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The Coming of the Greeks to Provence and Corsica: Y-Chromosome Models of Archaic Greek Colonization of the Western Mediterranean

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The Coming of the Greeks to Provence and Corsica: Y-Chromosome Models of Archaic Greek Colonization of the Western Mediterranean King et al. BMC Evolutionary Biology 2011, 11:69 http://www.biomedcentral.com/1471-2148/11/69 RESEARCHARTICLE Open Access The coming of the Greeks to Provence and Corsica: Y-chromosome models of archaic Greek colonization of the western Mediterranean Roy J King1, Julie Di Cristofaro2, Anastasia Kouvatsi3, Costas Triantaphyllidis3, Walter Scheidel4, Natalie M Myres5, Alice A Lin1, Alexandre Eissautier2, Michael Mitchell6, Didier Binder7, Ornella Semino8, Andrea Novelletto9, Peter A Underhill1, Jacques Chiaroni2* Abstract Background: The process of Greek colonization of the central and western Mediterranean during the Archaic and Classical Eras has been understudied from the perspective of population genetics. To investigate the Y chromosomal demography of Greek colonization in the western Mediterranean, Y-chromosome data consisting of 29 YSNPs and 37 YSTRs were compared from 51 subjects from Provence, 58 subjects from Smyrna and 31 subjects whose paternal ancestry derives from Asia Minor Phokaia, the ancestral embarkation port to the 6th century BCE Greek colonies of Massalia (Marseilles) and Alalie (Aleria, Corsica). Results: 19% of the Phokaian and 12% of the Smyrnian representatives were derived for haplogroup E-V13, characteristic of the Greek and Balkan mainland, while 4% of the Provencal, 4.6% of East Corsican and 1.6% of West Corsican samples were derived for E-V13. An admixture analysis estimated that 17% of the Y-chromosomes of Provence may be attributed to Greek colonization. Using the following putative Neolithic Anatolian lineages: J2a- DYS445 = 6, G2a-M406 and J2a1b1-M92, the data predict a 0% Neolithic contribution to Provence from Anatolia. Estimates of colonial Greek vs. indigenous Celto-Ligurian demography predict a maximum of a 10% Greek contribution, suggesting a Greek male elite-dominant input into the Iron Age Provence population. Conclusions: Given the origin of viniculture in Provence is ascribed to Massalia, these results suggest that E-V13 may trace the demographic and socio-cultural impact of Greek colonization in Mediterranean Europe, a contribution that appears to be considerably larger than that of a Neolithic pioneer colonization. Background olive oil [2]. The Greeks, at the beginning of the 1st mil- The collapse of the Late Bronze Age societies of the lennium BCE founded cities along the Asia Minor (Ana- Eastern Mediterranean (circa 1200 BCE) led to a cas- tolian) coast, divided into the Aeolian cities of cade of initial demographic retrenchment then expan- northwest Anatolia, the Ionian cities of central western sion, particularly among the Phoenicians of the coastal Anatolia and the Dorian cities of southwest Anatolia Levant and the Greeks of the Aegean Sea [1]. Both the [1,3]. Although the Greek colonies of Magna Graecia of Greeks and Phoenicians established a set of partitioned southern Italy and Sicily were established from a mix- colonies along the coast of Mediterranean Europe and ture of predominantly Dorian cities of the Aegean, the North Africa and engaged in extensive trade of a variety Peloponnesus and central Greece, the historical attesta- of goods including tin and other minerals, wine and tion of the Greek colonization of the western Mediterra- nean coastal regions of Provence, Spain and Corsica * Correspondence: [email protected] indicates a dominant influence from the Ionian city of 2Unité Mixte de Recherche 6578, Biocultural Anthropology, Centre National Phokaia (AKA Focia, Phocaea) (Figure 1) [4]. The Pho- de la Recherche Scientifique, Etablissement Français du S ang and Université kaian Greeks founded the city of Massalia circa 600 BCE Faculté de Médecine - Secteur Nord-Batiment A - Bd Pierre Dramard, 13344 Marseille Cedex 15, France at the location of the present city of Marseille and Alalie Full list of author information is available at the end of the article circa 560 BCE on the eastern coast of Corsica [4]. © 2011 King et al; licensee BioMed Central Ltd. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/2.