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Wednesday 26 April 2023

Ancient Greek ancestry in Greeks and Italians through G25 and qpAdm Analysis

Blog moved here: https://genesoftheancients.wordpress.com/

Related posts to read (click to open):

  1. Attempting to improve the qpAdm models from the new study: A genetic history of the Balkans from Roman frontier to Slavic migrations
  2. Did ancient Greeks leave a genetic impact on West Anatolia? qpAdm and G25 analysis
  3. Modeling the Neolithic ancestry of Modern and Ancient Europeans/West Asians using qpAdm, including ancient Greeks/Mycenaeans, Minoans, Hittites, Phoenicians, Yamnaya
  4. Modelling the Roman era sample from Marathon Greece on qpAdm and G25
  5. What is the DNA of the Cypriots?
  6. Ancient Greek skulls from Athens compared to modern populations

Updated 7/12/2023

Ancient Greek ancestry in Greeks and Italians through G25 and qpAdm Analysis

In this post, I will examine the genetic makeup of modern populations in comparison to the ancient Greeks, analyzing how their DNA differs and identifying any similarities that may exist.

Who is most similar genetically today to the ancient Greeks and Minoans? According to the study:
We estimated FST of Bronze Age populations with present-day West Eurasians, finding that Mycenaeans are least differentiated from populations from Greece, Cyprus, Albania, and Italy (Fig. 2), part of a general pattern in which Bronze Age populations broadly resemble present-day inhabitants from the same region (Extended Data Fig. 7).

Roman era Greek admixture of modern Greeks and Italians (updated 7/12/2023)

The Global25 admixture charts you see below illustrate the percentage mixtures needed to get the closest match to a modern target when comparing their DNA to ancient DNA samples. In simpler terms, they show how much mixing of different ancient DNA samples is required to match the genetic profile of present-day populations. This doesn't necessarily imply a direct descent from the exact population depicted; it could instead be from a population closely related or similar to the one shown.


Model without Armenian proxy because it seems to overfit on Dodecanese.
Italians

The models are focused on the direct ancestral profiles, that's why i use the Roman-era Aegeans. It's easier to trace indirect admixtures that way.

Source populations:
 

Balkan Thraco-Roman (Early Vlach and Early Albanian) vs Eastern Roman Aegean model:


 

Admixture of Thraco-Romans:

The Thraco-Romans are probably the ancestors of the Albanians and Aromanians/Vlachs. The fully Latinized ones are the Vlachs while the not fully Latinized ones are the Albanians.
 
Are the Vlachs Latinized or do they have admixture from Latin speaking people from Italy? 
The answer is not certain but Roman era Italy, from Rome to Sicily had a significant Latinized Greek population. Medieval Albania shows admixture from Roman era Greeks but it is hard to say if it came from Italy or Greece. The ALB_MA samples score some Republic era Latin admix but it is not certain if it's an overfit or real. I would need more samples to make a more precise model. The optimal distances in big sample averages must be around 1% but ALB_MA consists of only 3 samples and the distance is above 1.5%.

Multiple possible models with small variations.
 
 

Admixture of the Aegean Eastern Romans: 

Models need to be improved, it's harder to model them with precision with this model because of too few samples.

Admixture of the Hellenistic-Roman era Greeks:

Click this to see the models below recreated on qpAdm: Did ancient Greeks leave a genetic impact on West Anatolia? qpAdm and G25 analysis

The Roman era sample from mainland Greece Attica was identical to the Roman era Greeks of Mugla West Anatolia: Modelling the Roman era sample from Marathon Greece on qpAdm and G25

According to this G25 admix model they were:
37.6% Mycenaean
21.8% Pre-Greek West-Central Anatolian
25% East and South East Anatolian
15.6% North Levantine


This is a G25 neolithic chart so keep in mind its not as accurate as qpAdm. I will remake this chart with qpAdm.

