The G25 Proto-Indo-European master sheet

I'm putting together a G25 Proto-Indo-European master sheet that I'll be using in a future blog post about the Indo-European, or, if you wish, Indo-Anatolian homeland.

An early version of the master sheet is available HERE. Please note, the first pop label is my own while the second one is from the v66.2M.AADR dataset.

I'm aiming to expand it significantly and update it regularly. Below is a Vahaduo G25 West Eurasia PCA with all the samples. What am I missing? Feel free to let me know in the comments.

Spoiler alert! Here are a few of the revelations that you can expect in the aforementioned blog post about the Indo-European homeland:

- Yamnaya is almost certainly derived from Serednii Stih (aka Sredny Stog). I've been talking about this for years (see here) and you can probably pick it up without too much trouble in the PCA above. Finally, even Iosif Lazaridis, David Reich, Nick Patterson and friends are now on board with this idea as per their latest paper on the topic (see Lazaridis et al. 2025).

- However, I'm not convinced by the Lazaridis et al. hypothesis that the Caucasus–lower Volga (CLV) cline is intimately linked to the Indo-Anatolian and proto-Anatolian expansions. That's because the CLV cline is an artifact of isolation-by-distance working for thousands of years across a vast and highly diverse landscape, and it includes a wide range of populations that usually have nothing to do with each other.

- The question of who actually spoke Indo-Anatolian first will, in all likelihood, never be solved to everyone's satisfaction. But the number of true candidate groups is actually quite small, and I reckon they're almost certainly all highlighted in my PCA above. Wink, wink, nudge, nudge.

See also...

The Caucasus is a semipermeable barrier to gene flow

The PIE homeland controversy: December 2024 open thread

It seems like we're getting close to the moment when Iosif Lazaridis has to finally admit that the Proto-Indo-European (PIE) homeland was located in Eastern Europe, and also that the ancestors of the Hittites and other Anatolian speakers entered Anatolia via the Balkans.

Let's discuss.

However, please note that comments from total morons, trolls and/or mentally unstable people will not be approved.

See also...

Indo-European crackpottery

The story of the Khvalynsk people

I'm totally serious when I say that this video is more objective, informative and accurate than any peer-reviewed paper published to date when it comes to the genetic origins of the Khvalynsk people.

However, that's not to say it's perfect. I think it misses some important details. See here...

The Caucasus is a semipermeable barrier to gene flow

Dear David, Nick, Iosif...let's set the record straight

Understanding the Eneolithic steppe

Indo-European crackpottery

I've now had the chance to read and digest the following two papers in Science about the origin of Indo-European languages:

Language trees with sampled ancestors support a hybrid model for the origin of Indo-European languages, Heggarty et al.

The genetic history of the Southern Arc: A bridge between West Asia and Europe, Lazaridis, Alpaslan-Roodenberg et al.

The Heggarty et al. paper is pure fluff. It offers nothing useful or even remotely interesting.

For instance, the authors derive some Indo-European languages in Europe from Anatolian farmers and others from Caucasus hunter-gatherers (see here). This is not just exceedingly far fetched, but also obviously forced.

Wolfgang Haak and Johannes Krause, you should be deeply ashamed of yourselves.

I've already commented extensively about the Lazaridis, Alpaslan-Roodenberg et al. paper (for example, see here). But the one thing I need to add is that this paper is what it is due to the inherent bias of some of the lead authors to push the Indo-Anatolian homeland into West Asia. I won't even bother mentioning their names, because we all know who they are.

See also...

Crazy stuff

Dear David, Nick, Iosif...let's set the record straight

The story of R-V1636

The story of R-V1636

Who wants to bet against this map? Keep in mind that ART038 (~3000 calBCE) remains the oldest sample with the V1636 and R1b Y-chromosome mutations in the West Asian ancient DNA record. Ergo, there's nothing to suggest that V1636 or R1b entered Eastern Europe from West Asia.

See also...

A tantalizing link

How relevant is Arslantepe to the PIE homeland debate?

The Kura-Araxes people deserve better

When discussing the Kura-Araxes culture and its people it's important to understand these key points:

- there is Eastern European steppe ancestry in Kura-Araxes samples, and if you're not seeing it then you're not looking hard enough

- Armenian Kura-Araxes samples are mainly a mixture between three different groups currently best represented in the ancient DNA record by ARM_Areni_C, IRN_Hajji_Firuz_C and RUS_Darkveti-Meshoko_En

- ergo, most of the steppe ancestry in the Kura-Araxes population of what is now Armenia must have been mediated via local Chalcolithic groups like ARM_Areni_C

- Kura-Araxes samples show Mesopotamian-related ancestry, and this mustn't be ignored.

Oh, you don't believe it because you just read a big paper in Science claiming otherwise?

Well, the authors of that paper, Lazaridis, Alpaslan-Roodenberg et al., used distal mixture models to study the ancestry of their Kura-Araxes samples, and such models can miss important details.

Consider these three proximate mixture models for a relatively high quality and very homogenous Kura-Araxes sample set from the aforementioned paper. They were done with the qpAdm software

ARM_Kura-Araxes_Berkaber
ARM_Areni_C 0.239±0.068
IRN_Hajji_Firuz_C 0.379±0.068
RUS_Darkveti-Meshoko_En 0.382±0.054
P-value 0.285122 (Pass)
Full output

ARM_Kura-Araxes_Berkaber
IRN_Hajji_Firuz_C 0.569±0.051
RUS_Darkveti-Meshoko_En 0.363±0.058
RUS_Progress_En 0.068±0.020
P-value 0.20306 (Pass)
Full output

ARM_Kura-Araxes_Berkaber
IRN_Hajji_Firuz_C 0.531±0.060
RUS_Darkveti-Meshoko_En 0.469±0.060
P-value 0.0132579 (Fail)
Full output

Some caveats apply. For instance, the pass threshold (P-value ≥0.05) is arbitrary. But the point is that the models look much better with steppe-related and steppe reference populations (ARM_Areni_C and RUS_Progress_En, respectively).

Moreover, the unique and vital Darkveti-Meshoko population is represented by just one individual. I also have the genotypes of his brother and sister, but relatives aren't allowed in these sorts of tests.

