The East Asians and the Mongoloid race.

This group is dedicated to the Armenians but given that various subjects are discussed that are indirectly related to IE and thus Armenian origins so let's take a broader view on Eurasia. Another reason to discuss the East Asian genetics is related to our two neighbouring countries who speaks Turkic languages that came from the aforementioned region.

After it's initial spread in Eurasia (45.000years ago) humans divided into two major groups. East Eurasians living east of Himalayan mountains and West Eurasians.

Initially both groups were close genetically. Ancient DNA confirmed it. But after the Last Glacial Maximum, thus 20.000 years ago in East Asia new drifted group formed which had Mongoloid phenotype. It formed as a result of evolution and not migration. Today those people are known as Mongoloid race, while West Eurasians are known as Caucasoid or Europoid race.

The East Asian ancestry formed in northern central China / Amur region. The main Y DNA associated with them are the N, O and C2. See the attached maps. The haplogroup Q living in Paleolithic Siberia had West Eurasian origin and was initially associated with people having Caucasoid phenotype. The main population associated with them is known as ANE. Ancient North Eurasian which do not exist anymore.

Amerindians formed as a result of mixture of "caucasoid" ANE and old Mongoloids in Far East. Even though their most important Y DNA remained the "western" Q1. They settled in America some 15.000 years ago.

At some point number of important innovations occurred in East Asia providing a competitive advantage to them. One of oldest potteries are found in Amur region, north China dated between 12.000-18.000 years before present. Some 7000-8000 years ago the haplogroup N moved from Mongolia to "Caucasoid" Siberia starting a large migration and conquest which reached modern Finland and Europe. Today the highest level of haplogroup N is found in Uralic speakers, who's homeland was probably in east of Uralic mountains. Turkic speakers also have the N haplogroup but in a lesser degree. Haplogroup O expansion was more related to farming invention in China. Both in south (rice) and north (millet) China. They had large expansion to southern regions, replacing there Black people related to Aborigenes, Onge and Papua. And finally, the haplogroup C2 initially restricted solely to Amur region had a dramatic expansion in Current Era related to Mongols and Evenk (Tungusic) speakers.

Here is a comprehensive list of linguistic families that expanded from East Eurasia spreading genes associated with Mongoloid race in the rest of the world. You can see the autosomal impact in 4th map. It's probably a little bit exaggerated but overall give an impression of the scale of these expansions.

+ Uralic

+ Turkic

+ Mongolic

+ Evenks also known as Tungussic

+ Sino-Tibetan, the most famous member is the Chinese

+ Austro-Asiatic that reached India.

+ Austronesian stretched from Madagascar to Easter Island in Pacific via Indonesia.

I will dedicate separate threads to some linguistic families that are interesting from the Armenian history perspective

R1b-M269 and the Steppe Eneolithic.

In 2019 Wang et al. published three ancient samples from north Caucasian foothill steppe region. They got the label Steppe Eneolithic. The term "Steppe" is usually used in genetic papers to label DNA related or similar to Yamnaya genetic profile.

Those Steppe Eneolithic samples were remarkable because not only they have the genetic profile similar to Yamnaya but were also older than it. Back then they didn't get the attention they deserved because they were not from a famous culture. In 2024 Lazaridis et al. proposed a new name for them the Caucasus - Lower Volga cline and added new similar samples. Genetic calculations showed that genome wide Yamnaya got most of their ancestry from this CLV folks. Despite this advance the two most important Y markers were still not securely linked to this CLV population. The R1b-M269 and R1a-M417 which had a large expansion all over Eurasia spreading the Indo-European languages.

Ghalichi et al. 2024 filled this gap. Two new cases of R1b-M269 were found before the Yamnaya near North of Caucasus region. They had similar to other Steppe Eneolithic groups ancestry dated at around 3800BC. It's worth to note that there was another older M269 sample (I33307) from Kalmykia dated around 3700 but it's genetic profile is identic to Core Yamnaya raising the suspicion that it is younger than the radiocarbon dates show. Lazaridis et al. labeled this sample as Russia_Caspianinland_EBA_Yamnaya group that is why I didn't included it in this list of Steppe Eneolithic Y DNA (4700-3000BC. See the map for the geographic area)

R1b-V1636 -----9

R1b-M269. -----2

R1a. -----1

I2-L699. ------1

J2a ------1

J2b2a. ------1

Besides those cases another possible old R1b-M269 (around 4500BC) was found in Bulgaria in Varna culture burial. It's autosomes showed an obvious Steppe Eneolithic related ancestry. Which means that it was not related to local Neolithic farmers and was a migrant from the Caucasus Lower Volga region. Varna was a peculiar culture with highly hierarchical organization. Large number of gold items were found from Varna culture graves.

