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Friday, May 12, 2017

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...

All of the post-Middle Neolithic samples from the recent Mittnik et al. and Saag et al. preprints on the ancient population history of the Baltic region belonged to Y-chromosome haplogroup R1a. And most of them belonged to the R1a-M417 (R1a1a) subclade that makes up almost 100% of the R1a lineages in the world today. This is what the results look like in a table (the sample IDs are of my own design):

Earlier samples from the same region belonged to Y-haplogroups I2a and R1a, but this was a subclade of R1a defined by the YP1272 mutation that is extremely rare today even in Northeastern Europe.

And now shifting our focus west of Scandinavia: all but two of the post-Middle Neolithic samples from around the North Sea from the recent Olalde et al. preprint on the Bell Beaker phenomenon and ancient population history of Northwest Europe belonged to Y-chromosome R1b, and more specifically to the R1b-M269 (R1b1a1a2) subclade, which makes up almost 100% of the R1b lineages in the world today. Here's a table:

Earlier samples from the same region belonged to Y-haplogroups I2a, I, G2a and CF, and most of the instances of I and the CF would probably be classified as I2a if not for missing data.

Interestingly, despite the R1a vs R1b dichotomy between these post-Middle Neolithic obvious newcomers to the Baltic and North Sea regions, respectively, they were very similar in terms of overall genetic structure, obviously closely related, starkly different from Middle Neolithic Northern Europeans, and in all likelihood mainly derived from the same homeland that was not located in Northern Europe.

So can we locate this homeland with any degree of certainty, you might wonder? In fact, you might ask, isn't this a futile search for the time being, as we await ancient DNA from many prehistoric Eurasian populations?

Not at all, because when attempting to answer this question we're bounded by two key constraints: the exceptionally high frequencies of R1a and R1b in the post-Middle Neolithic Baltic and North Sea samples, and their close genetic affinity to earlier and contemporaneous populations from the Pontic-Caspian Steppe, part of which is due to significant Caucasus Hunter-Gatherer (CHG) admixture that was lacking in Middle Neolithic Northern Europeans.

Indeed, to date, the Pontic-Caspian Steppe is the only region where both R1a and R1b have been found in ancient remains from the same sites dating to the Mesolithic, Neolithic and Eneolithic. Here's a table based on results from Mathieson et al. 2015 and 2017. The R and R1 might really be R1a or R1b if not for missing data.

The Pontic-Caspian Steppe also abuts the Caucasus foothills, and we know that CHG admixture was a major feature of its inhabitants from at least the Eneolithic. So odds are, and make no mistake, these are indeed excellent odds, that the homeland we're looking for was on the Pontic-Caspian Steppe.

But of course I2a has also been recorded in prehistoric samples from the Pontic-Caspian steppe. So, you might ask, why did the populations migrating out of the steppe belong to R1a and R1b, and why did some of them seemingly carry only R1a while others only R1b? This can be explained by local founder effects on the steppe due to patrilocality. Moreover, it's possible that some groups moving out of the steppe did carry high frequencies of I2a, but they're yet to enter the ancient DNA record. [Edit: Maybe they already have? See here]

Now, the aforementioned post-Middle Neolithic newcomers to the Baltic and North Sea regions are most certainly in large part the direct ancestors of modern-day Northern Europeans, speaking languages belonging to the three daughter branches of Late Proto-Indo-European (PIE): Balto-Slavic, Celtic and Germanic. It's highly unlikely that languages ancestral to these present-day languages were spoken by Middle Neolithic farmers, nor introduced into Northern Europe after it was colonized by the migrants from the Pontic-Caspian Steppe.

What this strongly suggests is that the Pontic-Caspian Steppe was also the late PIE homeland.

But, you might argue, the Pontic-Casian Steppe may have just been the expansion point for some of the late PIE language branches. No, that won't work. For one, modern-day populations speaking languages belonging to all other late PIE branches, such as Armenian, Greek, Indo-Iranian and Italic, show signals of the same population expansion from the Pontic-Caspian Steppe that gave rise to modern-day Northern Europeans, in the form of Yamnaya-related genome-wide genetic admixture and appreciable frequencies of Y-chromosome haplogroups R1a-M417 and/or R1b-M269.