Including a singleton in the analysis means that I can't use the inbreed: YES option, which apparently can be a bad thing. Nevertheless, these models do look very solid.

Indeed, I can also model ARM_Kura-Araxes_Berkaber as practically 100% RUS_Maykop_Novosvobodnaya, perhaps with some excess ARM_Areni_C-related input.

ARM_Kura-Araxes_Berkaber
ARM_Areni_C 0.094±0.087
RUS_Maykop_Novosvobodnaya 0.906±0.087
P-value 0.284259 (Pass)
Full ouput

This makes good sense, because RUS_Maykop_Novosvobodnaya can also be modeled solidly as a mixture between IRN_Hajji_Firuz_C, RUS_Darkveti-Meshoko_En and RUS_Progress_En.

RUS_Maykop_Novosvobodnaya
IRN_Hajji_Firuz_C 0.614±0.056
RUS_Darkveti-Meshoko_En 0.307±0.064
RUS_Progress_En 0.080±0.022
P-value 0.141468 (Pass)
Full output

I don't know whether the genetic relationship between ARM_Kura-Araxes_Berkaber and RUS_Maykop_Novosvobodnaya shown in my model is due to Maykop ancestry in the former. It might just be a coincidence in the sense that the same or similar processes led to the formation of both groups. Feel free to let me know your thoughts about that in the comments.

The fact that the Kura-Araxes people harbored steppe ancestry might be very important in the debate over the location of the so called Indo-Anatolian homeland. For instance, it's possible that the proto-Anatolian language spread from the North Caucasus into Anatolia via the Kura-Araxes culture.

But, admittedly, such a solution doesn't have strong support from historical linguistics data, which suggest that the Indo-Anatolian homeland was located in what is now Ukraine and that Anatolian speakers entered West Asia via the Balkans:

Indo-European cereal terminology suggests a Northwest Pontic homeland for the core Indo-European languages

See also...

R-V1636: Eneolithic steppe > Kura-Araxes?

Dear Iosif...Yamnaya

But Iosif, what about the Phrygians?

Dear Iosif...Yamnaya

Even though the Yamnaya culture probably originated in what is now Ukraine, the earliest Yamnaya samples currently available are from the modern-day Samara region of Russia. They mostly date to around 3,000 BCE. I can analyze their ancestry using Principal Component Analysis (PCA) data.

Target: RUS_Yamnaya_Samara
Distance: 3.2816% / 0.03281581
81.0 RUS_Progress_En
14.4 UKR_N
4.6 HUN_Vinca_MN
0.0 ARM_Aknashen_N
0.0 ARM_Masis_Blur_N
0.0 AZE_Caucasus_lowlands_LN
0.0 BGR_C
0.0 BGR_Dzhulyunitsa_N
0.0 IRN_Ganj_Dareh_N
0.0 IRN_Hajji_Firuz_C
0.0 IRN_Seh_Gabi_C
0.0 IRN_Tepe_Abdul_Hosein_N
0.0 IRN_Wezmeh_N
0.0 RUS_Darkveti-Meshoko_En
0.0 RUS_Maykop
0.0 RUS_Maykop_Late
0.0 RUS_Maykop_Novosvobodnaya

The above results show exactly zero ancestry from West Asia. Admittedly, both RUS_Progress_En and HUN_Vinca_MN are European ancients with significant West Asian-related ancestry. However, this ancestry is very distantly West Asian-related, and, for instance, it almost certainly has no relevance to the Indo-Anatolian homeland debate.

The Afanasievo culture of Central Asia is regarded to have been an early offshoot of the Yamnaya culture. A good number of Afanasievo samples are available, so let's have a look if their results match those of the Yamnaya folks. And indeed they do, since BGR_C is very similar to HUN_Vinca_MN.

Target: RUS_Afanasievo
Distance: 3.4055% / 0.03405499
84.0 RUS_Progress_En
11.4 UKR_N
4.6 BGR_C
0.0 ARM_Aknashen_N
0.0 ARM_Masis_Blur_N
0.0 AZE_Caucasus_lowlands_LN
0.0 BGR_Dzhulyunitsa_N
0.0 HUN_Vinca_MN
0.0 IRN_Ganj_Dareh_N
0.0 IRN_Hajji_Firuz_C
0.0 IRN_Seh_Gabi_C
0.0 IRN_Tepe_Abdul_Hosein_N
0.0 IRN_Wezmeh_N
0.0 RUS_Darkveti-Meshoko_En
0.0 RUS_Maykop
0.0 RUS_Maykop_Late
0.0 RUS_Maykop_Novosvobodnaya

To try this at home, stick the PCA data in the text file here into the relevant fields here and cranck up the "Cycles" to 4X. You should see exactly zero ancestry from West Asia every time.

I can, more or less, reproduce these results with tools that are routinely used in peer reviewed papers. Below is a table of mixture models produced with the qpAdm software. I set the pass threshold to P ≥0.05, which is an arbitrary value, but the pattern is clear. The full output from each qpAdm run is available here.

Importantly, qpAdm needs to be fed the relevant "right pop" outgroups to be able to discriminate accurately between reference populations.

right pops:
CMR_Shum_Laka_8000BP
MAR_Taforalt
Levant_Natufian
IRN_Ganj_Dareh_N
Levant_PPNB
TUR_Marmara_Barcin_N
HUN_Starcevo_N
HUN_Koros_N
SRB_Iron_Gates_HG
Iberia_Southeast_Meso
RUS_Karelia_HG
RUS_West_Siberia_HG
RUS_Boisman_MN
MNG_North_N
TWN_Hanben
BRA_LapaDoSanto_9600BP

So, for instance, if one were to use in this role the modern-day Mbuti people, as opposed to, say, the ancient hunter-gatherers of Shum Laka, one might find that many models look statistically better than they should. And then one might also find that the Yamnaya samples carry significant West Asian ancestry.

Actually, I'm not opposed to the idea of some West Asian ancestry in Yamnaya. Indeed, considering the extraordinary mobility of the Yamnaya people and their Eneolithic predecessors on the Pontic-Caspian steppe, it would be unusual if they didn't come into close contact and mix, to some degree, with their neighbors from West Asia.

