the Carpathian Basin.

See, that's probably why it's hard to root Germanic on a tree of Indo-European languages; it's kind of both eastern and western.

I'm just speculating though.

Krefter said...

@Matt,
"The Siberian late Bronze Age / Iron Age in the Keyser paper gives dates of 3400 YBP to 2800 YBP for LBA and 3000 YBP to 1900 YBP for Iron Age, so it sounds like the same selective story as in Europe"

The Siberians already had modern-like pigmentation in 1,400-1,800BC, while Unetice in Germany a few hundred years earlier had slightly more dark alleles. Maybe it happened in Siberia earlier than in Europe.

Think about it. Andronovo's ancestors less than a thousand years before were probably as dark as west Asians.

Krefter said...

Does amyone know exactly what this is, and if there's any new and or useful information?

https://www.academia.edu/11530946/%D0%92%D1%82%D0%BE%D1%80%D0%B0%D1%8F_%D0%B2%D0%BE%D0%BB%D0%BD%D0%B0_%D0%BC%D0%B8%D0%B3%D1%80%D0%B0%D1%86%D0%B8%D0%B8_%D0%B5%D0%B2%D1%80%D0%BE%D0%BF%D0%B5%D0%BE%D0%B8%D0%B4%D0%BE%D0%B2_%D0%B2_%D0%AE%D0%B6%D0%BD%D1%83%D1%8E_%D0%A1%D0%B8%D0%B1%D0%B8%D1%80%D1%8C_%D0%B8_%D0%A6%D0%B5%D0%BD%D1%82%D1%80%D0%B0%D0%BB%D1%8C%D0%BD%D1%83%D1%8E_%D0%90%D0%B7%D0%B8%D1%8E_%D0%BF%D1%80%D0%B5%D0%B7%D0%B5%D0%BD%D1%82%D0%B0%D1%86%D0%B8%D1%8F_2015_-_The_Second_Wave_of_Caucasoid_Migration_to_Southern_Siberia_and_Central_Asia_PowerPoint_2015_._Please_download_the_presentation--it_gets_badly_distorted_in_the_preview_mode

Matt said...

Krefter: The Siberians already had modern-like pigmentation in 1,400-1,800BC, while Unetice in Germany a few hundred years earlier had slightly more dark alleles. Maybe it happened in Siberia earlier than in Europe.

Sure, could be. There's no reason why the sweep of the variant couldn't have got to a higher frequency a few hundred years earlier in South Siberia than North-Central Europe (or even earlier). There's be enough room for like a 0.1-0.2 frequency difference at the late Bronze Age / early Iron Age, like between the modern day Italians and Spanish samples in Mathieson. Esp. as there are other genes in play that could've slowed down or speeded up selection on rs16891982 (like if pigmentation is the target selection on rs16891982 could be weaker if other light skin alleles like derived variants at TYRP1, MC1R, IRF4 were around in higher frequency).

It looks hard for me to tell what the frequencies are in the Keyser samples though - that paper gives "Eye color according to rs12913832", "Phenotype according to OCA2 diplotype" (giving all the samples "fair or medium skin") and then an overall ancestry score as European or Asian according to "rs1545397, rs16891982, rs2031526" (where rs1545397 and rs2031526 are East Asian predictive variants, while rs16891982 indicates European). So it's not 100% clear what the frequency of rs16891982 was at what time in these samples. The actual SNPs and frequencies might be in "Pigment phenotype and biogeographical ancestry from ancient skeletal remains: inferences from multiplexed autosomal SNP analysis" which was the paper directly before or after that one (but I can't find a copy online).

Grey said...

@Mike Thomas

"Simply: the Yamnaya pastoralists dervive from certain Eastern European late Neolithic groups. So it seems a bit arbitrary that you're assigning yamnaya as "point zero " when the trail clearly begins elsewhere and earlier."

option 1) farmers move onto the steppe and absorb the local HGs into the farmer culture.

option 2) farmers moved onto the steppe and (unusually) lose to the local HGs thus creating a new type of culture where there is a mounted hunter elite over herder/farmer commoners.

If it was option 1 you'd be right.

If it's option 2 then Yamnaya (or thereabouts) would be the point zero of this new cultural form.

(And it must be getting obvious to almost everyone by now that it's option 2.)

Grey said...

Unless I'm missing something it seems to me...

The various depigmentation genes seem to derive from a (very) roughly triangular region centered in the northern interior of Eurasia with its points at Baltic, Siberia, Himalayas.

There seems to be a bunch of them that developed independently in different places and the entirety of the phenotype is a product of all of them coming together.

LP existed on the steppe but at much lower frequencies than is found in western Europe.

Mike Thomas said...

Grey

I told you (and I think others have also)- show evidence; not wishy-washy speculations that you incessantly pull out of thin air.

The scenarios you imagine do not represent reality, and there was no "war of civilisations" (farmers vs pastaralists, of farmers vs hunter-gatherers).


capra internetensis said...

