By sequencing 727 ancient individuals from the Southern Arc (Anatolia and its neighbors in Southeastern Europe and West Asia) over 10,000 years, we contextualize its Chalcolithic period and Bronze Age (about 5000 to 1000 BCE), when extensive gene flow entangled it with the Eurasian steppe. Two streams of migration transmitted Caucasus and Anatolian/Levantine ancestry northward, and the Yamnaya pastoralists, formed on the steppe, then spread southward into the Balkans and across the Caucasus into Armenia, where they left numerous patrilineal descendants. Anatolia was transformed by intra–West Asian gene flow, with negligible impact of the later Yamnaya migrations. This contrasts with all other regions where Indo-European languages were spoken, suggesting that the homeland of the Indo-Anatolian language family was in West Asia, with only secondary dispersals of non-Anatolian Indo-Europeans from the steppe. Description Connecting genes and history Stories about the peopling—and people—of Southern Europe and West Asia have been passed down for thousands of years, and these stories have contributed to our historical understanding of populations. Genomic data provide the opportunity to truly understand these patterns independently from written history. In a trio of papers, Lazaridis et al. examined more than 700 ancient genomes from across this region, the Southern Arc, spanning 11,000 years, from the earliest farming cultures to post-Medieval times (see the Perspective by Arbuckle and Schwandt). On the basis of these results, the authors suggest that earlier reliance on modern phenotypes and ancient writings and artistic depictions provided an inaccurate picture of early Indo-Europeans, and they provide a revised history of the complex migrations and population integrations that shaped these cultures. —SNV A web of migrations between Anatolia, its neighbors, and the Steppe suggests a West Asian origin of Indo-Anatolian languages. INTRODUCTION For thousands of years, humans moved across the “Southern Arc,” the area bridging Europe through Anatolia with West Asia. We report ancient DNA data from 727 individuals of this region over the past 11,000 years, which we co-analyzed with the published archaeogenetic record to understand the origins of its people. We focused on the Chalcolithic and Bronze Ages about 7000 to 3000 years ago, when Indo-European language speakers first appeared. RATIONALE Genetic data are relevant for understanding linguistic evolution because they can identify movement-driven opportunities for language spread. We investigated how the changing ancestral landscape of the Southern Arc, as reflected in DNA, corresponds to the structure inferred by linguistics, which links Anatolian (e.g., Hittite and Luwian) and Indo-European (e.g., Greek, Armenian, Latin, and Sanskrit) languages as twin daughters of a Proto-Indo-Anatolian language. RESULTS Steppe pastoralists of the Yamnaya culture initiated a chain of migrations linking Europe in the west to China and India in the East. Some people across the Balkans (about 5000 to 4500 years ago) traced almost all their genes to this expansion. Steppe migrants soon admixed with locals, creating a tapestry of diverse ancestry from which speakers of the Greek, Paleo-Balkan, and Albanian languages arose. The Yamnaya expansion also crossed the Caucasus, and by about 4000 years ago, Armenia had become an enclave of low but pervasive steppe ancestry in West Asia, where the patrilineal descendants of Yamnaya men, virtually extinct on the steppe, persisted. The Armenian language was born there, related to Indo-European languages of Europe such as Greek by their shared Yamnaya heritage. Neolithic Anatolians (in modern Turkey) were descended from both local hunter-gatherers and Eastern populations of the Caucasus, Mesopotamia, and the Levant. By about 6500 years ago and thereafter, Anatolians became more genetically homogeneous, a process driven by the flow of Eastern ancestry across the peninsula. Earlier forms of Anatolian and non–Indo-European languages such as Hattic and Hurrian were likely spoken by migrants and locals participating in this great mixture. Anatolia is remarkable for its lack of steppe ancestry down to the Bronze Age. The ancestry of the Yamnaya was, by contrast, only partly local; half of it was West Asian, from both the Caucasus and the more southern Anatolian-Levantine continuum. Migration into the steppe started by about 7000 years ago, making the later expansion of the Yamnaya into the Caucasus a return to the homeland of about half their ancestors. CONCLUSION All ancient Indo-European speakers can be traced back to the Yamnaya culture, whose southward expansions into the Southern Arc left a trace in the DNA of the Bronze Age people of the region. However, the link connecting the Proto-Indo-European–speaking Yamnaya with the speakers of Anatolian languages was in the highlands of West Asia, the ancestral region shared by both. Many partings, many meetings: How migration and admixture drove early language spread. Westward and northward migrations out of the West Asian highlands split the Proto-Indo-Anatolian language into Anatolian and Indo-European branches. Yamnaya pastoralists, formed on the steppe by a fusion of newcomers and locals, admixed again as they expanded far and wide, splitting the Proto-Indo-European language into its daughter languages across Eurasia. Border colors represent the ancestry and locations of five source populations before the migrations (arrows) and mixture (pie charts) documented here.

