Genetic History of Anatolia during Holocene

[eupator]

Goodness knows what Herodotus meant by Persians, though. Persians as in straight from Persia, or Persians as in people in the Persian Empire, which would have included Anatolians?

Ethnography was not exactly his forte, as we have discovered in his writings about the Etruscans.

Also, the spread was pretty extreme, from ten or less to a few with 20%, and the Caucasus/Iran Neo in Iranians looks like 80% in some of them, so I guess it depends on what you mean by kinship.

Kinship as in there were blood relations between them.

20% top end means it's almost 1/4 grandparent. It's quite significant, evident also in the J uniparentals (and G for the Caucasus).

Anatolia stands in between the Iranian Plateau/Caucasus and the Aegean coastline in Greece, there is a gradient that is visible in autosomals.

The Pelasgian substratum is of Anatolian (major) + Irano-Caucasian (minor) extraction, then you add the EHG signal from the north, Herodotus is being vindicated almost to a tee in what he wrote about his contemporaries in Greece.

 

[torzio]

it still a T you should be happy
i see in this paper again there is no E
even the southern arc paper had some :unhappy:

of course .............same as the 28000 year old T's found in western China ..............all count

 

[eupator]

As for who the Persians were, Herodotus mentions that they were comprised of many tribes that were united by Cyrus the Great.

A lot of them came from the Medes, who were called the Aryans, then the Pasargádai, Maráphioi, Máspioi, Panthialaîoi and Germánioi.

Cyrus himself, the uniter of the Persian ethnos, was the founder of the Achaemenids, whose patrilineal genealogical trace went back all the way to Perses and his father Perseas, the founding father of the Mycenaeans.

[1.125.3] ἔστι δὲ Περσέων συχνὰ γένεα, καὶ τὰ μὲν αὐτῶν ὁ Κῦρος συνάλισε καὶ ἀνέπεισε ἀπίστασθαι ἀπὸ Μήδων· ἔστι δὲ τάδε, ἐξ ὧν ὧλλοι πάντες ἀρτέαται Πέρσαι· Πασαργάδαι, Μαράφιοι, Μάσπιοι· τούτων Πασαργάδαι εἰσὶ ἄριστοι, ἐν τοῖσι καὶ Ἀχαιμενίδαι εἰσὶ φρήτρη, ἔνθεν οἱ βασιλέες οἱ Περσεῖδαι γεγόνασι· [1.125.4] ἄλλοι δὲ Πέρσαι εἰσὶ οἵδε· Πανθιαλαῖοι, Δηρουσιαῖοι, Γερμάνιοι·

 

[Dianatomia]

Am I to understand that there were people in the Aegean area with higher steppe content than the Mycenaeans? And that they came through the Southern route while we think that the Greek tribes were later arrivals through the Balkans?

EDIT: Never mind about EHG coming through Anatolia. I just reread the summary.

No, the Greek speakers came earlier than assumed. And they did come from the North. Why Mycenaeans had less steppe is also a puzzle to me. Perhaps people with less steppe also arrived in Greece. From the islands perhaps? Or some natives fled to the mountains/islands when the first newcomers arrived, after which some came back after a few centuries. Who knows? Or the the specimens we have so far are not representative for Mycenaean Greeks. The Italian Greeks may have Sicilian admixture. And the Greek from Empuries was intermixed with people from Asia Minor.

In any case, these new specimens might be more closer to Log 1 and Log 2 further North.

 

[Angela]

^^Did you look at the PCA? A good number of them still plot with Aegean Neolithic, which is almost indistinguishable from Anatolian Neolithic. They're the ones from the Peloponnese, from which the Mycenaean culture radiated.

The one sample which pulls away is from Thessaly.

As with the Slavic admixture, the steppe admixture seems to be on a north/south cline.

I'm sure we'll see something similar with Classical Greeks.

The heartland of the illustrious Greek civilization and its accomplishments seems to be more "southern".

 

[bigsnake49]

No, the Greek speakers came earlier than assumed. And they did come from the North. Why Mycenaeans had less steppe is also a puzzle to me. Perhaps people with less steppe also arrived in Greece. From the islands perhaps? Or some natives fled to the mountains/islands when the first newcomers arrived, after which some came back after a few centuries. Who knows? Or the the specimens we have so far are not representative for Mycenaean Greeks. The Italian Greeks may have Sicilian admixture. And the Greek from Empuries was intermixed with people from Asia Minor.