@Nirjhar

Well it isn't a 30 year old paper. It's a 45 year old book.

That's not what I had in mind, but thanks anyway.

@Grey

I'm afraid it isn't obvious to everyone that mounted hunter-gatherers formed an elite over farmer serfs, no. More like one of any number of possible but unsupported explanations we could pull out of our butts.

Grey said...

@Mike Thomas

We're using the same limited evidence - you're just pretending it's weighted to your preferred default hypothesis when it clearly isn't.

.

There's only a limited number of logical alternatives:

1) farmers moved onto the steppe and mostly erased the foragers
2) farmers moved onto the steppe but were mostly erased by the foragers
3) farmers moved onto the steppe and they merged somehow with the foragers

we now know it wasn't (1) or (2)

(and I dare say if I could be bothered to read back through all the threads there'd be people saying before the Samara data that it was option (1))

anyway option (3) has a number of logical alternatives also:

3a) the two merged with farmers dominant
3b) the two merged with foragers dominant
3c) the two merged with neither dominant

add the ydna into that and there's another three options
x) only the farmers were R1b
y) only the foragers were R1b
z) both were R1b
and we know (x) isn't true because of the old R1b forager

(and I dare say if I read back through the old threads there'd be people saying the steppe foragers were definitely not R1b before the Samara data)

anyway
so we're left with:
option 3a (farmers dominant)
- requires both R1b

option 3b (foragers dominant)
- either forager only R1b
- or both R1b

option 3c (equal merger)
- requires both R1b

So which of those default hypotheses is the most parsimonious and doesn't require new data?

Not yours.

.

As it happens I think the farmers are quite likely to have been R1b also so time will tell but at the moment the weight is on the side of a forager dominant merger.

.

@capra

"I'm afraid it isn't obvious..."

Sure but it's as obvious to me as the opposite is obvious to Mike Thomas so I'd say that was square.

"any number of possible"

I'd say there was a limited number of options (although with very wide gradation within each option).


Mike Thomas said...

Grey

I'll explain one last time, out of good faith

Its not about options, but facts and hard data. Almost all your comments consist of baseless speculations, cliches like "domino effect" & "big fish eats little fish", and infantile arithmetic algorithms.

Now these are facts:

* west Yamnaya had over 60% of faunal bones to be cattle and pigs - which clearly resemble the faunal assemblages of late Neolithic central/ southeastern Europe

* the mesolithic steppe was scantily populated until the Yamnaya (and immediate pre-Yamnaya) period. By contrast, the cucuteni-tripolye culture had a population excess with hundreds of sites and 'mega-cities' which eclipsed even contemporary Mesopotamian sites. In no small way, archaeologists proposed that at the final stage of this, the population spilled over onto the steppe.

* this is now confirmed by ancient mtDNA

What I did not say was there was some kind of epic battle between farmers and foragers, or that the farmers had special sheep who shot magic arrows out their arse, and that the farmers then enslaved the foragers, and promptly chained and gagged them .

Rather there was complex interaction at local and regional levels.

capra internetensis said...

@Grey

All those options are subsets of one mechanism

1) R1b became dominant through social dominance of the original carrying group.

There are other options:

2) It became dominant through drift in a mixed group

3) It became dominant through selection in a mixed group

Anyway, assuming the R1b was from the foragers and did become modal through social dominance, there are quite a few ways this could have happened besides the formation of a mounted forager elite.

Mike Thomas said...

Capra

That's what I have pondered also
But Davidski, and others, point out - why did R1b become dominant in all Western Europe if it were due to localised processes of drift and/or selection (social or natural)?


Hence the continued appeal of "mass-migrations". Although I personally think the former scenario is certainly possible؛ & what we think of as a mass R1b rise was a more protracted process of specific lineage selection/ drift (?)

capra internetensis said...

@Mike Thomas

I don't think the structure of Western European R1b-L51, with a huge number of branches coming off at almost the same phylogenetic level, can be explained by local processes of drift or selection. It really does seem to be a marker of some kind of rapid large-scale expansion. However, it could have resulted from the expansion of a successful minority, and only later increased in frequency to its present ridiculously high levels as a result of selection, rather than being imposed all at once.

But I was only thinking of R1b on the steppe. When farmers and foragers mixed to become the new steppe population drift and/or selection within the mixed population could have resulted in dominance of R1b *locally* - that is, within the population that became the source for westward migrants.

Mike Thomas said...

Agreed
Thanks Capra

Simon_W said...

@ Mike Thomas

If the West Asian ancestry in Corded people is just from regular early European farmers, like Cucuteni-Tripolye and the like, then why do they have so little of the orange farmer component (in the ADMIXTURE analysis in Haak et al., at K = 16 – 19)? Why do they have a whole lot of the teal West Asian component instead?

And why do they have more than 20% of the Gedrosia component, when Karelian EHG had only 8.6% Gedrosia and all early and middle Neolithic farmers had 0%?