Some of these signals are certainly due to fairly recent admixture from Northern Europeans, like in much of Greece as a result of the Slavic expansions during the Early Middle Ages, but most cannot be explained in this way.

Secondly, Balto-Slavic, Celtic and Germanic are not more closely related to each other than to some of the other late PIE branches. For instance, Balto-Slavic is considered far more closely related to Indo-Iranian than to Celtic, which is generally seen as a sister branch to Italic. Therefore, if Balto-Slavic and Celtic derive from a homeland on the Pontic-Caspian Steppe, then logically this is also where we should look for the origins of Indo-Iranian and Italic.

So as far as the late PIE homeland is concerned, thanks to ancient DNA, the debate is now practically over. But the PIE homeland debate is still wide open, or so we're told.

Apparently, Mathieson et al. 2017 aren't comfortable with putting the PIE homeland on the Pontic-Caspian Steppe because they can't find any evidence in their ancient DNA dataset of a significant migration through the Balkans that would potentially bring Anatolian languages from the Pontic-Caspian Steppe to Anatolia. From the paper:

One version of the Steppe Hypothesis of Indo-European language origins suggests that Proto-Indo European languages developed in the steppe north of the Black and Caspian seas, and that the earliest known diverging branch – Anatolian – was spread into Asia Minor by movements of steppe peoples through the Balkan peninsula during the Copper Age around 4000 BCE, as part of the same incursions from the steppe that coincided with the decline of the tell settlements. [51] If this were correct, then one way to detect evidence of it would be the appearance of large amounts of characteristic steppe ancestry first in the Balkan Peninsula, and then in Anatolia. However, our genetic data do not support this scenario. While we find steppe ancestry in Balkan Copper Age and Bronze Age individuals, this ancestry is sporadic across individuals in the Copper Age, and at low levels in the Bronze Age. Moreover, while Bronze Age Anatolian individuals have CHG/Iran Neolithic related ancestry, they have neither the EHG ancestry characteristic of all steppe populations sampled to date [20] , nor the WHG ancestry that is ubiquitous in southeastern Europe in the Neolithic (Figure 1A, Supplementary Data Table 2, Supplementary Information section 1). This pattern is consistent with that seen in northwestern Anatolia [11] and later in Copper Age Anatolia [23], suggesting continuing migration into Anatolia from the East rather than from Europe.

And this...

On the other hand, our data could still be consistent with the Steppe-Balkans-Anatolia route hypothesis model, albeit with constraints. It remains possible that populations dating to around 1600 BCE in the regions where the Indo-European Luwian, Hittite and Palaic languages were spoken did have European hunter-gatherer ancestry. However, our results would require that such ancestry was not ubiquitous in Bronze Age Anatolia, and was perhaps tightly linked to Indo-European speaking groups. We predict that additional insight about the genetic origins of the potential speakers of early Indo-European languages will be obtained when ancient DNA data become available from additional sites in this key period in Anatolia and the Caucasus.

But I'd say the authors are taking that one particular version of the Steppe Hypothesis way too seriously. They might even be implying things that the creator(s) of the said hypothesis never posited.

Why do they seemingly expect a massive surge of steppe admixture into the Balkans during the Copper Age? If the steppe people are just shooting through the Balkans on their way to Anatolia, why would they leave a lot of admixture along the way? And if the locals are abandoning their tell settlements and running for the hills as far away from the oncoming steppe invaders as they can, how exactly would they acquire steppe admixture? Osmosis or what?

The Balkans is not Northern Europe, and the hypothesized migration of the proto-Anatolians from the Pontic-Caspian Steppe to Anatolia through the Balkans was never, as far as I know, meant to parallel the massive Corded Ware expansion across Northern Europe. In other words, why should all of the early Indo-European expansions have been of the same character, especially considering that they moved into such starkly different areas of Eurasia?