However, based on everything I've seen, from uniparental markers to different types of autosomal genetic tests, it's clear to me that there's no substantial West Asian ancestry in any Yamnaya samples, except for an outlier female from modern-day Ozera, Ukraine (see here).

Admittedly, ancient DNA does have a habit of throwing curveballs, so I'm eagerly awaiting new Eneolithic samples from the Pontic-Caspian steppe, particularly those associated with the Yamnaya-like Sredni Stog culture, to help finally settle this issue.

Believe it or not, a contact recently sent me a supposedly unpublished female sample from a ~4,200 BCE Sredni Stog burial in modern-day Igren, east central Ukraine. So what the hell, let's assume for the time being that this sample is genuine. This is how Miss Sredni Stog behaves in my PCA mixture test.

Target: UKR_Sredni_Stog
Distance: 4.0769% / 0.04076877
75.6 RUS_Progress_En
17.8 UKR_N
6.6 HUN_Vinca_MN
0.0 ARM_Aknashen_N
0.0 ARM_Masis_Blur_N
0.0 AZE_Caucasus_lowlands_LN
0.0 BGR_C
0.0 BGR_Dzhulyunitsa_N
0.0 HUN_Vinca_MN
0.0 IRN_Ganj_Dareh_N
0.0 IRN_Hajji_Firuz_C
0.0 IRN_Seh_Gabi_C
0.0 IRN_Tepe_Abdul_Hosein_N
0.0 IRN_Wezmeh_N
0.0 RUS_Darkveti-Meshoko_En
0.0 RUS_Maykop
0.0 RUS_Maykop_Late
0.0 RUS_Maykop_Novosvobodnaya

Wow, just wow. Have we actually found Miss Proto-Yamnaya? What does qpAdm have to say in the matter?

UKR_Sredni_Stog
HUN_Vinca_MN 0.034±0.028
RUS_Progress_En 0.796±0.045
UKR_N 0.170±0.034
P-value 0.41088

Again, this is an excellent match with the results from my PCA test, especially if we take into account the standard errors. However, with qpAdm it's also possible to model this individual's ancestry as part West Asian.

UKR_Sredni_Stog
AZE_Caucasus_lowlands_LN 0.056±0.039
RUS_Progress_En 0.761±0.061
UKR_N 0.183±0.036
P-value 0.465667

As I pointed out above, it's plausible for such people to harbor some West Asian ancestry, but I'm very sceptical that this is really the case here, despite the rather solid qpAdm statistical fit. That's because UKR_Sredni_Stog is not a high quality sample, and, from my experience, qpAdm often has problems analyzing fine scale ancestry in singletons or even small groups that show excess DNA damage and/or offer much less than a million markers.

See also...

Dear Iosif, about that ~2%

But Iosif, what about the Phrygians?

Dear Iosif, about that ~2%

The debate over the location of the so called Indo-Anatolian homeland won't be decided by the persistence of any type of genetic ancestry in ancient Anatolia.

It'll be decided by a multidisciplinary study on the interactions between the ancient peoples of the North Pontic steppe, the eastern Balkans, and western Anatolia.

If such a study finds a pulse of steppe-related gene flow from the Balkans into Anatolia sometime during the early metal ages, it'll corroborate the linguistic hypothesis that a language ancestral to Hittite, Luwian and related tongues moved into Anatolia from Eastern Europe.

Why do we only need a pulse of gene flow, you might ask? Obviously, because:

- language and genetic ancestry can start with a strong association but, since they're not linked, they can eventually follow very different trajectories

- the dilution of genetic ancestry is an important factor, especially in ancient West Asia, and it must be taken into account in models of language spread, rather than ignored in favor of simple, elegant models that do not reflect reality.

Here's my favorite quote from the recent Lazaridis, Alpaslan-Roodenberg et al. paper, because, probably unbeknownst to the authors, it's exceptionally revealing about the spread of a wide range of Indo-European speakers into Anatolia.

However, in individuals from Gordion, a Central Anatolian city that was under the control of Hittites before becoming the Phrygian capital and then coming under the control of Persian and Hellenistic rulers, the proportion of Eastern hunter-gatherer ancestry is only ~2%, a tiny fraction for a region controlled by at least four different Indo-European–speaking groups.

Indeed, this is exactly what the Lazaridis, Alpaslan-Roodenberg et al. paper should've been about. That is, the authors should've given us a painstaking account of the spread of different ancient Indo-European speaking groups into Anatolia and explained how, overall, their DNA was rapidly diluted to a trace amount.

However, instead they treated us to a make-believe tale about a so called Indo-Anatolian homeland in what is now Armenia.

See also...

Dear Iosif...Yamnaya

But Iosif, what about the Phrygians?

Dear Iosif...

Dear Iosif #2

Dear Iosif #3

But Iosif, what about the Phrygians?

A paper in Science co-authored by around 200 scientists from some of the world's top academic institutions surely must mean something, right? Not necessarily.

In this short blog post I'll try to explain, as simply as I can, why the Lazaridis, Alpaslan-Roodenberg et al. paper doesn't get us any closer to solving the riddle of the so called Indo-Anatolian homeland.

However, it must be said that the paper does include many interesting and valuable samples. I'll be using six of these samples, labeled TUR_C_Gordion_Anc, to argue my case.

The TUR_C_Gordion_Anc sample set is from Gordion, the capital of ancient Phrygia, and thus, in all likeliness, it represents Phrygian speakers.

Phrygian is an Indo-European language and the leading hypothesis is that it originated in the Balkans.

In terms of fine scale ancestry, TUR_C_Gordion_Anc can be reliably divided into two genetic clusters. In the Principal Component Analysis (PCA) below these clusters are labeled TUR_C_Gordion_Anc1 and TUR_C_Gordion_Anc2.

Note that TUR_C_Gordion_Anc1 is obviously pulling away from TUR_C_Gordion_Anc2 towards samples from the Balkans. I've used ancient samples from what is now North Macedonia, labeled MKD_Anc, to represent the Balkans. To see an interactive version of the plot, paste the PCA coordinates from here into the relevant field here.