In any case, these new specimens might be more closer to Log 1 and Log 2 further North.

From Googling the toponyms, I have located the them to Thessaly for Theopetra, Corinthia in Peloponnese for Perachora and around Heraklion in Crete for Sarakinos. Does anybody have more accurate locations?Is there a map in the Thesis?

Never mind it is Figure 5 in the pdf.

 

[Angela]

As for who the Persians were, Herodotus mentions that they were comprised of many tribes that were united by Cyrus the Great.

A lot of them came from the Medes, who were called the Aryans, then the Pasargádai, Maráphioi, Máspioi, Panthialaîoi and Germánioi.

Cyrus himself, the uniter of the Persian ethnos, was the founder of the Achaemenids, whose patrilineal genealogical trace went back all the way to Perses and his father Perseas, the founding father of the Mycenaeans.

Yes, I know. I can read too.

It was a rhetorical question, as in, some of the "Persians", or people in the Persian confederation were indeed pretty similar to the Greeks, and some most decidedly were NOT. With such a general statement, people could get confused.

 

[bigsnake49]

^^Did you look at the PCA? A good number of them still plot with Aegean Neolithic, which is almost indistinguishable from Anatolian Neolithic. They're the ones from the Peloponnese, from which the Mycenaean culture radiated.

The one sample which pulls away is from Thessaly.

As with the Slavic admixture, the steppe admixture seems to be on a north/south cline.

I'm sure we'll see something similar with Classical Greeks.

The heartland of the illustrious Greek civilization and its accomplishments seems to be more "southern".

It was always more southern whether it was the Minoan civilization, the Cycladic civilization, the Mycenaean civilization or the classical Athenian civilization. Not to diminish the material civilization of the more northern tribes.

 

[Angela]

Kinship as in there were blood relations between them.

20% top end means it's almost 1/4 grandparent. It's quite significant, evident also in the J uniparentals (and G for the Caucasus).

Anatolia stands in between the Iranian Plateau/Caucasus and the Aegean coastline in Greece, there is a gradient that is visible in autosomals.

The Pelasgian substratum is of Anatolian (major) + Irano-Caucasian (minor) extraction, then you add the EHG signal from the north, Herodotus is being vindicated almost to a tee in what he wrote about his contemporaries in Greece.

As someone with no dog in the hunt, as they say, I see 2/7 Bronze Age samples with about 1/4 CHG, 1/7 with about 15%, and 4 with none.

Again, it's structured by area.

Whether with time it got more homogeneous I don't know. We would need Dark Age and Classical Era samples from lots of parts of Greece to figure that out.


I find it interesting that there's Turkic in Anatolian samples already in the late Roman period. I wonder how widespread that was?

 

[Dianatomia]

^^Did you look at the PCA? A good number of them still plot with Aegean Neolithic, which is almost indistinguishable from Anatolian Neolithic. They're the ones from the Peloponnese, from which the Mycenaean culture radiated.

The one sample which pulls away is from Thessaly.

As with the Slavic admixture, the steppe admixture seems to be on a north/south cline.

I'm sure we'll see something similar with Classical Greeks.

The heartland of the illustrious Greek civilization and its accomplishments seems to be more "southern".

They are still pulled towards Mycenaeans, who were approx 15% steppe, but these ones are more like 18% to 25%. I think Crete_Armenoi (also higher in Steppe) was somewhat neglected in the earlier study. Clearly there were some Mycenaeans with higher Steppe levels. There was considerable variation between Greeks in the Mycenaean Age. The sample which pulls away is not from Thessaly, it's from Boeotia which is South of Thessaly and not too far from Attica. Achilles who was described to be blonde (in whatever context) was from Phthia, Thessaly. He could have been light haired due to higher Steppe ancestry in his region during that time. Seems that Homer didn't make it up.

You are right. The heartland of the Greek civilization is more Southern. Could be due to the fact that it was connected to Crete and also due to a better climate. The area which is suspected to be proto-Hellenic (Thessaly, Epirus, part of Macedonia) seems to be more Steppe admixed. Even during the Early Bronze Age.

 

[Philjames100]

It's interesting that the sample from Theopetra with steppe ancestry had Y-DNA I2a2a1b.

I2a2a1b1b1 was also found in two samples with steppe ancestry from the Swat valley c.1000 BC, published in Narasimhan et al. 2019.