This all points to one conclusion: There was a new West Asian population at play which was not derived from early European farmers. And this has probably expanded from the Transcaucasus, it was the same population that triggered the Maykop phenomenon in the North Caucasus.

Davidski said...

ADMIXTURE works in mysterious ways. The unusually high level of ANE among the Corded Ware might be depressing their membership in the EEF component, and inflating it in the teal component.

The teal component is obviously packed with ANE. But it peaks too far south because the only ANE sample in the whole dataset is MA1. If there were more ANE samples the teal component wouldn't exist.

Simon_W said...

Yes, that's something worth considering. But it has to be said that no sample shares more drift with MA-1 than the EHG, and they had only slight membership in the teal component. But, with farmer admixture...

Some CT admixture in the western Yamnaya had to be expected, but the eastern steppe was out of reach for these CT farmers. Not just Samara, but even the majority of the Yamnaya area wasn't old CT farming ground. And where the Corded Ware originated is still a moot point, but from the genetic point of view it was rather not too far in the west. And possibly it was without a farming substrate, too.

Alberto said...

@Simon_W

"If the West Asian ancestry in Corded people is just from regular early European farmers, like Cucuteni-Tripolye..."

Also we don't know exactly if those farmers from CT were the same as the samples we have from Hungary and Germany. Maybe they already had ANE from Ukrainian HGs, or maybe some Caucasus-like population had already entered the Balkans before they entered the steppe.

We also don't know if places like Poland or Belarus had ANE before CWC.

Still too many gaps in the data. But I agree with Mike that it won't be a simple East to West mass migration (even if as a whole there are many eastern genes coming into Europe). We still need to find out all the details of how this happened. But I'd still expect a surprise or two when more genomes are sampled.

Simon_W said...

Well Alberto, we may assume that CT farmers mixed with local HGs like central European and other early farmers did: Slowly and in modest amounts. Until the middle Neolithic it had summed up to, I don't know, 20% HG on top of the HG that was already in LBK_EN. Also, if there was Caucasus admixture in the Balkans it would show up as Caucasus component in Dodecad K12b, not as Gedrosia. Most European farmers had quite a strong Caucasus component in K12b, but always 0% Gedrosia. HGs in Belarus most probably had quite a lot of ANE, and Belarus went from Comb-pitted HG straight to Corded Ware without an intermediate farmer stage. Polish HGs possibly had some ANE. But the Swedish TRB had none even as late as 3000 BC, and these were surrounded by SHG with ANE. Gok2 had quite a lot of HG admixture, but without ANE, and the roots of this Swedish TRB were probably in Poland. I think Mike Thomas puts more weight on the archeological evidence than on the genetic one, as many archeologists believe that the Corded Ware originated in Poland, from local people. But, there are also other opinions in the archeological community, and unfortunately the C14 dates of this period have also a very wide range of uncertainty. So I don't think it's that wise to put so much weight on controversial archeological evidence, when the genetic evidence makes a massive immigration quite obvious.

Simon_W said...

Davidski, I've thought about it, it doesn't work out. I could cite the f-stats in Haak et al. pointing to an Armenian-like, not an ENF-like population that admixed with EHG. But even according to your very own K8 PCA of the data, it's obvious that you can't get Yamnaya from a mixture of EHG and ENF. If you extend a line from the EHG via Yamnaya long enough, you end up somewhere in the Caucasus and (in one case) in Tadjiks. And there is really no reason why Cucuteni-Tripolye should have been an Armenian-like population. CT was partly descended from LBK, partly it had somewhat younger influences from the Balkans.

To me the most reasonable explanation for the teal component in Yamnaya and Corded people is a Copper Age expansion of ANE admixed Transcaucasians on the steppe. Presumably it was them who brought along R1b-P297 and R1b-L23. This would explain how these haplogroups can make up ~30% of some Armenian groups although the latter have extremely low northern hunter-gatherer ancestry. Incidentally my view is almost completely in line with Maciamo's:
http://cache.eupedia.com/images/content/R1b-migration-map.jpg

Davidski said...

Transcaucasians and Armenians have ancestry from the same Neolithic Near Eastern population as early European farmers, which is why they also show the EEF component in Haak's ADMIXTURE run. So saying that the Near Eastern admixture in the Yamnaya is Armenian-like and/or from the Caucasus doesn't explain anything here. There must be another reason why the Yamnaya lack the EEF component.

And Maciamo's model is obsolete. That's because we now know Eastern European hunter-gatherers carried R1, including R1b. I'm betting you'll fall off your chair when you see an ancient genome from pre-Bronze Age Armenia.

Simon_W said...

You've made a point there. Even Iranians and Tajiks have it; strange...

Sure, EHG had R1a1 and R1b1, but this doesn't prove that they had R1b1a. Otherwise you could also say that Iberian farmers had R1b1a. That there was R1b1 in Samara EHG may be a sign that R1b1a developped there. Or it may be a coincidence, a consequence of a generally higher incidence of R derived haplogroups in the east.

