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• W1963647637 abstract "The origin of the Etruscan people has been a source of major controversy for the past 2,500 years, and several hypotheses have been proposed to explain their language and sophisticated culture, including an Aegean/Anatolian origin. To address this issue, we analyzed the mitochondrial DNA (mtDNA) of 322 subjects from three well-defined areas of Tuscany and compared their sequence variation with that of 55 western Eurasian populations. Interpopulation comparisons reveal that the modern population of Murlo, a small town of Etruscan origin, is characterized by an unusually high frequency (17.5%) of Near Eastern mtDNA haplogroups. Each of these haplogroups is represented by different haplotypes, thus dismissing the possibility that the genetic allocation of the Murlo people is due to drift. Other Tuscan populations do not show the same striking feature; however, overall, ∼5% of mtDNA haplotypes in Tuscany are shared exclusively between Tuscans and Near Easterners and occupy terminal positions in the phylogeny. These findings support a direct and rather recent genetic input from the Near East—a scenario in agreement with the Lydian origin of Etruscans. Such a genetic contribution has been extensively diluted by admixture, but it appears that there are still locations in Tuscany, such as Murlo, where traces of its arrival are easily detectable. The origin of the Etruscan people has been a source of major controversy for the past 2,500 years, and several hypotheses have been proposed to explain their language and sophisticated culture, including an Aegean/Anatolian origin. To address this issue, we analyzed the mitochondrial DNA (mtDNA) of 322 subjects from three well-defined areas of Tuscany and compared their sequence variation with that of 55 western Eurasian populations. Interpopulation comparisons reveal that the modern population of Murlo, a small town of Etruscan origin, is characterized by an unusually high frequency (17.5%) of Near Eastern mtDNA haplogroups. Each of these haplogroups is represented by different haplotypes, thus dismissing the possibility that the genetic allocation of the Murlo people is due to drift. Other Tuscan populations do not show the same striking feature; however, overall, ∼5% of mtDNA haplotypes in Tuscany are shared exclusively between Tuscans and Near Easterners and occupy terminal positions in the phylogeny. These findings support a direct and rather recent genetic input from the Near East—a scenario in agreement with the Lydian origin of Etruscans. Such a genetic contribution has been extensively diluted by admixture, but it appears that there are still locations in Tuscany, such as Murlo, where traces of its arrival are easily detectable. The Etruscan culture developed and prospered in the 9th century b.c. over a region in central Italy named “Etruria”—usually referred to in Greek and Latin sources as “Tyrrhenia.” Etruria encompassed what is now known as “Tuscany,” “northern Latium,” and “western Umbria.” The culture of the Etruscan people was not only extremely advanced but also distinctively different from the cultures of the surrounding populations of Italy. Because of this, the origin of the Etruscan people has been a source of controversy for the past 2,500 years, with two major scenarios being disputed. The first is that the Etruscans were an autochthonous population of Italy; the alternative hypothesis is that they were direct descendants of an immigrant group of Aegean/Anatolian ancestry.1Beekes RSP The origins of the Etruscans. Nederlandse Akademie van Wetenschappen, Amsterdam2003Google Scholar Etruria was dominant in the Italian peninsula after 650 b.c., when Etruscans began to expand toward both the north (the Po River Valley) and the south. Etruscan kings conquered and ruled Rome for 100 years, until 509 b.c., when the last Etruscan king, Lucius Tarquinius Superbus, was removed from power and the Roman Republic was established. From that moment, the expansion period of the Etruscans ended and was followed by a slow process of conquest and