Visually, this is not an especially dramatic outcome, but it's an incredible result nonetheless, because it shows that even a few ancient samples can help to solve an age old mystery.

Across many dimensions of genetic variation, the shift in the PCA from TUR_C_Gordion_Anc1 to TUR_C_Gordion_Anc2 represents about 20% admixture from the Balkans, and about 8% from the Eastern European steppe. That's plenty enough to corroborate the linguistic hypothesis that the Phrygians originated in the Balkans, and that some of their ancestors came from the steppe. The mixture models below were done with the tools here.

Target: TUR_C_Gordion_Anc1
Distance: 1.6634% / 0.01663373
40.6 Kura-Araxes_ARM_Kaps
22.2 Anatolia_Barcin_N
21.8 MKD_Anc
13.6 Levant_PPNB
1.4 IRN_Ganj_Dareh_N
0.4 Han

Target: TUR_C_Gordion_Anc1
Distance: 1.7109% / 0.01710904
40.2 Kura-Araxes_ARM_Kaps
37.8 Anatolia_Barcin_N
12.4 Levant_PPNB
8.0 Yamnaya_RUS_Samara
1.2 IRN_Ganj_Dareh_N
0.4 Han

Target: TUR_C_Gordion_Anc2
Distance: 2.0293% / 0.02029339
51.0 Kura-Araxes_ARM_Kaps
26.8 Anatolia_Tepecik_Ciftlik_N
17.6 Anatolia_Barcin_N
4.6 Levant_PPNB

Surprisingly, Lazaridis, Alpaslan-Roodenberg et al. didn't have much to say about this topic. This quote basically sums it up:

However, in individuals from Gordion, a Central Anatolian city that was under the control of Hittites before becoming the Phrygian capital and then coming under the control of Persian and Hellenistic rulers, the proportion of Eastern hunter-gatherer ancestry is only ~2%, a tiny fraction for a region controlled by at least four different Indo-European–speaking groups.

I have no doubt that Lazaridis, Alpaslan-Roodenberg et al. can run a very decent PCA, and then blow it up to a size big enough to show that the Gordion samples represent two genetically somewhat distinct groups. I'm also sure that, if they really try, they can locate significant levels of proximate and relevant European ancestry in some of these samples.

They don't have to use my methods; they can use any methods they like. My point is that they won't find much if they're just looking for genetic signals from the Upper Paleolithic or Mesolithic.

Now, considering the way that the Phrygian question was treated by Lazaridis, Alpaslan-Roodenberg et al., despite the fact that they managed to sequence a few likely Phrygian speakers from none other than the Phrygian capital, let's not pretend that their paper brought us any closer to understanding the genetic origins of Anatolian speakers or pinpointing their ancestral homeland.

In order to even try to solve these problems with ancient DNA, we need a wide range of samples from Hittite, Luwian and other key sites where Anatolian languages were spoken. And then we must analyze them properly.

I'm guessing that Lazaridis, Alpaslan-Roodenberg et al. went out of their way to get such samples, but for one reason or another they failed. If so, that's OK, but I have a feeling that even if they got them, they wouldn't know what to do with them, because at best these samples would only show ~2% Eastern hunter-gatherer ancestry. Haha.

For what it's worth, I believe that the ancient data in the Lazaridis, Alpaslan-Roodenberg et al. paper point to the North Pontic steppe as the Indo-Anatolian homeland, and I'll lay out my arguments in an upcoming blog post.

See also...

Dear Iosif...Yamnaya

Dear Iosif, about that ~2%

Dear Iosif...

Dear Iosif #2

Dear Iosif #3

Dear Iosif #3

Back in 2016 I made this prediction about the origins of the Yamnaya people (Steppe_EMBA):

But here's my prediction: Steppe_EMBA only has 10-15% admixture from the post-Mesolithic Near East not including the North Caucasus, and basically all of this comes via female mediated gene flow from farming communities in the Caucasus and perhaps present-day Ukraine.

The relevant blog post is still here. Looking back, my analysis is a bit sloppy and I didn't articulate my ideas too well. But that was a pretty good prediction for its time, and I believe it still has a chance of being confirmed, more or less.

On the other hand, the widely publicized hypothesis that the Yamnaya population is a ~50/50 mixture between indigenous Eastern European hunter-gatherers and Near Eastern or West Asian migrants never looked right to me. So I'm glad that it's now dead and buried.

Those of you not up to date with this topic, all you need to know is that the Yamnaya genotype existed in Eastern Europe at least a thousand years before Yamnaya, and, moreover, the Yamnaya people are largely derived from Eastern European foragers already rich in Near Eastern-related ancestry. The relevant ancient genomes are on the way (for instance, see here).

Nevertheless, the narrative that waves of Near Eastern migrants moved into prehistoric Eastern Europe, leading to the emergence of the Yamnaya culture and even the Proto-Indo-European language, is still being pushed by some notable scientists working with ancient DNA.

My hope is that, considering the latest revelations about the genetic origins of the Yamnaya people, these scientists can embrace a more nuanced view. How about something like this?

- people moved around, and they were especially mobile on the Eastern European steppe from the Eneolithic onwards

- when they made contact they sometimes mixed, so there was admixture between far flung steppe groups

- since population densities on the steppe were low until the Yamnaya period, minor admixture that entered the steppe during the Neolithic and Eneolithic wasn't dilluted easily.

See also...

Dear Iosif #2

But Iosif, what about the Phrygians?

Dear Iosif, about that ~2%

Dear Iosif...Yamnaya

Dear Iosif...

Update 29/08/22: Dear Iosif #2

...


I'm skimming through the Lazaridis, Alpaslan-Roodenberg et al. paper that just came out at Science. And I feel like someone punched me in the face.

Nevertheless, I'll try to be diplomatic. Suffice to say, for now, that there's some rather strange stuff in this paper.

The main problem is that the authors are attempting to study fine scale ancestry with a somewhat rough distal model. As a result, they miss important details.

For instance, this quote is from the paper's supplementary PDF file, freely available here.

However, the complete lack of association of R-haplogroup descendants and EHG ancestry in either Armenia or Iran is consistent with either a massive dilution of EHG ancestry in these populations resulting in the dissociation of Y-chromosome lineages from autosomal ancestry over time, or with a scenario in which R-M269 was not associated with substantial EHG ancestry to begin with.

