Paeonians Y DNA from ancient North Macedonia

Ancient Iron Age Y DNA from modern North Macedonia. Two outliers are not included. Based on Lazaridis 2022 paper data.

Those people were known as Paeonians in Greek records. G2, C1 and J2a-Z6055 are from Neolithic period. R1b is from Yamnaya. The E1b is V13 related to Thracian expansion from Carpathian Mountains around 1000BC.

Based on this data Paeonians were less likely to be related to Illyrians, given the absence of J2b-L283. They were probably related to Phrygian and ancient Macedonians. A connection with Thracians is also possible but less likely.

An obsolete theory proposed a Paei> Hai shift to prove a migration of Proto Armenians from Balkans. But now we know that it was a wrong theory.

In any case Armenian and Paeonians are related to each other as part of Indo-European family. Also based on shared Neolithic ancestry from genetic point of view.

See also

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

The haplogroup G

 The haplogroup G is the third most frequent in Armenia after R1b and J2.

G was prominent in early Neolithic farmers, especially in those that moved to Europe. Despite its popularity in West Caucasus Rootsie 2012 analyzed the modern diversity of G and made this conclusion about its homeland.

..by evaluating 1472 haplogroup G chromosomes belonging to 98 populations ranging from Europe to Pakistan. Although no basal G-M201* chromosomes were detected in our data set, the homeland of this haplogroup has been estimated to be somewhere nearby eastern Anatolia, Armenia or western Iran, the only areas characterized by the co-presence of deep basal branches as well as the occurrence of high sub-haplogroup diversity. ...

In the last decade ancient DNA supported this homeland theory with a slight shift to more southern regions toward Fertile Crescent.

Three branches are frequent in Armenia.

• G2a2b-M406 more than 3.5%
• G2a2b-P303 around 3%
• G2a2a-PF3147 around 2.5%
  

Other branches are less frequent.

• G2a1
• G1 prominent in Hamshen Armenians
• G2b - The genetic history 
• Over time there will be reviews about those branches.

The Genetic History of J2a-Z6065

J2-Z6065 is the third most frequent branch of J2a in Armenia, after M67 and L25. Its initial homeland was almost certainly situated in historic Armenia. Currently, the oldest Z6065 sample comes from ancient Armenia, specifically the Neolithic site of Masis Blur. It was absent from early Anatolian and European farmers and even missing or rare in ancient Minoans, suggesting that Z6065 has a slightly more eastern homeland compared to M67 and M319, with which it shares a common ancestor.
J2-Z6065 has two important sub-branches:

**Y13341 / FGC15782**: Like most haplotypes that formed in historic Armenia, Z6065 benefited from expansions during the Chalcolithic period, moving westward towards Anatolia. It was found at the Ilipinar site in Anatolia and in the Urmia basin at Dinkha Tepe. However, its most successful expansion began in the Middle Bronze Age (4300 year ago). The subclade Z6065>YP879 was apparently part of the Van-Urmia culture and expanded with it and the affiliated Karmir-berd culture. YP879 was found in the ancient Urartian city of Sardurihinili and in Late Bronze Age Keti, modern Armenia. Today, YP879 constitutes 2.5% of the Y-DNA of modern Armenians.

**Y7687**: This sub-branch likely has a similar history to the first one, but its two successful subclades are related to different events. The Y7687>Z43661 is probably related to Bronze Age Anatolia, and there is a strong possibility that it was a Hittite-Luwian branch. Ancient DNA from the Bronze Age Anatolian site in Ovaoren supports this theory. This branch is well represented in Turkey and Armenia. 
M47 has a very different distribution, being prevalent among Gulf Arabs and Iranians. M47 expanded during the Bronze Age but also has more recent expansions. The historic events related to this expansion are unknown due to the absence of ancient DNA. One possible theory is that it was initially a Kassite lineage that later became part of Iranian tribes and Semites.

See also 

• The haplogroup J2

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

Ancient Kartvelians and the Formation of Iberia

The possible steppe connection and Indo-European affiliation of the Trialeti-Vanadzor culture has been known for a long time. However, its partial presence in the lands which in antiquity became known as Iberia and Caucasian Albania was a hindrance to accepting and promoting that theory. Today, ancient DNA allows us to better understand population shifts in the past and resolves these apparent contradictions.

