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Origins of East Caucasus Gene Pool: Contributions of Autochthonous Bronze Age Populations and Migrations from West Asia Estimated from Y-Chromosome Data
Abstract
The gene pool of the East Caucasus, encompassing modern-day Azerbaijan and Dagestan populations, was studied alongside adjacent populations using 83 Y-chromosome SNP markers. The analysis of genetic distances among 18 populations (N = 2216) representing Nakh-Dagestani, Altaic, and Indo-European language families revealed the presence of three components (Steppe, Iranian, and Dagestani) that emerged in different historical periods. The Steppe component occurs only in Karanogais, indicating a recent medieval migration of Turkic-speaking nomads from the Eurasian steppe. The Iranian component is observed in Azerbaijanis, Dagestani Tabasarans, and all Iranian-speaking peoples of the Caucasus. The Dagestani component predominates in Dagestani-speaking populations, except for Tabasarans, and in Turkic-speaking Kumyks. Each component is associated with distinct Y-chromosome haplogroup complexes: the Steppe includes C-M217, N-LLY22g, R1b-M73, and R1a-M198; the Iranian includes J2-M172(×M67, M12) and R1b-M269; the Dagestani includes J1-Y3495 lineages. We propose J1-Y3495 haplogroup’s most common lineage originated in an autochthonous ancestral population in central Dagestan and splits up ~6 kya into J1-ZS3114 (Dargins, Laks, Lezgi-speaking populations) and J1-CTS1460 (Avar-Andi-Tsez linguistic group). Based on the archeological finds and DNA data, the analysis of J1-Y3495 phylogeography suggests the growth of the population in the territory of modern-day Dagestan that started in the Bronze Age, its further dispersal, and the microevolution of the diverged population.
Keywords: East Caucasus; gene pool; Y-chromosome; Bronze Age populations; Dagestan; Azerbaijan; migrations; West Asia
1. Introduction
The East Caucasus, which spans modern-day Dagestan and Azerbaijan, is an important land bridge connecting Europe to West Asia. One of the gateways through the Caucasus is the Big Caucasian Pass, a strip of coastal land that runs between the East Caucasus mountains and the Caspian Sea all the way from Derbent to Sumgait.
The early humans of the Oldowan culture dispersed to the East Caucasus about 2 million years ago [1]. The ancient Chokh settlement site in today’s Dagestan is the key monument of the Mesolithic, Neolithic, and Bronze Age cultures that used to exist in the mountainous areas of the Northeast Caucasus. Evidence from recent radiocarbon dating suggests that domestication of animals, crop cultivation, and pottery making in this region began no later than the late 7th or early 6th millenniums BC [2]. Agriculture was booming; according to Nikolai Vavilov, the mountainous parts of Dagestan were the center of terraced farming [3]. The economic development during the Neolithic period (crop farming and animal husbandry, the advent of pottery, weaving, stone grinding, and polishing techniques) led to rapid population growth and colonization of new territories. In Azerbaijan, advanced Neolithic societies emerged at the dawn of the 6th millennium BC as a succession from the Neolithic societies of Southwest Asia [4,5]. The discovery of metals in the Eneolithic period (~7 kya) spurred the development of technology. The Bronze Age (6–4 kya) was marked by the emergence of new metalwork techniques, economic and social changes (social inequality, tribal confederations), and trade relations with the populations of Southeastern Europe and West Asia. In the Middle Bronze Age, nomads from the Eurasian steppe arrived in the East Caucasus, introducing the kurgan burial custom into the region.
In the 7th–6th centuries BC, the south of the region was infiltrated via the Derbent Pass by the Iranian-speaking Scythians who formed a military-political alliance known as Ashguza on the lands of today’s West Azerbaijan and Northwest Iran. Later, this territory was annexed by the Median Kingdom. In the late 1st millennium BC, the state of Caucasian Albania emerged from a tribal confederation in the northern part of the region. It had its own writing system, and its population adopted Christianity in the 4th century AD. Caucasian Albania was largely made up of autochthonous Caucasian tribes that spoke the Lezgic languages of the Nakh-Dagestanian family, although its reign extended over some Iranian tribes too. The territory of modern-day Dagestan was a cradle to such early states as Lakz, Tabarseran, Haidak, Sarir, Gumik, etc., which came into being in the first half of the 1st millennium AD.
