Armenian Highland population prehistory

A new preprint at bioRxiv claims that some sort of large-scale population movement resulted in the spread of Sardinian-like ancestry into both the Armenian Highland and East Africa during or just after the Middle-Late Bronze Age. See Hovhannisyan et al. here.

In all seriousness, my suggestion is that the authors should familiarize themselves with the scientific concept of the sanity check and then try again.

For what it's worth, here's a brief outline of the population history of the Armenian Highland based on what I've learned about the topic from ancient DNA in recent years:

- overall, the Neolithic populations of the Armenian Highland were surely very similar to the Caucasus_lowlands_LN samples from what is now Azerbaijan from the recent Skourtanioti et al. paper (see here)

- Chalcolithic era migrations from the Pontic-Caspian steppe and/or the North Caucasus introduced steppe ancestry to the Armenian Highland, bringing at least some of its populations closer genetically to those of Eastern Europe (a somewhat outdated but still useful blog post about this subject is found here)

- population expansions during the Early Bronze Age associated with the Kura-Araxes cultural phenomenon, which may have originated in what is now Armenia, resulted in a resurgence of indigenous Caucasus hunter-gatherer (CHG) ancestry across the Caucasus, as well as its spread to many other parts of West Asia (see here)

- another significant pulse of Eastern European admixture affected the Armenian Highland during the Middle-Late Bronze Age and Early Iron Age (see here)

- it's not yet completely clear what happened in the Armenian Highland during the Iron Age in terms of significant genetic shifts, due to the lack of ancient human samples from the region dating to this period, but it's still possible that the speakers of proto-Armenian arrived there from the Balkans at this time

- the present-day Armenian gene pool is the result of the processes described above, as well as later events, such as those associated with the Urartian and Ottoman Empires.

Indeed, it's probably not a coincidence that present-day Armenians cluster more or less between the prehistoric populations from the Armenian Highland and surrounds in the Principal Component Analysis (PCA) below.

To see a more detailed and interactive version of the plot, copy paste the data from the text file here into the relevant field at the Vahaduo Globabl25 PCA Views here.

Citation...

Hovhannisyan et al., AN ADMIXTURE SIGNAL IN ARMENIANS AROUND THE END OF THE BRONZE AGE REVEALS WIDESPREAD POPULATION MOVEMENT ACROSS THE MIDDLE EAST, bioRxiv, Posted June 24, 2020, doi: https://doi.org/10.1101/2020.06.24.168781

See also...

Armenian confirmation bias

Perhaps a hint of things to come

Understanding the Eneolithic steppe

Like three peas in a pod

One of the most interesting questions still waiting to be answered by ancient DNA is where exactly did the ancestors of the present-day European and South Asian bearers of Y-haplogroup R1a part their ways? Indeed, the answer to this question is likely to be informative about the place and time of the split between the Balto-Slavic and Indo-Iranian language families.

I was doing some reading today and discovered that the peoples associated with the Bronze Age Fatyanovo-Balanovo and Unetice archeological cultures shared strikingly similar metalwork, despite being separated by well over two thousand kilometers of forest and steppe. Apparently, this similarity is especially pronounced in the metalwork of the Unetice culture from what is now Slovakia (see Ancient Metallurgy in the USSR: The Early Metal Age, page 136).

S11953 is currently the only sample from Slovakia associated with the Unetice culture (Sirak et al. 2020). There are no Fatyanovo-Balanovo samples available yet. However, as far as I can tell, I0432 from Samara, Russia, should be a decent stand in (Mathieson et al. 2015).

Of course, both S11953 and I0432 belong to Y-haplogroup R1a. Moreover, S11953 belongs to a typically Balto-Slavic subclade of R1a, while I0432 belongs to a closely related subclade that is dominant nowadays among the Indo-Iranian speakers of Asia.

S11953 is younger than I0432, but this doesn't necessarily mean that his ancestors arrived in East Central Europe from deep in Russia during the Bronze Age. Indeed, the opposite is more likely to be true. That is, I0432 is probably the recent decedent of migrants from somewhere near the North Carpathians, because he shows elevated European Neolithic farmer ancestry compared to earlier ancients from the Samara region (see here).