Does anyone know anything more about this haplogroup?

 

[Leopoldo Leone]

^^Did you look at the PCA? A good number of them still plot with Aegean Neolithic, which is almost indistinguishable from Anatolian Neolithic. They're the ones from the Peloponnese, from which the Mycenaean culture radiated.

The one sample which pulls away is from Thessaly.

As with the Slavic admixture, the steppe admixture seems to be on a north/south cline.

I'm sure we'll see something similar with Classical Greeks.

The heartland of the illustrious Greek civilization and its accomplishments seems to be more "southern".

Why some people are surprised that early proto-greeks were steppe heavy is something that escape me, since it is to be expected that the first IE arrivals would have had higher steppe ancestry than subsequent ones for the latter were the result of the former mixing with pre-IE speaking groups.
As for the idea that during the classical age Thessalians were logkas-like, it is impossible since even Thracians were Mycenaean-like (and I'd specify to the most steppe rich end of the mycenean samples, as classical Greeks were), which means that it was the common genetic profile in the area independent of ethnic affiliations ("Paeonians", as the samples from today's FYROM are addressed on these sites, do seem like an intrusive element and they likely are), though one could expect a very minor north-south gradient in steppe ancestry.
How I see it Mycenaeans primarily formed in north Greece between 2200 and 1900/1800 BC, when logkas-like people settled in the area and mixed with either pechora-like people or Minoan-like people, then during 1900/1800 BC (a little before the shaft grave period) they descended on the rest of Greece largely replacing whomever had been living there.
In support for this model, I can cite the last paper on Himera, since they model the Himera_1 group with around 20% steppe, which is similar to the Mycenaean samples but to the steppe-rich end rather than the centre (around 10%); indeed they modelled them also using Greece_LBA (which I mean they used the average) and it turns out that in such model they need a 20% Balkan_IA admixture; since the idea of a migration from the Balkan at the end of the bronze age into Greece has been discarded recently by archaeology, I prefer the idea that mostly people on the higher end of steppe ancestry (15-20%) left more offspring during the Mycenaean period, likely because they were the ones that conquered the area.

 

[Leopoldo Leone]

They are still pulled towards Mycenaeans, who were approx 15% steppe, but these ones are more like 18% to 25%. I think Crete_Armenoi (also higher in Steppe) was somewhat neglected in the earlier study. Clearly there were some Mycenaeans with higher Steppe levels. There was considerable variation between Greeks in the Mycenaean Age. The sample which pulls away is not from Thessaly, it's from Boeotia which is South of Thessaly and not too far from Attica. Achilles who was described to be blonde (in whatever context) was from Phthia, Thessaly. He could have been light haired due to higher Steppe ancestry in his region during that time. Seems that Homer didn't make it up.

You are right. The heartland of the Greek civilization is more Southern. Could be due to the fact that it was connected to Crete and also due to a better climate. The area which is suspected to be proto-Hellenic (Thessaly, Epirus, part of Macedonia) seems to be more Steppe admixed. Even during the Early Bronze Age.

It is really doubtful that higher steppe in Greece correlates with higher rates of blondism, since it does correlate in the rest of Europe because steppe ancestry correlates with bell beaker admixture who had higher rates of blondism but oddly enough thanks to their globular amphora admixture (EEF); it is why indirectly steppe admixture in the rest of Europe correlates with higher rate of blondism but according to genetic studies EEF ancestry directly correlates with blondism in Estonians (furthermore blondism in north Europe continued to be selected for in the following centuries).
In Greece steppe ancestry was mediated by individuals who were the result of catacomb mixing with local balkanite farmers (serbian according to what I've seen on g25), who contrary to global amphora didn't develop high rates of blondism.

 

[enter_tain]

It is really doubtful that higher steppe in Greece correlates with higher rates of blondism, since it does correlate in the rest of Europe because steppe ancestry correlates with bell beaker admixture who had higher rates of blondism but oddly enough thanks to their globular amphora admixture (EEF); it is why indirectly steppe admixture in the rest of Europe correlates with higher rate of blondism but according to genetic studies EEF ancestry directly correlates with blondism in Estonians (furthermore blondism in north Europe continued to be selected for in the following centuries).
In Greece steppe ancestry was mediated by individuals who were the result of catacomb mixing with local balkanite farmers (serbian according to what I've seen on g25), who contrary to global amphora didn't develop high rates of blondism.