The first Y-DNA chart is based on the data from the Skourtanioti 2024 paper. A single Y-DNA sample from western Georgia was not included in the chart.

The LBA-EIA (1500-800 BC) period is known in eastern Georgia as the Lchashen-Tsitelgori culture. The Y-DNA distribution from this period is very similar to that from Armenia, with a predominance of R1b and I2, which expanded during the Middle Bronze Age Trialeti-Vanadzor culture period. The two J2 samples are from the Bazaleti site, which was north of Mtskheta, close to the foothills of the Greater Caucasus range. One of them also had low steppe ancestry in autosomes, indicating the northern border of Trialeti-Vanadzor culture's impact. This corresponds to the geography of the region, dividing lowlands and highlands. (see the map)

A single G2a1 in the LBA suggests that the Kura-Araxes culture of central regions of Georgia could have a different Y-DNA structure than those from Kakhetia, who were predominantly J1.

In the Iron Age 2 (800-600 BC), the data size is small, with only one Y-DNA sample, which is G2a1.

The Early Antique period (600-300 BC) roughly corresponds to the formation of the Achaemenid Empire. R1a appears in both Georgia and Armenia, and we see a significant change in the Y-DNA distribution. G2a1 and an as-yet unknown branch of J2 (most likely a subclade of CTS900) are now the most important haplotypes. Unlike in Armenia, which during this period shifted south, in Georgia we can expect a different shift. Unfortunately, the qpAdm models from the paper do not have enough resolution to understand the possible direction of the shift. A familial grave from Bragdzor near the Shnogh village close to modern Armenian-Georgian border can provide a hint about the nature of the population that expanded in Iron Age east Georgia.

**The Bragdzor Family**

Three samples from Bragdzor were related to each other. The father and his brother had the same Y and mito DNA, J2 and K1a4c respectively. The Bragdzor father had very different autosomes compared to the contemporary Lchashenians. He is marked as an outlier in the AADR dataset. He had high CHG and virtually no steppe ancestry, similar to modern West Georgians, with no admixture from the Etiuni-Lchashen people. This indicates that he arrived in Bragdzor during his lifetime, around 800 BC. Incidentally, this date corresponds to the proposed age of separation of Georgian and Zan. Two languages usually separate when the geographic distance between them increases, typically as a result of migration.

Another indirect piece of evidence that the Bragdzor father and his brother were new migrants is the fact that his son's mother (most probably his wife buried in the same grave) was an Etiuni woman, suggesting that they did not have women from the same ethnic group when they arrived. In any case, the Bragdzor family did not leave a lasting genetic impact in northern Lori. Numerous samples from the same region in succeeding periods show no evidence of significant change.

On the other hand, the kinsmen of the Bragdzor father left an important impact in eastern Georgia, laying the foundation of Proto-Iberia, which became known as Iberia in the Hellenistic period. There are two other samples outside of Georgia with possible Kartvelian affiliation having similar autosomes to Bragdzor: one from Keti (G2a1, ~650BC) and another from Hellenistic-era Samsun. This latter was in most likelihood related to Zan people. All of them have high CHG, differentiating them from possible Caucasian Albanian sample who had Zagros Neolithic ancestry and lower CHG.

**Conclusion**

In conclusion, steppe migrants that arrived from north of the Caucasus 4500 years ago left an important impact in the South Caucasus that lasted until the Iron Age. In the Iron Age, multiple unrelated events resulted in significant changes. Near the Greater Caucasus range, mountainous people moved to the lowlands, and Iberia and Caucasian Albania formed. Meanwhile, in historic Armenia, Urartu's and the Orontid (Yervandunis) efforts to create centralized states resulted in a different change in the genetic profile. Most likely, the Achaemenid Empire also played a role in these genetic shifts in the South Caucasus, but the scant data do not permit to have definitive conclusions.

See also

• The possible homeland of Kartvelian languages

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

This is the first paper with a large number of ancient DNA samples from Georgia. The bulk of them are from eastern Georgia, within the Kur-Araxian basin, which permitted the authors to compare the genetic histories of modern Armenia and Georgia. The time transect starts from the Early Bronze Age and the Kura-Araxes culture. Genome-wide, the Kura-Araxes samples from eastern Georgia are similar to those found in Armenia. The Y-DNA of all three samples is J1 (Z1842). We now have 12 Y-DNA samples from Kura-Araxes, and 9 of them (75%) are J1, which further reinforces the idea that J1-Z1842 expanded with the Kura-Araxes culture and was probably the most important Y-DNA in the northeastern parts of the Kura-Araxes horizon.

