Genetic study A genetic history of the Balkans from Roman frontier to Slavic migrations

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Reich et al.

I've been waiting for a comprehensive analysis of the ancient Balkans for some time now.

This study analyzed ancient DNA from 136 individuals from the Balkan region dating from the 1st to 10th centuries CE to understand population changes during the Roman Empire and the Migration Period. Here's a summary of the key findings:

  • Limited Italic/Roman ancestry: Despite Roman influence and administration, there was surprisingly little genetic contribution from people of Italic origin in the Balkans.
  • Influx of Anatolian ancestry: During the Roman Imperial period (1st-3rd centuries CE), there was a significant influx of people with Anatolian ancestry, particularly in urban centers. This influx was male-biased. This ancestry persisted in later populations, averaging 23%.
  • African ancestry: The study found evidence of individuals with North and East African ancestry, highlighting the reach of the Roman Empire and long-distance mobility. One individual from Viminacium had East African ancestry and a diet indicating consumption of marine protein, suggesting he spent his early life in East Africa.
  • Central/Northern European and Steppe ancestry: From the 3rd/4th centuries CE, individuals with ancestry related to Central/Northern Europe and the Pontic-Kazakh Steppe appeared. This suggests migration or integration of "barbarian" groups, possibly including Goths. These groups appear to have already been admixed before arriving in the Balkans. The Y-chromosome analysis showed a significant shift with the appearance of haplogroups R1a-Z93, I1, and R1b-U106, associated with these groups. However, this ancestry significantly decreases after 700 CE.
  • Slavic migrations: After 700 CE, a new ancestry component related to modern Eastern European Slavic-speaking populations appeared. This migration had a major demographic impact, contributing 30%-60% of the ancestry of present-day Balkan populations. This migration included both men and women. The study found evidence of intermarriage and integration between the newcomers and existing populations.
  • Present-day Balkan populations: Present-day Serbs, Croats, Bulgarians, and Romanians show a similar ancestral composition to the ancient individuals from the later period of the study (after 900 CE), with a significant Eastern European/Slavic component.
In short, the study reveals a complex history of migrations and admixture in the Balkans during the Roman and post-Roman periods. While Roman cultural influence was strong, genetic input from Italy was limited. Instead, the region saw significant influxes from Anatolia, Africa, Central/Northern Europe, and finally, Eastern Europe, with the Slavic migrations having the most lasting demographic impact on present-day Balkan populations.

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There are lots of E-V13 and G2a already from the Roman period and seem to make up the bulk of male lineages before the migration period. J2b is the third most common local Illyrian lineage.

In total I counted :
  • 2x E-L618
  • 15x E-V13 (Roman and Medieval but especially Roman period)
  • 1x E-V12 (Roman period)
  • 2x E-M123 (Roman period)
  • 6x G2a-P303 (including 5x L140)
  • 4x G2a-PF3148 (Roman and Medieval)
  • 3x I1 (Late Roman period)
  • 4x I2a-L621 (all from 775 CE onwards)
  • 1x I2c-L596 (Roman period)
  • 2x J1-Z2217 (Roman and Medieval)
  • 3x J2a-L26 (Roman and Medieval)
  • 7x J2b (Roman and Medieval)
  • 2x Q1a (700 to 1000 CE)
  • 6x R1a-Z282 (from 8th century onwards)
  • 3x R1a-Z93>Z94>Z2124 (3rd to 6th century)
  • 1x R1b-P312>DF27 (Roman period)
  • 1x R1b-P312>L21(Roman period)
  • 2x R1b-P312>DF99 (9th-10th century)
  • 2x R1b-P312>U152>L2 (Roman period)
  • 1x R1b-U106 (1st or 2nd century Viminacium)
  • 5x R1b-Z2103 (Roman and Medieval)
  • 1x R1b-PF7562 (15th century Albania)
  • 3x T1a2 (one from Roman Serbia, the other two from Greece and Albania)

So the Slavic R1a-Z282 and I2a-L621 start appear from about 775 CE onwards, together with some Q1a.

The Germanic I1 shows up from the late Roman period, but not the lone R1b-U106 who was already there in the Early Imperial period.
 
There are lots of E-V13 and G2a already from the Roman period and seem to make up the bulk of male lineages before the migration period. J2b is the third most common local Illyrian lineage.