It's not a 1:1 correlation for looks vs. steppe:eef, but the true ancient Attic/Ionic Greeks (ones with no or little steppe) would have looked like Middle Eastern people today. Colloquially, they would not really be considered "white" even though they were European genetically.

Obviously as Dorian/Dorian-like and Illyrian DNA sipped into Greeks it would increase incidence of light hair. This would even more so be the case for Slavic people, especially in Greek Macedonia.

 

[Francesco]

It's not a 1:1 correlation for looks vs. steppe:eef, but the true ancient Attic/Ionic Greeks (ones with no or little steppe) would have looked like Middle Eastern people today. Colloquially, they would not really be considered "white" even though they were European genetically.

Obviously as Dorian/Dorian-like and Illyrian DNA sipped into Greeks it would increase incidence of light hair. This would even more so be the case for Slavic people, especially in Greek Macedonia.

Just to clarify the obvious: we don't have many samples of Attc/Ionic Greek from the Iron Age, but their predecessor from the bronze age looked, not surprisinlgy, southern european. Doubt the Doric invasion brought so much change, since the few samples from Iron age Greeks are pretty similar to their bronze age ancestors.

https://www.eupedia.com/forum/threa...ransect-(Mesolithic-to-Medieval)-from-Biomuse

 

[real expert]

It is really doubtful that higher steppe in Greece correlates with higher rates of blondism, since it does correlate in the rest of Europe because steppe ancestry correlates with bell beaker admixture who had higher rates of blondism but oddly enough thanks to their globular amphora admixture (EEF); it is why indirectly steppe admixture in the rest of Europe correlates with higher rate of blondism but according to genetic studies EEF ancestry directly correlates with blondism in Estonians (furthermore blondism in north Europe continued to be selected for in the following centuries).
In Greece steppe ancestry was mediated by individuals who were the result of catacomb mixing with local balkanite farmers (serbian according to what I've seen on g25), who contrary to global amphora didn't develop high rates of blondism.

Well, according to Reich, Hanel, and Carlberg and recent papers about selection and admixture in Ancient Eurasians and Neolithic Europeans, the Steppe people introduced blond hair to Europeans too. But another source for light hair was the EEF ancestry. Moreover, the earliest known individual with the genes for blond hair was an ANE female from 16130-15749 BC. Reich concludes that the KITLG gene for blond hair was brought to Europe in a population migration wave from the Steppe. Hanel and Carlberg also suggest that Steppe folks were responsible for contributing this gene to Europeans. And Mathieson, argued that the genes for blond hair were derived from ANE population, too. With that being said, blond hair does not by default correlate with high Steppe ancestry. Hence someone with low Steppe input can of course exhibit blond or light brown hair too. Anyway, as the Hanel&Carlberg study showed the strong spread of Steppe pastoralists likely caused the rapid selective sweep in European populations towards light skin and hair.

For instance, among the few who were blond in this current Classical period Greek army study, was this Baltic-like individual who happened to be pale, blue-eyed, and blond:

“This observation of the derived allele in I10943/W0396 agrees with his genetic ancestry, with the greatest affinity to BA and IA Eastern Baltic groups. Notably, this individual also has an uncommon physical appearance among the studied samples, with HirisplexS estimating highest probabilities for a combination of blond hair, blue eyes and pale skin. This phenotype also rose in frequency in the Eastern Baltic in the 1st millennium BCE”

 

[Silesian]

Well, according to Reich, Hanel, and Carlberg and recent papers about selection and admixture in Ancient Eurasians and Neolithic Europeans, the Steppe people introduced blond hair to Europeans too. But another source for light hair was the EEF ancestry. Moreover, the earliest known individual with the genes for blond hair was an ANE female from 16130-15749 BC. Reich concludes that the KITLG gene for blond hair was brought to Europe in a population migration wave from the Steppe. Hanel and Carlberg also suggest that Steppe folks were responsible for contributing this gene to Europeans. And Mathieson, argued that the genes for blond hair were derived from ANE population, too. With that being said, blond hair does not by default correlate with high Steppe ancestry. Hence someone with low Steppe input can of course exhibit blond or light brown hair too. Anyway, as the Hanel&Carlberg study showed the strong spread of Steppe pastoralists likely caused the rapid selective sweep in European populations towards light skin and hair.

