Domestic horses were introduced into Anatolia and Transcaucasia during the Bronze Age (Guimaraes et al. 2020)

Over at Science Advances at this LINK. This is a very important paper because it basically eliminates West Asia as the source of the modern domestic horse lineage, which leaves the Pontic-Caspian steppe in Eastern Europe as the only viable option.

It also corroborates the linguistic theory that the Proto-Indo-European homeland was located on the Pontic-Caspian steppe. That's because the horse is a key animal in the Proto-Indo-European pantheon, and it appears in Indo-European mythology in intricate roles. This suggests that the speakers of Proto-Indo-European weren't just familiar with the horse but also managed to domesticate it. From the paper:

Abstract: Despite the important roles that horses have played in human history, particularly in the spread of languages and cultures, and correspondingly intensive research on this topic, the origin of domestic horses remains elusive. Several domestication centers have been hypothesized, but most of these have been invalidated through recent paleogenetic studies. Anatolia is a region with an extended history of horse exploitation that has been considered a candidate for the origins of domestic horses but has never been subject to detailed investigation. Our paleogenetic study of pre- and protohistoric horses in Anatolia and the Caucasus, based on a diachronic sample from the early Neolithic to the Iron Age (~8000 to ~1000 BCE) that encompasses the presumed transition from wild to domestic horses (4000 to 3000 BCE), shows the rapid and large-scale introduction of domestic horses at the end of the third millennium BCE. Thus, our results argue strongly against autochthonous independent domestication of horses in Anatolia.

Guimaraes et al., Ancient DNA shows domestic horses were introduced in the southern Caucasus and Anatolia during the Bronze Age, Science Advances 16 Sep 2020: Vol. 6, no. 38, eabb0030, DOI: 10.1126/sciadv.abb0030

See also...

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

R1b-M269 in the Bronze Age Levant

The new Harvard genotype datasets that I blogged about recently include a couple of potentially very useful samples from the Levant dated to 1400-1100 BCE. Search for IDs I2062 and I1934 in the anno files here. They're both from an archeological paper about a Late Bronze Age (LBA) burial site in what is now Israel that was published back in 2017 (see here).

Surprisingly, individual I2062 is listed in the anno files as belonging to Y-haplogroup R1b1a1a2, which is also known as R1b-M269. The reason that this is a surprise to me is because R1b-M269 is closely associated with the Bronze Age expansions of pastoralists from the Pontic-Caspian steppe in Eastern Europe, and these expansions didn't impact the Levant in any direct or significant way.

The Y-haplogroup assignment may or may not be correct. Sometimes the Y-haplogroups in these sorts of datasheets are indeed wrong. Unfortunately, as far as I know, the BAM file for I2062 isn't available anywhere online, so I can't check whether he does really belong to R1b-M269. But, intriguingly, his autosomes do show a subtle signal of Yamnaya-related ancestry from the Pontic-Caspian steppe that is missing in earlier ancients from the Levant.

To characterize his genome-wide ancestry, I first ran a series of unsupervised and supervised analyses with the Global25/nMonte3 method (using this datasheet). For the sake of simplicity, I narrowed things down to the mixture models below based on three reference populations each. Levant_ISR_C is made up of Chalcolithic samples from Israel. The identities of the other reference sets should be obvious to most readers. If confused, feel free to ask for more details in the comments below.

Levant_ISR_MLBA:I2062
Levant_ISR_C,66.8
IRN_Seh_Gabi_C,27
Yamnaya_RUS_Samara,6.2
[1] distance%=1.8905

Levant_ISR_MLBA:I2062
Levant_ISR_C,66.2
Kura-Araxes_ARM_Kaps,30.2
Yamnaya_RUS_Samara,3.6
[1] distance%=2.0856

Levant_ISR_MLBA:I2062
Levant_ISR_C,67.8
Kura-Araxes_RUS_Velikent,31.8
Yamnaya_RUS_Samara,0.4
[1] distance%=2.1738

To further confirm the reliability of my models, I tested them with the formal statistics-based qpAdm software. As far as I can tell, the output from qpAdm looks very solid across the board.

