Iron Age Y-DNA from North Macedonia

Iron Age Y-DNA from North Macedonia

This dataset shows Iron Age Y-DNA samples from the territory of modern North Macedonia, based on data from Iosef Lazaridis et al. (2022). Two outlier individuals are not included in the chart.

These populations were identified in Greek historical records as the Paeonians.

The haplogroups present in these samples reflect several different historical layers:

• G2, C1, and J2a-Z6055 derive from Neolithic populations.

• R1b reflects ancestry connected with Yamnaya culture expansions.

• E1b-V13 is associated with Thracian-related expansions from the Carpathian region around 1000 BCE.

Based on the current data, the Paeonians were unlikely to have been closely related to the Illyrians, since the characteristic J2b-L283 haplogroup, commonly associated with Illyrian populations, is absent.

Instead, they may have been closer to populations such as the Phrygians and the Ancient Macedonians. A connection with the Thracians is also possible, though somewhat less likely.

An older and now obsolete theory proposed a Paei → Hai sound shift as evidence for a migration of Proto-Armenians from the Balkans. However, this interpretation is no longer supported by current historical and genetic evidence.

Nevertheless, Armenians and Paeonians remain related in a broader sense, both as members of the Indo-European languages, and through shared Neolithic ancestry reflected in the genetic record.

See also

• Mycenaean Greek Y DNA from continental part of Greece and a Minoan Y DNA from Crete

Dragon Stone Burial from Lchashen

Dragon Stone Burial from Lchashen

The G25 coordinates for the Dragon Stone burial from Lchashen are now available (see the comments). The data come from Bobokhyan et al. 2024.

This unique kurgan burial contained the remains of two newborn children and a woman, who is presumed to have been their mother. A dragon stone (vishapakar) was placed above the tomb.

The autosomal DNA of the individuals is similar to that of other populations living in the territory of the modern Armenia during the same period.

The burial is dated to approximately 1550 BCE, corresponding to the Middle Bronze Age.

The J2 haplogroup has a complex structure.

The J2 haplogroup has a complex structure. For those who want to understand it better here is a tree made by Rozhansky more than 6 years ago. We have already four detailed topics related to each major subbranch. Top four of most popular J2 branches in Armenia.

• The history of J2-M67
• The history of J2-L25
• The genetic history of J2-Z6065
• The genetic history of J2b

Other interesting branches are the

• +Z6049 found in Caucasian hunters. Popular today in Caucasus. A review about one of its subclade.
• +PF5197 found in India, Iran and Gulf region. Also, in ancient Neolithic Armenia.
• +M319 Popular in ancient Crete in Minoans. But also, in Eneolithic steppe.

The populations from left to right are Nakh, Balkar, Ossetian, Georgian, Armenian.

Updated in 2024 July

J2-Z6065 in the Armenian Highlands

J2-Z6065 in the Armenian Highlands

J2-Z6065 is the third most frequent branch of J2a in Armenia, after M67 and L25. Its initial homeland was almost certainly located in historic Armenia.

At present, the oldest known Z6065 sample comes from the Neolithic site of Masis Blur in Armenia. Notably, this lineage is absent among early Anatolian and European farmers and is rare or missing in ancient Minoan populations, suggesting that Z6065 had a slightly more eastern origin compared to related branches such as M67 and M319, with which it shares a common ancestor.

J2-Z6065 has two major sub-branches:

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1. Y13341 / FGC15782

Like many lineages that formed in the Armenian Highlands, Z6065 appears to have expanded during the Chalcolithic period, moving westward into Anatolia. It has been identified at sites such as Ilipinar in Anatolia and Dinkha Tepe in the Urmia basin.

However, its most significant expansion occurred during the Middle Bronze Age (around 4300 years ago).

• The subclade Z6065 > YP879 was likely associated with the Van–Urmia cultural sphere and expanded alongside it, as well as with the related Karmir Berd culture.
• This lineage has been identified in the Urartian city of Sardurihinili and in Late Bronze Age Keti (modern Armenia).

Today, YP879 accounts for approximately 2.5% of the Y-DNA among modern Armenians.

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2. Y7687

This sub-branch likely followed a broadly similar early trajectory but diverged through different historical processes.

• The subclade Z43661 is most likely associated with Bronze Age Anatolia, and there is a strong possibility that it represents a Hittite–Luwian–related lineage.
• Ancient DNA evidence from the Bronze Age site of Ovaören supports this hypothesis.

Today, this branch is well represented in both Turkey and Armenia.

A distinct lineage within this branch, M47, shows a very different geographic distribution, being most prevalent among Gulf Arab and Iranian populations.