Below is a Principal Component Analysis (PCA) showing how S11953 and I0432 compare to each other in the context of ancient West Eurasian genetic variation. Obviously, they're sitting in the same part of the plot, which suggests that they harbor very similar ratios of ancient genetic components and probably share relatively recent ancestry. The relevant PCA datasheet is available here.

I've also highlighted myself, Davidski, on the plot. That's because I share the same Balto-Slavic-specific subclade of R1a with S11953 and, in terms of overall ancestry, I'm similar to both S11953 and I0432. Moreover, I'm the speaker of Polish, which is a Balto-Slavic language. What are the chances that we're dealing here with a remarkable string of coincidences? Indeed, was the North Carpathian region perhaps the homeland of the language ancestral to both Balto-Slavic and Indo-Iranian?

However, please note that there's nothing unusual or remarkable about my ancestry. The vast majority of people of Central, Eastern and Northern European origin - that is, mostly the speakers of Balto-Slavic, Germanic and Celtic languages - would also land in this part of the plot.

See also...

On the doorstep of India

Y-haplogroup R1a and mental health

The mystery of the Sintashta people

Maykop ancestry in Copper Age Arslantepe

At least four individuals from the Late Chalcolithic (LC) burial site of Arslantepe show ancestry typical of the population associated with the contemporaneous Maykop culture in the North Caucasus. They are ART018, ART020, ART027 and ART039 from the recent Skourtanioti et al. paper. I've labeled them TUR_Arslantepe_LC_Maykop in my qpAdm mixture model below:

TUR_Arslantepe_LC_Maykop
RUS_Maykop_Novosvobodnaya 0.318±0.041
TUR_Arslantepe_LC 0.682±0.041
chisq 9.969
tail prob 0.533159
Full output

Considering the tight statistical fit, I think it's even possible that some of these people harbor direct ancestry from Maykop Novosvobodnaya. Here's a Principal Component Analysis (PCA) showing why my qpAdm model works so well. It was produced with the data in the text file here and the Vahaduo PCA tools here.

Moreover, one of the Arslantepe males, ART038, belongs to Y-haplogroup R1b-V1636 (R1b1a2). This is clearly a marker of paternal steppe ancestry, because it's been reported in two Eneolithic samples from the southernmost part of the Pontic-Caspian steppe near the North Caucasus foothills (see here). These individuals are dated to ~4,200 calBCE, so they lived about a thousand years earlier than ART038.

ART038 probably lacks steppe and Maykop-related ancestries on his autosomes. Nevertheless, my point about his Y-haplogroup stands, because autosomal admixture can be bred out and disappear completely within a couple hundred years, or about 6 to 8 generations.

Interestingly, Skourtanioti et al. argued against the possibility of significant steppe and Maykop-related ancestries in the Arslantepe LC samples. They also didn't see R1b-V1636 as an obvious signal of paternal steppe ancestry. I find this very puzzling indeed, because to me it seems way off the mark.

See also...

Perhaps a hint of things to come

An early Mitanni?

Perhaps a hint of things to come

It's still a mystery how the Hittites and other Anatolian speakers ended up in the Near East. However, the leading theory is that their ancestors migrated from the steppes of Eastern Europe to western Anatolia via the Balkans sometime during the Copper Age.

Consider the qpAdm mixture models below, made possible thanks to some of the ancient samples published recently along with Skourtanioti et al. 2020. The key ancients are described in a text file available here.

TUR_Barcin_C
AZE_Caucasus_lowlands_LN 0.471±0.094
RUS_Vonyuchka_En 0.148±0.040
TUR_Barcin_N 0.381±0.069
chisq 12.874
tail prob 0.116261
Full output

TUR_Barcin_C
RUS_Vonyuchka_En 0.107±0.029
TUR_Buyukkaya_EC 0.893±0.029
chisq 12.107
tail prob 0.207331
Full output

I'd say it's quite clear now that TUR_Barcin_C harbors minor ancestry from the Pontic-Caspian (PC) steppe. The reason this isn't widely accepted yet is because demonstrating it convincingly hasn't been possible without a proximate Anatolian ancestry source for TUR_Barcin_C, precisely like TUR_Buyukkaya_EC.