In total I counted :
  • 2x E-L618
  • 15x E-V13 (Roman and Medieval but especially Roman period)
  • 1x E-V12 (Roman period)
  • 2x E-M123 (Roman period)
  • 6x G2a-P303 (including 5x L140)
  • 4x G2a-PF3148 (Roman and Medieval)
  • 3x I1 (Late Roman period)
  • 4x I2a-L621 (all from 775 CE onwards)
  • 1x I2c-L596 (Roman period)
  • 2x J1-Z2217 (Roman and Medieval)
  • 3x J2a-L26 (Roman and Medieval)
  • 7x J2b (Roman and Medieval)
  • 2x Q1a (700 to 1000 CE)
  • 6x R1a-Z282 (from 8th century onwards)
  • 3x R1a-Z93>Z94>Z2124 (3rd to 6th century)
  • 1x R1b-P312>DF27 (Roman period)
  • 1x R1b-P312>L21(Roman period)
  • 2x R1b-P312>DF99 (9th-10th century)
  • 2x R1b-P312>U152>L2 (Roman period)
  • 1x R1b-U106 (1st or 2nd century Viminacium)
  • 5x R1b-Z2103 (Roman and Medieval)
  • 1x R1b-PF7562 (15th century Albania)
  • 3x T1a2 (one from Roman Serbia, the other two from Greece and Albania)

So the Slavic R1a-Z282 and I2a-L621 start appear from about 775 CE onwards, together with some Q1a.

The Germanic I1 shows up from the late Roman period, but not the lone R1b-U106 who was already there in the Early Imperial period.
Basically the modern population of Albania and Kosova is similar to the late antiquity population of the region before the slavic migration, but the authors do not state this obvious fact.
 
Basically the modern population of Albania and Kosova is similar to the late antiquity population of the region before the slavic migration, but the authors do not state this obvious fact.
Yes, although there is more R1b in modern Albania and less G2a than in Roman-age Serbia and Croatia.
 
Yes, although there is more R1b in modern Albania and less G2a than in Roman-age Serbia and Croatia.

That G2a presence is wild and unexplainable. E-V13 appears more frequently than Iron Age due to cremation bias, the population started inhumating the remains.
 
That G2a presence is wild and unexplainable. E-V13 appears more frequently than Iron Age due to cremation bias, the population started inhumating the remains.
Probably there could be some Jewish input through the G2a. The synagogue in Saranda/Albania is considered to be of the 4th-5th AD and there might be others dispersed in the Balkans area.
 
There are lots of E-V13 and G2a already from the Roman period and seem to make up the bulk of male lineages before the migration period. J2b is the third most common local Illyrian lineage.

In total I counted :
  • 2x E-L618
  • 15x E-V13 (Roman and Medieval but especially Roman period)
  • 1x E-V12 (Roman period)
  • 2x E-M123 (Roman period)
  • 6x G2a-P303 (including 5x L140)
  • 4x G2a-PF3148 (Roman and Medieval)
  • 3x I1 (Late Roman period)
  • 4x I2a-L621 (all from 775 CE onwards)
  • 1x I2c-L596 (Roman period)
  • 2x J1-Z2217 (Roman and Medieval)
  • 3x J2a-L26 (Roman and Medieval)
  • 7x J2b (Roman and Medieval)
  • 2x Q1a (700 to 1000 CE)
  • 6x R1a-Z282 (from 8th century onwards)
  • 3x R1a-Z93>Z94>Z2124 (3rd to 6th century)
  • 1x R1b-P312>DF27 (Roman period)
  • 1x R1b-P312>L21(Roman period)
  • 2x R1b-P312>DF99 (9th-10th century)
  • 2x R1b-P312>U152>L2 (Roman period)
  • 1x R1b-U106 (1st or 2nd century Viminacium)
  • 5x R1b-Z2103 (Roman and Medieval)
  • 1x R1b-PF7562 (15th century Albania)
  • 3x T1a2 (one from Roman Serbia, the other two from Greece and Albania)

So the Slavic R1a-Z282 and I2a-L621 start appear from about 775 CE onwards, together with some Q1a.

The Germanic I1 shows up from the late Roman period, but not the lone R1b-U106 who was already there in the Early Imperial period.
When do you plan to update the threads on R1a, including M458 trees etc?

Theres a plethora of aDNA from BA, IA, LA, EMA for M458, and they are not all Slavic.

I am happy to provide the studies and sample IDs in question for their eventual update on Eupedia if such is planned.
 