Next, in the Middle Bronze Age period (2200-1500 BC), there is an increase in steppe ancestry. The Y-DNA R1b-Z2103 and I2a2b appear associated with the Trialeti-Vanadzor culture. Apparently, those steppe pastoralists had a lower impact on northeastern parts of Georgia, likely due to the geography and high mountains. We see a preponderance of J2 there, while the southeastern regions of Kakhetia and regions bordering Armenia were more strongly affected by the steppe. This situation persists until the Iron Age 2 and Early Antique era (800-300 BC), when in the Urartian-Orontid period, we see a drop in steppe ancestry in Armenia. In Georgia, the drop in steppe ancestry was not abrupt but slow. This difference is another argument favoring the idea expressed in Petrosyan 2023 that this drop in Armenia is related to Urartu activities and post-Urartian events.

Starting from the Iron Age, the genetic histories of Armenia and Georgia begin to diverge. Georgia witnesses' migrations from the south in the Late Antique period to Early Medieval periods (1-400 AD). The authors link those migrants with the spread of Christianity but given that some of them appear in the 1st century AD (NAT004, 1950 ybp), we can assume they were related to the political events of that period. In the 1st century AD, Rome was trying to reinforce its control of Greater Armenia and the South Caucasian kingdoms of Iberia and Albania. Meanwhile, the Parthian Arsacids were trying to counteract them. As a result of these wars, the Arsacids became the rulers of Armenia and tried to impose their cadet branch in Iberia as well. Armenia on the other side gets a Levantine influence in Hellenistic period. From historic point of view, it is quite possible but given the small sample size and issues with models where various periods are not compared with the same source populations, there is a need of extra data to confirm this.

Starting from 400 AD, eastern Georgia witnesses new sporadic waves of migrants from the steppe, this time associated with the Huns and Sarmatians. My feeling is that the Maskut tribes' invasion is also associated with this event. These steppe intruders had a practice of artificial deformation of the skull, which the locals borrowed. The sample size from Armenia is small from those periods, which is probably why we don't see them in Armenia. Another possibility is that they didn't reach Armenia in large number. After the Middle Ages, there are no remarkable events.

Overall, this is a good paper that describes the genetic history of the region. This comparison would be more comprehensive if we had dense sampling of Armenia and Azerbaijan from the relevant periods. We should not forget that the sample size in the post-Urartian period Armenia is incomparable to what we have now from Georgia.

Besides the positive side there are some issues mentioned by other commentators:

- The choice of Iran_Chl as a source population instead of Iran_Neo. This choice can mask the Levant N ancestry present in the South Caucasus since the Neolithic period.

- Mentions of obsolete papers like Pagel 2013, where Pagel and Atkinson calculated the ages of divergence of different linguistic groups in Eurasia.

It must be noted that current genetic data do not forbid the deep origins of Kartvelian from CHG. Given that there was a resurgence of CHG in the Bronze Age Georgia corresponding to the age of divergence of Proto-Kartvelian.

The deep origin of PIE from local farmers is also still on the table. However, the most frequent Y-DNA of Kartvelian people (G2a1 from Fertile Crescent) and those in the CLV cline (R1b, from EHG) create other possibilities also. More data will obviously help us better understand the deep origins of Kartvelian and PIE.

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

Lernagog 1 - a new Neolithic site in Republic of Armenia near Talin.

New details are emerging about the Lernagog 1 site which was discovered in 2017 by an Armeno-Japanese team. Radiocarbon dates show that it is the oldest Neolithic site in South Caucasus and can be qualified as Early or Middle Neolithic (8-7th millennium BCE) Until the discovery of this site the advent of food producing in South Caucasus was mostly dated after 6200 BCE, known as Shulaveri- Aratashen- Shomutepe culture.