Indeed, as Mathieson et al. 2017 point out in the quote above, the evidence for the fleeting presence of steppe peoples in the Copper Age Balkans is in their dataset. For instance, in their Varna 1 sample set from Bulgaria, three out of the five individuals show significant steppe admixture. One of these individuals is almost 50% Yamnaya-like. Surely, there's really no need to expect anything more than that when looking for signals of a proto-Anatolian migration from the Pontic-Caspian Steppe to Anatolia.

In fact, even though I do appreciate the incredible work these guys are doing and the data they're making available to myself and everyone else, I suspect that there's a little bit of, shall we say, schadenfreude going on here.

They sequenced all of three Early Bronze Age Anatolians of obscure origin (are they actually suspected Anatolian speakers, like Luwians?), and apparently it's a big deal that they can't find any steppe admixture in Early Bronze Age Anatolia. Come on.

And then we're offered just three Yamnaya samples from the Pontic Steppe in Ukraine. One happens to be a massive outlier towards the Caucasus. Wow, what are the chances of that? And guess what, all three of these Yamnayans are females, so of course we're left wondering about the Y-haplogroups of the Yamnaya males on the Pontic Steppe. What happened to the males? Next paper, that's what.

Update 19//05/2017: Please note that the authors are not holding back any Yamnaya males from Ukraine for a future paper, as per my claim in the last paragraph above. They used what they had for the time being.

Update 21/05/2017: Actually, I suspect that we already have a population from the Bronze Age steppe in the ancient DNA record with a high frequency of Y-haplogroup I2a. See here.

See also...

Eastern Europe as a bifurcation hotspot for Y-hg R1

Globular Amphora people starkly different from Yamnaya people


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Alberto said...


does looking for steppe masks CHG?

I don' think so in this case. They're checking using Yamnaya (which is EHG + CHG) against Anatolia Neolithic, so if those samples had any CHG admixture it would show up as a shift towards Yamnaya. But the signal is very weak, so either there is nothing or very little.

We'll wait for North Africa, but who knows how long that will take. Probably we'll see Shulaveri and VNSP or similar before.

Matt said...

@Azra.... hmmm... nice to see that tried; still I think the proportions of qpAdm lack much specific to be compelling (don't distinguish well between Iberia+Central Europe) and not sure about mixes using one population outliers. Time will tell with what Davidski and others actually model with qpAdm.

@OM: "But that is the opossite of what this papers are stating. so according to "them", Bell beaker germany as 25% Iberian chalc is something they do not see as possible."

Well, they only tested whether it was better modelled as 50% Iberia Chalc and 50% Yamnaya or 50% Globular Amphora and 50% Yamnaya, and found that Bell Beaker Central Europe did not model well as if Chalcolithic Iberians and steppe mixed without any ancestry from any of the populations between them. A model of Yamnaya landing like space aliens in Iberia, mixing with the local people and then massively replacing other populations in Central Europe is what they tested and found not to work (and that's not a total waste of time, as some people had proposed it in the past, I believe).

Whether a model like CordedWareGermany+FranceMN at 75:25 or CordedWareGermany+FranceMN+IberiaCA at 75:13:12 would actually fit for Central Europe Bell Beaker even better still is, I guess, something they either didn't test for reasons, or is beyond the capabilities of the data.

@ Kristiina, not so much I don't not believe you, I'm sure you're right, but what you have said adds complication to what I thought was a simple way of checking which autosome in ADMIXTURE mapped to which burial and ancient individual. Seems better to just ask the authors to correct than guess based on complicated speculation based on modern distributions of pretty rare groups! So I left a comment and will leave the topic there to see what they say.

Arza said...

@ Matt

I know, I know. I'm more surprised with CWC. I expected that nMonte will pick up a little bit of everything and produce a "perfect CWC". Could this mean that the population that Yamnayan-like CWC mixed in was very different from the populations that we have in this paper?

For me outliers are the most important samples. Sign of distant connections or unsampled yet populations. Best example - Srubnaya Outlier. The second one - RISE568 - Behemoth forced radiocarbon dating - instead of BB Czech we have an early Slav, which at least partly solves the mystery (RISE568 ancestry in all Balto-Slavs).

