Post by Admin on Jun 27, 2023 17:35:50 GMT
Discussion
Our genomic transect of the population of Albania from the Neolithic to the modern era reveals fluctuations in genetic ancestry over a period of 8000 years. In contrast to the southeastern Balkans, where the arrival of Pontic-Caspian steppe ancestry and the associated Indo-European cultural package during the EBA did not lead to a lasting genetic turnover (4), we show that contemporary populations in Albania were genetically transformed both in autosomal and paternal ancestry (Fig. 8; Table S5). We find that more than a millennium later, BA-IA Balkan populations with high levels of steppe ancestry (30-40%) formed a distinct genetic cluster that extended from northwestern Greece, North Macedonia and the Adriatic coast (including Albania) and transcended archaeological and linguistic boundaries (Fig. 4A). This genetic continuum was broken down across the Balkans during the Roman and Migration period (Fig. 5A), due to mass settlement of Germanic and Slavic-speaking groups in the region.
However, in agreement with linguistic studies, we find that Albanians likely descend from a surviving West palaeo-Balkan population that experienced significant demographic increase approximately between 500-800 CE (Fig. 10), perhaps after a population bottleneck. We show that in contrast to the rest of the Balkans, the Medieval samples from both North and South Albania experienced little to no contribution from surrounding Slavic populations (Fig, 6B-C; Tables S12-S13, S15) and maintained high levels of BA-IA West Balkan ancestry. Remarkably, the same genetic profile persisted 500-800 years later in most of the post-Medieval samples from Bardhoc, as shown both by the PCA (Fig. 5), qpAdm analyses (Tables S16-S18), and IBD data (Table S20), which indicate significant genetic continuity from the Medieval populations of Albania. However, qpAdm models cannot exclude the possibility of additional admixture with currently unsampled neighbouring late Roman-early Medieval palaeo-Balkan groups with a similar ancestry profile. Based on linguistic data, the area of modern Kosovo and southeastern Serbia may have been such a source (15, 33, 34).
Despite being largely unaffected by the demographic changes that took place during the Migration period, the historical Albanians did not emerge in isolation. At the peak of the Migration Period, the Medieval population of Albania displayed genetic links as far as Pannonia (Tables 20-21), while in post-Medieval times we detected the presence of individuals likely related to modern Roma people (Fig. 5A). Furthermore, two of the post-Medieval samples exhibit significant admixture with South Slavic populations (Tables S16-S18), and modern Albanians display highly variable levels of Slavic ancestry (Fig. 5B, Tables S16-S18). This indicates complex historical interactions with South Slavic populations, as suggested by toponymy and linguistics (23, 35).
We reveal that a significant proportion of the paternal ancestry of modern Albanians derives from groups ultimately descending from the BA-IA West Balkans (Tables S33-S34), including those traditionally known as “Illyrians” (Figs. S9, S11), which reflects our findings on autosomal ancestry. However, inferring the language spoken by the Medieval samples from Albania is challenging, as Greek, South Slavic and West Balkan Romance are the only recorded languages of the region (14, 15, 25), while there is no indication of the survival of “Illyrian” following the first centuries of Roman rule (7, 8). Furthermore, Albanian displays Latin loans from both the Western and Eastern Balkans (85), which attests to linguistic influences beyond the confines of modern Albania. Testing the Messapic hypothesis for Albanian (7, 19, 30, 32) was not possible due to the low coverage of said samples (12). Although the presence of haplogroups J2b-L283, I-M223, and R1b-Z2103 among the Messapians (Table S30) suggests a West Balkan origin, whether a related language persisted in the Balkans during Medieval times is unknown.
Even though Eastern Roman historians were unfamiliar with Albanians (22), we cannot exclude the possibility that proto-Albanians interacted with populations speaking Greek, Aromanian, or Slavic in what is now southern Albania during Medieval times. Given that genetic data strongly suggest a predominantly local origin for Albanians, their Medieval ancestors may have inhabited a geographically restricted area [possibly the region of Mat in central Albania (14)], only occasionally venturing towards the south. These movements may have increased in scale over time, finally attracting the attention of Greek-speaking historians in the 11th century (22).
While the quest for the origins of the Albanian language will certainly continue, we expect that the present study will shape these debates and provide the necessary framework for more extensive research on the genetic ancestry of the ancient and modern inhabitants of Albania.