During the first few centuries AD, other Iranian tribes, including Sarmatians, Maskut, and Alans, were arriving in the Caucasus. The South Caucasus was raided by Turkic-speaking Huns, Sabirs, and Khazars who came from the North via the Derbent Pass in the early Middle Ages (between the late 4th and 8th centuries) [6]. In the 7th century AD, the north of lowland Dagestan was occupied by the rising Khazar Khanate, whereas the south of the region was colonized by the Sasanian Empire in the 4th century. In the 7th–9th centuries, the East Caucasus was controlled by the Arab Khalifate, which was pursuing an aggressive migration policy that resulted in the Islamization of the local population. Incursions of Turkic-speaking Seljuks, the founders of the powerful empire spanning Central and West Asia and the South Caucasus, began in the 11th century as the invaders continued their expansion from Central Asia into the East Caucasus, including Derbent. Turkic peoples from the Cumania (Desht-i-Kipchak) migrated to the South Caucasus in the 11th–12th centuries; their descendants may have contributed to the emergence of the subethnical Azebaijani-Karapapakh group [7]. Other Turkic-speaking peoples of the East Caucasus are Kumyks and Nogais. There is no consensus on the origin of Kumyks. The prevailing hypothesis traces their descent to the local autochthonous population that had close ethnocultural contacts with foreign Turkic tribes like Sabir, Khazar, or Kipchak. In turn, Nogais were nomads from a late migration wave; most of them are settled in Dagestan and are represented by the Karanogai population.
After the Tatar-Mongol expansion that started in the 14th century, Islamic influence regained its primacy in Dagestan. The local medieval principalities of that time existed until the 19th century when they were absorbed by the Russian Empire. The southern regions of the East Caucasus were under Iranian influence during the 15th–19th centuries; they were incorporated by the Russian Empire in the aftermath of the Russo-Persian wars when the peace treaties were signed in 1813 and 1828. Later, these territories became the state of Azerbaijan.
www.mdpi.com/2073-4425/14/9/1780
Abstract
The gene pool of the East Caucasus, encompassing modern-day Azerbaijan and Dagestan populations, was studied alongside adjacent populations using 83 Y-chromosome SNP markers. The analysis of genetic distances among 18 populations (N = 2216) representing Nakh-Dagestani, Altaic, and Indo-European language families revealed the presence of three components (Steppe, Iranian, and Dagestani) that emerged in different historical periods. The Steppe component occurs only in Karanogais, indicating a recent medieval migration of Turkic-speaking nomads from the Eurasian steppe. The Iranian component is observed in Azerbaijanis, Dagestani Tabasarans, and all Iranian-speaking peoples of the Caucasus. The Dagestani component predominates in Dagestani-speaking populations, except for Tabasarans, and in Turkic-speaking Kumyks. Each component is associated with distinct Y-chromosome haplogroup complexes: the Steppe includes C-M217, N-LLY22g, R1b-M73, and R1a-M198; the Iranian includes J2-M172(×M67, M12) and R1b-M269; the Dagestani includes J1-Y3495 lineages. We propose J1-Y3495 haplogroup’s most common lineage originated in an autochthonous ancestral population in central Dagestan and splits up ~6 kya into J1-ZS3114 (Dargins, Laks, Lezgi-speaking populations) and J1-CTS1460 (Avar-Andi-Tsez linguistic group). Based on the archeological finds and DNA data, the analysis of J1-Y3495 phylogeography suggests the growth of the population in the territory of modern-day Dagestan that started in the Bronze Age, its further dispersal, and the microevolution of the diverged population.
Keywords: East Caucasus; gene pool; Y-chromosome; Bronze Age populations; Dagestan; Azerbaijan; migrations; West Asia
1. Introduction
The East Caucasus, which spans modern-day Dagestan and Azerbaijan, is an important land bridge connecting Europe to West Asia. One of the gateways through the Caucasus is the Big Caucasian Pass, a strip of coastal land that runs between the East Caucasus mountains and the Caspian Sea all the way from Derbent to Sumgait.