Obviously, EHG means Eastern European Hunter-Gatherer. But why focus on EHG? Surely, this makes little sense when looking at the genetic prehistory of West Asia, because no one ever argued that this region was settled by EHG populations. It was widely settled by Yamnaya-related groups, with already heavily diluted EHG ancestry, during the metal ages.

OK, so the authors are actually aware about the potential dilution of EHG ancestry, but they don't really do anything about it.

If we're looking at the origins of West Asian R1b-M269, and using its association with autosomal DNA components as a guide, then we should be focusing on Yamnaya-related ancestry.

For instance, here's a fine scale ancient ancestry model based on Principal Component Analysis (PCA) data. It shows the ancestry proportions of two relatively high coverage Iron Age males from two different sites in Iran from the Lazaridis, Alpaslan-Roodenberg et al. dataset. Both belong to R1b-M269 and both show significant Yamnaya-related ancestry.

Target: IRN_HajjiFiruz_IA:I2327_all
Distance: 2.2930% / 0.02292994
39.6 Kura-Araxes_ARM_Kaps
24.2 IRN_Ganj_Dareh_N
18.2 Levant_PPNB
12.4 Yamnaya_RUS_Samara
4.4 Anatolia_Tepecik_Ciftlik_N
1.2 Han

Target: IRN_Hasanlu_IA:I4232_all
Distance: 2.5179% / 0.02517895
26.0 IRN_Ganj_Dareh_N
25.6 Kura-Araxes_ARM_Kaps
24.4 Anatolia_Tepecik_Ciftlik_N
15.8 Yamnaya_RUS_Samara
7.6 Levant_PPNB
0.6 IRN_Shahr_I_Sokhta_BA2

As a control, here's an earlier, Chalcolithic sample bearing Y-haplogroup J2b from the same region. Not surprisingly, this individual totally lacks the Yamnaya-related signal.

Target: IRN_HajjiFiruz_ChL:I4241_all
Distance: 2.7938% / 0.02793782
32.6 Kura-Araxes_ARM_Kaps
25.6 IRN_Ganj_Dareh_N
23.6 Anatolia_Tepecik_Ciftlik_N
18.2 Levant_PPNB

Overall, these results make perfect sense. I could probably locate very minor signals of EHG ancestry in the Iron Age samples, but that would be more difficult and much less certain, so I won't bother.

Soon I'll be able to rerun these analyses with Bronze Age samples from Dagestan and surrounds. That should bump up the levels of Yamnaya-related ancestry and improve the statistical fits (wink, wink, nudge, nudge).

Disappointingly, Lazaridis, Alpaslan-Roodenberg et al. go so far as to suggest that R1b-M269 may have originated in West Asia.

However, considering the scores of ancient Eastern European populations rich in R1b-M269 and many near and far related subclades of R1b, this makes no sense whatsoever.

Indeed, contemplating nowadays that R1b-M269 might be native to West Asia, where R1b only starts showing up in the ancient DNA record during the Copper Age, is about as stupid as claiming that gravity doesn't exist.

Largely due to their distal model approach, Lazaridis, Alpaslan-Roodenberg et al. also argue that the Indo-Anatolian homeland was located in what is now Armenia and surrounds. I'm far from convinced that this solution will stand the test of time.

In terms of the more widely accepted theory that the Indo-Anatolian homeland was located on the Pontic-Caspian steppe in Eastern Europe, the most important samples in the paper are the three Bronze Age individuals from Yassitepe in western Anatolia. That's because they're from a region that is traditionally seen as the entry point of Indo-Anatolian speakers into Anatolia from the European steppe via the Balkans.

Interestingly, individual I5737, dated to 2035-1900 calBCE or the Middle Bronze Age, belongs to Y-chromosome haplogroup I2a-P78, which surely must be a signal of European ancestry. I see this as a significant result.

Here's how the trio from Yassitepe look in my fine scale ancient ancestry model. Minor Yamnaya-related ancestry does show up, although, admittedly, it might just be noise in individual I5735.

Target: TUR_Aegean_Izmir_Yassitepe_MBA:I5737
Distance: 2.7507% / 0.02750748
58.4 Anatolia_Barcin_N
20.4 Kura-Araxes_ARM_Kaps
9.2 Anatolia_Tepecik_Ciftlik_N
5.6 IRN_Ganj_Dareh_N
3.8 Yamnaya_RUS_Samara
2.6 Levant_PPNB

Target: TUR_Aegean_Izmir_Yassıtepe_EBA:I5733
Distance: 2.7969% / 0.02796887
52.0 Anatolia_Barcin_N
27.2 Kura-Araxes_ARM_Kaps
8.6 Levant_PPNB
6.2 Yamnaya_RUS_Samara
6.0 IRN_Ganj_Dareh_N

Target: TUR_Aegean_Izmir_Yassıtepe_EBA:I5735
Distance: 3.1270% / 0.03127009
36.0 Kura-Araxes_ARM_Kaps
32.4 Anatolia_Tepecik_Ciftlik_N
26.0 Anatolia_Barcin_N
2.8 IRN_Shahr_I_Sokhta_BA2
1.2 Yamnaya_RUS_Samara
1.0 Levant_PPNB
0.6 MAR_Taforalt

This isn't much, especially considering it's already late 2022, but it's better than nothing. Fortunately, more samples from Bronze Age western Anatolia are on the way (wink, wink, nudge, nudge).

However, I'm not done with the Lazaridis, Alpaslan-Roodenberg et al. dataset yet. I'm planning to spend much more time on this blog in the coming weeks and months and will be using their samples in a wide range of analyses.

Citation...

Iosif Lazaridis, Songül Alpaslan-Roodenberg et al., The genetic history of the Southern Arc:A bridge between West Asia and Europe, Science 377, eabm4247 (2022)

See also...

Dear Iosif #3

But Iosif, what about the Phrygians?

Dear Iosif, about that ~2%

Dear Iosif...Yamnaya

David Reich on the origin of the Yamnaya people (!?)

Harvard's David Reich is doing a talk next month about the genetic history of West Asia and nearby parts of Europe. This is a quote from an online abstract of the talk (found here).