Another important finding in the Steppe Eneolithic context is off course the J2b2a-L283. Its autosomes were more Caucasus shifted which is understandable. But overall, it was part of that emerging kurgan Eneolithic groups from which the Yamnaya and Corded Ware Culture will stem few centuries later.

We haven't yet found the R1a-M417 in the forementioned context. But given that one of oldest R1a-M198 (close to M417) was found in Middle Volga region we can assume that M417 will be found in Steppe Eneolithic / CLV region also. Currently one of oldest R1a-M417 is from Yamnaya layer in Balkans.

Based on this data I will restate my scepticism about the possibility that Yamnaya descend from Sredny Stog located in Ukraine expressed on some boards. Highest diversity of Z2103+ and negative to Z2106 is found in Near East which means that the region from which Z2103 started its expansion must be close to Caucasus. More ancient DNA will hopefully solve this question also.

PS. I didn't include in that Eneolithic Y DNA list samples from Steppe Maykop who have a different Central Asian origin. Also, one of V1636 is dated at 2800BC which is later than the 3000BC. But its autosomes shows that it was from preceding period and not from Core Yamnaya.

See also

• Qpadm models about the Core Yamnaya origins.
• The Yamnaya culture and it's legacy.

The Kura-Araxes culture. An open thread.

Ghalichi et al. 2024 have published 7 new KA ancient samples from Georgia. Site Dzedzevbi near Dmanisi. And two more samples from Velikent Dagestan.

The low coverage didn't permit to find the deep subclades of those samples. But apparently two male samples from Dzedzevbi are J2b2. While the other is J1 almost certainly from the Z1842 branch. The Velikent is also J1.

Overall, there is now 16 Y DNA from known Kura-Araxes layers. 11 of them are J1 from the Z1842 branch. Most cases of J1 are found in regions that are geographically close to eastern Great Caucasian range. Dagestan, Kakhetia, Berkaber in Tavush and one case near Dmanisi. The strong prevalence of J1 is almost certainly the result of founder effect, because in earlier Neolithic periods the J1 was rare.

Moving away from the eastern parts of Caucasian range we see other haplotypes. Like R1b-V1636 from Sevan basin, G2b from Kaps Shirak. J2-M92 from Doghlauri central Georgia. And now two cases of J2b2 from Dzedzevbi. According the Genarchivist activists one of them is from the J2b-FT3464 a minor branch found today in Near East and Europe. While the other J2b was J2b2b-Z2453. An old Neolithic branch found in Shulaveri culture and Hajji Firuz tepe.

What can we deduce from this distribution? As I have already noted the J1 in northeastern parts of Kura-Araxes horizon can be associated with North-East Caucasian (NEC) speakers. But the whole KA horizon couldn't have been NEC speakers otherwise this would have left linguistic traces. For which there is no evidence. The rapid shift in Y DNA distribution when moving away from eastern Caucasus is another strong argument that there was another ethnic group (or groups) in Kura-Araxes.

Who could be this other group(s)?

The two main candidates are the Anatolian speakers and Hurro-Urartians. Currently the data is still too small to connect the dots between South Caucasus and Anatolia/Levant/Mesopotamia where Anatolian Hurro-Urartian languages were spoken. But some patterns are already visible.

See also

• Here is an excerpt from the Petrosyan 2023 paper about the origins of Kura-Araxes culture.
• 
  

I2a2b-Y16419 a possible Ukraine hunter gatherer lineage in Yamnaya/Catacomb?

Lazaridis et al. 2024 had discovered that Yamnaya has some Ukraine Neolithic hunter gatherer ancestry. UNHG. This term can be confusing but it's a result of different naming convention in ex-Sovietic countries, when a hunter-gatherer community gets the Neolithic label if they had a pottery.

Ghalichi et al. 2024 confirmed the presence of some 15% of UNHG in Yamnaya. The I2-L699 regularly found in steppe was derived from UNHG for example.