G25 genetic distance heatmaps to modern populations:


Distance to Roman era Greeks 2000 years ago from South Greece and West Anatolia

Distance to Mycenaeans from South West Peloponnese. They were the same genetically as Mycenaeans from Attica.
Distance to Mycenaeans from North West Peloponnese.
 
What do i use to represent Roman-era Greeks?
I used this:
South_Greece/West_Anatolia(Roman-era_1-200AD),0.1087389,0.1527357,-0.0335888,-0.0700049,0.0004309,-0.0225901,0.0002193,-0.0049076,-0.0057949,0.0243225,0.0046767,0.0034469,-0.0066204,0.0031103,-0.0139793,-0.0054362,0.0047721,0.0004729,0.0040811,-0.0070116,-0.0044421,0.0023824,-0.0023993,0.0029722,0.0008222

The G25 coordinate above is the average of these samples: https://pastebin.com/c9QYBQm7

The average consists of DNA samples from West Anatolia and Imperial Rome that are 2000 years old. I used 30% of the Imperial Rome samples that were 95-100% identical to the samples from West Anatolia and Marathon Southern Greece dated to around 300 AD. 

The Marathon sample's coordinates are not good (high distances and noise) which is why i have to use the other samples from Imperial_Rome and West Anatolia as a proxy for Late Hellenistic and Roman-era Greek ancestry. But i managed to get a good model for Marathon without overfits, as you can see in the chart below its identical to the Greeks from West Anatolia and Rome.



Click on this link to read about Greek ancestry in West Anatolia: Did ancient Greek leave a genetic impact on West Anatolia? qpAdm and G25 anaylisis

Contrary to the misconception by some people that Greeks have never established settlements in Western Anatolia, historical evidence reveals not only widespread Greek colonization in the region but also mixing between Greeks and Anatolians, even in areas beyond Greek control.

According to Strabo, Carians, of all the "barbarians", had a particular tendency to intermingle with the Greeks,
"This was particularly the case with the Carians, for, although the other peoples were not yet having very much intercourse with the Greeks nor even trying to live in Hellenic fashion or to learn our language ... yet the Carians roamed throughout the whole of Greece serving on expeditions for pay. ... and when they were driven thence [from the islands] into Asia, even here they were unable to live apart from the Greeks, I mean when the Ionians and Dorians later crossed over to Asia." (Strabo 14.2.28)

I won't get into too many details on archeology so here's a part from the wiki on archeology of Caria:

Throughout the 1950s, J.M. Cook and G.E. Bean conducted exhaustive archaeological surveys in Caria.[18] Cook ultimately concluded that Caria was virtually devoid of any prehistoric remains. According to his reports, third millennium finds were mostly confined to a few areas on or near the Aegean coast. No finds from the second millennium were known aside from the Submycenean remains at Asarlik and the Mycenaean remains at Miletus and near Mylasa. Archaeologically, there was nothing distinguishing about the Carians since the material evidence so far only indicated that their culture was merely a reflection of Greek culture.[19]

During the 1970s, further archaeological excavations in Caria revealed Mycenean buildings at Iasus (with two "Minoan" levels underneath them),[20] as well as Protogeometric and Geometric material remains (i.e. cemeteries and pottery).[21] Archaeologists also confirmed the presence of Carians in SardisRhodes, and in Egypt where they served as mercenaries of the Pharaoh. In Rhodes, specifically, a type of Carian chamber-tomb known as a Ptolemaion may be attributed to a period of Carian hegemony on the island.[22] Despite this period of increased archaeological activity, the Carians still appear not to have been an autochthonous group of Anatolia since both the coastal and interior regions of Caria were virtually unoccupied throughout prehistoric times.[23]

As for the assumption that the Carians descended from Neolithic settlers, this is contradicted by the fact that Neolithic Caria was essentially desolate.[24][failed verification] Though a very small Neolithic population may have existed in Caria,[25] the people known as "Carians" may in fact have been of Aegean origin that settled in southwestern Anatolia during the second millennium BC.[26] 

https://www.jstor.org/stable/581122?origin=crossref
https://www.jstor.org/stable/501620
https://www.journals.uchicago.edu/doi/10.2307/501620
https://www.jstor.org/stable/581114?origin=crossref

This is more evidence that Southern Greeks before mixing with Slavs were like the Marathon sample and West Anatolians. Using genoplot i removed 15.2% Slavic from modern Maniots, the least mixed Greeks, to simulate their pre-Slavic genetic profile. As you can see below it's about the same as Roman era West Anatolians.