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The genetic history of the Southern Arc: A bridge between West Asia and Europe

Genetic association data from national biobanks and large-scale association studies have provided new prospects for understanding the genetic evolution of complex traits and diseases in humans. In turn, genomes from ancient human archaeological remains are now easier than ever to obtain, and provide a direct window into changes in frequencies of trait-associated alleles in the past. This has generated a new wave of studies aiming to analyse the genetic component of traits in historic and prehistoric times using ancient DNA, and to determine whether any such traits were subject to natural selection. In humans, however, issues about the portability and robustness of complex trait inference across different populations are particularly concerning when predictions are extended to individuals that died thousands of years ago, and for which little, if any, phenotypic validation is possible. In this review, we discuss the advantages of incorporating ancient genomes into studies of trait-associated variants, the need for models that can better accommodate ancient genomes into quantitative genetic frameworks, and the existing limits to inferences about complex trait evolution, particularly with respect to past populations.

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Quantitative Human Paleogenetics: What can Ancient DNA Tell us About Complex Trait Evolution?

Significance By studying genome-wide data from 54 individuals from eighth- to fifth-century Sicily, we gain insights into the composition of Classical Greek armies (ca. fifth c. BCE) and the populace of a Greek colony. The presence of mercenaries in Greek armies fighting in the Mediterranean, as early as 480 BCE and with origins as far away as northern Europe and the Caucasus, is absent from historical texts and thus so far underappreciated in ancient classical scholarship. Our interdisciplinary study both underlines the value of integrating genetic studies to complement archaeological and historical research and highlights the importance of warfare in facilitating continental-scale human mobility, cultural contact, and cooperation in the Mediterranean of the Classical period.

The diverse genetic origins of a Classical period Greek army

Literary and archaeological sources have preserved a rich history of Southern Europe and West Asia since the Bronze Age that can be complemented by genetics. Mycenaean period elites in Greece did not differ from the general population and included both people with some steppe ancestry and others, like the Griffin Warrior, without it. Similarly, people in the central area of the Urartian Kingdom around Lake Van lacked the steppe ancestry characteristic of the kingdom’s northern provinces. Anatolia exhibited extraordinary continuity down to the Roman and Byzantine periods, with its people serving as the demographic core of much of the Roman Empire, including the city of Rome itself. During medieval times, migrations associated with Slavic and Turkic speakers profoundly affected the region. Description Connecting genes and history Stories about the peopling—and people—of Southern Europe and West Asia have been passed down for thousands of years, and these stories have contributed to our historical understanding of populations. Genomic data provide the opportunity to truly understand these patterns independently from written history. In a trio of papers, Lazaridis et al. examined more than 700 ancient genomes from across this region, the Southern Arc, spanning 11,000 years, from the earliest farming cultures to post-Medieval times (see the Perspective by Arbuckle and Schwandt). On the basis of these results, the authors suggest that earlier reliance on modern phenotypes and ancient writings and artistic depictions provided an inaccurate picture of early Indo-Europeans, and they provide a revised history of the complex migrations and population integrations that shaped these cultures. —SNV Polities of the ancient Mediterranean world preserved contrasts of ancestry since the Bronze Age but were linked by migration.

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Eleni Kalliga

Papers

The genomic history of the Aegean palatial civilizations