The presence of I2 in Yamnaya related cultures is an important subject for the Armenian ethnogenesis given the presence of I2a2b in Trialeti-Vanadzor culture.

Till now we didn't have any extra information about this haplotype. The closest European sample to it was found in Eneolithic Croatia with a common ancestor reported to live at 8200BC by the FTDNA. This age is amply sufficient for this lineage to be present both in European farmers and in UNHG.

Ghalichi et al. 2024 had a new sample which probably can help to understand how I2a2b could end up in early Yamnaya communities as a rare lineage. The paper reported a single female sample from near Azov and Black Sea joining region labeled as "Steppe Eneolithic outlier west". KHB dated at 4000BC near Taman peninsula. This sample has large amount of UNHG ancestry (43%). In most likelihood it infiltrated there from Ukraine.

If this outlier represents a regular population living in that region, then this raises the possibility that the I2a2b-Y16419 was also living, there since the Eneolithic and with the formation of Yamnaya/Catacomb became part of those cultures as a rare lineage. Later it moved to south and had a great luck to expand with Trialeti-Vanadzor culture.

For verifying this theory more ancient DNA is needed from the aforementioned region in particular and from south Ukraine in general.

Shulaveri-Aratashen-Shomutepe (SAS) culture. 7000/6200-5300BC.

 Shulaveri-Aratashen-Shomutepe (SAS) culture. 7000/6200-5300BC.

We have five good quality new samples from Shulaveri (SAS) culture in Georgia (Aruchlo). They had YDNA H2, J2a1a and R. This latter is from the R2 haplogroup according Genarchivist activists.

On the PCA three of five farmers plot close to related Neolithic samples from Armenia and Azerbaijan while modern people close them are the Armenians in G25. It's now obvious that this was the main genetic profile in SAS/Shulaveri culture. Those were the first farmers in South Caucasus and their ancestry was largely derived from central regions of Fertile Crescent hence the reason that occasionally we call them Central farmers. Ghalichi 2024 used also the term East Anatolian farmers. Given some archaeological data from Van region we can assume that most of historic Armenia (except probably the most western and southwestern regions) was inhabited by this type of farmers.

Besides this "Central"/"Armenian like" type there were also farmers with higher CHG ratio. First we have seen them in Aknashen from Armenia. Now we have another similar CHG shifted sample from Georgia plotting close to modern Georgians. It's not exactly identic to Aknashen but rather plots close to Darkveti-Meshoko ( labeled as Caucasus Eneolithic ) raising the possibility that Darkveti-Meshoko culture formed as a mixture of Shulaveri and CHG. The Darkveti culture is remarkable because genome wide it's genetic profile looks a good candidate for being Pre-Proto-Kartvelian. High CHG and very low Steppe. But the absence of G2a1a there and scant sampling from west Caucasus makes those suggestions still speculative.

Another sample from Aruchlo/Georgia plots close to CHG hunters. The ARO006 with YDNA R2. Making it a hunter who learned farming without having any significant admixture from those farmers. The presence of such hunter related genetic profile in Shulaveri culture means that there were at last two different languages in SAS. One derived from the first farmers and another derived from the hunters who learned farming. It's remarkable that archaeology supports this dualistic nature of Shulaveri culture. Two different potteries were made in Shulaveri. One of them was Chaff-tempered. The other one was Grit-tempered. Chaff-tempered was almost certainly made by the first farmers who came from southern regions of historic Armenia. While the Grit-tempered was made by groups derived from the local hunters.

It's interesting that both pottery traditions continued in ancient South Caucasus and historic Armenia after the Neolithic period. Grit-tempered was prominent in Sioni (also found in Adablur and Guinchi) culture which evolved in Early and Middle Chalcolithic (5300-4300BC). . We can conjecture that they were CHG shifted. Offcourse this is a just a prediction based on archaeology which can be wrong, given that currently there are no samples from this period. While Chaff-tempered pottery became prominent in Late Chalcolithic period (4300-3600BC). The Late Chalcolithic period DNA both from Armenia and Azerbaijan shows that they were mostly derived from the first Neolithic farmers having some extra new admixtures.