Levant_ISR_MLBA_I2062
IRN_Seh_Gabi_C 0.193±0.052
Levant_ISR_C 0.710±0.038
Yamnaya_RUS_Samara 0.098±0.026
chisq 9.304
tail prob 0.67676
Full output

Levant_ISR_MLBA_I2062
Kura-Araxes_ARM_Kaps 0.249±0.076
Levant_ISR_C 0.681±0.051
Yamnaya_RUS_Samara 0.071±0.035
chisq 11.101
tail prob 0.52032
Full output

Levant_ISR_MLBA_I2062
Levant_ISR_C 0.661±0.042
Kura-Araxes_RUS_Velikent 0.339±0.042
chisq 7.979
tail prob 0.844942
Full output

Admittedly, even though I2062 can be modeled with Yamnaya-related admixture, he doesn't need to be. Indeed, his ratio of this type of ancestry varies significantly between the models, from around 10% to nothing. This appears to be dependent on the geography of the non-Levant and non-Yamnaya reference populations; the closer they are to the Pontic-Caspian steppe, the smaller the ratio of Yamnaya-related ancestry in I2062. I'd describe this as an artifact of the isolation-by-distance phenomenon, and it totally makese sense, but it prevents me from confirming beyond any doubt that I2062 does harbor genome-wide steppe ancestry. Unfortunately, individual I1934 doesn't offer enough data to be analyzed with the same methods.

Samples associated with the Kura-Araxes or Early Transcaucasian culture are particularly strong references for the eastern ancestry in I2062. This probably isn't a coincidence, and it might also explain his Y-haplogroup, because, at its maximum extent, the territory occupied by the Kura-Araxes culture stretched all the way from the Pontic-Caspian steppe to the southern Levant. The map below is from Wilkinson 2014.

See also...

Downloadable genotypes of present-day and ancient DNA data

Early chariot riders of Transcaucasia came from...

R-V1636: Eneolithic steppe > Kura-Araxes?

Early chariot riders of Transcaucasia came from...

I'm finding it increasingly difficult nowadays to fully appreciate all of the ancient DNA samples that are accumulating in my dataset. But it's not entirely my fault.

Among the hundreds of ancient samples published last year there was a couple of Middle Bronze Age (MBA) individuals from what is now Armenia labeled "Lchashen Metsamor" (see here). I wasn't planning to do much with these samples because, even after reading the Nature paper that they came with a couple times over, I didn't have a clue what they were about. But after some digging around, I now know that their people, those associated with the Lchashen Metsamor archeological culture, were among the earliest in Transcaucasia, and indeed the Near East, to use the revolutionary spoked-wheel horse chariot. How awesome is that?

The invention of the spoked-wheel chariot is generally credited to the Middle Bronze Age Sintashta culture of the Trans-Ural steppe in Central Asia, and its rapid spread is often associated with the early expansions of Indo-European languages deep into Asia. On the other hand, some have argued that this type of chariot was first developed in the Near East, and directly derived from solid-wheeled wagons pulled by donkeys.

It's now obvious, thanks to ancient DNA, that the Sintashta people were by and large migrants to Central Asia from somewhere in Eastern Europe, and that they didn't harbor any recent ancestry from the Near East. So if chariot technology spread into the steppes from the Near East, then it did so without any accompanying gene flow, which is possible but not entirely convincing. This begs the question of whether the Lchashen Metsamor population was of Sintashta-related origin, because if it was, then this would corroborate the consensus that spoked-wheel chariots were introduced into Transcaucasia from the steppes to the north.

Below is a Principal Component Analysis (PCA) of West Eurasian genetic variation. It does suggest that the Lchashen Metsamor pair (labeled Armenia_MBA_Lchashen), as well as most of the other currently available samples from what is now Armenia dating to the Middle to Late Bronze Age (MLBA), harbor some steppe ancestry. That's because they appear to form a cline between samples associated with the Sintashta and Kura-Araxes cultures. Of course, the Kura-Araxes culture was a major Early Bronze Age (EBA) archeological phenomenon centered on Transcaucasia and surrounds, so its population can be reasonably assumed to have formed the genetic base of most subsequent populations in the region. The relevant PCA datasheet is available here.

