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 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 Nalchik surprise

If, like Iosif Lazaridis, you subscribe to the idea that the Yamnaya people carry early Anatolian farmer-related admixture that spread into Eastern Europe via the Caucasus, then I've got great news for you.

We now have a human sample from the Eneolithic site of Nalchik in the North Caucasus, labeled NL122, that packs well over a quarter of this type of ancestry (see here). Below is a quick G25/Vahaduo model to illustrate the point (please note that Turkey_N = early Anatolian farmers).

Target: Nalchik_Eneolithic:NL122
Distance: 2.1934% / 0.02193447
60.8 Russia_Steppe_Eneolithic
26.2 Turkey_N
13.0 Georgia_Kotias

On the other hand, if, again like Iosif Lazaridis, you subscribe to the idea that the Indo-European language spread into Eastern Europe via the Caucasus in association with this early Anatolian farmer-related admixture, then I've got terrible news for you.

That's because NL122 is apparently dated to a whopping 5197-4850 BCE (see here). This dating might be somewhat bloated, possibly due to what's known as the reservoir effect, because the Nalchik archeological site is generally carbon dated to 4840–4820 BCE.

However, even with the younger dating, this would still mean that early Anatolian farmer-related ancestry arrived in the North Caucasus, and thus in Eastern Europe, around 4,800 BCE at the latest. That's surprisingly early, and just too early to be relevant to any sort of Indo-European expansion from a necessarily even earlier Proto-Indo-Anatolian homeland somewhere south of the Caucasus.

This means that NL122 effectively debunks Iosif Lazaridis' Indo-Anatolian hypothesis. Unless, that is, Iosif can provide evidence for a more convoluted scenario, in which there are at least two early Anatolian farmer-related expansions into Eastern Europe via the Caucasus, and the expansion relevant to the arrival of Indo-European speech came well after 5,000 BCE.

I haven't done any detailed analyses of NL122 with formal stats and qpAdm. But my G25/Vahaduo runs suggest that it might be possible to model the ancestry of the Yamnaya people with around 10% admixture from a population similar to NL122.

Target: Russia_Samara_EBA_Yamnaya
Distance: 3.4123% / 0.03412328
72.6 Russia_Progress_Eneolithic
18.2 Ukraine_N
9.2 Nalchik_Eneolithic

However, I don't subscribe to the idea that the Yamnaya people carry early Anatolian farmer-related admixture that spread into Eastern Europe via the Caucasus (on top of what is already found in Progress Eneolithic). Based on basic logic and a wide range of my own analyses, I believe that they acquired this type of ancestry from early European farmers, probably associated with the Trypillia culture. For instance...

Target: Russia_Samara_EBA_Yamnaya
Distance: 3.2481% / 0.03248061
80.2 Russia_Progress_Eneolithic
13.6 Ukraine_Neolithic
6.2 Ukraine_VertebaCave_MLTrypillia
0.0 Nalchik_Eneolithic

Another way to show this is with a Principal Component Analysis (PCA) that highlights a Yamnaya cline made up of the Yamnaya, Steppe Eneolithic and Ukraine Neolithic samples. As you can see, dear reader, there's no special relationship between the Yamnaya cline and Nalchik_Eneolithic. The Yamnaya samples, which are sitting near the eastern end of the Yamnaya cline, instead seem to show a subtle shift towards the Trypillian farmers.

Indeed, I also don't exactly understand the recent infatuation among many academics, especially Iosif Lazaridis and his colleagues, with trying to put the Proto-Indo-Anatolian homeland somewhere south of the Caucasus. Considering all of the available multidisciplinary data, I'd say it still makes perfect sense to put it in the Sredny Stog culture of the North Pontic steppe, in what is now Ukraine.

Please note that all of the G25 coordinates used in my models and the PCA are available HERE.

See also...

The Caucasus is a semipermeable barrier to gene flow

The Caucasus is a semipermeable barrier to gene flow

The scientists at the David Reich Lab are a clever bunch. But they're not always on top of things, and this can be a problem.

For instance, they fail to understand that the Caucasus has effectively stymied human gene flow between Eastern Europe and West Asia through the ages. That is, the Caucasus is a semipermeable barrier to human gene flow.

Until they accept and understand this fact, they won't be able to accurately characterize the ancestry of the ancient human populations of the Pontic-Caspian (PC) steppe, including the Yamnaya people.

In turn, they also won't be able to locate the Indo-Anatolian homeland.

Now, the Caucasus isn't a barrier to gene flow because it's difficult to cross, and, indeed, many human populations have managed to cross it since the Upper Palaeolithic. As a result, the peoples of the North Caucasus are today genetically more similar to the populations of the Near East than Europe.

In fact, the clear genetic gap between most West Asian and Eastern European populations through the ages is actually caused by the extreme differentiation between the mountain ecology of the Caucasus and the steppe ecology of the PC steppe.

That is, the Caucasus is ecologically so different from the PC steppe that it has been practically impossible for human populations to make the transition from one to the other.

Indeed, it's important to understand that there's no reliable record of any prehistoric human population successfully making the transition from the mountain ecology to the steppe ecology in this part of the world.

In other words, contrary to claims by people like David Reich and David Anthony, there's no solid evidence of any significant prehistoric human migration from the Caucasus, or from south of the Caucasus, to Eastern Europe by hunter-gatherers, farmers or pastoralists.

But, you might ask, how on earth did the Yamnaya people get their significant Caucasus/Iranian-related admixture if not via a mass migration from the Caucasus and/or the Iranian Plateau, as is often argued by the above mentioned scholars and their many colleagues?

Well, obviously, the diffusion of alleles from one population to another can happen without migration. All that is needed is a contact zone between them.

The ancient DNA and archeological data currently available from the Caucasus and the PC steppe suggest to me that there was at least one such contact zone in this area bringing together the peoples of the mountain and steppe ecoregions. This allowed them to mix, probably gradually and over a long period of time, by and large without leaving their ecoregions.

Once the Caucasus alleles entered the steppe, they were spread around by local hunter-gatherers and pastoralists who were highly mobile and well adapted to the steppe ecology.

Someone should write a paper about this.

See also...

The Nalchik surprise

Understanding the Eneolithic steppe

Matters of geography

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 Yassitepe challenge

This is about the only successful qpAdm model that I can find for the pair of Early Bronze Age (EBA) females from Yassitepe, Turkey, using a decent set of outgroups and markers. I wouldn't take it too literally, but it does suggest a potentially significant level of European ancestry, including some steppe ancestry, in these Yassitepe individuals.

TUR_Aegean_Yassitepe_EBA
AZE_Caucasus_lowlands_LN 0.565±0.054
ROU_N 0.387±0.041
RUS_Progress_En 0.048±0.022
P-value 0.103248
Full output

If anyone reading this can find a better, more convincing solution then I'd love to see it. Feel free to share it in the comments below.

Obviously, both of the Yassitepe samples are from the recent Lazaridis, Alpaslan-Roodenberg et al. paper. Their EBA dating suggests that they might be relevant to the debate over the origins of Anatolian speakers, such as the Hittites and Luwians.

See also...

Dear Iosif, about that ~2%

The precursor of the Trojans

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...

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...