The impermeability of Anatolia to exogenous migration contrasts with our finding that the Yamnaya had two distinct gene flows, both from West Asia, suggesting that the Indo-Anatolian language family originated in the eastern wing of the Southern Arc and that the steppe served only as a secondary staging area of Indo-European language dispersal.

If this is actually what David Reich is going to claim then I'd say his team has a lot of work to do before they put out their paper on the topic.

First of all, Yamnaya did not have two distinct gene flows from West Asia. I don't even know what that means exactly, but there's no way that this statement is correct no matter how one interprets it.

In fact, the Yamnaya population formed on the Pontic-Caspian steppe from earlier groups native to this part of Eastern Europe, such as the people associated with the Sredny Stog culture.

That is, there were no migrations from West Asia into Eastern Europe that can be claimed to have been instrumental in the emergence of the Yamnaya population. On the other hand, Yamnaya may have been significantly influenced by cultural impulses from West Asia, but this is nothing new.

In terms of deep population structure, the Yamnaya genotype can be described as a mixture between Eastern European and West Asian-related genetic components. However, these Asian-related components were already in Europe thousands of years before Yamnaya came into existence.

Indeed, soon to be published ancient DNA shows that hunter-gatherers very similar to the Yamnaya people, packing quite a lot of West Asian-related ancestry, lived in the Middle Don region (just north of the Pontic-Caspian steppe) well before 5,000 BCE (see here).

So, did the West Asian ancestors of these Middle Don hunter-gatherers speak Proto-Indo-European, or, as David Reich calls it, Indo-Anatolian? Keep in mind that most linguists put the birth of Indo-Anatolian around 4,000 BCE, which is actually the Sredny Stog period.

Moreover, in underlining Anatolia's supposed impermeability to exogenous migration, David Reich is arguing against things that no one worth their salt ever claimed. That's because the spread of Indo-Anatolian speakers into Anatolia has never really been described by archeologists and linguists as a massive migration, but rather as an infiltration into lands already heavily populated by the Hattians (for instance, see here).

We may have already seen the genetic evidence of this infiltration in the presence of steppe Y-chromosome haplogroup R-V1636 in a Chalcolithic burial at Arslantepe (see here and here). Let's wait and see what else crops up over the next few years as many more ancient Anatolian genomes are sequenced by David Reich and colleagues.

See also...

The PIE homeland controversy: February 2022 status report

I think we'll see the emergence of two main competing proto-Indo-European (PIE) homeland theories over the next few years:

- a homeland in the Eneolithic North Caucasus, and the spread of Anatolian languages into West Asia with Maykop-related ancestry

- a homeland in the North Pontic region, possibly within the Eneolithic Sredny Stog archeological culture, and the spread of Anatolian languages into West Asia via the Balkans.

Both theories have support from ancient DNA. Some of it has already been published (for instance, see here).

At this point, I can see myself firmly in the North Pontic camp, even if it turns out that North Pontic-related ancestry only made a fleeting impact on Bronze Age Anatolia.

After all, there's no direct relationship between genes and languages, so to prove that Anatolian languages came from the North Pontic, there's no need for North Pontic-related ancestry to persist in Anatolia, as long as we have solid evidence that people with this type of ancestry moved there at the right time.

In my mind, for now, the Maykop culture provides an excellent explanation for non-Indo-European influences in PIE, and there's no need to make it Indo-European speaking, let alone PIE speaking.

See also...

The PIE homeland controversy: June 2021 status report

R1a vs R1b in third millennium BCE Central Europe (Papac et al. 2021)

R1a-M417 and R1b-L51 are by far the most important Y-chromosome haplogroups in Europe today. More precisely, R1a-M417 dominates in Eastern Europe, while R1b-L51 in Western Europe.

It's been obvious for a while now, at least to me, that both of these Y-haplogroups are closely associated with the men of the Late Neolithic Corded Ware culture (CWC). Indeed, in my mind they're the main genetic signals of its massive expansion, probably from a homeland somewhere north of the Black Sea in what is now Ukraine.

I'm still not exactly sure how the east/west dichotomy between R1a and R1b emerged in Europe, but, thanks to a new paper by Papac et al. at Science Advances, at least now I have a working hypothesis about that. Below is a quote from the said paper, emphasis is mine:

Our results suggest that the Y-lineage diversity in early CW males was supplanted by a nonrandom process [selection, social structure, or influx of nonlocal R1a-M417(xZ645) lineages] that drove the collapse in Y-chromosomal diversity. A simultaneous decline of Y-chromosomal diversity dating to the Neolithic has been observed across most extant Y-haplogroups (64), possibly due to increased conflict between male-mediated patrilines (65). We view that changes in social structure (e.g., an isolated mating network with strictly exclusive social norms) could be an alternative cause but would be difficult to distinguish in the underlying model parameters.

Right, so even though the CWC was clearly a community of closely related groups, there must have been some competition between its different clans. And since these clans were highly patriarchal and patrilineal, this competition probably led to different paternal lineages dominating different parts of the CWC horizon, with M417 becoming especially common in the east and L51 in the west.

Of course, the expansions of post-Corded Ware groups, such as the M417-rich Slavs in Eastern Europe and L51-rich Celts in Western Europe, were also instrumental in creating Europe's R1a/R1b dichotomy, but obviously these groups were in large part the heirs of the CWC.

By the way, most of the samples from Papac et al. are already in the Global25 datasheets linked here. Look for the labels listed here. Below is a plot made from the Global25 data courtesy of regular commentator Matt.

Citation: L. Papac, M. Ernée, M. Dobeš, M. Langová, A. B. Rohrlach, F. Aron, G. U. Neumann, M. A. Spyrou, N. Rohland, P. Velemínský, M. Kuna, H. Brzobohatá, B. Culleton, D. Daněček, A. Danielisová, M. Dobisíková, J. Hložek, D. J. Kennett, J. Klementová, M. Kostka, P. Krištuf, M. Kuchařík, J. K. Hlavová, P. Limburský, D. Malyková, L. Mattiello, M. Pecinovská, K. Petriščáková, E. Průchová, P. Stránská, L. Smejtek, J. Špaček, R. Šumberová, O. Švejcar, M. Trefný, M. Vávra, J. Kolář, V. Heyd, J. Krause, R. Pinhasi, D. Reich, S. Schiffels, W. Haak, Dynamic changes in genomic and social structures in third millennium BCE central Europe. Sci. Adv. 7, eabi6941 (2021).