Admittedly, though, the statistical fits in my models aren't all that great. I suspect the problem lies with RUS_Vonyuchka_En, which is likely to be a rather poor stand in for the people who brought steppe ancestry, and possibly early Anatolian speech, to western Anatolia.

So let's see what happens when I try a more proximate reference for the steppe ancestry in TUR_Barcin_C. How about Yamnaya_BGR, an individual of mixed Balkan and steppe origin from what is now Bulgaria?

TUR_Barcin_C
AZE_Caucasus_lowlands_LN 0.518±0.075
TUR_Barcin_N 0.203±0.056
Yamnaya_BGR 0.279±0.067
chisq 10.602
tail prob 0.225269
Full output

TUR_Barcin_C
TUR_Buyukkaya_EC 0.749±0.058
Yamnaya_BGR 0.251±0.058
chisq 9.687
tail prob 0.376414
Full output

That's a little better. Unfortunately, the problem now is that the models are anachronistic, because TUR_Barcin_C is about a thousand years older than Yamnaya_BGR. Clearly, we need more Copper Age samples from the western edge of the PC steppe, the eastern Balkans, and especially northwestern Anatolia.

The Principal Component Analysis (PCA) below effectively illustrates why my qpAdm models work. It was produced with Global25 data using the Vahaduo PCA tools freely available here. Note that TUR_Barcin_C is shifted away from the essentially perfect cline formed by AZE_Caucasus_lowlands_LN, TUR_Barcin_N and TUR_Buyukkaya_EC towards samples from ancient Eastern Europe, including Yamnaya_BGR.

See also...

Steppe invaders in the Bronze Age Balkans

An early Mitanni?

I've updated my Global25 datasheets with most of the ancients from the new Skourtanioti et al. paper. Here's a Principal Component Analysis (PCA) based on the data. It was produced with the Vahaduo PCA tools freely available here and the text file here.

Note that one of the Bronze Age females from Alalakh, labeled ALA019, appears to have ancestry from Turan and the Eurasian steppe. She may well have been a Mitanni of Indo-Aryan origin.

Interestingly, a Copper Age male from Arslantepe, ART038, belongs to Y-haplogroup R1b1a2 aka R1b-V1636. This is an unusual find, because R1b hasn't yet been reported in any Copper Age or earlier samples from outside of Europe and the Eurasian steppe.

As far as I can tell, this individual doesn't harbor any genome-wide ancestry from north of the Caucasus. However, R1b-V1636 is a rare lineage that is first attested in Eneolithic samples from the North Caucasus Piedmont steppe, so ART038's Y-chromosome might be the first evidence of the presence of steppe ancestry in Copper Age Anatolia.

I've also added most of the ancients from the new Agranat-Tamir et al. paper to the Gobal25 datasheets. The PCA below is based on the text file available here.

The Megiddo samples include a trio of interesting outliers dated to 1600-1500 BCE with significant ancestry from the steppe. One of these individuals is a male, I2189, who belongs to Y-haplogroup R and probably R1a. So he might also be of Indo-Aryan origin.

Another Megiddo male, S10768, belongs to R1b-M269 and probably shows a few per cent of steppe ancestry. I've already discussed how R1b and steppe ancestry may have ended up in the Bronze Age Near East in a couple of my previous posts:

R1b-M269 in the Bronze Age Levant

How did steppe ancestry spread into the Biblical-era Levant?

R-V1636: Eneolithic steppe > Kura-Araxes?

Understanding the Eneolithic steppe

Archeologist David Anthony has teamed up with Harvard's David Reich Lab to work on a paper about the Eneolithic period on the Pontic-Caspian steppe.