Simon_W said...

Alberto, I think the odds against an early presence of modern Caucasian-like farmers in Ukraine are considerable: Hungary isn't that far from the Ukraine, and it's on the same geographical latitude. The Ukraine can't be said to be somehow closer to Anatolia. Yet, if you study the HungaryGamba samples, you'll note that the affinity to modern Caucasus populations did not at all increase during the Neolithic and Copper Age. BR1 at 2000 BC showed no such affinity either. Only BR2 at 1200 BC does. The same can be said about the yDNA. In all the Hungarian samples, the earliest recorded presence of the West Asian haplogroup J2 is around 1200 BC.

I would also mention Dienekes' Rolloff analyses of the Greeks and Bulgarians, according to which the ANE-rich West Asians mixed into the Greeks around 1840 BC and into Bulgarians around 1670 BC.

Together this points to a Bronze Age arrival of ANE-rich West Asians in Southeastern Europe.

Alberto said...

@Simon

I actually agree with you in the general picture:

"This all points to one conclusion: There was a new West Asian population at play which was not derived from early European farmers. And this has probably expanded from the Transcaucasus, it was the same population that triggered the Maykop phenomenon in the North Caucasus. "

Also in the likelihood of this population being R1b (though I'm open to any possibility regarding this).

But we're still missing a lot of data. When you say:

"So I don't think it's that wise to put so much weight on controversial archaeological evidence, when the genetic evidence makes a massive immigration quite obvious."

I'd also say that the genetic evidence is still very spotty and paints a simplistic picture (for example, a HUGE impact of Yamnaya in half Eurasia). I think that more data points will show a more complex process, different routes, different populations involved...

By the way, where would you place the origin of such mass migration? Do you think that the South Caucasus could be the origin of a populations whose genes have extended in high percentages from India to Ireland? Or do you think they arrived to the South Caucasus from somewhere else in the middle/Late Neolithic?

Mike Thomas said...

Good convo points chaps
I agree
Colin, my point was that this mtDNA data shows directly movement from CE to the steppe
I know that haaks PCA paints yet a different picture - a very Central Asian one

But ultimately , I have to agree with Alberto . The supposed yamnaya impact is overestimated , and what it's really showing is that yamnaya itself was part a larger process of intermixture

Mike Thomas said...

@ Colin

" So I don't think it's that wise to put so much weight on controversial archeological evidence, when the genetic evidence makes a massive immigration quite obvious."

Perhaps. But I think we still need more aDNA to make the genetic evidence unequivocal.
Moreover I never suggested that CT farmers colonised all the way to samara
Nor do I think we can definitely argue that they should be the same as german farmers

Finally; what id call Epi-Neolithic groups survived in the Baltic and down through mainland poland (ie residual foragers). Along with SHG people from Denmark etc; they'd have made a small but sure constribution to CWC, and it's ANE rich profile

Mike Thomas said...

@ Colin

" Presumably it was them [i.e. cis-Caucasians],who brought along R1b-P297 and R1b-L23."

I've also pondered this possibility ; whilst aware that R1b1 was already present in Samara HG

Davidski said...

A simple two-way admixture event between ancient eastern Anatolians or Transcaucasians and Eastern European foragers certainly doesn't explain the genome-wide structure of the Samara Yamnaya.

The only other samples that show the teal component without any EEF-related admixture are the Kalash. So either there was a migration from the Hindu Kush to the steppe, or from some other place that populated both the Hindu Kush and the steppe.

I think the latter is much more likely. But this wouldn't have been a Copper Age expansion, nor the Proto-Indo-European expansion. Rather, it was probably the same population movement that influenced present-day Pakistan during the early Neolithic, and possibly gave rise to sites like Mehrgarh. See here...

http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0095714

In other words, it looks like the Near Eastern admixture in the Yamnaya comes from a place that lacked EEF-related ancestry during the Neolithic, at least for a time. I'd say this was probably early Neolithic Iran. There aren't many other options.

Mike Thomas said...

Davidski

"But this wouldn't have been a Copper Age expansion, nor the Proto-Indo-European expansion. "

Well I think more data will pin down the chronology ; and as for if they were "indo-europeans" - that I think is a rather moot point since we have no way of telling what languages were spoken by distant societies. As I've argued ad nauseum before; linguistic palaeoantoligists have had too much confidence with what they claim to prove; and have largely operated within the dictum of what they set out to prove in the first place

Mike Thomas said...

"it looks like the Near Eastern admixture in the Yamnaya comes from a place that lacked EEF-related ancestry during the Neolithic, at least for a While "

To throw a curve ball: could it be somehow possible that the EEF were largely Mesolithic southern- Central Europeans ? Hence the apparent lack in some parts of central- west Asia

either that or Iran had a largely "indigenous " development of agriculture

Krefter said...