Basically the modern population of Albania and Kosova is similar to the late antiquity population of the region before the slavic migration, but the authors do not state this obvious fact.
This though is a bit of a coincidence. There is an Anatolian component in Albanians that shifted the Illyrian basis in southern direction and then in early Medieval times the Slavic component shifted the population in northern direction which coincidentally resembles the old Illyrian structure. Same that happened in Italy, just with Germanic infiltration in the Medieval period.
 
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You can use the Medieval Foggia samples + Slavic to model modern Albanians. The proportion of Slavic ancestry matches what the paper determined, which is around 30%.
 
There are lots of E-V13 and G2a already from the Roman period and seem to make up the bulk of male lineages before the migration period. J2b is the third most common local Illyrian lineage.

The "bulk" of E-V13 is concentrated in Viminacium (Moesia Superior, today Eastern Serbia) and Timacum Minus (Southeastern Serbia). It is too geographically limited to be considered the bulk of male lineages before the migration period. As far as I could see, there are only three E-V13 samples from Croatia (Trogir and Gardun in Dalmatia; Nustar in Eastern Croatia). One E-V13 in Serbia is from Jakovo which is part of the wider region of Syrmia like Nustar in Croatia. The two E-M123 samples are both from Viminacium.

I'm not sure what you mean by stating that J2b is the third most common local Illyrian lineage. The bulk of E-V13 is concentrated in pretty much two locations in Eastern Serbia (Viminacium and Timacum) which is irrelevant to the areas settled or associated with Illyrians and the Roman province of Illyricum. You cannot paint a larger picture from such quantitatively and geographically limited data.
 
If I'm understanding Figure C correctly, modern ethnic Albanians are very similar to the populations from bronze age and iron age Albania and that Greeks from Macedonia and the Peloponnese have been shifted away from the the populations that resided there from the bronze age and iron age, is this correct?

How much have the populations that have entered the Balkans genetically contributed to the ancestry of Albanians and Greeks today? It is kind of shocking that Anatolian male migration into the Balkans had a 23% influence without some sort of resistance from the natives, if I'm understanding this correctly.
 
If I'm understanding Figure C correctly, modern ethnic Albanians are very similar to the populations from bronze age and iron age Albania and that Greeks from Macedonia and the Peloponnese have been shifted away from the the populations that resided there from the bronze age and iron age, is this correct?

How much have the populations that have entered the Balkans genetically contributed to the ancestry of Albanians and Greeks today? It is kind of shocking that Anatolian male migration into the Balkans had a 23% influence without some sort of resistance from the natives, if I'm understanding this correctly.
Medieval Albanians cluster with Central and South Italians (Apulians from Zollino). Then were pulled to their current position due to Slavic enrichment. Later I will post an interactive PCA when one can observe this first hand. Bronze Age Albanians cluster with Northern Italians/Tuscans.
 
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Medieval Albanians cluster with Central and South Italians (Apulians from Zollino). Then were pulled to their current position due to Slavic enrichment. Later I will post an interactive PCA when one can observe this first hand. Bronze Age Albanians cluster with Northern Italians.
eoeaeAg.png

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That G2a presence is wild and unexplainable. E-V13 appears more frequently than Iron Age due to cremation bias, the population started inhumating the remains.
Quite likely. The Hellenistic and Roman Era was a time when some ancient cultures probably had gone extinct. And so did some religious practices.

I have my doubts of a large influx from North Africa. As this would have probably been accompanied by other haplogroups other than E-V13. Especially from the 1st to 3D century AD.
 
Quite likely. The Hellenistic and Roman Era was a time when some ancient cultures probably had gone extinct. And so did some religious practices.

I have my doubts of a large influx from North Africa. As this would have probably been accompanied by other haplogroups other than E-V13. Especially from the 1st to 3D century AD.

Didn't quite understand your reply because arguably E-V13 arose in Europe, but it's parent E-L618 or the para-version of it came from North Africa specifically Mesolithic Egypt in Late Mesolithic/Early Neolithic times.

Anyway, i don't even think G2a is present at all in North Africa to make such a presence in Balkans.

The Carpathian-Balkan sphere of culture even Central Balkans were heavy cremation users with proper Thracians resulting in biritual practices, i do think these were precursors of E-V13 heavy populations.
 
