https://www.biorxiv.org/content/10.1101/2022.03.13.483276v1
Hannah M Moots et al.
doi: https://doi.org/10.1101/2022.03.13.483276
This article is a preprint and has not been certified by peer review
Abstract
The Iron Age saw the expansion of Phoenician and Greek colonies across the Mediterranean and the rise of Carthage as the major maritime power of the region. These events were facilitated by the ease of long-distance travel following major advances in seafaring. We know from the archaeological record that trade goods and materials were moving across great distances in unprecedented quantities, but it is unclear how these patterns correlate with human mobility. To investigate population mobility and interactions directly, we sequenced the genomes of 30 ancient individuals from Carthaginian and Etruscan port cities around the central Mediterranean, in Tunisia, Sardinia, and central Italy. At all three locations, there is a meaningful contribution of autochthonous populations (from Bronze Age North Africa, Sardinia, and Italy, respectively), as well as highly heterogeneous ancestry including many individuals with ancestry from other parts of the Mediterranean region. These results highlight both the role of autochthonous populations and the extreme interconnectedness of populations in the Iron Age Mediterranean. By studying these trans-Mediterranean neighbors together, we explore the complex interplay between local continuity and mobility that shaped the Iron Age societies of the central Mediterranean.
Characterizing the Genetic Heterogeneity at Kerkouane At Kerkouane, a Carthaginian town on the Cap Bon peninsula in Tunisia (see extended description in Materials), we observe a highly heterogeneous population, spanning across the PCA space in Fig. 3 from modern Mozabite populations to modern Sicilian populations, consisting of three primary genetic clusters.
One of the genetic groups we identified includes four individuals who have genetic continuity with preceding Maghrebi neolithic farmers, suggesting that these individuals represent an autochthonous North African population (Fig. 4). One individual, R11778, can be modeled in qpAdm with 100% Morocco Late Neolithic farmer ancestry, while three individuals, R11746, R11755, R11790, can be modeled predominantly with this component, along with the addition of 15 - 20% Steppe-related ancestry. A second cluster, visible in PCA (Fig. 3 and Fig. S5) and identified in qpWave (Fig. 5), contains seven individuals who are genetically similar to Bronze Sicilian and central Italian populations, as well as some individuals from the Hellenistic Iberian Greek colony of Emp ries (14, 23). For R11759, who projects near modern Mozabite and Moroccan populations in PCA space, there were no working distal qpAdm models with the original set of 5 distal source populations (Fig. 5). We replaced Morocco Late Neolithic with Morocco Early Neolithic farmers and a hunter-gatherer individual from Ethiopia from ~4500 BP (24), both of which produced working models. Using competition modeling (where possible sources are rotated to the outgroup), the best model uses ~70% Morocco Early Neolithic ancestry and ~30% Anatolia Neolithic (Fig. 4, Fig. S6). When compared to other ancient individuals using qpWave analysis (Fig. 5), this individual forms a clade with ancient Canary Island inhabitants thought to be representative of the original founding population (25). The Canary Islands were originally settled in the 1st millennium BCE by a population genetically ancestral to today’s Amazigh populations of Saharan
Africa (26).
Discussion
...
These results indicate that autochthonous North African populations contributed substantially to the genetic makeup of Kerkouane. The contribution of autochthonous North African populations in
Carthaginian history is obscured by the use of terms like “Western Phoenicians”, and even to an extent, “Punic”, in the literature to refer to Carthaginians, as it implies a primarily colonial population and diminishes indigenous involvement in the Carthaginian Empire. As a result, the role of autochthonous populations has been largely overlooked in studies of Carthage and its empire. Genetic approaches are well suited to examine such assumptions, and here we show that North African populations contributed substantially to the genetic makeup of Carthaginian cities. The high number of individuals with Italian and Greek-like ancestry may be due to the proximity of Kerkouane to Magna Graecia, as well as key trans-Mediterranean sailing routes passing by Cap Bon (1, 28). Yet, surprisingly, we did not detect individuals with large amounts of Levantine ancestry at Kerkouane. Given the roots of Carthage and its territories as Phoenician colonies, we had anticipated we would see individuals with ancestry similar to Phoenician individuals, such as those published in (12). One possible explanation is that the colonial expansion of Phoenician city-states at the start of the Iron Age did not involve large amounts of population mobility, and may have been based on trade relationships rather than occupation. Alternatively, this could potentially be due to differential burial practices (although Phoenician burial practices were thought to have shifted from cremations to interments in the central and western Mediterranean around 650 BCE (29), predating the individuals in the study), or to a disruption in connections between Carthaginian territories and the Eastern Mediterranean, after the fall of the Phoenician city-states to Babylon.
