A map based mainly on Urartian and other inscriptions.

A map based mainly on Urartian and other inscriptions, from Grekyan (2023), The Urartian Onomasticon: A Prosopographic Study.

Ten Years Since the First Yamnaya Ancient DNA

Ten Years Since the First Yamnaya Ancient DNA

Ten years ago, in 2015, the first ancient DNA samples from the Yamnaya archaeological culture were published in the study by Haak et al. The discovery of R1b-Z2103 in Yamnaya caused a major sensation, as many commentators at the time expected to find R1a, whose correlation with the spread of Indo-European languages seemed self-evident.

Last year, Lazaridis et al. (2024) largely clarified the origins of Yamnaya autosomes. However, the precise origin of its main Y-DNA lineage, Z2103, remains unresolved. Yediay (2024) addressed this issue (see picture 2), noting that virtually all known cases of Z2103 found in the Eurasian steppe belong to the Z2106+ branch, while all currently known cases of basal Z2103* are found in the ancient South Caucasus.

How can this apparent contradiction be explained?

At the current stage of research, we know that Z2103 experienced a dramatic founder effect during the spread of the Yamnaya culture and was practically absent from the Pontic–Caspian steppe before the emergence of that culture around 3300 BCE. As a result, 41 out of 51 Y-DNA samples extracted from Yamnaya individuals are Z2103-positive (more than 80%). Therefore, the original homeland of Z2103 must have been close to the region where the Yamnaya culture first emerged.

Yamnaya autosomes are largely derived from the preceding CLV (Caucasus–Lower Volga) cline. The genetically closest known sample comes from near the Azov region (Krivianskiy). However, in addition to CLV ancestry, Yamnaya populations also carried extra Ukraine Neolithic Hunter-Gatherer (UNHG) ancestry and some Caucasian farmer–related ancestry. These two components significantly narrow down the possible homeland.

Previously, an infiltration of UNHG ancestry into the Krasnodar region of Russia has been observed, while the additional Caucasian farmer–related ancestry was also present in the same region. Based on this evidence, it is reasonable to assume that Z2103 began its expansion somewhere between the Azov and Caucasus regions.

Such a homeland helps explain several observations:

• The reason why all three basal Z2103* branches (Y4362, L584, PF331) are found in West Asia. Their proximity to the Caucasus likely facilitated southward migrations.

• Z2106 may also have formed near the Caucasus. Its main subbranch, R-Y503415, has been identified both south of the Caucasus and north of it.

• An initial spread from west to east would also explain why the Afanasievo culture appears almost simultaneously with Yamnaya in the Altai region. Most Afanasievo males belong to Z2108, a rare subclade today found in China. Some Z2108 lineages also remained in the Pontic–Caspian steppe, where the subclade KMS67 later expanded.

• Finally, after filling the Pontic–Caspian steppe with their settlements, Yamnaya groups moved into the Balkans, where the subbranch Z2110 is found. However, in the Balkans the most successful founder effect appears to have occurred in J2b2a-L283, another minor Yamnaya lineage originating from South Caucasian farmers.

Although this principle is not universal, a potential homeland should also be associated with a diversity of related archaeological cultures. Near the Krasnodar region we find two other cultures related to Yamnaya: Novotitorovka and the North Caucasus culture. The Novotitorovka culture is particularly noteworthy because it yielded numerous wagons similar to those discovered in Lchashen in Armenia. This further raises the possibility that the region between the Azov area and the North Caucasus is precisely where we should expect to find Y-DNA lineages ancestral to those later found in West Asia, including the still elusive I2a2b.

Moesia and the Question of Its Namee

Moesia and the Question of Its Name

Moesia was a province of the Roman Empire. It was located in what is now Bulgaria, Serbia, and partly Romania, extending in the west as far as the borders of modern Albania.

Its name has often been compared with that of the Mysians, a people who lived in the northwestern region of Anatolia. The presence of a similar name in the Balkans has sometimes been used as an argument for the migration of the Mushki from that region.

However, Boteva questions the origin of the name Moesia and argues that it was likely a fabrication of the Roman Imperial period. No known source mentions this term in connection with that region before the 1st century AD. According to this view, the name itself was derived from Mysia and written as Moesia in order to conform to older Latin orthographic conventions.

The reason the Romans began using this term for the region was probably ideological and political. Western Moesia was largely inhabited by the Dardani, a tribal name that played an important role in Roman epic traditions concerning Trojan origins. In these traditions, the Roman founding ancestors were linked to the Dardani. However, when the Romans conquered the Balkans, they came into conflict with the actual Dardani people. In Roman sources, these Dardani were portrayed as barbarians attacking Roman territories, creating a contradictory narrative: in one case the Dardani were noble ancestors, while in another they appeared as hostile enemies.

To avoid referring to their new enemies by the prestigious name Dardani, Roman authorities may have chosen to designate their lands with a new name. Thus, the term Musia or Moesia was adopted, borrowing a name from Anatolia. In this way, the name Dardani became obscured and gradually faded from use in reference to the region.

If this interpretation is correct, then the argument that the name Moesia reflects a migration of the Mysians or Mushki from the Balkans is no longer tenable.