And finally we can now say with high degree of certitude that the CHG shifted genetic profile of Kura-Araxes culture (3600-2400BC) had local origins. When farmers settled all over Kur and Arax river valleys the forest-mountain zone between those two valleys also known as Lesser Caucasus became a sort of refugium where they preserved the initial hunter gatherer ancestry in higher proportion. This is the reason why the oldest radiocarbon dated Kura-Araxes sites are found in north Armenia (Gegharot) and south Georgia.

But this is not the whole story. Apparently Kura-Araxes also had two potteries and genetic profiles. Which could mean that for at last 4000 years two genetic profiles were competing in South Caucasus starting from the Neolithic period till the end of Early Bronze Age.

PS. There was in reality six samples from Aruchlo. But one of them has so little coverage that it was not included in the calculations.

Ghalichi et al. 2024 and the Indo-European question.

 Ghalichi et al. 2024 and the Indo-European question.

A new paper with more than 100 samples provided new details about the interaction of Eastern European populations and West Asians over Caucasus.

A new Mesolithic sample from north Caucasus (Satanae cave SJG001) dated at 6100BC yielded an interesting genetic profile. It was practically identic to other Eastern European hunter gatherers (EHG). The sample had the R1a haplogroup, from a minor branch (not the same as M417) that in most likelihood is related to later IE migrations.

This EHG result from north Caucasus means that the genetic profile of Yamnaya didn't existed since the immemorial times but formed later than the 6000BC. Yamnaya has ancestry both from EHG and from south of Caucasus. The event that triggered the formation of Yamnaya better known as Steppe genetic profile was the apparition of farmers in South Caucasus. The Shulaveri-Aratashen-Shomutepe (SAS) culture starts no later than 6200BC. And starting from this period farmers mixed both with Caucasian hunter gatherers and made attempts to colonize the steppe in north of Caucasus.

A new case of J2b2a-L283 was found in steppe zone of North Caucasus (the ZO1002 on the map). It was dated at 3800BC, older than Maykop and Kura-Araxes. And already had some EHG admixture which means that it was probably a lineage descending from Shulaveri farmers trying to settle in steppe. Later this haplotype will become a rare Yamnaya lineage and will move to Balkans where it will have an important expansion.

Despite this finding the most predominant Y DNA in Eneolithic steppe pastoralistic groups had local Eastern European origin (R1 and I2) rather than from south of Caucasus. We have another Y DNA from Nalchik farmers and this time once more it's R1b. From the V1636 branch. We have also now one of oldest R1b-M269 from Konstantinovka dated at 3800BC. KST001.

So two important question arises..

Why despite having large amount of West Asian/Caucasian ancestry the Eneolithic steppe groups and Yamnaya has so little cases of J haplogroup.

And the second question is linguistic. Who's language was the Proto Indo Anatolian? Those from Shulaveri culture farmers or those from EHG / R1 side.

Before answering the first question we should note that this phenomenon of mismatch between autosomes occurred not only in East Europe but also in West Europe. After the initial success of G2 haplogroup , later there was a widespread resurgence of local I2 haplogroup. In British islands farmers for instance had 100% of I2 haplogroup. Yet in autosomes they had 70% of Anatolian farner ancestry.

There can be numerous theories to explain this. I am personally inclined to believe that social factors played a key role. Neolithic farmers of West Asia had more egalitarian or even matriarchal culture. A recent paper from Catalhoyuk showed an obvious matrilineal culture. While eastern European hunters could have more patriarchal culture. When a matriarchal and patriarchal cultures meet each other this affects in different ways the uniparental markers. Y DNA of one side will prevail and mtdna of the other side will also prevail.

There are other theories like mass killing of males. Those theories are popular in internet. But they can't explain one important thing. Why Scythians, Saka and Turkic tribes who are known to have a warrior culture and where frequently killing all males of the competing clan had one of most diverse Y DNA. The answer is self evident. Because during a conflict it's not granted that always one haplogroup prevails while others always are defeated. The outcome of the clans conflict can be very random which will create a very random distribution of haplogroups.

When farmers moved to north of Caucasus they were the technologically advanced side. It would be strange that they lost all battles without wining any of them and expanding their lineage. So wars and mass killings can't explain the emergence of this pattern. There can be other explanations. I just demonstrated one possible.

As for the linguistic part I will dedicate another topic

According to the 12th-century historian Vardan Areveltsi, the original name of Van city was Yervandavan.