assimilation by the Romans, which culminated at the time of the “Social War” (90–88 b.c.), with the attribution of Roman citizenship to all Etruscans. Rapidly, the Etruscan culture and language disappeared,2Barker G Rasmussen T The Etruscans. Blackwell, Oxford, United Kingdom1998Google Scholar, 3Rasmussen T Urbanization in Etruria.in: Osborne R Cunliffe B Mediterranean urbanization 800-600 BC. Oxford University Press, Oxford, United Kingdom2004Google Scholar possibly also because, in the following decades, Etrurian lands were often distributed to Roman veterans and were partially repopulated by poor Roman citizens. However, despite a potentially extensive dilution of the ancestral Etruscan gene pool with that from surrounding Italic populations, there is no evidence that such a gene pool had been completely replaced.4Piazza A Cappello N Olivetti E Rendine S A genetic history of Italy.Ann Hum Genet. 1988; 52: 203-213Crossref PubMed Scopus (147) Google Scholar In addition, part of the process of initial assimilation might have been mainly male mediated, since incoming Roman veterans often married women from local communities. Thus, some populations of modern Tuscany should have retained at least a fraction of such an ancestral gene pool—particularly the exclusively maternally inherited mtDNA pool—possibly at a variable extent, given the differential degree of geographical and genetic isolation of the different Tuscan communities during the past 25 centuries. To evaluate the nature and extent of mtDNA variation in modern Tuscans, the mtDNA from 322 subjects from three areas of Tuscany was analyzed: 86 from Murlo, a rather isolated town of Etruscan origin in the Siena province; 114 from Volterra, a former major Etruscan city in the province of Pisa; and 122 from the Casentino Valley in the province of Arezzo, which was part of historical Etruria. mtDNA profiles in the three populations were determined by sequencing ∼750 bp from the control region for each subject (from nucleotide position [np] 16024 to np 210, thus including the entire hypervariable segment [HVS]–I [nps 16024–16383] and part of the HVS-II [nps 57–372]). This was followed by a hierarchical survey of haplogroup and subhaplogroup diagnostic markers in the coding region,5Carelli V Achilli A Valentino ML Rengo C Semino O Pala M Olivieri A Mattiazzi M Pallotti F Carrara F et al.Haplogroup effect and recombination of mitochondrial DNA: novel clues from the analysis of Leber hereditary optic neuropathy pedigrees.Am J Hum Genet. 2006; 78: 564-574Abstract Full Text Full Text PDF PubMed Scopus (148) Google Scholar which allowed the classification of mtDNAs into 39 haplogroups (table 1). Most haplogroups that are typical of modern European populations are present,6Torroni A Huoponen K Francalacci P Petrozzi M Morelli L Scozzari R Obinu D Savontaus M-L Wallace DC Classification of European mtDNAs from an analysis of three European populations.Genetics. 1996; 144: 1835-1850PubMed Google Scholar, 7Richards M Macaulay V Hickey E Vega E Sykes B Guida V Rengo C Sellitto D Cruciani F Kivisild T et al.Tracing European founder lineages in the Near Eastern mtDNA pool.Am J Hum Genet. 2000; 67: 1251-1276Abstract Full Text Full Text PDF PubMed Scopus (745) Google Scholar, 8Palanichamy Mg Sun C Agrawal S Bandelt H-J Kong Q-P Khan F Wang C-Y Chaudhuri TK Palla V Zhang Y-P Phylogeny of mitochondrial DNA macrohaplogroup N in India, based on complete sequencing: implications for the peopling of South Asia.Am J Hum Genet. 2004; 75: 966-978Abstract Full Text Full Text PDF PubMed Scopus (277) Google Scholar and a few East Asian (D4g1)9Kong Q-P Bandelt H-J Sun C Yao Y-G Salas A Achilli A Wang C-Y Zhong L Zhu CL Wu S-F et al.Updating the East Asian mtDNA phylogeny: a prerequisite for the identification of pathogenic mutations.Hum Mol Genet. 2006; 15: 2076-2086Crossref PubMed Scopus (307) Google Scholar and sub-Saharan African (L3d and L1b)10Kivisild T Shen P Wall DP Do B Sung R Davis K Passarino G Underhill PA Scharfe C Torroni A et al.The role of selection in the evolution of human mitochondrial genomes.Genetics. 