Varna, Trypillia and Yamna Ukraine outliers have a great potential for some breakthroughs. Especially the last one. It plots in a place where Proto-Indo-Iranians should be expected. Not the ones from the steppes, but rather from BMAC.

Quick thought - what if Yamna-Iran-ASI model is wrong? Maybe something from Caucasus rolled into steppe, produced this Yamna Outlier... and then they moved directly to India?

We can only wait for the files. There should be a rule on Biorxiv - no Global 10 coordinates (at least), no publication.

Olympus Mons said...

In reich paper, one of the Iberia bell beaker shows similar GREEN (CHG) as the rest of the Central Europeans. do we know which one is it?
Actually in the PCA it is near all others central europeans...
page 22 Extended Data Figure 1. Population structure

Olympus Mons said...

@alberto and Matt,
Thanks. always very instructive.

Olympus Mons said...

...and I know I should insist... but:
Its just that is annoying. Cova da Moura is natural cave outside the "circle of bell beaker" (Zambujal, Vnsp and Leceia) and has inhumations from 3700 - 2200 BCE. And all fragmented. So they dont really know what is the context. it can be a guy from around 3700BC....

Arza said...

@ OM
Probably this one(s) (sample on the left also have very tiny green part):
We find that the great majority of Beaker Complex
individuals outside of Iberia derive a large portion of their ancestry from Steppe populations
(Fig. 2a), whereas in Iberia, such ancestry is absent in all sampled individuals, with the
exception of two (I0461 and I0462) from the Arroyal I site in northern Spain.
Page 5

Olympus Mons said...

and shouldn't insist,
But even the Cisterna girls... were bell beakers just because the were found near V-perforated ivory buttons. thats it. nothing else, not even pottery, was there. So a bell beaker passes by, bangs a chick, leaves his coat... Jesus...!
And them they go and compare with samples, full of beakers, arrows, arches , daggers.... shit. not even a palmela point or something. I want the guys buried with large useless daggers, palmela points, copos, bell beakers.

Even a bigger shit. If they have that adn in Perdigoes for 3000 bc with Mtdna U4, H and U6, why in hell didnt at least look into those?!

capra internetensis said...


The Cova da Moura sample has a radiocarbon date, I assume it is actually for that sample and they didn't just pick one at random.

Olympus Mons said...

Like I always said, 2800BC in Portugal, 2700 BC in Northern spain, 2650 BC jump over the Pyrenees. Something like that.

and this one, a first true bell beaker burial. so, no real bell beakers in Portugal and the first real BB burial, with a bell beaker and a carinated one (so, a family from the south) turns out with CHG. humm lets keep an eye on this girl as well...I0462

Olympus Mons said...

Yes. by AMS. tks. - nevertheless nothing with him was bell beaker... the place has 1500 years of burials.

Algan mardi said...

"Before the hittites , the north-Levant and all of Anatolia except coastal black sea anatolia was IE peoples" could you extend this a little?
in wiki about Kültepe " It is the site of discovery of the earliest traces of the Hittite language, and the earliest attestation of any Indo-European language, dated to the 20th century BC"
"differentiation of Hittite and Luwian, within the Anatolian group, already ca. 2000 BC, in the documents of Kültepe, what means that COMMON ANATOLIAN MUST BE MUCH OLDER
Matt link this where the oldest branch is hittite.
¿ could not be a signal, even a little, about a possible PIE homeland?

Davidski said...

@Algan mardi

Could not be a signal, even a little, about a possible PIE homeland?

No, because people travel.

If even some of the Hittites, in particular those from elite burials, are shown to be genetically different from earlier West Asian peoples, like Hattians, then the Hittite and other Anatolian languages need not be native to Anatolia.

Algan mardi said...

Yes,people travel, acording with wiki (sorry, i don´t like this kind of link but is the faster one, " The Hattians spoke Hattic, a non-Indo-European language of uncertain affiliation. Hattic is now believed by some scholars to be related to the Northwest Caucasian language group", maybe neither need be native from abnatolia

Davidski said...

Yes, and the three Early Bronze Age Anatolians from the upcoming Lazaridis et al. paper may well be Hattians.