@Matt,
"Good convo points chaps
I agree
Colin, my point was that this mtDNA data shows directly movement from CE to the steppe
I know that haaks PCA paints yet a different picture - a very Central Asian one "

Are you saying the FST statistics from Wilde. 2015 are evidence of a migration from Central Europe into the Steppe?

If you look at the samples one by one you see Neolithic Central Europeans were totally differnt from Steppe people mtDNA wise. Formal stats with mtDNA, especially low coverage samples, can reveal little in my opinion.

Modern West Asians, not EEFs, have fairly similar mtDNA to Steppe(in terms of ENF lineages, not overall). Also, most modern west Asian's maternal lineages separated from EEF's maternal lineages a very very very long time ago, probably in west Asia.

My very nubby opinion is that the ENF ancestry of Samara Yamna mostly came directly from west Asia, not southeast Europe.

Mike Thomas said...

Krefter

You're right, for eastern Yamnaya samples at least

David

Can you possibly do some K15s for the 4 ancient Thracian samples please ?

Davidski said...

They don't have enough markers for the K15. Very poor overlap. So the results are meaningless.

Mike Thomas said...

Righto, no problem.

Nirjhar007 said...

Mike and others
As you said its impossible to know what language a distant society spoke and it is only actually be provable by Ancient Texts and deciphered scripts anyway i would like to have your and others opinions on this phenomenon-

http://en.wikipedia.org/wiki/Seima-Turbino_Phenomenon
What is you guys guesstimate on the Language that they ''may'' have spoken, Do you think they were related to Pre-Scythians who migrated West Wards towards Volga and NE Europe from their ''origin'' Around Mongolia or they were some Uralic Type Tribe or Indo-Iranian?

Chad Rohlfsen said...

Mike,

There won't be ANE in Danish hunters. They're part of TRB, and show no sign of excess eastern affinity.

Mike Thomas said...

Chad
I'm not saying that TRB will be "ANE"; but there were unincorporated Mesolithic residual groups which persisted until the start of Bronze age , in the Baltic and North sea rim , scandinavia , central Poland even

If we ever find such skeletal samples , they'd be worth sampling

Alberto said...

Yes, good points made above.

David, yes, I think that the "West Asian" expansion was probably not a single late Neolithic movement. It doesn't make sense that large populations like the IVC was replaced by any other in the late Bronze Age.

There are connection from West Iran (Zagros), South Caspian, North East Iran border with Turkmenistan (Jeitun), North Afghanistan... From there it could go north through east Tajikistan and Kyrgyzstan up to Tian Shan (in South East Kazakhstan).

From Zagros it could also extend south east to Mehrgarh, and from Afghanistan south to Pakistan.

Along the Neolithic, probably all of Iran, Central Asia and S-C Asia became very "West Asian". Not sure about the Caucasus. They might have moved there early too (from South Caspian) or be a later movement.

Mike posted a link to this book a while ago:

http://unesdoc.unesco.org/images/0009/000944/094466e.pdf

@Mike

"To throw a curve ball: could it be somehow possible that the EEF were largely Mesolithic southern- Central Europeans ? Hence the apparent lack in some parts of central- west Asia"

It's a possibility, but I think that there is a clear connection with at least the Eastern and Northern parts of the Fertile Crescent. I recently read some paper that EEF arrived to Greece by sea from there (and not by land from Western Anatolia). The question is: Were the people from Mesopotamia the same as from further east? Or were they more "West Asian"?

About the Thracian samples, I saw some analysis of them and they are certainly very noisy:

https://genetiker.wordpress.com/2014/06/11/analyses-of-iron-and-bronze-age-bulgarian-genomes/

Simon_W said...

Another argument against the „solution“ that some possible EHG admixture in CT transformed these EEF into an Armenian-like population: Effectively this would be the same as deriving Yamnaya from a mix of EEF and EHG. If CT = EEF + EHG and this admixed with EHG to produce Yamnaya, the equation would be Yamnaya = EHG + EEF + EHG, in other words a simple two way mix of EEF and EHG. And we know from PCA and f-stats that this doesn't work out.

Likewise, Armenians don't seem to be a simple mix of basal Eurasians + EHG, as again indicated by PCA and f-stats.

Obviously, the reason for all this being that there is more than one type of ANE. Or rather, the MA-1 related ancestry drifted and evolved differently in different populations. It's natural, because MA-1 is a rather old sample, compared to EEF, EHG and Neolithic West Asians etc.

I think it's possible that the teal component is to some extent a composite of basal Eurasians and northern HGs. But it seems to include a lot of south central Asian ANE as well, which cannot be explained as a mix of basal + northern HG.

@ Davidski

I would agree that there was something coming from Iran into Yamnaya, the question is: On which way, east or west of the Caspian? The lack of the EEF component in Samara might suggest that it came on an eastern route. But I'm not sure if that's archeologically feasible. I would also consider that some smaller part of the teal component is EEF + EHG.

Simon_W said...