Medieval Albanians cluster with Central and South Italians (Apulians from Zollino). Then were pulled to their current position due to Slavic enrichment. Later I will post an interactive PCA when one can observe this first hand. Bronze Age Albanians cluster with Northern Italians.
eoeaeAg.png

Medieval Albanians cluster with Central and South Italians (Apulians from Zollino). Then were pulled to their current position due to Slavic enrichment. Later I will post an interactive PCA when one can observe this first hand. Bronze Age Albanians cluster with Northern Italians/Tuscans.
I added all of the albanian samples
 
Didn't quite understand your reply because arguably E-V13 arose in Europe, but it's parent E-L618 or the para-version of it came from North Africa specifically Mesolithic Egypt in Late Mesolithic/Early Neolithic times.
Given that E-V13 arose in Europe, I am cautious about a significant North (East) African influx in the Balkans in the 1-3 century A.D. But after a closer look, this paper does not claim that. They just want to emphasise that there was mobility because they found some evidence of East African ancestry.

On another note, I don’t understand the BA+IA Greeks cluster. I am not aware of any research with any significant number of IA mainland Greeks. I am saying this because it is possible that the Anatolian admixture in Greece increased during the IA. And certainly during the Classical Age.

The first option is the so called Dorian invasion hypothesis. Some historians have postulated that there is evidence that Dorians came from farther East. Archaeological evidence supports that as well.

Besides that, we also know that the Greek world starting the Late Bronze Age included Anatolia as well. So internal mobility within the Greek world would have increased Anatolian ancestry in Greeks. This ancestry may have impacted the Balkans as well.
 
Given that E-V13 arose in Europe, I am cautious about a significant North (East) African influx in the Balkans in the 1-3 century A.D. But after a closer look, this paper does not claim that. They just want to emphasise that there was mobility because they found some evidence of East African ancestry.

On another note, I don’t understand the BA+IA Greeks cluster. I am not aware of any research with any significant number of IA mainland Greeks. I am saying this because it is possible that the Anatolian admixture in Greece increased during the IA. And certainly during the Classical Age.

The first option is the so called Dorian invasion hypothesis. Some historians have postulated that there is evidence that Dorians came from farther East. Archaeological evidence supports that as well.

Besides that, we also know that the Greek world starting the Late Bronze Age included Anatolia as well. So internal mobility within the Greek world would have increased Anatolian ancestry in Greeks. This ancestry may have impacted the Balkans as well.

Well, the paper has been around forever, they never mentioned E-V13 with East Africa, there has been a sample found from there (East African Roman mercenary) in Viminacium i think, and we were aware.

Otherwise i think even original E-L618 from Egypt would be different from East Africans in having heavy Mesolithic Egyptian HG = ANA + Dzudzuana-like.

In addition the Bronze Age origin of E-V13 has nothing to do with Anatolia as well, they migrated and invaded North-West Anatolia during Late Bronze Age. This is well known in archaeological circles related to archaeological cultures from where E-V13 was found in one of them in big numbers during EIA, namely Psenicevo in Kapitan Andrevo. This was a Balkan-Carpathian Culture, an Urnfield sub-variant.
 
The "bulk" of E-V13 is concentrated in Viminacium (Moesia Superior, today Eastern Serbia) and Timacum Minus (Southeastern Serbia). It is too geographically limited to be considered the bulk of male lineages before the migration period. As far as I could see, there are only three E-V13 samples from Croatia (Trogir and Gardun in Dalmatia; Nustar in Eastern Croatia). One E-V13 in Serbia is from Jakovo which is part of the wider region of Syrmia like Nustar in Croatia. The two E-M123 samples are both from Viminacium.

I'm not sure what you mean by stating that J2b is the third most common local Illyrian lineage. The bulk of E-V13 is concentrated in pretty much two locations in Eastern Serbia (Viminacium and Timacum) which is irrelevant to the areas settled or associated with Illyrians and the Roman province of Illyricum. You cannot paint a larger picture from such quantitatively and geographically limited data.

Viminacium is not neccessarily related to Illyrians but Timacum Minus has an Illyrian relation seeing the J-L283 and R1b found with some Illyrian profiles, the E-V13 there is probably from Thracian mixed with Illyrian, Some E-V13 profiles found there are shifted towards Illyrians, some shift towards Southern Thracians. Moesia was a territory that had all groups. E-V13 during the Roman period was not only located in Viminacium or Timacum Minus. E-V13 made up also 20% of the Y-dna samples in Roman Dalmatia so far.
 
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