Tthe youngest hunter-gatherer date obtained in foragers cave (Kmlo 2) was 7400 BCE in Armenia. Thus, there was a hiatus of more than 1000 years between the end of the last forager and the start of farming and pastoralism. The Lernagog 1 site fills this hiatus. Research shows that they were making clay houses and were mostly pastoralists. No human bone is found yet

There are two possibilities. One is that Lernagog were local hunters who learned the pastoralism. If so, they would be closer to Aknashen sample. Another possibility is that they were migrants from Fertile Crescent "core area" (see the map). This latter scenario would mean that they were closer to Masis blur sample we have.

Qpadm models about the Core Yamnaya origins.

We have one sample from north Caucasus PG2004 which is similar to so called BP group of Caucasus Lower Volga cline.

Modeling Yamnaya as a three-way mixture of CLV_BP, Aknashen and Ukraine Neolithic hunter gatherers was very easy, as in the paper.

BP group 66.5%

Aknashen 19.5%

UNHG. 14.0%

p value 0.17

It didn't require from me any extra effort to find the correct "settings" (outgroups). Adding western Ukraine Trypllia farmers didn't made the model better. Aknashen was still wanted while Trypllia not. I tried various ways to make Trypllia wanted, but it didn't work. Finally, I removed Aknashen and forced the model to rely solely in Trypllia. I got a p value lower than 0.05. So practically a failure. Nikitin 2024 dedicates a special chapter to this subject. Their conclusion is that Yamnaya probably do have some Trypllia but it's very low.

My conclusions are the same. Aknashen ancestry in Yamnaya is real. It is supported by Y DNA of preceding period, and I am sure that the scrupulous analysis of mtDNA will show the same result. Finally, if I had access to the Nalchik farmer data then the percentage would be twice higher.

Then I tried the same in G25 and here a surprise. The raw unscaled models behaved in the same manner as qpadm albeit with a different proportion. But the scaled G25 showed a clear a preference for Trypllia. I said in this group many times that tools and settings matters. And this an excellent example why the use of scaled models can lead to misleading results. I want to remind that scaled coordinates are artificially altered numbers.

Keeping this in mind let's see what means CLV-steppe ancestry in Bronze Age Anatolia that Lazaridis 2024 supposedly has found but Lazaridis 2022 didn't detected. I don't have Cayonu files to reproduce their models in qpadm but what we know from G25 behavior is that:

+ When Anatolia N and CHG are used in models as source they mask the steppe ancestry and show a lower number. This was done in Lazaridis 2022 which not only didn't find any steppe in BA Anatolia but even in Van Urartu.

+ When a Mesopotamian or Levantine and Iran Neo are used as sources and CHG and Anatolia_N are ommitted then this exaggerates the steppe ancestry in northwest Asia. In Lazaridis 2024 they used Cayonu Neolithic which was a north Mesopotamian population. This exaggerated the steppe ancestry in Bronze Age Anatolia where Hittite lived. If the same Cayonu was used for Minoans, they would find steppe even in Minoans and probably Alalakh Hurrians and Semites also. Which doesn't make sense. Uniparental markers do NOT support such high level of CLV-steppe ancestry in BA Anatolia. Quite contrary they speak about very low or virtual absence of it. Just one R1b-V1636 in the midst of more than dozen local haplotypes.

My conclusions are that based on this genetic data it is not possible to consider IE homeland issue fully solved. We are very close it. And the broad picture. But the exact details are not still there, and linguistics also can be helpful.

What is needed now is to have more than 50 Y DNA from Bronze Age Anatolia. Also, a large number of Y DNA is needed from South Caucasus and historic Armenia LC-EBA period to see what happened to those Late Chalcolithic migrants from north. How much they left an impact and why their autosomes were diluted after the Areni C. It is also important to have samples from LC kurganic burials like Aknalich and Soyuq Bulaq.

See also

• When the tools and settings matters. Part 1 and Part 2

The Kurgan builders.

The Caucasus Lower Volga cline (CLV) having both south Caucasian and Eastern European foragers ancestry is a genetic term. The material cultures behind this term were variable but they had one important common feature. Virtually all the samples from CLV cline were found from kurgans. Kurgan is an artificial mound, a tumulus on top of the grave. A kurgan could harbor a single or multiple graves. Given that it requires a lot of manpower it was usually built for elite persons. Smaller kurgans also existed. Other prominent features of kurganic burials in Pontic Caspian steppe were the red ochre, the raised knee position etc. The origins of this tradition are uncertain, and the dates of the oldest kurgans debated but what is well known now that it expanded with Eneolithic (is equal to Chalcolithic) pastoralists (4500-3500bc) who had the CLV ancestry, replacing older local hunter's flat grave tradition. Besides kurgans, we know that those people had a patrilocal and exogamic culture.