See also...

On the origin of the Corded Ware people

Understanding the Eneolithic steppe

Conan the Barbarian probably belonged to Y-haplogroup R1a

On the origin of the Corded Ware people

There's been a lot of talk lately about the finding that the peoples associated with the Corded Ware and Yamnaya archeological cultures were genetic cousins (for instance, see here). As I've already pointed out, this is an interesting discovery, but, at this stage, it's difficult to know what it means exactly.

It might mean that the Yamnayans were the direct predecessors of the Corded Ware people. Or it might just mean that, at some point, the Corded Ware and Yamnaya populations swapped women regularly (that is, they practiced female exogamy with each other).

In any case, I feel that several important facts aren't being taken into account by most of the interested parties. These facts include, in no particular order:

- despite being closely related, the Corded Ware and Yamnaya peoples were highly adapted to very different ecological zones - temperate forests and arid steppes, respectively - and this is surely not something that happened within a few years and probably not even within a couple of generations

- both the Corded Ware and Yamnaya populations expanded widely and rapidly at around the same time, but never got in each others way, probably because they occupied very different ecological niches

- despite sharing the R1b Y-chromosome haplogroup, their paternal origins were quite different, with Corded Ware males rich in R1a-M417 and R1b-L51 and Yamnaya males rich in R1b-Z2103 and I2a-L699

I suppose it's possible that the Corded Ware people were overwhelmingly and directly derived from the Yamnaya population. But right now my view is that, even if they were, then the Yamnaya population that they came from was quite different from the classic, R1b-Z2103-rich Yamnaya that spread rapidly across the steppes.

Indeed, perhaps what we're dealing with here is a very early (proto?) Yamnaya gene pool located somewhere in the border zone between the forests and the steppes, that then split into two main sub-populations, with one of these groups heading north and the other south?

I do wonder what David Anthony would say if he was made aware of the above mentioned facts? Then again, perhaps he's already aware of them, and simply chose to ignore them when formulating his latest theory about the origin of the Corded Ware people?

See also...

The PIE homeland controversy: June 2021 status report

Archeologist David Anthony has made several appearances online recently to promote his theories about the origins of the Corded Ware and Yamnaya cultures and peoples.

In a clip on Youtube he reiterated his theory that the so called Iranian-related ancestry in the Yamnaya people actually came from what is now Iran, and, more precisely, that it was carried by hunter-gatherers who travelled relatively rapidly from the South Caspian region into the Volga Delta in what is now Russia.

It's still a complete mystery to me as to why a group of hunter-gatherers from the South Caspian would undertake such a migration, instead of, say, expanding their range gradually over thousands of years, first into the Caucasus and eventually into Eastern Europe.

But there's a more serious problem with Anthony's theory: it contradicts the currently available ancient DNA. That's because the so called Iranian-related ancestry in the Yamnaya people is most closely related to the Kotias and Satsurblia hunter-gatherers from what is now Georgia, and these hunter-gatherers form a separate clade from the earliest samples from what is now Iran. For instance, see here and here.

Also, in a podcast on Razib's blog, Anthony doubled down on his theory that Y-chromosome haplogroup R1a was closely associated with Yamnaya plebs who were excluded from Kurgan burials, and, as a result, their remains haven't yet been sampled.

At least this theory isn't yet contradicted by ancient DNA, but it's more complicated and less parsimonious than my theory, which posits that R1a, or rather R1a-M417, was simply a very rare lineage in the Yamnaya population, and that it only became a common and widespread marker thanks to the Corded Ware expansion (see here).

Intriguingly, my understanding is that there are several unpublished R1a samples from the Caspian and Volga steppes at Harvard's David Reich Lab that have been classified by its scientists as Yamnaya outliers. Of course, Anthony is collaborating on at least one major paper with this lab (see here).

Ergo, I strongly suspect that Anthony's theory is in part based on these Yamnaya outliers. However, I also believe that these samples are wrongly dated and probably represent Scythians and/or Sarmatians. I'll be able to look into that if they're ever published.

Speaking of the David Reich Lab, its leading scientists, David Reich and Nick Patterson, have also made appearances online recently, on Youtube and Razib's blog, respectively, to reveal that the Corded Ware and Yamnaya peoples aren't just very similar genetically, but in fact close cousins.

This is a very interesting finding. Apparently it's based on a relatively high level of Identity-by-Descent (IBD) segment sharing between Corded Ware and Yamnaya samples, but that's all I know. I'm guessing that the relevant paper is coming soon (that is, within the next five years).

However, the long-standing question that the readers of this blog want to see answered is not whether the Corded Ware and Yamnaya peoples are close cousins, but whether Yamnaya migrants founded the Corded Ware culture. The obvious way to prove that they did is to find at least one ancient population unambiguously classified as part of the Yamnaya horizon that is rich in the typically Corded Ware Y-haplogroups R1a-M417 and R1b-L151.

See also...

On the origin of the Corded Ware people

The PIE homeland controversy: January 2019 status report

The PIE homeland controversy: August 2019 status report

Balto-Slavic drift

A few years ago I began using the term "Balto-Slavic genetic drift" to describe the fine-scale genetic signal that is shared by the speakers of Baltic and Slavic languages to the exclusion of Europeans without significant Balto-Slavic ancestry.

As a result, nowadays, many people online use the term "Balto-Slavic drift" when referring to this phenomenon.

The easiest way to prove that Balto-Slavic drift exists is to run a fine-scale Principal Component Analysis (PCA) of European genetic variation with a lot of Balto-Slavic samples in the mix. Indeed, my Global25 PCA analysis does a great job of illustrating the impact of Balto-Slavic drift on the population structure of Europe both in PCA plots and mixture models (for instance, see here).