In the Hellenistic-Roman era, a significant migration occurred from the Levant/East Anatolia to West Anatolia and Greece and then from Greece/West Anatolia and the Greek colonies to Italy. This resulted in the majority of the average citizens of the city of Rome being predominantly Greek around 2000 years ago. Specifically, 30% of the sampled population in Rome was fully Greek while the average scores 69% Greek.`
"The Roman Empire did, however, stimulate demographic change in the Balkans. In this early period, ∼1/3 of the individuals (15 of the 45) fall beyond the Balkan clines in PCA (Figures 1C and S4) but close to Near Easterners and can be modeled as deriving their ancestry predominantly from Roman/Byzantine populations from western Anatolia and, in one case, from Northern Levantine groups (Figure 2A; Data S2, Table 6). Most of these individuals were excavated at four different Viminacium necropolises, but we also found them at other urban centers such as Tragurium (Trogir) and Iader (Zadar). A very strong demographic shift toward Anatolia is also evident in Rome and central Italy during the same period15,25 and demonstrates long-distance mobility plausibly originating from the major eastern urban centers of the Empire such as Ephesus, Corinth, or Byzantium/Constantinople, and our results show that these migrants had a major demographic impact not only on the Imperial capital but also on other large towns on the Empire’s northern periphery. Our data also provide insights concerning the social dynamics of this demographic process."
G25 Model of all samples from imperial Rome averaged:

There are even Roman satirical poems written about the "bad Greek immigrants from the east" by the Romans, for example Juvenal's Satires.
"I now proceed to speak of the nation specially favoured by our wealthy compatriots, one that I shun above all others. I shan’t mince words. My fellow Romans, I cannot put up with a city of Greeks; yet how much of the dregs is truly Achaean? The Syrian Orontes has long been discharging into the Tiber, carrying with it its language and morals and slanting strings, complete with piper, not to speak of its native timbrels."


G25 distance chart showing the most similar modern people genetically to the ancient Greeks and Minoans:

The numbers below indicate how close they plot on the G25 PCA, so the smaller the number the more similar genetically to the target on top "Distance to". I used G25 because it has more DNA samples and less noise because its a PCA that is meant to differentiate populations that are very similar genetically to each other. Fst has too much noise (Standard Errors) to tell exactly which South East European region is the closest genetically because they are all relatively as close to the Mycenaeans.

G25 genetic distance heatmaps and charts to modern populations

Distance to Late Hellenistic and Roman era Greeks.


Distance Bronze Age Greeks:

The Bronze Age southern Greeks (Mycenaaeans from Peloponnese and Attica) are most similar genetically to modern Southern Greeks, Greek islanders, Greek Cypriots and South Italians
Genetically closest modern Greeks to the Mycenaeans from Peloponnese.

Distance to Mycenaeans from Central Greece.

While we don't have ancient Macedonian samples yet, based on the samples near Macedonia they were likely most similar genetically to modern Thessalian Greeks, Albanians and Central/North Italians. 

G25 distance heatmap to modern populations:




Before vs after the Slavic and Turkic invasions:

Before Slavic and Turkic invasions
After Slavic and Turkic invasions


Assessing the performance of qpAdm: a statistical tool for studying population admixture 


qpAdm models:

Outgroups/reference pops: https://pastebin.com/hKu6GSZ7

Aegean 1-200AD samples used https://pastebin.com/KdHE0uwT

Extra models:

Total Yamnaya ancestry from paleo Balkans vs Slav.




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