Kurganic people moved to north, toward middle Volga region and the Khvalynsk culture (after 4500bc) emerged there. In middle Volga region various branches of haplogroup Q1 were integrated into early steppe pastoralist communities. Q-L939 the branch of Georgian Bagrationi. Q-YP1669, probably Q-F26062 and some others to be found.

They also moved toward the Balkanian peninsula where they mixed with local European farmers where various kurganic cultures emerged (Cernavoda, Usatovo, Suvorovo). This migration has been proposed to be the source of Anatolian languages. Currently there is no evidence that they reached deep into Anatolia. Nevertheless it's possible that some obscure ethnicities like Mysians in Turkish Thrace and north west Anatolia were derived from those migrants. Although a later Yamnayan origin is also possible for them. Yamnayans themselves also had kurgans.

The new "finding" in Lazaridis 2024 paper is about the migration of those kurganic CLV people to south Caucasus. I use brackets because the presence of steppe ancestry in Areni cave was known since 2016. But only now it got an interpretation. This migration apparently occurred at 4300BC a period known as late Chalcolithic which is associated with Chaff faced ware. Chaff ware was not from steppe but had a local Neolithic origin. A syncretic culture emerged. A variant of which is known as Leila tepe culture in what is now Azerbaijan. We have already seen that those chaff groups moved to west toward Anatolia. The R1b-V1636 is associated with this event. We have now three cases of V1636 stretched from Aintab to Sevan basin.

The new proposal of Lazaridis is that they were the IE Anatolians and based on this they propose that the CLV is the place were Indo-Europeans emerged. This is the main difference of this paper from the Lazaridis 2022 in which they placed the homeland of PIE ( or Indo-Anatolians ) in south of Caucasus.

My next post will be about the Indo European origins based on the linguistics and how those known migrations fits into the great picture. Also I will discuss the real number of steppe ancestry in Bronze Age Anatolia and will show that the current data size is very small for having a good understanding of the situation.

** The first picture is a reconstruction of a kurgan .They are usually eroded over time. This one reconstructed. The second picture is the tumulus of Lydian king Alyattes. This is one of the largest known kurgans.

Pastoralism in East Europe

We have now quite large number of samples before the Yamnaya period in East Europe ( before 3300BC) to have some conclusions about the origins of pastoralism in Pontic Caspian steppe and forest steppe regions.

If we keep aside local hunter gatherer lineages and those associated with Maykop culture in North Caucasus, then we are left with haplotypes that are associated with South Caucasian farmers. Shulaveri Aratashen culture.

• J2-M319 - an obvious farmer lineage, which usually is associated with Minoans but it's deeper origins are without doubt in historic Armenia. Absent in European farmers.
• J2b2b from Eneolithic Moldova associated with migrations from east. This branch was never found in any Euro-Anatolian farmer site but was found in Mentesh tepe in northwest of Azerbaijan.
• J2b2a1-L283 currently the oldest sample from this branch is from Yamnaya but there is little doubt that it was present in Eneolithic steppe also. It became part of Yamnaya communities, moved to west Balkans and had a successful founder effect there. It's parallel branch the J2b2a2 is a lineage found in farmers from Central Asia who from there moved to India. Thus, indirect evidence supports that it was a south Caucasian farmer lineage. Its hunter gatherer origin is less likely.
• J1b - it was a west Caucasian hunter lineage. It could be a CHG lineage in steppe. But a farmer lineage can't be ruled out completely given that this branch was found in many farmers sites stretched from Pakistan to Crete. In steppe it was found in Volga region and Usatovo cultures.
• J1-CTS1026 - similar to J1b uncertain affiliation. It's south Caucasian origin in steppe is without doubt.

Besides those cases there were also J1* in Karelia in hunter gatherer context. Those were almost certainly from CHG.