Our genomic transect of the population of Albania from the Neolithic to the modern era reveals fluctuations in genetic ancestry over a period of 8000 years. In contrast to the southeastern Balkans, where the arrival of Pontic-Caspian steppe ancestry and the associated Indo-European cultural package during the EBA did not lead to a lasting genetic turnover (4), we show that contemporary populations in Albania were genetically transformed both in autosomal and paternal ancestry (Fig. 8; Table S5). We find that more than a millennium later, BA-IA Balkan populations with high levels of steppe ancestry (30-40%) formed a distinct genetic cluster that extended from northwestern Greece, North Macedonia and the Adriatic coast (including Albania) and transcended archaeological and linguistic boundaries (Fig. 4A). This genetic continuum was broken down across the Balkans during the Roman and Migration period (Fig. 5A), due to mass settlement of Germanic and Slavic-speaking groups in the region.
However, in agreement with linguistic studies, we find that Albanians likely descend from a surviving West palaeo-Balkan population that experienced significant demographic increase approximately between 500-800 CE (Fig. 10), perhaps after a population bottleneck. We show that in contrast to the rest of the Balkans, the Medieval samples from both North and South Albania experienced little to no contribution from surrounding Slavic populations (Fig, 6B-C; Tables S12-S13, S15) and maintained high levels of BA-IA West Balkan ancestry. Remarkably, the same genetic profile persisted 500-800 years later in most of the post-Medieval samples from Bardhoc, as shown both by the PCA (Fig. 5), qpAdm analyses (Tables S16-S18), and IBD data (Table S20), which indicate significant genetic continuity from the Medieval populations of Albania. However, qpAdm models cannot exclude the possibility of additional admixture with currently unsampled neighbouring late Roman-early Medieval palaeo-Balkan groups with a similar ancestry profile. Based on linguistic data, the area of modern Kosovo and southeastern Serbia may have been such a source (15, 33, 34).
Despite being largely unaffected by the demographic changes that took place during the Migration period, the historical Albanians did not emerge in isolation. At the peak of the Migration Period, the Medieval population of Albania displayed genetic links as far as Pannonia (Tables 20-21), while in post-Medieval times we detected the presence of individuals likely related to modern Roma people (Fig. 5A). Furthermore, two of the post-Medieval samples exhibit significant admixture with South Slavic populations (Tables S16-S18), and modern Albanians display highly variable levels of Slavic ancestry (Fig. 5B, Tables S16-S18). This indicates complex historical interactions with South Slavic populations, as suggested by toponymy and linguistics (23, 35).
We reveal that a significant proportion of the paternal ancestry of modern Albanians derives from groups ultimately descending from the BA-IA West Balkans (Tables S33-S34), including those traditionally known as “Illyrians” (Figs. S9, S11), which reflects our findings on autosomal ancestry. However, inferring the language spoken by the Medieval samples from Albania is challenging, as Greek, South Slavic and West Balkan Romance are the only recorded languages of the region (14, 15, 25), while there is no indication of the survival of “Illyrian” following the first centuries of Roman rule (7, 8). Furthermore, Albanian displays Latin loans from both the Western and Eastern Balkans (85), which attests to linguistic influences beyond the confines of modern Albania. Testing the Messapic hypothesis for Albanian (7, 19, 30, 32) was not possible due to the low coverage of said samples (12). Although the presence of haplogroups J2b-L283, I-M223, and R1b-Z2103 among the Messapians (Table S30) suggests a West Balkan origin, whether a related language persisted in the Balkans during Medieval times is unknown.
Even though Eastern Roman historians were unfamiliar with Albanians (22), we cannot exclude the possibility that proto-Albanians interacted with populations speaking Greek, Aromanian, or Slavic in what is now southern Albania during Medieval times. Given that genetic data strongly suggest a predominantly local origin for Albanians, their Medieval ancestors may have inhabited a geographically restricted area [possibly the region of Mat in central Albania (14)], only occasionally venturing towards the south. These movements may have increased in scale over time, finally attracting the attention of Greek-speaking historians in the 11th century (22).
While the quest for the origins of the Albanian language will certainly continue, we expect that the present study will shape these debates and provide the necessary framework for more extensive research on the genetic ancestry of the ancient and modern inhabitants of Albania.