@ Alberto

I'd say the south central Asian type of ANE arrived in West Asia quite late, from the east. Dienekes' rolloff analysis of the Lezgins as a mix of Sardinians + Burusho yielded an admixture date of 3750 BC. That's perfectly in line with the archeological findings here:
http://dienekes.blogspot.com/2013/05/origins-of-maykop-phenomenon.html

And the fact that the EEF had a lot of the Dodecad K12b Caucasus component while lacking the K7b West Asian and the K12b Gedrosia component, quite in contrast to modern Caucasus populations, may suggest that the Caucasus is a mix of an old component related to EEF and another, ANE-rich component that came in from the east.

Of course I've also had doubts that the Indo-Europeans may have sprung from a small mountain area. It would seem more believable that large masses of herders in the wide open steppe were at the origin of IE. But on the other hand, if a group of PIE from the Transcaucasus had managed to assimilate the EHG on the steppe, this would be half the battle. We may also think what would have become of IE if they hadn't assimilated the steppe EHG. IE would be a rather small language family now, with Anatolian and Tocharian extinct, there would only Greek, Albanian and Armenian be left – if these were from West Asia, as I'm suspecting.

Simon_W said...

@ Mike Thomas

You confused me with Colin. :D

Iran today doesn't lack the EEF component.

And while we don't know what pre-Neolithic southeastern Europeans were like, possibly similar to Neolithic Anatolians, those in central Europe were certainly like Loschbour and KO1, and not EEF-like.

Davidski said...

Roloff doesn't give migration dates, it gives mixture dates, and they always seem to be a bit late, probably because of continuing mixing and minor admixture events. So a date of 3750 BC might actually mean a migration around 4500 BC or much earlier. The Yamnaya seem to have been a fairly stable blend by 3300 BC, so I'd say 3750 BC is too late.

Alberto said...

@Simon

Yes, something around 4000 BC looks reasonable to me as to when the "West Asians" moved east into the Southern Caucasus, though it could have been earlier. See:

http://dienekes.blogspot.com/2012/09/rolloff-analysis-of-french-as-mixture.html

"Of course I've also had doubts that the Indo-Europeans may have sprung from a small mountain area. It would seem more believable that large masses of herders in the wide open steppe were at the origin of IE."

But when did the open steppe had "large masses of herders"? There was probably a population boom with Yamnaya, but that implies the new arrival of people from somewhere else, and therefore Yamnaya being just a part of a larger scale process (as Mike said).

For languages I don't make any strong case, because they are subject to certain degree of randomness (as Y-DNA). Why the Caucasus people are non-IE speakers? Probably for the same reason as Basques. Or for the same reason as Hungarians speak Hungarian. Or Burusho people speak Burushaki.

But I see difficult to relate IE to EHG. Usually HGs from anywhere have left as their biggest legacy their genes, and not much more. They are by definition small scattered groups, that lack cohesion and many of the features that we define as a culture. When a superior population comes (superior technically, socially, culturally, demographically an economically) it's difficult to imagine strange scenarios where HGs (Western, Eastern or from anywhere) impose their language and culture onto the incoming population.

Genetically, EHG are strongly related to Uralic speakers, and not so strongly to Altaic ones. The genetic connection to IE speakers is quite weak. But as I said, strange things can happen, so who knows...

Nirjhar007 said...

Well guys i think by the end of this year we may get the solution of the delicious PIE riddle...

Mike Thomas said...

@ Simon

I do apologise

"And while we don't know what pre-Neolithic southeastern Europeans were like, possibly similar to Neolithic Anatolians, those in central Europe were certainly like Lobschbour and KO1, and not EEF-like."

True. But that's for central Europe-Carpathian Basin- northern Balkans.

Thrace and Greece were probably Anatolian-like, who show contacts with Antalya region and even further to Natufian culture.

Some parts of the Ukrainian LGM refuge extended to eastern Romania

So, that is why the Balkans Mesolithic was likely a mixed place.

Grey said...

@Davidski

"So either there was ... (snip) ... early Neolithic Iran."

My vote would currently go for the option you outlined there.

(I prefer apple valley (Almaty / Kazakh) as the origin but not for logical reasons. I just like the idea of an apple/pear forest and goats drunk on fermented fruit self-domesticating with Borat Bacchus the fruit forager.)

.

@Alberto


"Usually HGs from anywhere have left as their biggest legacy their genes, and not much more."

For things to turn out differently than they usually did the local HGs would have needed a significant and unusual home advantage.

.

It would be an incredible fluke but given that LP in ancient dna is currently very rare and given the very high rate of LP in western Europe and its overlap with R1b distribution does R1b "need" to have been a mass migration?

I'm not saying it wasn't but does it need to have been?

.

random thought of the day

Before the Persian Royal Road Was the fastest overland route between the Indus and Greece by going north of the Caspian rather than crossing all those mountains?

http://www.lhs.rcs.k12.tn.us/teachers/crumbyj/The%20Persian%20Empire_files/slide0142_image023.gif

Alberto said...