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. King et al. BMC Evolutionary Biology 2011, 11:69 Page 2 of 8 http://www.biomedcentral.com/1471-2148/11/69 Rhone river Monoikos Nikaia Massalia Antipolis Basques Taureis Olbia Alalia (Corsica ) T1 Nea Nikomedeia Sesklo and Dimini Phokaia T7 Smyrna Franchthi Cave T8 T6 Alpes de Vaucluse Haute Provence Bouches Du Rhone Var Figure 1 Map showing locations of the Asia Minor Greek cities, Phokaia and Smyrna, in blue, and their western Mediterranean colonies, in red, as well as source populations studied in Greece and Turkey (in yellow). Putative source populations including Basques, central Anatolians, and Mediterranean Anatolians are designated with black circles. Inset shows the 4 districts in Provence from which samples were collected. Here the Phokaians encountered and interacted with the sub-Haplogroups of J2 have also been associated with indigenous Celto-Ligurian populations, as evidenced by the Neolithic colonization of mainland Greece, Crete large caches of wine amphora, which the local tribes dis- and southern Italy [12]. On the other hand, E-V13 tributed along the Rhone River and the Mediterranean appears to have originated in Greece or the southern coast [5]. Balkans [13,14] and then spread to Sicily at high fre- Few genetic studies have explored the Greek contribu- quencies with the Greek colonization of the island. tion to the modern populations of Italy and France. E-V13 is also found at low frequencies on the Anatolian A recent study of Y-chromosome haplogroups in a Sici- mainland [13] and thus may be useful in teasing apart lian population [6] showed a major impact of presump- the relative contributions of Greek colonization (E-V13) tive Greek immigration to the island estimated by from Early Neolithic colonization (J2) to Western Eur- admixture analysis to be about 37% using a localized ope. In this report, a sampling of individuals whose Balkan/Greek marker E-V13. This level of admixture ancestry traces to the Ionian Greek city of Phokaia will was higher than that predicted by classical demographic be compared through Y-chromosome genotyping to studies. Previous Y-chromosome genetic studies of samples from the Aeolian/Ionian city of Smyrna and a Phoenician colonization have demonstrated that hap- set of samples from Provence. These data will reveal logroup J2 frequency was amplified in regions contain- genetic patterning characteristic of 1) the Ionian founda- ing the Phoenician colonies of Iberia and North Africa tion of Phokaia versus the Aeolian/Ionian foundation of in comparison to areas not containing Phoenician colo- Smyrna. 2) the relative Y chromosome contributions of nies [7]. However, these studies did not address the role Phokaian Greeks and local Anatolian/Neolithic and/or of either Greek colonization or early Neolithic coloniza- central Anatolian populations in these two Asia Minor tion of Western Europe. Greek city-states and 3) the contribution of Greek and/ Y-chromosome studies have investigated the contribu- or Neolithic Y-chromosomes to the demographic pat- tion of various Y haplogroups to the spread of farming tern of Provence. from the Near East to Europe [8-10]. Haplogroup J2 fre- quency has been correlated with aspects of the symbolic Results material culture of the Neolithic in Europe and the Near The phylogenetic relationships and haplogroup frequen- East (painted pottery and ceramic figurines) [11] and cies for the data from the two sites in Asia Minor: King et al. BMC Evolutionary Biology 2011, 11:69 Page 3 of 8 http://www.biomedcentral.com/1471-2148/11/69 Phokaia and Smyrna, three mainland Greek sites, the demonstrated that both language/religion and geography four regions from Turkey and the Neolithic sites in discriminated the sample groups (Table 3). Provence are given in Figure 2. Phokaia and Smyrna MDS analyses show that both mainland Greek and have just subtle differences in their haplogroup compo- Phokaia separate from the Turkish samples while sition. The dominant haplogroups in both Phokaia and Smyrna positions between mainland Greeks and the Smyrna are E-V13 (19.4% and 12.1%) and R1b-M269 Turks (Figure 3). Since the Phokaian and Smyrnian sam- (22.6% and 27.8%) respectfully. In addition, J2a is also plescouldnotbedistinguishedfromeachotherin common, attaining a frequency of 9.7% in Phokaia and terms of Fst, they were aggregated for the subsequent 15.5% in Smyrna. This high frequency of haplogroup admixture analyses. J2a-Page55 (formerly DYS413≤ 18) in Smyrna is charac- The dominant haplogroup of Provence is R1b-M269 teristic of non-Greek Anatolia. Table 1 describes popu- at 58.8% (Figure 2). Also found in Provence is hap- lations analyzed in this study. The AMOVA (Table 2) logroup E-V13 (3.9%) and J2a-DYS445 = 6 (7.8%). All showed no significant distinction between Phokaia and the V13 derived samples are from western Provence Smyrna, whereas Smyrna was significantly differentiated along the Rhone, while all the J2a-DYS445 = 6 are from from central Anatolia and Phokaia from western Anato- Var in eastern Provence. lia. Smyrna also differed from both the Sesklo/Dimini The admixture analysis (Table 4) indicates a high level samples from Thessaly and the Lerna/Franchthi Cave of indigenous Basque admixture throughout Provence samples from the Peloponnese. The AMOVA analysis (70-90%). Also detected
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