Elamites

Elamites

The paper by Amjadi et al. (2025) published ancient DNA from the Chalcolithic period of the South Zagros, dated to around 4600 BCE. The sample comes from the Gol Afshan Tepe site, associated with the Bakun cultural period, which immediately precedes the so-called Lapui period (4100–3500 BCE), usually linked to Proto-Elamite developments. Assuming that there was no dramatic population change between the Bakun and Lapui periods, we may conjecture that the Proto-Elamite language originated among a population with a large amount of Zagros/Iran Neolithic ancestry.

Consistent with its autosomal profile, the sample carried Y-DNA haplogroup R2. This lineage was common among ancient Zagros farmers and today is found mainly in South Asia and in some parts of West Asia—for example, among Armenians in Sasun (see the second map).

The Elamite civilization flourished in the southwestern Iranian plateau. The Elamite language was widely used in the Achaemenid Empire as an administrative language. Persians referred to the Elamite region as Hujiya > Khuzi, from which the name of the modern Khuzestan province is derived. Elamite may have still been spoken in the medieval period, as Arab sources report the presence of an incomprehensible Khuzi language in that region, distinct from Persian. The Armenian word khuzhan, meaning hooligan or wild person, is derived from the name of that region.

If further research confirms a connection between the Elamite language and Zagros/Iran Neolithic populations, several implications may follow:

• Sumerian: Since Sumerian appears unrelated to Elamite, it may instead be associated with another population known as the Central Iranian farmers rather than with Zagros farmers.

• Northern Iran: Northern Iran seems to have had a different genetic history. A contemporary Chalcolithic DNA sample from the Urmia Basin (Hajji Firuz Chl) is strongly shifted westward and plots close to Central Iranian farmers. Their migration eastward may have diluted Zagros Neolithic ancestry in northern Iran, where other ethnic groups are known to have lived during the Bronze Age. The most prominent among them were the Kassites. The Kassite language has no known connection with Elamite, although a possible link between Kassite and Hurrian has been proposed.

• Elamite–Dravidian hypothesis: A theory connecting Elamite with Dravidian languages of South Asia has been proposed, though it is not widely accepted. If Elamite is indeed linked to Zagros/Iran Neolithic populations, this could provide additional support for the hypothesis. However, Dravidian-speaking populations also carry a substantial component of South Asian-specific ancestry related to Andamanese hunter-gatherers.

• Administrative languages and ethnicity: The widespread use of Elamite in the administration of the multiethnic Achaemenid Empire may offer insight into the status of the Urartian language in the kingdom of Biainili. We now know that Biainili was not solely an Urartian state; other ethnic groups, most prominently Armenians, were also part of the kingdom. Elamite inscriptions from the Achaemenid period have been discovered in Armavir, Armenia. This again illustrates that the presence of written texts in a particular language in a given region does not necessarily imply a widespread presence of its speakers there.

The western Iranians.

 The western Iranians.

Three major ancient western Iranian groups were the Persians, Medes and Parthians. Until now we didn't have ancient DNA samples that could be securely linked to this groups. Iron Age samples from south of Urmia could be related partly to Medes but given their genetic profile and dating such linking was remaining speculative.

Amjadi et al. 2025 published first historic western Iranian ancient DNA samples that were either Parthians, either eastern Medes. The location (Marsin Chai, Vestermin) of those samples was in the border of Medes and Parthian satrapies so there is some incertitude how they were identified in their lifetime. (See the map).

Liarsangbon samples could be related to Medes given their more western location. It's remarkable that some of them had also slightly more western genetic profile. J1-620(xP58)>FGC6069 was found in Liarsangbon. Without much doubt of local origin. Another undefined J2a was also uncovered from that site in Parthian empire period.

Genome wide Achaemenid period samples from Marsin Chai were grosso modo looking similar to Bronze Age samples from the same region (TepeHissar, Shahtepe) nevertheless they are not identical.

Looking closer on the PCA it's clear that they are slightly more "northern" shifted. And in some cases slightly "western" shifted. The slight western shift could be of local origin but could be also from Sintashta/Andronovo cultures in Central Asia that had substantial Anatolian Farmer related ancestry.

Despite this no R1a was found from those ancient Iranian remains, only J2b2a2. This can mean that R1a was low but also can be due to sampling issue. Given that many samples were relatives and had the same Y DNA.

Overall those ancient Parthians/Medes have quite high Zagros/Iran Neolithic ancestry with low EHG/ANE. Also they have some South Asian related Onge/Andaman island Hunter Gatherer ancestry.

They are different from Urmia basin Iron Age samples which are more western and who plot close to antique era samples from Armenia. This difference can be due to geography but can be also related to different ethnic background. Ancient Medes samples from core Media are needed to understand this difference.

Returning to the R1a issue it must be noted that the R1a appear both in Armenia and Georgia at Achaemenid period. Even more interesting samples from Meskhetia region in Georgia yielded surprising results. Out of seven Y DNA from Sassanian period Meskhetia here were 2 cases of R1a and 2 cases of Q2 which almost certainly was introduced there by western Iranians. Another J1-P58>Z1853 from Meskhetia was also of non local origin. All this show that western Iranians must have more diverse Y DNA pool and more sampling is needed to understand their diversity.