On the other hand, there was no single European farmers lineage found in Pontic Caspian steppe. Archaeologists believed that west Ukraine farmers played an important role in the introduction of pastoralism, but the genetic data do not support that idea. In west Ukraine Cucuteni Trypllia farmers we have G2a2a, G2a2b, E1b-L618, C1a2, I2a2a etc. None of them is found in steppe.

An unpublished sample from Nalchik proves that farmers from South Caucasus moved to north. Despite this southern input, the Eneolithic steppe was predominantly R1b-V1636 and I2-L699. This latter was a Ukraine hunter gatherer lineage in its deep origin.

What was the reason of the mismatch between autosomes and Y DNA is hard to say but a similar scenario occurred in main Europe where after an initial success, the G2 farmers lost their positions and I2 became more frequent. While the autosomes didn't change in a significant manner.

Those genetic results is supported by the linguistics data also. Sahala notice that the Sumerian word gud, gu meaning ox, bull is a good parallel with PIE *gou(s) meaning cow, ox. From which the Armenian word kov is derived.

Matasovic notice that the reconstructed PIE language morphology shares features with North Caucasian languages.

PS You can see the Caucasus Lower Volga cline on the map which had both CHG and South Caucasian farmers ancestry mixed with local hunter gatherers

The Yamnaya culture and it's legacy

Yamnaya was an Early Bronze Age culture (3300-2700BC) that expanded in Pontic Caspian steppe in East Europe. It is remarkable due to its homogenous population which contrasts to the diversity of the preceding Eneolithic (means Chalcolithic) cultures. In just 1000 years genes associated with Yamnaya culture reached what is now Portugal and western Mongolia, Britain and South-Central Asia. The scale of this expansion can be considered as one of the most important demographic events in West Eurasia after the invention of farming in Fertile Crescent. In 2022 Lazaridis et al. announced that all modern living IE nations languages can be derived from the Yamnaya.

In 2015 the first ancient DNA of Yamnayans was published and it became clear that they do not directly descend from Eastern European hunter gatherers. Roughly the half of their ancestry was from south, and the "Armenian like" term was used to label this southern part. Then in late 2015 the Caucasian hunter gatherers (CHG) were discovered. In 2016 Iranian Zagros Neolithic farmers were published and since then both those populations were used to model the Yamnayans.

Despite those statistical models the formation of Yamnayans gene pool was remaining an obscure subject. Finally in April 2024 two large papers dedicated to this subject provided more clarity. Various Neolithic and Eneolithic groups living in East Europe had both shared ancestry and differences. One group who was living from North Caucasus foothills to Lower Volga (site Berezhnovka ) was labeled as "Caucasus Lower Volga cline". CLV had both southern Caucasian hunters and farmers (Aknashen) ancestry.

I will dedicate a special topic to describe what is CLV but at this stage it's sufficient to say that Yamnayans had up to 80% of their ancestry from CLV. Not only Yamnayans but all Kurganic cultures (Sredni Stog, Khvalynsk, Cernabida, Usatovo) of eastern Europe and Balkans had the CLV ancestry mixed with various local European populations. For the case of Yamnayans the second part of their ancestry was the Ukraine hunter gatherers (UNHG), which made them similar to Sredni Stog. Even though the most important Y DNA's were missing in Sredni Stog which could mean that Proto Yamnaya's exact immediate geographic location is not found yet but shouldn't be outside from Don - Volga - Caucasus triangle. With more sampling we will learn the exact origin of the two most successful lineages. R1b-M269 and R1a-M417.

The samples from the new paper are not available yet but we have two samples from the forementioned CLV cline. They are from the Progress - 2 site in the North Caucasus (4500-4000BC, see the last map for the location). I used them to model various Yamnaya samples and got the same result as in Lazaridis 2024. More than 75-80% of ancestry in regular Yamnaya samples is from the Progress / CLV. Some outliers have extra ancestries which is normal given their location.

You may wonder how "Hittites from east, from historic Armenia" and "Yamnaya from CLV" are connected to each other? After all there is no evidence that Hittites and other Anatolian speakers descend from Yamnaya culture. What is the common shared ancestry between IE Anatolians and Yamnaya?

Another topic will be dedicated to this subject.