A couple of other labs are also preparing papers on similar topics, and they've already sequenced and analyzed many of their ancient samples (for instance, see here). However, I don't have a clue when these papers will be published. My guess is that we'll have to wait a year or so.

Needless to say, knowing what happened on the Pontic-Caspian (PC) steppe and surrounds during the Eneolithic is crucial to understanding the origins of the present-day European gene pool. It's also likely to be highly relevant to the debate about the location of the Proto-Indo-European (PIE) homeland.

In this blog post I'll explain what I've learned about the Eneolithic peoples of the PC steppe based on already published data.

If we ignore Steppe Maykop samples, the currently available Eneolithic individuals from the eastern part of the PC steppe form an essentially perfect cline in my Principal Component Analysis (PCA) of ancient West Eurasian genetic variation.

The cline runs from the Mesolithic hunter-fishers of the Eastern European forest zone to those of the Eneolithic sites of Progress 2 and Vonyuchka in the North Caucasus foothills. Let's call this the Khvalynsk cline, because three of the samples are from a burial site in the Volga River valley associated with the Khvalynsk culture. The relevant datasheet is available here.

The reason that these samples form the cline is because they carry different ratios of admixture related to Caucasus hunter-gatherers (CHG) from what is now Georgia. Moreover, the Khvalynsk individuals appear to be relatively recent mixtures between sources rich and poor in this type of ancestry.

I also marked a Maykop cline on the plot. This cline is made up of individuals associated with the Maykop and Steppe Maykop cultures from the Caucasus Mountains and nearby parts of the PC steppe, respectively. The Maykop culture is dated to the Early Bronze Age (EBA) period, but the PC steppe was still part of the Eneolithic world at the time.

The Maykop cline is more complicated than the Khvalynsk cline, because some of the Maykop individuals carry genetic components that the others lack. These genetic components are closely related to the aforementioned CHG, as well as Anatolian Neolithic farmers (ANF) and Western Siberian hunter-gatherers (WSHG).

Note that the two clines intersect, but this isn't because any of the Khvalynsk cline samples harbor Maykop-related ancestry. It's largely because the Steppe Maykop individuals carry high levels of Vonyuchka-related ancestry.

So unless we're dealing here with a remarkable string of coincidences, then the Vonyuchka hunter-fisher must be a decent proxy for the people who spread significant levels of CHG-related ancestry north of the Caucasus.

The important question, therefore, is where and when exactly did this population form? And it's a question that the authors of the aforementioned upcoming papers should be aiming to answer comprehensively.

In my view, it was the result of interactions between the hunter-fishers of the North Caucasus and the southernmost parts of the PC steppe during the Neolithic period, perhaps around 6,000 BCE, just before significant ANF-related ancestry spread across the Caucasus during the Eneolithic. That's because the Progress 2/Vonyuchka samples lack ANF-related ancestry, or at least an obvious signal of it, and are dated to ~4,200 BCE. And when I say Neolithic in this context, I don't mean the Near Eastern type of Neolithic with well developed farming, but rather the local type of Neolithic still based on hunting and fishing.

Now, obviously, the people of the Corded Ware and Yamnaya cultures were the children of the Eneolithic PC steppe. So you might be wondering how they fit into all of this. I still don't know, and apparently neither do the scientists at Harvard (see here). However, I'd say that the Maykop cline isn't relevant to this question. The Khvalynsk cline might be relevant, but even if it is, this doesn't necessarily mean that the Yamnaya people are by and large derived from the Khvalynsk people.

Here's the same PCA plot as above, but this time with early Corded Ware and Yamnaya samples also highlighted. Note that, apart from a few outliers, they form a rather tight cluster that is shifted slightly away from the Khvalynsk cline, but probably not in the direction of the Maykop cline.

A couple of the Yamnaya outliers are shifted towards the "eastern" end of the Khvalynsk cline, and thus near the Progress 2/Vonyuchka samples. This isn't surprising because these Yamnaya individuals are from burial sites close to the North Caucasus and probably harbor significant levels of local ancestry.