It's also possible to tease out Balto-Slavic drift with formal statistics. I showed this indirectly in a recent blog post about Greek population structure (see here). In this post I'm going to demonstrate how to explicitly and formally test for Balto-Slavic drift both in ancient and present-day samples.

To do this we need to find stats that basically split Baltic and Slavic speakers from other Europeans, such as f4(Outgroup,Test;Bell_Beaker_NDL,Baltic_LVA_BA). In this f4-stat, Baltic_LVA_BA is the ancient reference population with an unusually high level of Balto-Slavic drift, while Bell_Beaker_NDL is a fairly similar population overall in terms of ancient ancestry components, but with practically zero Balto-Slavic drift.

Note that the statistics with the most significant Z scores (>3) involve populations that speak Baltic or Slavic languages, or their neighbors who plausibly harbor significant Baltic and/or Slavic ancestry. Among the ancient, mostly Scandinavian, populations (from Margaryan et al. 2020 and marked with the VK2020 prefix), significant Balto-Slavic drift only appears in the more easterly and/or later groups from the Viking Age (VA).

Unfortunately, one of the problems with this analysis is that Baltic_LVA_BA and Bell_Beaker_NDL aren't identical in terms of their ancient ancestry proportions. For one, the latter has significantly more Neolithic farmer ancestry. No wonder then, that Greeks, who are mostly of early farmer stock, don't show a significant Z score, despite probably packing a significant amount of Balto-Slavic ancestry dating to the Middle Ages.

In the near future, as more ancient samples become available, it might be possible to find better reference populations for the job and create more accurate, finer-scaled tests.

See also...

Uralian genes

That old chestnut: Northeast vs Northwest Euros

The Uralic cline with kra001 - no projection this time

A whole lot of nonsense was posted online, often by people who should've known better, after I claimed that kra001 was a solid proxy for a proto-Uralic genome (see here).

For those of you who still don't get it, below are three Principal Component Analysis (PCA) plots featuring Uralic speakers and other present-day Eurasians. Kra001 is also there. These graphs are based on genotype data not reprocessed Global25 data. The relevant datasheet is available here.

Compared to my previous PCA with kra001, here I included a bigger range of East Eurasian populations to help mitigate the effects of extreme genetic drift in some of the Siberian groups, at least on the first few Principal Components (PCs). Moreover, kra001 wasn't projected onto PCs computed with modern-day samples, so he was free to influence the outcome of the PCA.

Note the east to west clines made up largely of Uralic speaking groups on the first two plots. These plots are based on PCs 1/2 and 1 /3, respectively. The third plot, based on PCs 1/4, is more complex and thus more difficult to interpret, but it also manages to isolate many of the Uralic populations from the others.

The Uralic-specific clines do intersect with the clines and clusters formed by the other linguistic groups. However, based on the three plots, the Yeniseian-speaking Kets are the only Asian group that can plausibly be confused for Uralic speakers.

Importantly, apart from the Kets, kra001 is the only Asian individual who shifts his position on all three plots as if he were a Uralic speaker. This might well be a coincidence, and we'll never know what language was spoken by kra001, but it does suggest to me that his genome is a solid proxy for a proto-Uralic genome.

See also...

First taste of Early Medieval DNA from the Ural region

The BOO people: earliest Uralic speakers in the ancient DNA record?

Fresh off the sledge

Finally, a proto-Uralic genome

Obviously, genes don't speak languages, people do. But sometimes it's possible to associate a linguistic group with a very specific genetic signature.

A while ago many of us in the blogosphere spotted an uncanny connection between the Uralic language family, Y-haplogroup N-L1026 and Nganasan-like genome-wide genetic ancestry.

As a result, we expected a Nganasan-like population rich in N-L1026 to eventually appear in the ancient DNA record, probably somewhere in Siberia and in burials from a likely proto-Uralic archeological culture. This hasn't happened yet, but we now have direct evidence that such a population must have existed somewhere deep in Siberia as early as the Bronze Age.

Kra001, whose genome was published recently along with Kilinc et al., belongs to a pre-N-L1026 lineage and, at least in terms of genome-wide genetic structure, could well be from a population directly ancestral to present-day Nganasans. Of course, the Nganasan language is part of the Samoyedic branch of Uralic.

Below is a series of Principal Component Analyses (PCA) featuring kra001. He's labeled RUS_Krasnoyarsk_BA, after the location and age of his burial. Note the obvious Uralic cline running across the plots. That is, from west to east. Kra001 is positioned at the end of this cline very close to a small cluster of Nganasans. To see interactive versions of the plots, paste the Global25 coordinates here into the relevant field here.

Admittedly, there's no way of knowing whether this individual spoke proto-Uralic or not. Indeed, he may have spoken something totally unrelated. The important point is that the very specific genetic signature shared by almost all present-day Uralic speakers, except perhaps Hungarians, is now finally represented in the ancient DNA record. And I can reveal to you that we'll soon be seeing many more ancients very similar to kra001 in upcoming papers.

See also...

The Uralic cline with kra001 - no projection this time

Fresh off the sledge

Looking forward to a post-Covid world

I was hoping that the Covid-19 pandemic wouldn't have an immediate impact on the publication of ancient DNA papers and new data, but considering how much things have slowed down in this respect, it seems that I was fooling myself.

So let's take a break until early next year, and then see what happens.

Trust me, we've got a lot to look forward to in the post-Covid-19 world. Based on what I've heard from various sources, here are some predictions about what we might see:

- the search for the Proto-Indo-European homeland will shift west to the North Pontic steppe

- on the other hand, the search for the Proto-Uralic homeland will move deep into Siberia

- the key role of the Single Grave (westernmost Corded Ware) culture in the population history of Western Europe will finally get some attention

- following on from the above, Y-haplogroup R1b-L51 will be revealed as a Single Grave marker

- the idea that the Pontic-Caspian steppe was colonized by migrants from Mesopotamia during the Bronze Age will be forgotten, and, ironically, we'll instead learn that there was a significant influx of steppe ancestry into ancient Mesopotamia

- Old Kingdom Egyptians will come out less Sub-Saharan African than present-day Egyptians.

I probably shouldn't blab everything out, so that's all you're getting from me for now. You'll just have to wait for the rest until next year, or perhaps even the year after that.

See also...