If so, their Caucasus-like ancestry would make perfect sense. Now we need to see some samples from elite Hittite graves.

Algan mardi said...

we will wait

Olympus Mons said...

Nope. they were Hittites and with CHG loaded up from the fact they might descend from SHulaveri stock?

... now, Shulaveri , north Africa, Iberia. looks bad. unless something new comes out (because samples they had, Reich and Rui, were just too lousy to even be on a paper about Bell beakers).

The jury is still out, but does not look good.

Davidski said...

I did some reading about that Early Bronze Age site at Isparta and checked out some maps.

Those three Anatolians weren't Hittites, unless on a holiday, and they just happened to die on their holiday. Not Luwians either; neither the date nor the location fits.

Folker said...

This thread is already far too long, but as we come back to Anatolian languages...

I looked at archelogical sites in North Western Anatolia, and didn't thing anything for a probable route from the Balkans to Central Anatalia. Especially, the land of Pala, where people spoke Palaïc before the conquest made by the Kaskas in the XVth century is totally blank. If we postuled a late arrival from the Balkans, just before 2000 BC, of some already differenciated IE groups, it is a clear difficulty to not have any sample to test form this part of Anatolia.

I'm completely agree with Dave, given what we know of the samples tested by Laradis, give their age, none could be attributed to Hittites, but all to Hattians, a non IE culture. So their results will be unconclusive in regards of IE languages and migrations.

epoch2013 said...


Hittites were renowned for not dying on their holidays back in the days, making them the absolute darlings of travel agencies back then. Not a very well known fact, mind you.

Gaspar said...

@algan mardi

Anatolia had IE languages ~4000BC , long before any hittites as I stated.

These IE languages must have come via the homeland of PIE in north caucasus area prior to 4000BC

I do not see what your issue is

Hittites absorbed some hatti languages when creating Hittite language......they did have a language before absorbing some Hatti, don't you agree that they spoke an IE language ?

Grey said...

coming back to what advantage cattle herders had over farmers in certain regions

if it wasn't LP from the beginning (but over time LP provided a selective advantage within the same context) then what might it have been?

if the root problem for the farmers was low crop yields in certain regions (for whatever reason) then people could compensate with calories from other sources like butter or cheese but the farmers had butter and cheese also so maybe it was just quantity?

iirc oxen were originally used primarily as draft animals but i assume at some point separate breeds of oxen vs milch cows had started to be developed but maybe the cattle herders had taken that process further than the farmers so their cows produced a lot more milk?

(for cheese and butter)

i wonder if that is provable from dna?

Grey said...

meant to link this paper on dominance of dairying in neolithic Britain

Samuel Andrews said...

This is for laughs. So I posted links to the new ancient DNA paper on Maju's blog.

Even after reading these papers Maju still argues R1b P312 originated in Neolithic France and that Corded Ware got its R1a from native Neolithic or Mesolithic Eastern Europeans. And he thinks elevated WHG ancestry is being confused as Steppe ancestry....

Some Maju quotes...

"In my work hypothesis: R1a1 arrived to Central-North Europe at low frequencies, maybe with Neolithic expansion....probably before Corded Ware"

"Very tentatively, I would suggest a South-to-North expansion of R1b-S116 with Artenacian first and BB later"

But there is progress. Because of overwhelming ancient DNA data from Britain he agrees R1b P312 came to Britain from the continent with lots of Steppe admixture. But recall that last year Maju argued R1b P312 arrived in Britain from France with farming.

jv said...

@ Kristiina,
THANK YOU for the mtDNA comparison posts! Really appreciate your research.( all 3 H6a1a's mentioned were female)

zardos said...

@grey: Being on the move is also more healthy. You can escape plagues and foes, while having your food always with you. Just look at the dirt of a tell settlement, dirt of generations of settlers piling up.

By the way, there is a good paper about the first Eastern European cattle breeders

The Eastern European economy was largely based on cattle and horse breeding, while the LBK and Balkan economy had much more pigs and smaller animals in their herds. In Eastern Europe, Kotova argues, horses exceeded pigs, while sheep and goats were not suitable for the forests. Especially in the forest steppe there was a specialisation on cattle and horses.