The most extreme Yamnaya outlier, from a site in what is now Ukraine, is clearly shifted towards the Maykop cline, and even towards the Caucasus Maykop cluster. However, this is a female with no grave goods and she may have been a foreign bride or captive, possibly from a late Maykop settlement. It's also possible that her 3095-2915 calBCE dating is wrong.

I'm pretty sure that when we find out why the Yamnaya cluster is so deliberately shifted away from the Khvalynsk cline, we'll also discover how the early Corded Ware and Yamnaya populations formed. For now, I strongly suspect that this has something to do with gene flow from the western edge of the PC steppe and the ethnogenesis of the Sredny Stog culture, which was located just west of the Khvalynsk culture.

By and large, the PC steppe is still seen by historical linguists and archeologists as the most sensible place to put the PIE homeland.

However, a theory that the PIE homeland was located somewhere south of the Caucasus, and that instead the PC steppe was the late or nuclear PIE dispersal point, has gained popularity in recent years, largely thanks to the apparent lack of PC steppe ancestry in a handful of samples from Hittite era Anatolia. In this scheme, the Maykop culture took PIE into Eastern Europe and the Yamnaya culture subsequently spread late/nuclear PIE from the PC steppe, while Proto-Anatolian, the ancestor of Hittite, was introduced into Anatolia from the east along with Maykop-related ancestry.

This is possible, in the sense that almost anything is possible, but it doesn't strike me as the most parsimonious interpretation of the facts.

Even before ancient DNA, it was known that the Maykop culture colonized parts of the PC steppe, at least for a short time, and probably had contacts with the Yamnaya people and/or their antecedents. But it was generally seen as the vector for Caucasian and other non-Indo-European influences in PIE.

Moreover, not only were the Maykop and Yamnaya populations of fundamentally different genetic origins, but apparently the Yamnaya people didn't absorb any perceptible Maykop ancestry as they expanded into the North Caucasus region at the tail end of the Maykop period.

That's really difficult to explain if we assume that these groups were close linguistic relatives, and much easier to reconcile with the assumption that they were derived from different worlds culturally and linguistically.

Another important question is what happened to the Steppe Maykop people, because right now it looks like they vanished almost without a trace, essentially as if they were pushed out or even erased by the Yamnaya expansion. If they were indeed pushed out or erased, then it's likely that their language was as well.

As for the lack of PC steppe ancestry in Hittite era Anatolians, I honestly can't see this is as a significant obstacle to a PIE homeland on the steppe, especially if we consider that the most widely accepted Indo-European phylogenies show the Anatolian family as the most basal node.

In the opinion of the vast majority of experts, it's the most basal node because the Proto-Anatolian speakers were the first to leave the PIE homeland. And if they were indeed the first to leave the homeland, then why should we expect their descendants to harbor significant ancestry from the homeland? In my view, such an assumption would contradict the most widely accepted Indo-European phylogenies.

See also...

The Caucasus is a semipermeable barrier to gene flow

Aesch25

During the early 3rd Millennium BC much of Central and Northern Europe was being infiltrated by pioneer herders, often young men, from the east associated with the Corded Ware culture (CWC).

In some important ways, this expansion may have been very similar to the European colonization of the more remote parts of the Americas during the 16th and 17th centuries.

For instance, the European newcomers weren't always able to dominate the indigenous peoples, and, sometimes, instead of trying to impose their culture on them, they accepted theirs.

I suspect that Aesch25, an ancient sample from the recent Furtwängler et al. paper on the social and genetic structure of the prehistoric populations of the Swiss Plateau, represents a similar case.

Aesch25 wasn't buried with grave goods so he wasn't given a cultural context in the said paper. However, dated to 2864-2501 calBC, he's the earliest individual in this part of Europe with the originally Eastern European Y-haplogroup R1b-M269 and a CWC-like genome-wide genetic structure.

Indeed, the other fourteen samples from the same burial site, dated to more or less the same period as Aesch25, are overwhelmingly of local Neolithic farmer origin.