@Grey

"I prefer apple valley (Almaty / Kazakh) as the origin but not for logical reasons"

I think that there might have been a connection between the South Caspian to Tian Shan, more or less following what are the modern capitals of those countries (Tehran, Ashgabat, Kabul, Dushanbe, Tashkent, Bishkek, Almaty). Kind of a predecessor of the Silk Road.

The forests, mountains, valleys, rivers, fauna and flora that seem to be present in PIE vocabulary could come from that area you mention.

Since you have that thing for horses, here's a paper that shows some evidence of early horse domestication in south east Uzbekistan, pre-dating other known possible places by maybe 2 millennia:

http://www.academia.edu/2765711/A_PROBLEM_OF_THE_EARLIEST_HORSE_DOMESTICATION._DATA_FROM_THE_NEOLITHIC_CAMP_AYAKAGYTMA_THE_SITE_UZBEKISTAN_CENTRAL_ASIA


"It would be an incredible fluke but given that LP in ancient dna is currently very rare and given the very high rate of LP in western Europe and its overlap with R1b distribution does R1b "need" to have been a mass migration?

I'm not saying it wasn't but does it need to have been?"

I don't think that LP specifically is the reason, because it doesn't seem to originate in R1b populations as has been speculated. For example, in the Basque Country, some site from 3000 BC showed 31% LP, while in Yamnaya samples it's absent:

http://www.nature.com/ejhg/journal/v20/n7/abs/ejhg2011254a.html

Also LP is not such a big advantage in Mediterranean environments, and Spain has about 70% R1b.

But you are right that some other advantage might have caused this haplogroup to become so prevalent in WE. We don't know many things about those times. But for example we know from historical times that the Black Death killed almost half of the European population in a matter of a few years. So any kind of better resistance to any kind of health issue from the Bronze Age or later could have had a big influence over time.

Grey said...

@Alberto

If the Atlantic coast provided an extreme selective advantage for LP and before that it was very rare then LP further inland in Western Europe should at least partially correlate with Atlantic coast ydna and/or mtdna.

For example the mtdna H that apparently spread dramatically out of Iberia with BB.

It would be interesting if there was any data from further inland which had both ydna, mtdna and LP status.

http://dienekes.blogspot.co.uk/2013/04/mtdna-haplogroup-h-and-origin-of.html

Simon_W said...

@ Alberto

You said: „4000 BC looks reasonable to me as to when the "West Asians" moved east into the Southern Caucasus“. In fact I was thinking of the opposite direction, i.e. that they moved west.

But I agree that the French date looks strangely early. Maybe really something that happened in CT, if the result is correct at all.

As for the large masses of herders, I was thinking of a lecture by Mallory where he stressed the large number of burials found in Khvalynsk, accompanied by plenty of cattle and sheep, pointing to a booming, well-fed population, a thousand years before the CT collapse. This obviously against the idea that only farming cultures can be populous.

Imho the reason why non-IE languages are common in the Caucasus is that it's a perfect refugium. That applies also to haplogroup G2a. Basque survived because it's on the western fringe of Europe, far from the PIE homeland, and also in a mountainous area. The Hungarian case looks indeed quite random, undoubtedly the result of elite dominance, though they are not completely without a genetic legacy of the early Magyars.

Well, David seems to think that the West Asians were peaceful weak matriarchalic farmers, like in Marija Gimbutas' scenario of EEFs, and hence an easy prey for warlike dominant patriarchalic EHG. But it's not self-evident that everything from West Asia must be like this. (Strictly speaking EEFs were not completely peaceful either.) We don't even have to invoke IE West Asians like the Hittites. The Assyrians were warlike killers too, and nowadays the IS and similar fanatics cannot be called submissive either, and in general women don't have a very high social standing in these cultures. In fact they had a better standing in some old IE cultures, like the Celtic one, and afaik it's not typical for Slavs to suppress their women, except for the machismo found in Balkan Slavic cultures.

Simon_W said...

A few points regarding Wilde's paper:

As I had suspected, the Yamnaya samples are almost all from the western part of the Yamnaya world, where there was undeniable CT farmer influence. From the Volga near Samara she has only samples from the later (Bronze Age) Poltavka culture. Three Yamnaya samples are from Manych and Kuban in the Caucasus foreland, the rest is from the Dnieper and further west.

Also, the paper's „main dish“ is the mtDNA analysis. yDNA is not considered, and from autosomal DNA only a number of genes are analyzed, not hundreds of thousands of SNPs. If the CT legacy shows itself most of all in the mtDNA, it's natural that the western Yamnaya groups show low Fst to EEF populations.