For instance, among the few who were blond in this current Classical period Greek army study, was this Baltic-like individual who happened to be pale, blue-eyed, and blond:

Upper Palaeolithic genomes reveal deep roots of modern Eurasians

https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4660371/#S1
Yamnaya--Afanasievo(R1b-Z2109+) are a combination of EHG(low in SLC24A5 light skin) and CHG(tend to have black hair, light skin). They scored relatively the same in light hair and light(HERC2/SLC24A5) skin compared to Anatolian samples. All things be equal, oldest blonde is from Steppe-ANE female from 16130-15749 BC.. HERC2/SLC24A5 can be found in CHG

.... Direct AMS dating of the bone yielded an estimated date of 13,300 BP for the age of the bone...The Satsurblia individual is genetically closest to an ancient individual, dating to around 9,700 BP, found at the Kotias Klde rock shelter in Georgia. Together, they form a genetically distinct cluster referred to as Caucasus Hunter-Gatherer (CHG).^[8][9]... The ancient individual from Satsurblia was male with black hair and brown eyes; however, the individual is one of the earliest found to carry the derived HERC2 allele for blue eyes. The Satsurblia individual also likely had light skin, as he was found to carry the derived SLC24A5 allele for light skin. The Satsurblia individual belongs to mtDNA Haplogroup K3 and Y-DNA Haplogroup J1-Y6313*.^[10][11][8] About 1.7-2.4% of the Satsurblia individual's DNA was Neanderthal in origin.^[9]

//The derived threonine allele (Ala111Thr; also known as A111T or Thr111) represented 98.7 to 100% of the alleles in European samples, while the ancestral or alanine form was found in 93 to 100% of samples of Sub-Saharan Africans, East Asians and Indigenous Americans. The variation is a SNP polymorphism rs1426654, which had been previously shown to besecond among 3011 tabulated SNPs ranked as ancestry-informative markers. This single change in SLC24A5 explains between 25 and 38% of the difference in skin melanin index between peoples of sub-Saharan African and European ancestry.^[5]The SNP rs2470102 independently affects skin pigmentation variation among the South Asian population.^[16]
Furthermore, the European mutation is associated with the largest region of diminished genetic variation in the CEU HapMap population, suggesting the possibility that the A111T mutation may be the subject of the single largest degree of selection in human populations of European ancestry.^[5] It is hypothesized that selection for the derived allele is based on the need for sunlight to produce the essential nutrient vitamin D. In northerly latitudes, where there is less sun, greater requirement for body coverage due to colder climate, and frequently, diets poor in vitamin D, making lighter skin more suitable for survival.^[17]The earliest known sample of the threonine allele is 13,000 years old from Satsurblia Cave in Georgia.^[18] The allele was widespread from Anatolia to Ukraine and Iran at the beginning of the Neolithic.^[19][20][21]

 

[Angela]

Why some people are surprised that early proto-greeks were steppe heavy is something that escape me, since it is to be expected that the first IE arrivals would have had higher steppe ancestry than subsequent ones for the latter were the result of the former mixing with pre-IE speaking groups.
As for the idea that during the classical age Thessalians were logkas-like, it is impossible since even Thracians were Mycenaean-like (and I'd specify to the most steppe rich end of the mycenean samples, as classical Greeks were), which means that it was the common genetic profile in the area independent of ethnic affiliations ("Paeonians", as the samples from today's FYROM are addressed on these sites, do seem like an intrusive element and they likely are), though one could expect a very minor north-south gradient in steppe ancestry.
How I see it Mycenaeans primarily formed in north Greece between 2200 and 1900/1800 BC, when logkas-like people settled in the area and mixed with either pechora-like people or Minoan-like people, then during 1900/1800 BC (a little before the shaft grave period) they descended on the rest of Greece largely replacing whomever had been living there.
In support for this model, I can cite the last paper on Himera, since they model the Himera_1 group with around 20% steppe, which is similar to the Mycenaean samples but to the steppe-rich end rather than the centre (around 10%); indeed they modelled them also using Greece_LBA (which I mean they used the average) and it turns out that in such model they need a 20% Balkan_IA admixture; since the idea of a migration from the Balkan at the end of the bronze age into Greece has been discarded recently by archaeology, I prefer the idea that mostly people on the higher end of steppe ancestry (15-20%) left more offspring during the Mycenaean period, likely because they were the ones that conquered the area.

