E-V13 entered Greece with Illyrians and Dorian invasions

[Aspar]

Thanks for the post. I am interested in the bolded. Do you have any specific samples in mind that show this change? I would agree that by Roman Imperial times we start finding modern looking Greek Aegean island profiles, but I am not sure any samples of similar genetics have been found on the mainland from this era.

Modern mainland Greeks have less Eastern genetic input that Empuries_2

Target: Iberia_Northeast_Empuries2
Distance: 2.7372% / 0.02737175
74.2 Anatolia_Barcin_N
11.8 Yamnaya_RUS_Samara
9.8 GEO_CHG
2.6 IRN_Ganj_Dareh_N
1.6 Levant_Natufian

Target: Greek_Thessaly
Distance: 2.3201% / 0.02320123
57.6 Anatolia_Barcin_N
33.6 Yamnaya_RUS_Samara
2.8 IRN_Ganj_Dareh_N
2.2 Baltic_LVA_HG
2.0 Levant_Natufian
1.8 GEO_CHG

Target: Greek_Central_Macedonia
Distance: 2.5255% / 0.02525525
56.2 Anatolia_Barcin_N
34.2 Yamnaya_RUS_Samara
3.4 Baltic_LVA_HG
3.4 IRN_Ganj_Dareh_N
2.4 Levant_Natufian
0.4 GEO_CHG

Target: Greek_Peloponnese
Distance: 1.9206% / 0.01920593
59.8 Anatolia_Barcin_N
29.0 Yamnaya_RUS_Samara
5.6 IRN_Ganj_Dareh_N
2.4 GEO_CHG
2.4 Levant_Natufian
0.8 Baltic_LVA_HG

Well I do have one from "Population genomic analysis of elongated skulls reveals extensive female-biased immigration in Early Medieval Bavaria", albeit it's found in a land far from Greece and the Balkans however the general opinion circulating on the forums like this one was that it was a sample from Greece or the Balkans ultimately. I am thinking about STR_300, the one found in medieval Bavaria buried with artifacts associated with the Mediterranean.
From the study:

All males and females with normal skulls had estimated origins with north and central
Europe, apart from STR_300 and STR_502, with their most likely geographic origins being
Greece and Turkey respectively (Fig. S29).

Using the G25 tool I will try to model the sample in question using all the samples of the current spreadsheet with ancient scaled samples:

Target: DEU_MA_o
Distance: 3.2673% / 0.03267303 | ADC: 0.5x
56.6 BGR_IA
18.4 Iberia_Northeast_Empuries2
16.2 HUN_MA_Szolad_o1
7.2 TUR_Isparta_EBA
1.6 ARM_Areni_C

STR_300 clearly is overwhelmingly of Balkan origin, whether this is an ancient Greek or Thracian or something else is very difficult to determine because of the similarity of those samples. In this model, the sample str_300 not only can be modeled mostly as of Thracian origin but it's also closest to the Iron Age sample from Bulgaria:

Distance to: DEU_MA_o
0.03732499 BGR_IA
0.04089265 Iberia_Northeast_Empuries2
0.04558788 HUN_MA_Szolad_o1
0.05106671 ITA_Rome_Late_Antiquity
0.05364110 GRC_Mycenaean
0.05461303 ITA_Rome_Imperial
0.05645685 ITA_Collegno_MA_o1
0.05843966 ITA_Rome_MA
0.06062882 Levant_LBN_MA_o4
0.06143632 ITA_Sardinia_Late_Antiquity
0.06284889 BGR_Krepost_N
0.06415580 ITA_Sardinia_IA
0.06455031 Levant_Ashkelon_IA1
0.06464730 ITA_Sicily_LBA

What is interesting is that the model shows that str_300 harbored some 8.8% West Asian ancestry derived mostly from Anatolia and the Lesser Caucasus.
That the sample in question shifts towards West Asia when compared to his predecessors from the Balkans could be also demonstrated with PCA:



Of course, If you are picky you might say that the Levant and Anatolia-Caucasus are two different thing however I have no doubt that different samples could pick up different things including those coming from the Levant. It's pity that we don't have many samples from the Roman period Balkans and one sample could hardly prove a point however we can even cite the recent paper about Rome and in the authors in the paper came to the same conclusion that during the Imperial times there was a big shift of the Rome's population towards West Asia and the Middle East and we can't doubt that the the same didn't happen in the Balkans IMO. Not only because Greece and the Balkans are closer geographically to West Asia and the Middle East but also because the ancient dna profile of Empuries2 and BGR_IA still evident in the EM period witnessed by the dna profile of
HUN_MA_Szolad_o1 didn't survive in the modern populations of Greece and the Balkans and Slavic admixture only can't explain the current position on the PCA of these modern populations but you also need something West Asian or Levantine to effectively model them:

Target: Greek_Thessaly
Distance: 1.2934% / 0.01293420
40.4 Iberia_Northeast_Empuries2
37.8 HUN_Avar_Szolad
13.4 TUR_Isparta_EBA
8.4 Levant_LBN_Roman

Empuries2 Eastern genetic input is represented mostly by CHG which in turn comes from the BA native populations of Greece such as the Minoans. Empuries2 definitely has less "IRN_Ganj_Dareh_N" and "Levant_Natufian" than modern mainland Greeks who when you take in consideration that they have additional ancestry from people of Northern and Eastern Europe with little or none West Asian and MENA admixture then it comes clear that some West Asian population also took a part in the making of the modern genetic profile of the mainland Greeks.

[Sorcelow]

Well I do have one from "Population genomic analysis of elongated skulls reveals extensive female-biased immigration in Early Medieval Bavaria", albeit it's found in a land far from Greece and the Balkans however the general opinion circulating on the forums like this one was that it was a sample from Greece or the Balkans ultimately. I am thinking about STR_300, the one found in medieval Bavaria buried with artifacts associated with the Mediterranean.
From the study:



Using the G25 tool I will try to model the sample in question using all the samples of the current spreadsheet with ancient scaled samples:

Target: DEU_MA_o
Distance: 3.2673% / 0.03267303 | ADC: 0.5x
56.6 BGR_IA
18.4 Iberia_Northeast_Empuries2
16.2 HUN_MA_Szolad_o1
7.2 TUR_Isparta_EBA
1.6 ARM_Areni_C

STR_300 clearly is overwhelmingly of Balkan origin, whether this is an ancient Greek or Thracian or something else is very difficult to determine because of the similarity of those samples. In this model, the sample str_300 not only can be modeled mostly as of Thracian origin but it's also closest to the Iron Age sample from Bulgaria:

Distance to: DEU_MA_o
0.03732499 BGR_IA
0.04089265 Iberia_Northeast_Empuries2
0.04558788 HUN_MA_Szolad_o1
0.05106671 ITA_Rome_Late_Antiquity
0.05364110 GRC_Mycenaean
0.05461303 ITA_Rome_Imperial
0.05645685 ITA_Collegno_MA_o1
0.05843966 ITA_Rome_MA
0.06062882 Levant_LBN_MA_o4
0.06143632 ITA_Sardinia_Late_Antiquity
0.06284889 BGR_Krepost_N
0.06415580 ITA_Sardinia_IA
0.06455031 Levant_Ashkelon_IA1
0.06464730 ITA_Sicily_LBA

What is interesting is that the model shows that str_300 harbored some 8.8% West Asian ancestry derived mostly from Anatolia and the Lesser Caucasus.
That the sample in question shifts towards West Asia when compared to his predecessors from the Balkans could be also demonstrated with PCA:



Of course, If you are picky you might say that the Levant and Anatolia-Caucasus are two different thing however I have no doubt that different samples could pick up different things including those coming from the Levant. It's pity that we don't have many samples from the Roman period Balkans and one sample could hardly prove a point however we can even cite the recent paper about Rome and in the authors in the paper came to the same conclusion that during the Imperial times there was a big shift of the Rome's population towards West Asia and the Middle East and we can't doubt that the the same didn't happen in the Balkans IMO. Not only because Greece and the Balkans are closer geographically to West Asia and the Middle East but also because the ancient dna profile of Empuries2 and BGR_IA still evident in the EM period witnessed by the dna profile of
HUN_MA_Szolad_o1 didn't survive in the modern populations of Greece and the Balkans and Slavic admixture only can't explain the current position on the PCA of these modern populations but you also need something West Asian or Levantine to effectively model them:

Target: Greek_Thessaly
Distance: 1.2934% / 0.01293420
40.4 Iberia_Northeast_Empuries2
37.8 HUN_Avar_Szolad
13.4 TUR_Isparta_EBA
8.4 Levant_LBN_Roman

Empuries2 Eastern genetic input is represented mostly by CHG which in turn comes from the BA native populations of Greece such as the Minoans. Empuries2 definitely has less "IRN_Ganj_Dareh_N" and "Levant_Natufian" than modern mainland Greeks who when you take in consideration that they have additional ancestry from people of Northern and Eastern Europe with little or none West Asian and MENA admixture then it comes clear that some West Asian population also took a part in the making of the modern genetic profile of the mainland Greeks.

I agree with you that mainland Greece received ancestry from the Levant/Caucasus region that was lacking in the Mycenaean and Minoan samples, but I believe that your models are inflating the Slavic related ancestry by not including more northerly Balkan samples. We know that heavier amounts of steppe ancestry was lurking to the north of Greece and since geneflow in this part of Europe usually came from the north southward, I think steppe ancestry was continuing to trickle down into Greece after the Bronze Age to the point where the pre-Slavic migration inhabitants of Greece were Prenestini_O like. Prenestini_O contains a bit more steppe ancestry than the Mycenaeans and also shows subtle signs of ancestry from the near east.

If I use Prenestini_O as a proxy for mainland Greece prior to the migration, we have this tight model, which fits well for most mainland Greece. There may have been multiple migrations to Greece from the Near East during different periods of time, but for the sake of keeping the model simple, I'm using modern Syrians as a proxy for any near eastern input that entered Greece, considering that these migrants may have been from the Greek speaking Antioch region:

[1] "distance%=1.3114"

Greek_Central_Macedonia

ITA_Prenestini_tribe_IA_o,56
HUN_Avar_Szolad,32.6
Syrian,11.4

[Aspar]

I agree with you that mainland Greece received ancestry from the Levant/Caucasus region that was lacking in the Mycenaean and Minoan samples, but I believe that your models are inflating the Slavic related ancestry by not including more northerly Balkan samples. We know that heavier amounts of steppe ancestry was lurking to the north of Greece and since geneflow in this part of Europe usually came from the north southward, I think steppe ancestry was continuing to trickle down into Greece after the Bronze Age to the point where the pre-Slavic migration inhabitants of Greece were Prenestini_O like. Prenestini_O contains a bit more steppe ancestry than the Mycenaeans and also shows subtle signs of ancestry from the near east.

If I use Prenestini_O as a proxy for mainland Greece prior to the migration, we have this tight model, which fits well for most mainland Greece. There may have been multiple migrations to Greece from the Near East during different periods of time, but for the sake of keeping the model simple, I'm using modern Syrians as a proxy for any near eastern input that entered Greece, considering that these migrants may have been from the Greek speaking Antioch region:

[1] "distance%=1.3114"

Greek_Central_Macedonia

ITA_Prenestini_tribe_IA_o,56
HUN_Avar_Szolad,32.6
Syrian,11.4

Well yeah, my model was only put forward to show the general picture and not to go in details. I've used it to show that there is some ancestry in modern mainland Greeks and not only them but also other Balkan people that comes from West Asia. While the Hun_Avar sample in this case is not only representative for Slavic ancestry but also for other Northern ancestry. Of course there is no perfect model however we agree for the extra West Asian and Levantine in modern mainland Greeks. On the other hand, I think that your usage of the Prenestini outlier is eating from the Levantine admixture because this sample was harboring significant Levantine ancestry as shown by Davidski in his blog.

ITA_Prenestini_tribe_IA_o
ITA_Proto-Villanovan 0.679±0.068
Levant_ISR_Ashkelon_IA2 0.321±0.068
chisq 7.222
tail prob 0.89033
Full output

Anyway this has nothing to do with E-V13 who in turn would have brought in Greece more ancestry associated with the Northern people rather than with the Levantines.