And at the end here is the list of Y DNA found in Yamnaya at this stage. R1b-Z2103 (west Asia Balkans), R1b-L51 (west Europe), R1a-M417 (Asia - Europe), I2-L699, J2b-L283 (west Balkans), R1b-PF7562 (Balkans Mycenean ) Q-L939 and isolated cases of J1 and R1b-V1636.

See also

• Population dynamics in North Caucasus

Hittites from east.

Before commenting the deeper origins of Indo Europeans proposed in the new Lazaridis 2024 paper I want to speak about the origins of Anatolians. Since the advent of paleogenetic it became clear that they can't be native to plain Anatolia. They could be either from east or from north-west. Since 2018 the accumulating data was giving clear preference to the eastern origins. From historic Armenia. But a doubt was still persisting, because the Balkanian route was not fully investigated. In 2023 Penske et al. has found a migration from steppe to eastern Balkans before Yamna around 4000BC but they didn't even discuss the possibility that they could be related to Hittites. It's because their autosomes were rapidly diluted having a limited impact in Bulgaria yet alone in Anatolia. Also the Y DNA associated to this migration the I2-L699 was missing from inland ancient Anatolian samples.

While a R1b-V1636 was found in Bronze Age Gaziantep quite close temporaly and geographically to the kingdom Armi mentioned in Eblaite texts around 2500bc. Hittites names have been discovered in this kingdom recently.

The V1636 was popular in CLV cline stretching from Caucasus to Lower Volga and was found in ancient Malatya and Sevan basin also.

Besides this R1b other haplotypes found in Bronze Age Anatolia also show self-evident eastern affiliation.

The J1-Z1828 who's subbranch BY94 can be an Iron Age Luwian lineage like the J2- L70 which itself is derived from the L25 a haplotype from east.

A J2-Z6065 which was found in Masis Blue in Neolithic Armenia. We have also bunch of of J2-M67 and G2-M406, even though this latter is related to Hattic people so it can't be interpreted as a prove of Hittites from east.

In conclusion all ancient Y DNA that we have in inland Anatolia do have an eastern origin, from historic Armenia and around. One of them is further from north of Caucasus region. Their autosomes are also shifted to east compared to the preceding periods.

One exception is the I2-P78 from MBA Yassi Tepe near Izmir, which is clearly from Mycenean Greece, so this person was almost certainly a Greek people representative who probably settled in western Anatolian coasts after the 2000BC. They are mentioned in Hittite texts as Ahhiyawa.

In sum we can now say with high certitude that Indo-European languages were spoken in Armenian Highlands at minimum the last 6300 years. The 4300BC is not a random number it's the date of the start of Late Chalcolithic associated with Chaff faced ware which had a tremendous impact on southern Anatolia. But not much on northern Anatolia where Hattic related people were settled since more earlier times. Probably since 5800BC.

But that would require another thread which I will do the next time.

PS a proposed location of Armi. Another location is in the northeast of Ebla.

See also

• One possible way to know the language(s) spoken in Kur-Araxian culture is to examine the onomastics in those places where there was continuity

Samples of Anatolian/Aegean origin from the Roman empire city Vimminacium in what is now the eastern part of Serbia

Samples of Anatolian/Aegean origin from the Roman empire city Vimminacium (Olalde 2023) in what is now the eastern part of Serbia. A large military camp was present in the city.

• E-L791 is remarkable because it's very close to Napoleon's Y DNA. Another L791 from Roman era Croatia.

  Distance to: Croatia_Zadar_Roman_oLevant.SG:R3742.SG
  0.03003301 Greek_Cappadocia
  0.03236809 Greek_Central_Anatolia
  0.03434637 Greek_Crete_Heraklion
  0.03806465 Armenian_Gesaria
  0.03847296 Greek_Crete

• I2-Y16419 is ultimately from Trialeti-Vanadzor culture. Based on genetics he was probably from Kesaria/Kayseri.

• The presence of J2-L70 and J1-BY94 is without surprise. These two haplotypes expanded in Iron Age Anatolia most probably.

• And the T1a2 could be local but in most likelihood, he was also from Anatolia.

Also, a lot of local Balkanian specific haplotypes like E-V13 with some Germanic ones.

Another evidence that Roman Empire had a significant impact on the genetics of South Europe. Read in more detail in the link below.

 

•  In 313 AD, the Emperor Constantine issued the Edict of Milan, which accepted Christianity