 According to the 12th-century historian Vardan Areveltsi, the original name of Van city was Yervandavan. Historically, however, the city was first called Tushpa, likely pronounced as Tosp, which later named the surrounding Tosp canton where Van is located. Interestingly, Greek and Roman sources don’t unambiguously mention “Van,” though “Tosp” is well-documented.

Vardan’s claim may have rational roots: a nearby canton was named Yervandunik’ ("land of Yervanduni"), which was also found in the Araratian plain, representing hereditary lands of the Yervanduni dynasty. Tosp thus became their capital, possibly leading to a new name. After Yervanduni’s decline, the city’s name evolved from the local parallel Biaina term (pronounced as Vayn) into Van.

Tosp was the royal city of Biaina dynasty (term used by Grekyan), or better known as Urartu’s rulers. Based on king names the Yervanduni and Biaina dynasties were distinct groups. A power shift from the Biaina to Yervanduni dynasties occurred at some point, though details remain unclear due to a lack of records. Archaeology and paleogenetics in Tosp/Van may eventually clarify this dynastic shift, often mistakenly seen as a transition from Urartu to Armenia

While we are waiting for the publication of new ancient DNA paper about Caucasus with more than 100 samples here are some preliminary remarks.

 While we are waiting for the publication of new ancient DNA paper about Caucasus with more than 100 samples here are some preliminary remarks.

All 6 Y dna from the site Guinchi in Dagestan (Shamil region) are J1. Some with good quality are BY100. Which is the branch popular in northeastern Caucasus.

They are apparently from Kura-Araxes culture period. They have medium level steppe ancestry which they obviously got from the preceding Chalcolithic period.

This further strengthen the theory that Nakh-Dagestani speakers descend from the Kura-Araxes culture that expanded in northeast of Caucasus after the 3500BC. We can say that this is not anymore a theory but a proven fact.

Linguistically the age of Proto-NEC is close to that date. The moderate presence of steppe ancestry also explains why modern Daghestani people have steppe ancestry. This is because they had it from the start.

Nevertheless modern levels of steppe ancestry are twice higher than in Early Bronze Age. Which means that modern Daghestani people got extra steppe ancestry later also. The prime candidate is obviously the Catacomb culture which moved to south Caucasus after the 2500BC. Another source of steppe ancestry are the later Iranic and Turkic nomadic incursions in that region.

Despite this advance still some questions remain unsolved. Nakh branch has less steppe compared to Daghestani people. They are also predominantly J2. Which could mean that Nakh people region had an extra influx from south which plausibly introduced that J2.

The origins of Lezgin branch remains unsolved. Whether they moved from north to south or descend directly from the Kura-Araxes people who survived on the southern slopes of Great Caucasian range is unknown.

We can now say with certitude that Udi people linguistically related to Lezgin group are a genetic outlier in NEC family. They have too much Armenian Highlands farmer related ancestry which they most probably got from Armenians. Given their Christian faith was the same as Armenians.

And finally. Despite the close connection of NEC family with Kura-Araxes there is no evidence that the whole Kura-Araxes horizon was speaking NEC related languages

A list of Assyrian cuneiform texts where the term Muški is mentioned.

 A list of Assyrian cuneiform texts where the term Muški is mentioned. The oldest one is dated at the first quarter of 12th century BC. Hittite empire's last king Šuppiluliuma II ascended to the throne at 1207BC which means that there is virtually no time window for Mushki to start from Balkans, cross the whole Anatolia and reach Alzi region (near Sassun) to be attested by Assyrians. Even skipping the time window issue such a sprint running migration is nonsensical from the economic and political perspective.

In conclusion even without ancient DNA the Muški's original homeland in Balkans was untenable. It's now impossible with the advent of genetics.

The source Karen Radner's review

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.

Dragon stone burial from Lchashen has now the G25 coordinates. See the comments. Bobokhyan et al. 2024.

 Dragon stone burial from Lchashen has now the G25 coordinates. See the comments. Bobokhyan et al. 2024..

Two newborn kids and a woman ( supposedly their mother ) were found in an unique kurgan burial with Dragon stone deposed on the tomb. Their autosomes is similar to the contemporary people who lived in that period in ancient Republic of Armenia. The burial is dated around 1550BC, 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

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

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