To investigate the possibility of Sintashta-related admixture in Lchashen Metsamor with formal methods, I ran a series of mixture models with the qpAdm software. Here are the three statistically most sound outcomes that I was able to come up with for Lchashen Metsamor:

Armenia_MBA_Lchashen
CWC_Kuyavia 0.183±0.036
Kura-Araxes_Kaps 0.817±0.036
chisq 13.941
tail prob 0.378021
Full output

Armenia_MBA_Lchashen
Balkans_BA_I2163 0.193±0.045
Kura-Araxes_Kaps 0.807±0.045
chisq 14.780
tail prob 0.321267
Full output

Armenia_MBA_Lchashen
Kura-Araxes_Kaps 0.788±0.043
Sintashta_MLBA 0.212±0.043
chisq 14.871
tail prob 0.315451
Full output

I sorted the output by "tail prob", but the fact that Sintashta_MLBA is in third place isn't a problem because the stats in all of these models are basically identical. Indeed, CWC_Kuyavia (Corded Ware culture samples from present-day Kuyavia, North-Central Poland) and Balkans_BA_I2163 (a Bronze Age singleton from what is now Bulgaria) are both very similar and probably closely related to each other and to the Sintashta samples.

Interestingly, and, I'd say, importantly, ancients from the steppe that are closest to Lchashen Metsamor in both space and time, but not particularly closely related to the Sintashta people, don't work too well as a mixture source in such models.

Armenia_MBA_Lchashen
Kubano-Tersk 0.184±0.046
Kura-Araxes_Kaps 0.816±0.046
chisq 22.179
tail prob 0.0526526
Full output

A couple of months ago I suggested that populations associated with the Early to Middle Bronze Age (EMBA) Catacomb culture were the vector for the spread of steppe ancestry into what is now Armenia during the MLBA (see here). After taking a closer look at the Lchashen Metsamor samples, I now think that the peoples of the Sintashta and related cultures were also important in this process. If so, they may have moved from the steppe into Transcaucasia both from the west via the Balkans and the east via Central Asia, and brought with them spoked-wheel chariots. I don't have a clue what language they spoke, but I'm guessing that it may have been something Indo-European.

See also...

The mystery of the Sintashta people

A potentially violent end to the Kura-Araxes Culture (Alizadeh et al. 2018)

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

Armenians vs Georgians

Armenians and Georgians are ethnic groups that live side by side in the south Caucasus, or Transcaucasia. By all accounts, they've both been there since prehistoric times and they're very similar in terms of overall genetic structure.

However, they speak languages from totally unrelated families: Indo-European and Kartvelian, respectively. How did this happen and might the answer lie in the small genetic differences that do exist between them?

To investigate this issue, I ran a series of qpAdm formal mixture models of present-day Armenians and Georgians using tens of ancient reference populations. To come up with as straightforward and meaningful results as possible, I constrained myself to two-way models. I then discarded the runs that produced "tail probs" under 0.1 and retained less than 400K SNPs. Only a handful of models passed muster, including these two:

Armenian
Mycenaeans_&_Empuries2 0.233±0.041
Kura-Araxes_Kaps 0.767±0.041
chisq 18.422
tail prob 0.142151
Full output

Georgian
Globular_Amphora 0.071±0.025
Kura-Araxes_Kaps 0.929±0.025
chisq 18.419
tail prob 0.142266
Full output

At the most basic level, the results suggest that both Armenians and Georgians are overwhelmingly derived from populations of Bronze Age Transcaucasia associated with the Kura-Araxes archeological culture, albeit with minor ancestries from somewhat different sources from the west. As far as I can see, when using more than 400K SNPs and a wide range and large number of outgroups (or right pops), neither Armenians nor Georgians can pass perfectly for any one ancient population in my dataset.