• M47 expanded during the Bronze Age, but also shows evidence of later expansions.
• The historical context of these expansions remains unclear due to the lack of ancient DNA.
• One possible hypothesis is that M47 was initially associated with the Kassites, and later became integrated into Iranian and Semitic populations.

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Conclusion

Overall, J2-Z6065 appears to be a lineage deeply rooted in the Armenian Highlands, with multiple expansion phases:

• Chalcolithic dispersals toward Anatolia and adjacent regions
• Middle Bronze Age expansions tied to local cultural complexes (e.g., Van–Urmia)
• Later regional developments, especially in Anatolia and the Near East

Further ancient DNA discoveries will be essential to clarify the full historical trajectory of its subclades.

Trialeti–Vanadzor Culture and Its Genetic Legacy in the South Caucasus

Trialeti–Vanadzor Culture and Its Genetic Legacy in the South Caucasus

The possible steppe connection and Indo-European affiliation of the Trialeti-Vanadzor culture have been discussed for a long time. However, its partial presence in regions that later became known as Iberia and Caucasian Albania made this interpretation difficult to accept.

Today, ancient DNA allows us to better understand past population shifts and helps resolve these apparent contradictions.

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Y-DNA Patterns in Eastern Georgia

The first Y-DNA chart (based on Skourtanioti et al. 2024) excludes one sample from western Georgia.

During the Late Bronze Age–Early Iron Age (1500–800 BCE)—known in eastern Georgia as the Lchashen-Tsitelgori culture—the Y-DNA distribution closely resembles that of Armenia:

• A predominance of R1b and I2, lineages that expanded during the Middle Bronze Age with the Trialeti–Vanadzor horizon
• Two J2 samples from the Bazaleti site (north of Mtskheta), near the foothills of the Greater Caucasus

One of these J2 individuals shows low steppe ancestry, indicating the northern boundary of Trialeti–Vanadzor influence, consistent with the region’s lowland–highland geographic division.

A single G2a1 sample suggests that central Georgian Kura–Araxes-derived populations may have had a different Y-DNA structure compared to Kakhetian groups, which were predominantly J1.

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Iron Age Transformations

In the Iron Age II (800–600 BCE), data are limited, with only one G2a1 sample available.

The Early Antiquity period (600–300 BCE)—coinciding with the rise of the Achaemenid Empire—shows a major shift:

• Appearance of R1a in both Armenia and Georgia
• Increased importance of G2a1 and J2 lineages (likely subclades of CTS900)

Unlike Armenia, which experienced a southward genetic shift, eastern Georgia likely underwent a different directional change, though current qpAdm models lack sufficient resolution to define it precisely.

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The Bragdzor Family: Evidence of Migration

A key insight comes from a familial burial at Bragdzor.

Three individuals were closely related:

• The father and his brother shared Y-DNA (J2) and mtDNA (K1a4c)
• The father displayed autosomal DNA very different from local Lchashen populations

He had:

• High CHG (Caucasus Hunter-Gatherer ancestry)
• Virtually no steppe ancestry
• A profile similar to modern West Georgians

This indicates that he was a migrant, arriving in the region around 800 BCE.

This date coincides with the proposed linguistic separation of Kartvelian languages, including the divergence of Georgian and Zan, which typically occurs alongside geographic expansion.

Further evidence of migration:

• His son’s mother (likely his wife, buried in the same grave) was an Etiuni/Lchashen woman, suggesting incoming males without accompanying women

Despite this, the Bragdzor lineage did not leave a lasting genetic impact in northern Lori, as later samples show continuity with earlier populations.

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Formation of Iberia and Kartvelian Expansion

In contrast, related groups did leave a strong impact in eastern Georgia, contributing to the formation of Proto-Iberia, later known as Iberia in the Hellenistic period.

Additional supporting samples:

• A G2a1 individual from Keti (~650 BCE)
• A Hellenistic-era individual from Samsun, likely linked to Zan populations

These individuals share:

• High CHG ancestry
• A clear distinction from Caucasian Albanian-related samples, which show more Zagros Neolithic ancestry and lower CHG

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Conclusion

The steppe-derived populations that entered the South Caucasus around 2500 BCE (Trialeti–Vanadzor horizon) had a long-lasting genetic impact, visible until the Iron Age.

However, during the Iron Age:

• Mountain populations expanded into lowlands, contributing to the formation of Iberia and Caucasian Albania
• In historic Armenia, state formation under Urartu and the Orontid dynasty led to different genetic shifts
• The Achaemenid Empire likely also influenced population structure across the region

Due to the limited dataset, these interpretations remain provisional, and further ancient DNA evidence will be necessary for definitive conclusions.