A PCA with Ancient and Modern Samples from Iran, Armenia, Azerbaijan, and Iraqi Kurdistan

A PCA with Ancient and Modern Samples from Iran, Armenia, Azerbaijan, and Iraqi Kurdistan

The Caucasian Albanian sample from Late Antiquity shows a clear shift toward Iranian populations. It does not fall along the Udi–Armenia LIA–Lezgin cline.

Ancient Iranian groups such as the Medes and Parthians would likely plot slightly to the left and somewhat below Tepe Hissar. They would probably fall near the position of Shah Tepe BA, or slightly to the left and bottom of it.

It is not yet entirely clear, but modern Iranians appear to have a slight western or northern shift compared with ancient populations.

Among modern groups, Kurds are the most western. Persians from Khorasan and Tats show the highest steppe ancestry (a shift toward the left), while the Mazandarani display the highest Zagros Neolithic ancestry.

The Shamakhi Armenian singleton appears to shift more toward the Armenia_LIA / Tat / Lezgin cluster than toward any other Iranian population.

The Dinkha Tepe BA–IA 2 sample, associated with the Grey Ware horizon in the late Middle Bronze Age, could represent an early wave of Indo-Iranian populations.

P.S. I updated the PCA and added the Shah Tepe BA sample.

The Eastern-Side Armenians

The Eastern-Side Armenians

Hovhannisyan et al. (2024) published numerous DNA samples from the Artsakh, Syunik, and Gardman regions. In addition, a member of our group, Diana Sarkisova Vikutan, provided several coordinates from Shamakhi Armenians, as well as DNA samples with mixed ancestry from the Shamakhi and Artsakh regions. We are grateful to her for sharing these data.

All these datasets were combined into a single PCA and ancestry proportion chart together with other modern and ancient samples spanning the area from Syunik to Shamakhi. The analysis was performed using unscaled coordinates. Based on this, several observations can be made:

• Artsakh and Syunik Armenians: Artsakh Armenians are genetically very similar to Syunik Armenians. Both belong to the genetic cline of historic Armenia, which shows a strong affinity with the so-called “Central farmer” ancestry. Gardman Armenians are also close to this cluster, though with a slight shift toward Caucasus hunter-gatherer (CHG) ancestry.

• Distance from Armenia LIA samples: All Armenian subgroups plot away from the Armenia LIA samples from Syunik. This is due to a so-called “southern” shift that may have occurred in these eastern regions sometime after the fall of Urartu.

• Shamakhi Armenians: Shamakhi Armenians show higher steppe ancestry than any other known Armenian subgroup. The two samples from the Shamakhi region differ somewhat from each other. One of them (Sham01) displays more Zagros Neolithic ancestry. The source of this shift is currently unclear. It is worth noting that a Late Antiquity sample (3rd century AD) from the Shamakhi region already showed extra Zagros/Iran Neolithic ancestry, almost certainly due to Iranian influence in that region. Another possibility is that the Zagros shift in Sham01 reflects ancestry from a more recent non-Armenian population. Yet another possibility is that the shift derives from a still poorly sampled subgroup of Tat Armenians.

• Mixed Shamakhi–Artsakh samples: Three DNA samples show mixed Shamakhi and Artsakh ancestry. As expected, they plot between the Shamakhi and Artsakh clusters. However, their dispersion suggests that the real diversity among Shamakhi Armenians could be greater than what the currently available two samples indicate.

• Udi population: The Udi people plot far from their linguistic relatives, the Lezgins. Moreover, they are also shifted away from the Late Antiquity Shamakhi sample that may represent Caucasian Albanian ancestry. The formation of the Udi genetic profile is still unclear, but their proximity to Armenians—who themselves are shifted toward the southwest—suggests that Udis may have acquired Armenian ancestry either in antiquity or, more likely, during the medieval period. It is also noteworthy that some Azerbaijanis from the Shaki region plot close to Udis, indicating that the influence of Armenian-like ancestry was not limited to the Udi population.

Additionally, the Skourtanioti et al. (2024) paper noted an “Anatolian” shift in Iberia and linked it to the spread of Christianity. It is possible that similar processes occurred in Caucasian Albania during Late Antiquity and the Early Medieval period. Hopefully, with the publication of a temporal transect from the Kakhetia region, we will gain a better understanding of how the genetics of Caucasian Albania evolved during historical times.

In summary, the genetic history of the territory of the modern Azerbaijani Republic appears to have become quite complex after the Bronze and Iron Ages. Multiple influences shaped the genetic diversity of the region. The single Caucasian Albanian sample remains somewhat puzzling: it is still unclear whether it represents typical Albanian ancestry or an outlier. As usual, more modern and ancient samples from these regions are needed to better understand their genetic history.

If You are an Armenian and have ancestry from Gardman, Shaki and Shirvan including Shamakhi regions You can join the relevant FTDNA project. You can find the link below. The title of the project will change in the future.

https://www.familytreedna.com/groups/gardmank-gandzak/about