In any case, irrespective of his cultural affiliation and life story, Aesch25 represents an important data point in the search for the homeland of the so called Bell Beakers who spread across much of Europe during the Copper Age. That's because most Bell Beaker males belong to R1b-M269 and are very similar to Aesch25 in terms of overall genetic structure, apart from an excess of Neolithic farmer ancestry.

My view is that the Bell Beakers were an offshoot of the Single Grave culture (SGC), the westernmost variant of CWC. Of course, the SGC was centered on what is now northwestern Germany and surrounds, and didn't reach into the Swiss Plateau. However, in all likelihood it was founded by men closely related to Aesch25.

Below is a Principal Component Analysis (PCA) based on Global25 data featuring Aesch25 and several other individuals from the Furtwängler et al. paper. To view an interactive version of the plot, copy paste the data from the text file here into the relevant field here, then press Add to PCA. Also, you should copy paste each population separately to make sure that they don't form one grouping in the PCA key.

Aesch25 can easily pass for a CWC individual from what is now Germany (DEU_CWC_LN). On the other hand, the CWC samples from the Swiss Plateau (CHE_CWC_LN) are clearly shifted "south" relative to the German CWC cluster, which suggests that they harbor more Neolithic farmer ancestry. Indeed, they all belong to Y-haplogroup I2, which is especially closely associated with Middle Neolithic European farmers.

MX265, from Singen in southwest Germany, is the only sample in the Furtwängler et al. dataset that belongs to Y-haplogroup R1a. This is a somewhat unexpected outcome, because R1a is, overall, the most common Y-haplogroup in CWC males (see here).

Another surprise is that this individual is dated to just 763-431 calBC, which is a period that overlaps with the Hallstatt and La Tene cultures in Central Europe. Considering that these cultures are often associated with early Celts, was this person perhaps the speaker of a long lost Celtic language?

See also...

Single Grave > Bell Beakers

Dutch Beakers: like no other Beakers

Hungarian Yamnaya > Bell Beakers?

Hungarian Yamnaya predictions

The Battle Axe people came from the steppe

Ancient DNA vs Ex Oriente Lux

In recent years you may have read academic papers, books and press articles claiming that the Early Bronze Age Yamnaya culture of the Pontic-Caspian steppe was founded by migrants from the Caucasus, Mesopotamia or even Central Asia.

Of course, none of this is true.

The Yamnaya herders and closely related groups, such as the people associated with the Corded Ware culture, expanded from the steppe between the Black and Caspian seas, and, thanks to ancient DNA, it's now certain that they were overwhelmingly derived from a population that had existed in this region since at least the mid-5th millennium BCE (see here).

So rather than being culturally advanced colonists from some Near Eastern civilization, the ancestors of the Yamnaya herders were a relatively primitive local people who still largely relied on hunting and fishing for their subsistence. They also sometimes buried their dead with flint blades and adzes, but hardly ever with metal objects, despite living in the Eneolithic epoch or the Copper Age.

As far as I know, this group doesn't have a specific name. But in recent scientific literature it's referred to as Eneolithic steppe, so let's use that.

It's not yet clear how the Yamnaya people became pastoralists. Some scholars believe that they were basically an offshoot of the cattle herding Maykop culture of the North Caucasus. However, the obvious problem with this idea is that the Yamnaya and Maykop populations probably didn't share any recent ancestry. In fact, ancient DNA shows that the former wasn't derived from the latter in any important or even discernible way (see here).

On the other hand, Yamnaya samples do harbor a subtle signal of recent gene flow from the west that appears to be most closely associated with Middle to Late Neolithic European agropastoralists (see here). Therefore, it's possible that herding was adopted by the ancestors of the Yamnaya people as a result of their sporadic contacts with populations living on the western edge of the Pontic-Caspian steppe.

Eneolithic steppe is currently represented by just three samples in the ancient DNA record, and all of these individuals are from sites on the North Caucasus Piedmont steppe (two from Progress 2 and one from Vonyuchka 1).