I agree with the bolded comment.

However, I don't know who the people are who are surprised at the steppe in the early samples from this paper. As you say, early arrivals are going to carry more steppe, but also the number of new arrivals may diminish the further south they go, as was the case with the Slavic speakers. Nor do I think 15-25% is steppe "heavy". Northeast Europeans are steppe heavy, and early Corded Ware was steppe heavy. 15-25% are Southern European like levels.

Whether the original Greek speakers were Logkas like (if I'm understanding you) or not I don't know, but what I do know is that the Mycenaeans genetically were lower in steppe on average than Logkas, whether because there were fewer steppe admixed people the further south they went or there were more locals or a combination of both I don't know. I highly doubt there was a "replacement" of the Neolithic like people. There's way too much Neolithic/early Minoan like ancestry in the Mycenaeans. What we also know is that the Mycenaean "culture" clearly formed in the Peloponnese, either by absorbing local elements and/or early contact with the advanced civilization of the Minoans, and then radiated outwards from there.

Also, your idea that people on the higher end of steppe ancestry left more descendants because of a class system is questionable in my mind as it is contradicted by the results from the Lazaridis paper and even by the authors of this paper. The higher status samples among the Mycenaeans did not necessarily have higher steppe. This does not at all have the trademark signs of a "conquest", whatever the fantasies of Eurogenes and people of his ilk.

 

[Angela]

I am NOT going to go down the rabbit hole once again with Northern Europeans determined to believe that steppe people were blonde and brought blondism to the west. Believe what you want.

However, it just isn't true.

The steppe people were NOT blonde. We've had study after study showing they were predominantly dark haired and eyed and not particularly light skinned either. Catacomb people, in particular, were darker than ANY modern Europeans, to quote the author of the study. Do we really have to keep posting the study to prove it, or the one which shows that regardless of where blonde hair can be found in isolated areas in the past, the blonde hair and fair skin carried and spread by ADMIXED people like Corded Ware can be found on the NEOLITHIC EUROPEAN part of the genome, and were obviously therefore picked up from people of cultures like Globulara Amphora.

Look the papers up and read them again, for God's sake.

This is just one of those wrong-headed ideas which Northern Europeans and even some Nordicists in Southern Europe refuse to let go, no matter what the science shows. So, as I said, believe what you want. It's irrelevant to me. This hobby is full of people who refuse to accept the science in the papers because it goes against long held beliefs.

Also, if the ancients were anything like their modern descendants in Southern Europe, the definition of "blonde" is different here than it is in the north. If you have light brown hair, you're a "bionda". The presence of platinum hair a la the Targaryens is unheard of once early childhood has passed.

 

[Silesian]

I really like to look into the Anatolian-Hittite word for wine, since no connection can be made between Anatolia and early R1b samples Corded Ware in Poland(aka horseless ware) or Eastern Hunter Gatherers-both being darker phenotypes(Compare R1b-Z2109 Yamnaya/Afanasievo and by extension Dom2 horses in Catacomb-- Upper Palaeolithic genomes reveal deep roots of modern Eurasians.
Chapter 10 The Word for Wine in Anatolian, Greek, Armenian, Italic, Etruscan, Semitic and Its Indo-European Origin

https://brill.com/view/book/edcoll/9789004413122/BP000013.xml

Abstract

It can be shown that Hittite u̯ii̯an(a)- ‘grapevine; wine’ continues a Proto-Indo-European nomen agentis *u̯éi̯h1-on-/ *u̯ih1-n-́ ‘twiner, creeping plant, grapevine’, and that this stem formation provides the derivational base for further Indo-European words for ‘wine’ such as Latin vīnum < *u̯ih1-n-ó- and Greek οἶνος < *u̯ói̯n-o- < *u̯ói̯(h1)-n-o-, with the corresponding affiliation adjective preserved in Armenian gini ‘wine’ < *gu̯īníi̯o < *u̯oi̯n-ih2ó-. The morphological structure of the respective base word and its derivatives corresponds to regular Proto-Indo-European patterns of word formation and is therefore congruous with the assumption of an Indo-European origin of the lexemes in question. Accordingly, it is likely that the etymologically isolated Semitic *u̯ai̯nu ‘wine’ was borrowed from Mycenaean Greek and that Proto-Kartvelian *γu̯ini̯o ‘wine’ is a loanword from Proto-Armenian or its prestage.