[Sorcelow]

Well yeah, my model was only put forward to show the general picture and not to go in details. I've used it to show that there is some ancestry in modern mainland Greeks and not only them but also other Balkan people that comes from West Asia. While the Hun_Avar sample in this case is not only representative for Slavic ancestry but also for other Northern ancestry. Of course there is no perfect model however we agree for the extra West Asian and Levantine in modern mainland Greeks. On the other hand, I think that your usage of the Prenestini outlier is eating from the Levantine admixture because this sample was harboring significant Levantine ancestry as shown by Davidski in his blog.



Anyway this has nothing to do with E-V13 who in turn would have brought in Greece more ancestry associated with the Northern people rather than with the Levantines.

Yes, its eating up some of the near eastern admixture, but I'm not sure if it has as much as what is shown in Davidksi's analysis. G25 is showing it as roughly intermediate between Mycenaean and MJ12, with slight additional near eastern ancestry. Either way, I think most E-V13 entered Greece from the north.

[Michalis Moriopoulos]

I favor a Hellenistic origin for the lion share of post-Bronze Age Near Eastern ancestry in European Greeks, but I can't rule out earlier or later migrations.

It seems possible that Iron Age Greeks have differed regionally. All we've got right now to go on are Emporiotes. Maybe Greeks in the far north had more steppe ancestry while Greeks in southern areas already had some Near Eastern admixture. We have just learned that there was significant Aegean ancestry in Iron Age Phoenicia, despite no obvious archaeological explanation for why that would be. Gene flow between Mycenaean and Levantine/Anatolian type people might have been bidirectional; the Phoenicians were seafarers, after all. So we might see some Levantine ancestry already in Iron Age Magna Graecia, the Aegean, parts of Anatolia, and maybe even southern Greece itself (Peloponnesus or Attica) well before the time of Alexander.

Of course, all we can do is fantasize until we see the samples.

[Generalissimo]

It seems possible that Iron Age Greeks have differed regionally. All we've got right now to go on are Emporiotes.

Bronze Age Greeks differed regionally too.

The currently available Mycenaean samples are fairly representative overall, but you'll see more diversity when other Mycenaean samples are released, both with much more and less steppe ancestry.

[Sorcelow]

Bronze Age Greeks differed regionally too.

The currently available Mycenaean samples are fairly representative overall, but you'll see more diversity when other Mycenaean samples are released, both with much more and less steppe ancestry.

Any idea when we will see more Greek samples released?

[Johnny ola]

I favor a Hellenistic origin for the lion share of post-Bronze Age Near Eastern ancestry in European Greeks, but I can't rule out earlier or later migrations.

It seems possible that Iron Age Greeks have differed regionally. All we've got right now to go on are Emporiotes. Maybe Greeks in the far north had more steppe ancestry while Greeks in southern areas already had some Near Eastern admixture. We have just learned that there was significant Aegean ancestry in Iron Age Phoenicia, despite no obvious archaeological explanation for why that would be. Gene flow between Mycenaean and Levantine/Anatolian type people might have been bidirectional; the Phoenicians were seafarers, after all. So we might see some Levantine ancestry already in Iron Age Magna Graecia, the Aegean, parts of Anatolia, and maybe even southern Greece itself (Peloponnesus or Attica) well before the time of Alexander.

Of course, all we can do is fantasize until we see the samples.

The northern regions of Greece especially regions like Macedonia, Thrace and Northern Thessaly/Epirus were never part of the Mycenean culture. These regions were inhabit mostly by Thracian tribes.Greeks from Bronze Age and until the classical period have stayed pretty much the same. The later admixtures especially from Slavs-Vlachs and West Asians have to do mostly with Roman-Byzantium ages.Another possibility of the west Asian components could be also hellenization of locals from the west anatolian coast. The people there might have BA Anatolia shift, so its very likely intermarriages to gave a more east med vibe.

[Johnny ola]

Bronze Age Greeks differed regionally too.

The currently available Mycenaean samples are fairly representative overall, but you'll see more diversity when other Mycenaean samples are released, both with much more and less steppe ancestry.

The current Mycenean samples are a joke. Thought I will agree with you I am waiting for R1a in the upcoming samples!!!

Most people bet on R1b thought...

[Oreo]

I believe that old Greeks in the Classical Period had their regional differences but those differences were fairly minimal, if the Thracian sample that we have is the average.