As a result, it might be tempting to argue that cultural, if not genetic, impulses from the Caucasus did play an important role in the formation of the Yamnaya and related peoples. However, it's important to note that the North Caucasus Piedmont steppe was the southern periphery of Eneolithic steppe territory.

Below is a map of Eneolithic steppe burial sites featured in recent scientific literature. It's based on data from Gresky et al. 2016, a paper that focused on a specific and complex type of cranial surgery or trepanation often practiced by groups associated with this archeological culture (see here).

Incredibly, one of the skeletons from Vertoletnoe pole has been radiocarbon dated to the mid-6th millennium BCE. My suspicion, however, is that this result was blown out by the so called reservoir effect (see here). In any case, the academic consensus seems to be that the roots of Eneolithic steppe should be sought in the Lower Don region, rather than in the Caucasus foothills (see page 36 here).

Considering that nine Eneolithic steppe skulls from the Lower Don were analyzed by Gresky et al., I'd say it's only a matter of time before we see the publication of genome-wide data for at least of couple of these samples. Indeed, the paper's lead author is from the Deutsches Archäologisches Institut, which is currently involved in a major archaeogenetic project on the ancient Caucasus and surrounds. Unfortunately, the study is scheduled to be completed in about four years (see here).

But whatever happens, the story of Eneolithic steppe deserves to be investigated in as much detail as possible, because it obviously had a profound impact on Europe and its people.

In my estimation, at least a third of the ancestry of present-day Northern Europeans, all the way from Ireland to the Ural Mountains in Russia, is ultimately derived from Eneolithic steppe groups. It's also possible that R1a-M417 and R1b-L51, the two most frequent Y-chromosome haplogroups in European males today, derive from a couple of Eneolithic steppe founders. If so, that's a very impressive effort for such an obscure archeological culture from what is generally regarded as a peripheral part of Europe.

See also...

Did Caucasus hunter-gatherers ever live in what is now Iran?

Nope, they only lived in the Caucasus Mountains. See that's probably why they're called Caucasus hunter-gatherers, or CHG for short.

But what about the hunter-gatherers from the Belt and Hotu caves in northern Iran, you might ask? Well, what about them? They're not CHG, nor are they significantly more CHG-like than the early farmers of the Zagros Mountains.

To illustrate the point, below are a couple of TreeMix graphs. I'd say they're rather straightforward and self-explanatory.

However, please note that I combined the Belt and Hotu individuals into one sample to help keep the marker count at over 100K. Also keep in mind that CHG is represented by Kotias_HG.

See also...

A final note for the year

A note on Steppe Maykop

Did South Caspian hunter-fishers really migrate to Eastern Europe?

Graphing the truth

I haven't used TreeMix since qpGraph became freely available for Linux. Among other things, the latter offers greater control, reproducibility and transparency.

However, I'd say that in its current form qpGraph is not the most objective way to analyze data. That's because if you're really good with it, and you want a graph to work, then often you can make it work by tweaking whatever it is that needs to be tweaked.

It's not possible to do a lot of tweaking with TreeMix. Indeed, once the user picks the samples for the TreeMix run, the rest of the process can be totally unsupervised, and thus free from human interference. Obviously, that's not a guarantee of accuracy, but it can be useful.

I feel I need to run more unsupervised analyses, especially when exploring new data. So to that end, I've dusted off TreeMix and will be using it regularly again.

There's been some talk lately online about migrations from Central Asia giving rise to the Eneolithic populations of the North Caucasus Piedmont steppe. In my opinion, that sounds like nonsense. But let's see what TreeMix has to say on the matter. In the graphs below look for the samples labeled Progress_En and Vonyuchka_En, respectively.

As far as I can tell, both of these graphs essentially corroborate the results from my recent Principal Component Analyses (PCA) with many of the same ancients (see here). In other words, Progress_En and Vonyuchka_En can be described as mixtures of populations closely related to the hunter-gatherers of the Caucasus on one hand, and those of Eastern Europe on the other. How does Central Asia fit into this, you might ask? It doesn't, unless you really want it to.

See also...

Did South Caspian hunter-fishers really migrate to Eastern Europe?