The early humans of the Oldowan culture dispersed to the East Caucasus about 2 million years ago [1]. The ancient Chokh settlement site in today’s Dagestan is the key monument of the Mesolithic, Neolithic, and Bronze Age cultures that used to exist in the mountainous areas of the Northeast Caucasus. Evidence from recent radiocarbon dating suggests that domestication of animals, crop cultivation, and pottery making in this region began no later than the late 7th or early 6th millenniums BC [2]. Agriculture was booming; according to Nikolai Vavilov, the mountainous parts of Dagestan were the center of terraced farming [3]. The economic development during the Neolithic period (crop farming and animal husbandry, the advent of pottery, weaving, stone grinding, and polishing techniques) led to rapid population growth and colonization of new territories. In Azerbaijan, advanced Neolithic societies emerged at the dawn of the 6th millennium BC as a succession from the Neolithic societies of Southwest Asia [4,5]. The discovery of metals in the Eneolithic period (~7 kya) spurred the development of technology. The Bronze Age (6–4 kya) was marked by the emergence of new metalwork techniques, economic and social changes (social inequality, tribal confederations), and trade relations with the populations of Southeastern Europe and West Asia. In the Middle Bronze Age, nomads from the Eurasian steppe arrived in the East Caucasus, introducing the kurgan burial custom into the region.
In the 7th–6th centuries BC, the south of the region was infiltrated via the Derbent Pass by the Iranian-speaking Scythians who formed a military-political alliance known as Ashguza on the lands of today’s West Azerbaijan and Northwest Iran. Later, this territory was annexed by the Median Kingdom. In the late 1st millennium BC, the state of Caucasian Albania emerged from a tribal confederation in the northern part of the region. It had its own writing system, and its population adopted Christianity in the 4th century AD. Caucasian Albania was largely made up of autochthonous Caucasian tribes that spoke the Lezgic languages of the Nakh-Dagestanian family, although its reign extended over some Iranian tribes too. The territory of modern-day Dagestan was a cradle to such early states as Lakz, Tabarseran, Haidak, Sarir, Gumik, etc., which came into being in the first half of the 1st millennium AD.
During the first few centuries AD, other Iranian tribes, including Sarmatians, Maskut, and Alans, were arriving in the Caucasus. The South Caucasus was raided by Turkic-speaking Huns, Sabirs, and Khazars who came from the North via the Derbent Pass in the early Middle Ages (between the late 4th and 8th centuries) [6]. In the 7th century AD, the north of lowland Dagestan was occupied by the rising Khazar Khanate, whereas the south of the region was colonized by the Sasanian Empire in the 4th century. In the 7th–9th centuries, the East Caucasus was controlled by the Arab Khalifate, which was pursuing an aggressive migration policy that resulted in the Islamization of the local population. Incursions of Turkic-speaking Seljuks, the founders of the powerful empire spanning Central and West Asia and the South Caucasus, began in the 11th century as the invaders continued their expansion from Central Asia into the East Caucasus, including Derbent. Turkic peoples from the Cumania (Desht-i-Kipchak) migrated to the South Caucasus in the 11th–12th centuries; their descendants may have contributed to the emergence of the subethnical Azebaijani-Karapapakh group [7]. Other Turkic-speaking peoples of the East Caucasus are Kumyks and Nogais. There is no consensus on the origin of Kumyks. The prevailing hypothesis traces their descent to the local autochthonous population that had close ethnocultural contacts with foreign Turkic tribes like Sabir, Khazar, or Kipchak. In turn, Nogais were nomads from a late migration wave; most of them are settled in Dagestan and are represented by the Karanogai population.
After the Tatar-Mongol expansion that started in the 14th century, Islamic influence regained its primacy in Dagestan. The local medieval principalities of that time existed until the 19th century when they were absorbed by the Russian Empire. The southern regions of the East Caucasus were under Iranian influence during the 15th–19th centuries; they were incorporated by the Russian Empire in the aftermath of the Russo-Persian wars when the peace treaties were signed in 1813 and 1828. Later, these territories became the state of Azerbaijan.
www.mdpi.com/2073-4425/14/9/1780