2006; 172: 373-387Crossref PubMed Scopus (348) Google Scholar mtDNAs were also detected. This latter finding is not unexpected, since low frequencies of African and even East Asian mtDNAs are not uncommon in populations of southern Europe. However, what was not anticipated was the relatively high frequency, especially in the Murlo sample, of haplogroup HV lineages that are non-H and non-HV0, as well as haplogroups R0a (formerly known as “(pre-HV)1”)11Torroni A Achilli A Macaulay V Richards M Bandelt H-J Harvesting the fruit of the human mtDNA tree.Trends Genet. 2006; 22: 339-345Abstract Full Text Full Text PDF PubMed Scopus (333) Google Scholar and U7 and U3, which are typical of Near Eastern populations.Table 1Frequencies of mtDNA Haplogroups in the Three Tuscan PopulationsNo. (%) of Subjects by PopulationHaplogroupaHV0, HV0*, HV0a, R0, and R0a replace pre-V, pre*V1, pre*V2, pre-HV, and (pre-HV)1, respectively.Murlo (N=86)Volterra (N=114)Casentino Valley (N=122)Total (N=322)H:27 (31.4)53 (46.5)46 (37.7)126 (39.1) H*14 (16.3)31 (27.2)15 (12.3)60 (18.6) H14 (4.7)13 (11.4)19 (15.6)36 (11.2) H35 (5.8)……5 (1.6) H53 (3.5)3 (2.6)8 (6.6)14 (4.3) H61 (1.2)1 (.9)…2 (.6) H11…2 (1.8)1 (.8)3 (.9) H14…1 (.9)…1 (.3) H15……2 (1.6)2 (.6) H19…2 (1.8)1 (.8)3 (.9)HV0:5 (5.8)7 (6.1)5 (4.1)17 (5.3) HV0*1 (1.2)3 (2.6)1 (.8)5 (1.6) HV0a1 (1.2)1 (.9)…2 (.6) V3 (3.5)3 (2.6)4 (3.3)10 (3.1)HV (without H/HV0):6 (7.0)3 (2.6)7 (5.7)16 (5.0) HV*5 (5.8)3 (2.6)6 (4.9)14 (4.3) HV11 (1.2)…1 (.8)2 (.6)R0:1 (1.2)5 (4.4)2 (1.6)8 (2.5) R0a1 (1.2)5 (4.4)2 (1.6)8 (2.5)R2:…1 (.9)…1 (.3) R2*…1 (.9)…1 (.3)J:7 (8.1)7 (6.1)4 (3.3)18 (5.6) J16 (7.0)5 (4.4)2 (1.6)13 (4.0) J21 (1.2)2 (1.8)2 (1.6)5 (1.6)T:8 (9.3)15 (13.2)13 (10.7)36 (11.2) T11 (1.2)3 (2.6)4 (3.3)8 (2.5) T27 (8.1)12 (10.5)9 (7.4)28 (8.7)U:26 (30.2)14 (12.3)29 (23.8)69 (21.4) U11 (1.2)1 (.9)…2 (.6) U2e5 (5.8)1 (.9)1 (.8)7 (2.2) U35 (5.8)1 (.9)3 (2.5)9 (2.8) U41 (1.2)4 (3.5)1 (.8)6 (1.9) U5a3 (3.5)2 (1.8)2 (1.6)7 (2.2) U5b……6 (4.9)6 (1.9) U6a1 (1.2)…1 (.8)2 (.6) U74 (4.7)1 (.9)3 (2.5)8 (2.5) K16 (7.0)4 (3.5)12 (9.8)22 (6.8)N1:…4 (3.5)2 (1.6)6 (1.9) N1b1……1 (.8)1 (.3) N1c…1 (.9)1 (.8)2 (.6) I…3 (2.6)…3 (.9)N2:1 (1.2)…7 (5.7)8 (2.5) N2a……1 (.8)1 (.3) W1 (1.2)…6 (4.9)7 (2.2)X:3 (3.5)2 (1.8)3 (2.5)8 (2.5) X23 (3.5)2 (1.8)3 (2.5)8 (2.5)M:1 (1.2)…2 (1.6)3 (.9) M1……2 (1.6)2 (.6) D4g11 (1.2)……1 (.3)Sub-Saharan L:1 (1.2)3 (2.6)2 (1.6)6 (1.9) L3d1 (1.2)3 (2.6)1 (.8)5 (1.6) L1b……1 (.8)1 (.3)a HV0, HV0*, HV0a, R0, and R0a replace pre-V, pre*V1, pre*V2, pre-HV, and (pre-HV)1, respectively. Open table in a new tab To determine whether the overall haplogroup frequencies in the Tuscan population samples were indeed different from those of other Italian or European populations, we compared haplogroup distributions in the three Tuscan populations (table 1) with those of 55 western Eurasian populations (including 7 from other Italian regions) (table 2), by using principal-component analysis (PCA)—a dimension-reduction method that seeks to explain the variance of multivariate data by a smaller number of variables (the principal components [PCs]), which are linear functions of the original variables—in this case, the haplogroup frequencies.12Cavalli-Sforza LL Piazza A Menozzi P The history and geography of human genes. Princeton University Press, Princeton, NJ1994Google Scholar After reducing the variables (haplogroups) to PCs, we reported the coordinates of the observations for the 58 populations in a four-dimensional graphic representing the genetic landscape of western Eurasia (fig. 1). We observed that the first PC is primarily east-west, separating Europeans from Near Easterners, and it accounts for 22% of the variation. The Basque country falls at one pole, Iran at the other. Such a strong geographical pattern, with the Near Easterners forming a well-defined group and the eastern and central Mediterranean populations of Europe as their most closely related neighbors, is in perfect agreement with the patterns reported elsewhere for both classical12Cavalli-Sforza LL Piazza A Menozzi P The history and geography of human genes. 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