...
The Iron Age appears to be a key period for the formation of the current genetic structure of North Africa. Previous research suggests present-day central and western North African populations can be modeled as having four primary ancestry components: a local/autochthonous Maghrebi component derived from paleolithic hunter-gatherer populations in the region (20, 31, 32); a Near Eastern component thought to have been introduced with Arab rule of the region in the Medieval period (26); a sub-Saharan African component that was introduced in the last 5,000 years (16, 20); and a European component originally thought to have arrived due to historical population movements. While many papers have suggested the Near Eastern and European components resulted from recent historical movements, such as Arab rule in Medieval North Africa and trans-Mediterranean trade in the last 500 years, we see evidence for these
components being present in North Africa in the Iron Age, around 2,500 years ago. Fregel et al. 2018 show the European component is, at least partially, linked to the farming expansion and is similar to Anatolian and early European farmers (20). Additionally, we show that both Near Eastern and sub-Saharan African components were present in North Africa earlier than previously thought, reflecting the ongoing interconnectedness of North Africa to these regions for millennia (20, 33–35). The sub-Saharan ancestry we observe at Kerkouane may result either from direct contact or indirect contact through the nomadic populations of the Sahara. These nomadic groups, known to the Greeks as Numidians, are thought to be ancestral to Amazigh populations living in North Africa today. Trans-Saharan trade routes, made easier by a greener, less arid Sahara than today, had connected the communities of North Africa with their sub-Saharan counterparts since the Bronze Age (36, 37). Herodotus noted the coexistence of sedentary peoples and nomadic peoples in the land of the "libou" in the 5th century BCE (38). In addition to overland networks, these connections to sub-Saharan Africa also occurred by sea. Herodotus described Phoenician trade routes as extended far beyond the Mediterranean to the British Isles and West Africa via the Atlantic coast and even that a Phoenician and Egyptian expedition had circumnavigated Africa the previous century (1, 39). The Iron Age may have
been a key period for gene flow across the Sahara as well.
Hannah M Moots et al.
doi: https://doi.org/10.1101/2022.03.13.483276
This article is a preprint and has not been certified by peer review
Abstract
The Iron Age saw the expansion of Phoenician and Greek colonies across the Mediterranean and the rise of Carthage as the major maritime power of the region. These events were facilitated by the ease of long-distance travel following major advances in seafaring. We know from the archaeological record that trade goods and materials were moving across great distances in unprecedented quantities, but it is unclear how these patterns correlate with human mobility. To investigate population mobility and interactions directly, we sequenced the genomes of 30 ancient individuals from Carthaginian and Etruscan port cities around the central Mediterranean, in Tunisia, Sardinia, and central Italy. At all three locations, there is a meaningful contribution of autochthonous populations (from Bronze Age North Africa, Sardinia, and Italy, respectively), as well as highly heterogeneous ancestry including many individuals with ancestry from other parts of the Mediterranean region. These results highlight both the role of autochthonous populations and the extreme interconnectedness of populations in the Iron Age Mediterranean. By studying these trans-Mediterranean neighbors together, we explore the complex interplay between local continuity and mobility that shaped the Iron Age societies of the central Mediterranean.
Characterizing the Genetic Heterogeneity at Kerkouane At Kerkouane, a Carthaginian town on the Cap Bon peninsula in Tunisia (see extended description in Materials), we observe a highly heterogeneous population, spanning across the PCA space in Fig. 3 from modern Mozabite populations to modern Sicilian populations, consisting of three primary genetic clusters.