The best proxies for the minor but significant western ancestry in Armenians are Mycenaeans of the Bronze Age Aegean region and Greek colonists from Iron Age Iberia (Empuries2). Obviously, and perhaps importantly, these are both attested Indo-European-speaking groups. On the other hand, the very minor western ancestry in Georgians is best characterized as gene flow from Middle to Late Neolithic European farmers rich in indigenous European forager ancestry. It's practically impossible to say what language or languages these farmers spoke. How about something Kartvelian?

In any case, for me, the perplexing thing about present-day Armenians is that they harbor very little steppe ancestry. By and large, no more than a few per cent. Compare that to the currently available samples from what is now Armenia dating to the Middle to Late Bronze Age, which show ratios of steppe ancestry of up to 25%. For now, I'm guessing that what we're dealing with here is the classic bounce back of older ancestry layers that has been documented for different parts and periods of prehistoric Europe.

See also...

Early chariot drivers of Transcaucasia came from...

Catacomb > Armenia_MLBA

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

Catacomb > Armenia_MLBA

It's now clear, thanks to ancient DNA, that Transcaucasia and surrounds were affected by multiple, and at times significant, population movements from Eastern Europe during the Chalcolithic and Bronze Age periods. Based on the ancient samples from what is now Armenia, I'd say that this process peaked during the Middle Bronze Age. But who exactly were the people who perhaps swarmed south of the Caucasus at this time?

The most likely suspects are the various groups that occupied the southernmost parts of the Pontic-Caspian steppe throughout the Bronze Age. They were associated with the so called Catacomb, Kubano-Tersk and Yamnaya archeological cultures. Below is a Principal Component Analysis (PCA) that compares samples from these cultures with those from Middle to Late Bronze Age Armenia (labeled Armenia_MLBA). The relevant datasheet is available here.

Note that Armenia_MLBA forms a cline that appears to be stretching out towards the Catacomb, Kubano-Tersk, Yamnaya and other Bronze Age steppe groups, and this suggests that it harbors significant and probably recent steppe-related ancestry. But PCA plots based on just two dimensions of genetic variation can be misleading at times, so let's check this out with some formal mixture models using qpAdm.

Armenia_MLBA
Catacomb 0.234±0.028
Kura-Araxes_Kaps 0.766±0.028
chisq 10.723
tail prob 0.826248
Full output

Armenia_MLBA
Kubano-Tersk 0.254±0.030
Kura-Araxes_Kaps 0.746±0.030
chisq 13.535
tail prob 0.633284
Full output

Armenia_MLBA
Kura-Araxes_Kaps 0.768±0.028
Yamnaya_Kalmykia 0.232±0.028
chisq 14.454
tail prob 0.564954
Full output

Armenia_MLBA
Kura-Araxes_Kaps 0.762±0.029
Yamnaya_Caucasus 0.238±0.029
chisq 15.916
tail prob 0.458816
Full output

All of these models are statistically very sound, and even though I ranked the results by "tail prob", there's nothing in the output that clearly points to any one of the southern steppe groups as the obvious source of the steppe-related ancestry in Armenia_MLBA. But, interestingly, Catacomb tops the ranking, and it probably also makes the most sense based simply on Carbon-14 chronology. So, for now, I'm going with Catacomb.

I didn't get a chance yet to investigate this issue in detail with the Global25. Does it contradict the results from my PCA and qpAdm analyses? If anyone reading this would like to take a close look that'd be great. Feel free to post your findings in the comments below. And if the answer is indeed Catacomb, then what language did these Catacomb-derived migrants, or perhaps invaders, speak? If not proto-Armenian then what?

By the way, please be aware that the Kubano-Tersk samples in my analyses are the same individuals as those featured in Wang et al. 2019 under the label "North Caucasus".

See also...

Early chariot drivers of Transcaucasia came from...

Likely Yamnaya incursion(s) into Northwestern Iran

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