The haplogroups called „Late Neolithic/EBA“ in Brandt, Haak et al. 2013 were most of all T1, I and U2. These were nearly absent in the Early and Middle Neolithic groups. Since U2 is clearly from EHG, it would be interesting to elucidate the origin of the T1 and I in Yamnaya. And now it's interesting that in the Yamnaya samples from along the Dnieper, there is no U, but only H, T1 and I. Also, the southern two of these three sites, plus the Kuban samples, in other words the samples around the Sea of Azov up to the Dnieper knee, have a high Fst to the other Yamnaya samples. The Kuban samples were T and J2b1d, though, and geographically very close to Maikop.

Another interesting finding is indeed the high Fst of the Catacomb culture to Yamnaya and other cultures, and its affinity to the Pitted Ware. This clearly suggests a movement of northern people southwards, and further reduces the credibility of a West Asian origin of IE. I think this may have resulted in the replacement of yHg R1b through R1a.

FrankN said...

Don't know if anybody will still be reading this - nevertheless:
Have a look at Table 13.11 right at the bottom of the paper, which compares Fst Distances between archeological cultures, and with current populations, and pay attention to BEC (Bernburg Culture), a.k.a "late Funnelbeaker" or "Nordic Megalithic" (actually its southerly variant). Bernburg Culture is a/o interesting, as it signifies the later phase of the swift (1 km/y) southward expansion of northern Funnelbeakers, thereby first assimilating residual HG populations in Northern Germany, and then intruding into Neolithic territory. Between 3,300 and 3,100 BC, an archeologically well documented "militarized border", characterised by multiple fortifications and many "warrior burials" stretched along a line from Erfurt via Potsdam to Stettin (forgive me Dave, I always get lost with the Polish ortography here, no revisionism intended!), which seperated the Bernburg Culture from the Salmünde Culture to the South. The latter was an offspring of the equally expansionist Danubian Baden-Boleraz-Cucuteni culture, credited with the introduction of copper metalurgy into Central Europe, and Salzmünde proper having the earliest Central European horse burial found so far. Ultimately, around 3,100 BC, the Bernburg Culture destroyed the Salzmünde settlement.
[In fact, everything you need to build your IE stereotype can be found here - Bernburg/ late Funnelbeakers has the cart (first evidence more-less simultaneously 3,400 BC from near Kiel and near Cracow) and the burial mounds, Salzmünde the horse plus copper metalurgy.]

First, for an impression, the Fst-statistics (probability of population identity in brackets) related to the potential sources of the Bernburg culture's (female) genetic make-up:
- Mesolithic Central Europeans: 0.056 (3.1%)
- Northern HGs: 0.077 (0.8%)
- Eastern HGs: 0.037 (8.1%) [here come the first ANE!]
- LBK: 0.027 (7.4%)
- Rössem: 0 (60.6%)
- Late neolithic farmers, Blätterhöhle/ Westfalia: 0 (64.0%)
- Funnelbeakers (Scandinavia/ Mecklenburg): 0.16 (26.2%)
- Dnepr-Don (pre-Yamnaya): 0.038 (17,3%)
- Salzmünde Culture (the "enemy"): 0.024 (10.8%)
The strong similarity to Westfalia, and low resemblance to NHG is remarkable. What at first sight appears to be a Scandinavian/ West Baltic phenomenon is actually proof of archeologically based assumptions that the Funnelbeaker phenomenon emerged from strong interaction with the Michelsberg culture, which from 4,500 BC on rapidly expanded out of the Paris Basin (or even Brittany) towards the Elbe and into Southern Germany, thereby also neolithicising the Netherlands plus, on a different route, the British Isles. In addition, the Bernburg Culture also picked up EHG (somewhere from Pomerania?) and a good chunk from the Dnepr-Don culture.

Now a look at affinities to later cultures:
- Pitted Ware: 0.065 (4.8%)
- Corded Ware: 0 (66.0%)
- Bell Beaker: 0 (51%)
- Unetice: 0 (46.6%)
- Pontic eneolithic: 0 (57.5%)
- Yamnaya West: 0 (87.3%)
- Yamnaya East: 0.093 (2.3%)
- Poltava Culture: 0.017 (28.4%)
- Early Catacomb: 0.031 (14.3%)
- Andronovo: 0 (56.5%)
- Bronze-age Kazakhstan: 0.027 (20,1%)
Well, yDNA may of course tell another story, but this looks very much like an expansion from Central Europe into the Steppe. Yamnaya East statistically indistinguishable from Bernburg! That doesn't neccessary mean there wasn't any movement into the opposite direction - most likely there was - but this looks very much like populations mixing rather than like a violent incursion (which is, btw, also what archeology in the Elbe-Saale region indicates).

Finally, comparison of BEC with today's (female) population:
- Germany: 0.075 (18.6%)
- Russia: 0.0023 (33.8%)
- Poland: 0.0021 (35.7%)
- Lithuania: 0.0016 (37.7%)
- Bulgaria: 0 (45.0%)
- Ukraine: 0 (77.4%)

Any questions? If somebody could get his/her hands on the Bernburg data, admixture simulations from it might be interesting.