One of the genetic groups we identified includes four individuals who have genetic continuity with preceding Maghrebi neolithic farmers, suggesting that these individuals represent an autochthonous North African population (Fig. 4). One individual, R11778, can be modeled in qpAdm with 100% Morocco Late Neolithic farmer ancestry, while three individuals, R11746, R11755, R11790, can be modeled predominantly with this component, along with the addition of 15 - 20% Steppe-related ancestry. A second cluster, visible in PCA (Fig. 3 and Fig. S5) and identified in qpWave (Fig. 5), contains seven individuals who are genetically similar to Bronze Sicilian and central Italian populations, as well as some individuals from the Hellenistic Iberian Greek colony of Emp ries (14, 23). For R11759, who projects near modern Mozabite and Moroccan populations in PCA space, there were no working distal qpAdm models with the original set of 5 distal source populations (Fig. 5). We replaced Morocco Late Neolithic with Morocco Early Neolithic farmers and a hunter-gatherer individual from Ethiopia from ~4500 BP (24), both of which produced working models. Using competition modeling (where possible sources are rotated to the outgroup), the best model uses ~70% Morocco Early Neolithic ancestry and ~30% Anatolia Neolithic (Fig. 4, Fig. S6). When compared to other ancient individuals using qpWave analysis (Fig. 5), this individual forms a clade with ancient Canary Island inhabitants thought to be representative of the original founding population (25). The Canary Islands were originally settled in the 1st millennium BCE by a population genetically ancestral to today’s Amazigh populations of Saharan
Africa (26).
Discussion
...
These results indicate that autochthonous North African populations contributed substantially to the genetic makeup of Kerkouane. The contribution of autochthonous North African populations in
Carthaginian history is obscured by the use of terms like “Western Phoenicians”, and even to an extent, “Punic”, in the literature to refer to Carthaginians, as it implies a primarily colonial population and diminishes indigenous involvement in the Carthaginian Empire. As a result, the role of autochthonous populations has been largely overlooked in studies of Carthage and its empire. Genetic approaches are well suited to examine such assumptions, and here we show that North African populations contributed substantially to the genetic makeup of Carthaginian cities. The high number of individuals with Italian and Greek-like ancestry may be due to the proximity of Kerkouane to Magna Graecia, as well as key trans-Mediterranean sailing routes passing by Cap Bon (1, 28). Yet, surprisingly, we did not detect individuals with large amounts of Levantine ancestry at Kerkouane. Given the roots of Carthage and its territories as Phoenician colonies, we had anticipated we would see individuals with ancestry similar to Phoenician individuals, such as those published in (12). One possible explanation is that the colonial expansion of Phoenician city-states at the start of the Iron Age did not involve large amounts of population mobility, and may have been based on trade relationships rather than occupation. Alternatively, this could potentially be due to differential burial practices (although Phoenician burial practices were thought to have shifted from cremations to interments in the central and western Mediterranean around 650 BCE (29), predating the individuals in the study), or to a disruption in connections between Carthaginian territories and the Eastern Mediterranean, after the fall of the Phoenician city-states to Babylon.
...
The Iron Age appears to be a key period for the formation of the current genetic structure of North Africa. Previous research suggests present-day central and western North African populations can be modeled as having four primary ancestry components: a local/autochthonous Maghrebi component derived from paleolithic hunter-gatherer populations in the region (20, 31, 32); a Near Eastern component thought to have been introduced with Arab rule of the region in the Medieval period (26); a sub-Saharan African component that was introduced in the last 5,000 years (16, 20); and a European component originally thought to have arrived due to historical population movements. While many papers have suggested the Near Eastern and European components resulted from recent historical movements, such as Arab rule in Medieval North Africa and trans-Mediterranean trade in the last 500 years, we see evidence for these
components being present in North Africa in the Iron Age, around 2,500 years ago. Fregel et al. 2018 show the European component is, at least partially, linked to the farming expansion and is similar to Anatolian and early European farmers (20). Additionally, we show that both Near Eastern and sub-Saharan African components were present in North Africa earlier than previously thought, reflecting the ongoing interconnectedness of North Africa to these regions for millennia (20, 33–35). The sub-Saharan ancestry we observe at Kerkouane may result either from direct contact or indirect contact through the nomadic populations of the Sahara. These nomadic groups, known to the Greeks as Numidians, are thought to be ancestral to Amazigh populations living in North Africa today. Trans-Saharan trade routes, made easier by a greener, less arid Sahara than today, had connected the communities of North Africa with their sub-Saharan counterparts since the Bronze Age (36, 37). Herodotus noted the coexistence of sedentary peoples and nomadic peoples in the land of the "libou" in the 5th century BCE (38). In addition to overland networks, these connections to sub-Saharan Africa also occurred by sea. Herodotus described Phoenician trade routes as extended far beyond the Mediterranean to the British Isles and West Africa via the Atlantic coast and even that a Phoenician and Egyptian expedition had circumnavigated Africa the previous century (1, 39). The Iron Age may have
been a key period for gene flow across the Sahara as well.