I think the differences were very much ideological too, because we deal with two very different ways and approaches to life. The CW people used the environment they found, didn't alter it too much and were ready to be constantly on the move. The farmer population was more immobile and reluctant to move on or fight their way throuph foreign territory, even if the conditions were very bad at home.

The main difference might be mobility and the sheer quantity of cattle breeding. The CW pastoralists specialised on breeding huge herds while dropping or drastically reducing crop farming, which they couldn't do on the move anyway.

Its interesting that she connects the earlist forms of animals husbandry in the Ukraine, cattle from the start, with the Azov sea area and caucasus, rather than the Balkans.

Eastern European pastoralists had time to evolve before their successful expansion. They could specialise on pastoralism with large animals.
I'm pretty sure they made progress in doing so, including animal breeding for higher productivity and better processing of animal products.

Grey said...


"By the way, there is a good paper about the first Eastern European cattle breeders"


the other thing i wonder about is vitamin A deficiency - best sources of vitamin A:
- fish
- liver
- eggs
- cheese
- butter
- milk (if drinkable)

one of the symptoms is slow growth and bone development in kids which might be relevant to height difference

again, would be interesting if it was possible to see signs of Vitamin A deficiency in neolithic bone samples

Algan mardi said...

Thanks. I thought you had some data about that. My issue is posted above, some diffuse ideas about the homeland of PIE, or protoPIE or howsoever should be called, locating it in Anatolia-Fertile Crescent area

Algan mardi said...

This papers propose the possibility of an incipient form of pastoralism in mesolithic northwest iberia 9300 cal BP. Elba, mtDNA U5b1

Ric Hern said...

@ zardos

Interesting paper thanks.

If that dates of domestication is accurate then it surely seems to rule out Maykop and Balkan Neolithic Farmers as the source of cattle domestication within the Steppe. It also rules out Maykop as the source.

This only leaves Northeast Anatolian who were not far from the first proposed domestication site in the Taurus Mountains. If R1b(V88) can be connected with the spread of cattle domestication to Africa and maybe be part of the first domestication in the Taurus it is hard not to imagine other R1bs also being in Anatolia during the late Mesolithic.

Davidski said...

OK, please note that Broad MIT/Harvard are not holding back any Yamnaya males from Ukraine for future papers, as suggested by me in the blog entry. They published what they had. Sorry.

Ric Hern said...

@ Davidski


Nirjhar007 said...

But certainly they are holding back females! ...

Davidski said...

But certainly they are holding back females!

Obviously not since all three Yamnaya individuals and the singleton Eneolithic individual from Ukraine are all females.

The really interesting thing you're ignoring is that one of these Yamnaya females from Ukraine is by far the most West Asian shifted of the Yamnaya samples currently available. And the Eneolithic female is also heavily West Asian shifted compared to the Neolithic samples from Ukraine rich in R1a and R1b.

What this suggests is that my Caucasian bride theory might be correct.

Nirjhar007 said...

Hehe I was kidding bro.

Samuel Andrews said...

The CHG-heavy Yamnaya female is definitely supports the Caucasian bride theory.

Because in the supplementary info it is mentioned that a mail order bride receipt was found in her grave. Apparently she was sold for 5 goats. I think the CHG men riped Yamnaya off with that price but that's another discussion.

But seriously if her X-Chromsome is overwhelmingly CHG and if archaeological data indicates was born somewhere else than that's really good evidence.

Ric Hern said...

I think the Ancient Shamans(Doctors) and Elders had an idea what inbreeding can do to a population through thousands of years of accumulated knowledge by observation.

So I think it is quite possible that the Shamans and Elders encouraged some breeding with outside populations.

Nick Patterson (Broad) said...


<"location of PIE homeland still uncertain"

<They already know it's the steppes, they want to make it look like its up for debate <until they release all the papers.

If "They" includes me, I sure don't know the location of the homeland, and
certainly won't unless and until we understand the origin of the Hittites.

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