See also

• The possible homeland of Kartvelian languages

Remarks on Skourtanioti 2024 paper on the Genetic History of South Caucasus

Elamites

Amjadi et al. (2025) published ancient DNA from the Chalcolithic period in the South Zagros, dated to around 4600 BC. The sample comes from the Gol Afshan Tepe site and is associated with the Bakun period, which immediately precedes the so-called Lapui period (4100–3500 BC), often linked to the Proto-Elamites.

Assuming there was no major population shift between the Bakun and Lapui periods, we can reasonably suggest that the Proto-Elamite language developed among populations with a strong Zagros/Iran Neolithic genetic component. This is supported by autosomal data, as well as the presence of Y-DNA haplogroup R2, which was common among ancient Zagros farmers and today is found mostly in South Asia and parts of West Asia (for example, among some Armenians from Sasun).

The Elamite civilization flourished in the southwest of the Iranian plateau. The Elamite language was widely used as an administrative language in the Achaemenid Empire. The Persians referred to the region as Hujiya > Khuzi, from which the modern name Khuzestan derives. Interestingly, Elamite may have survived into the medieval period—Arab sources mention an incomprehensible “Khuzi” language distinct from Persian in that region.

There is even a linguistic echo in Armenian: the word khuzhan (խուժան), meaning “hooligan” or “wild person,” is derived from that regional name.

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Possible implications

If future research confirms a strong link between Elamites and Zagros Neolithic populations, several important conclusions may follow:

• Sumerian is likely unrelated to Elamite, suggesting it originated from a different population, often referred to as “Central farmers,” rather than Zagros farmers.
• Northern Iran had a different genetic trajectory.
  A Chalcolithic sample from the Urmia basin (Hajji Firuz) is shifted toward western populations and clusters closer to Central farmers. This suggests eastern migrations diluted Zagros ancestry in northern Iran, where other groups—such as the Kassites—later appeared. Kassites are not clearly linked to Elamites, though connections with Hurrians have been proposed.
• Elamo-Dravidian hypothesis gains some support.
  The idea that Elamite is related to Dravidian languages is still debated, but a Zagros Neolithic origin could strengthen this theory. That said, Dravidian populations also carry strong South Asian-specific ancestry (including Andaman-related hunter-gatherers).
• Language vs. population mismatch in empires.
  The widespread administrative use of Elamite in the Achaemenid Empire shows that a language’s presence in inscriptions does not necessarily reflect the dominant spoken language of a region.
  This is relevant for understanding Urartian (Biainili) as well: inscriptions alone don’t prove linguistic dominance. For example, Elamite inscriptions have been found in Armavir (Armenia), yet that doesn’t imply Elamite-speaking populations lived there in large numbers.

See also

• Ancient Kartvelians and the Formation of Iberia

Two different stories in Anatolia

After the Neolithic period migrations from east changed the genetic landscape of plain Anatolia. Those migrations were not an one time event but two major events dated to different periods. It must be noted that in the current state of archaeologic knowledge there are no Neolithic sites in northern Anatolia. Food producing appear there in the Chalcolithic period which starts after the 5800 BCE. See the dividing line on the map.

Currently we have two Early Chalcolithic samples from Hattusa (Buyukkaya) in the north and Tell Kurdu in the south (the green circles on the map). They are dated to the same period after the 5800 BCE yet they have different shifts to east. The northern sample has a strong shift to east close to the Late Chalcolithic samples from the same place (Camlibel Tarlasi). While the southern site (Tell Kurdu) has a very little shift. In my previous thread dedicated to Aintab history I showed that in southern Anatolia the main migration from the east occurred at Late Chalcolithic, thus more than 1500 years later than in the north. Given that that those two events have a different archaeologic background then it's safe to assume that they are related to different ethnic groups.

Based on the currently available data we can link the Late Chalcolithic migrations in the south to the Minoans and IE Anatolians. While the northern Early Chalcolithic migration can be associated with Hattic people. This theory is supported by the strong presence of G2-M406 in Hattusan sites. While the same M406 was absent or rare in the Crete and Minoan civilization sites.

It's worth to note that in western Caucasus and Georgia the Neolithic settlements appear roughly in the same period. We can assume that the same impulse that introduced the food producing to north Anatolia moved also to western Georgia were a peculiar Neolithic culture emerged, different from the Shulaveri-Aratashen related sites in Kur-Araxian basin. Later those western Georgian farmers moved to north triggering the emergence of mountainous pastoralist culture known as Darkveti-Meshoko (after 4500BCE). We have samples from this latter culture. They are from a subbranch of J2-M67>CTS900. What linguistic group is related to the introduction of food producing to the west Caucasus is a more complex subject, which will be discussed later