*Correspondence and requests for materials should be addressed to D.M.B. (
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1 The genetic variation in the R1a clade among the Ashkenazi Levites' Y chromosome Doron M Behar* 1,2, Lauri Saag 1, Monika Karmin 1, Meir G. Gover 3, Jeffrey D. Wexler 4, Luisa Fernanda Sanchez 2, Elliott Greenspan 2, Alena Kushniarevich 1,5, Oleg Davydenko 5, Hovhannes Sahakyan 1,6, Levon Yepiskoposyan 6, Alessio Boattini 7, Stefania Sarno 7, Luca Pagani 1,8, Shai Carmi 9, Shay Tzur 9,1, Ene Metspalu 1,11, Concetta Bormans 2, Karl Skorecki 1,12, Mait Metspalu 1, Siiri Rootsi 1, Richard Villems 1,11 1 Estonian Biocentre, Tartu 511, Estonia 2 Genomic Research Center, Gene by Gene, Houston 778, Texas, USA 3 Independent Genetic Genealogy Researcher, Savyon 5695, Israel 4 levitedna.org, Los Angeles 945, California, USA 5 Institute of Genetics and Cytology, National Academy of Sciences of Belarus, 2272, Minsk, Belarus 6 Laboratory of Ethnogenomics, Institute of Molecular Biology of National Academy of Sciences, Yerevan, 14, Armenia 7 Department of Biological, Geological and Environmental Sciences, University of Bologna, Bologna, 4126, Italy 8 APE Lab, Dept. of Biology, University of Padova, 35121, Padova, Italy 9 Braun School of Public Health and Community Medicine, The Hebrew University of Jerusalem, Jerusalem, , Israel 1 Rambam Health Care Campus, Haifa 31961, Israel 11 Department of Evolutionary Biology, Institute of Molecular and Cell Biology University of Tartu, Tartu 511, Estonia 12 Ruth and Bruce Rappaport Faculty of Medicine, Technion-Israel Institute of Technology, Haifa, 31961, Israel *Correspondence and requests for materials should be addressed to D.M.B. ( behardm@genebygene.com)
2 R1a M42 M11784 Z645 R1a2 Z93 M17/M198 R1a1 Z283 R1a3 CTS4385 M582 Y YP268 GRC Ashkenazi Jew, Israelite GRC Ashkenazi Jew, Israelite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC12117 Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC949 Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC GS1545-ASM Ashkenazi Jew, Unknown GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC18273 Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC14189 Jewish, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Israelite GRC Ashkenazi Jew, Levite GRC Jew, Levite GRC4194 Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC11771 Ashkenazi Jew, Israelite GRC11348 Ashkenazi Jew, Levite GRC Ashkenazi Jew, Israelite GRC Turkish Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GS12263-ASM Ashkenazi Jew, Unknown GRC Turkish Jew, Levite GRC27612 European, Unspecified GRC9631 Ashkenazi Jew, Levite GRC Israeli Jew, Levite GRC12895 Ashkenazi Jew, Levite GRC1462 Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC24718 Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC24915 Ashkenazi Jew, Levite GS1627-ASM Ashkenazi Jew, Unknown GRC1622 Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Algerian Jew, Levite GRC Iraqi Jew_Israelite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GS14998-ASM Ashkenazi Jew, Unknown GRC European, Unspecified GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GS12262-ASM Ashkenazi Jew, Unknown GRC Ashkenazi Jew, Levite GRC Israeli Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC Ashkenazi Jew, Levite GRC917 Ashkenazi Jew, Levite GRC Ashkenazi Jew, Cohen GRC Iranian Kerman GRC Y GRC Iranian Azeri GRC Iranian Kerman GRC Spanish GRC Iranian Azeri GRC Arab, Israel GRC Arab, Israel GRC Balkar, North GRC Balkar, North GRC25392 Turkish, North GRC Scottish GRC Polish GRC Balkar, North GS15893-ASM Mixed GS13749-ASM Assyrian GS35237-ASM Azerbaijani GS16188-ASM Rushan Vanch, Tajikistan GS16276-ASM Bashkir GS13758-ASM Shugnan, Tajikistan GS35114-ASM Circassian, North Caucasus GS16198-ASM Iraqi Jew, Israelite GRC Iraqi Jew, Israelite GRC Iraqi GS16171-ASM Balkar, North Caucasus GS16974-ASM Mumbai Jew, Israelite GS22439-ASM Altaian, Siberia GS13757-ASM Kyrgyz GS16191-ASM Kyrgyz GS16196-ASM Ishkasim, Tajikistan GS14417-ASM Rushan Vanch, Tajikistan GS16972-ASM Iraqi Jew, Israelite GS1681-ASM Bengal, West GS14415-ASM Kazakh NA ASM Gujarati, India NA ASM Gujarati, India GS16799-ASM Gupta, India GS16966-ASM Kurmi, India GS1688-ASM Orissa, India GS16189-ASM Ishkasim, Tajikistan GS35239-ASM Cossack GS16875-ASM Iranian GS16193-ASM Kyrgyz GS16187-ASM Kyrgyz GS16174-ASM Altaian, Siberia GS16967-ASM Altaian, Siberia GS14329-ASM Komi GS3518-ASM Komi GS13756-ASM Russian, North GS13765-ASM Karelian GS1697-ASM Karelian GS16796-ASM Russian, West GS35177-ASM Ukrainian, West GS15891-ASM Ukrainian GS1327-ASM Ukrainian GRC Estonian GS16916-ASM Estonian GS16797-ASM Russian, Central GS35121-ASM Ukrainian, East GRC Estonian GRC Belarusian GS35113-ASM Circassian, North Caucasus GS1494-ASM USA, Unspecified GRC Belarusian GS35238-ASM Cossack GS253-DNA_B1_2_37-ASM Mixed GS1374-ASM Tatar GS1185-ASM USA, Unspecified GS3519-ASM Swede GS3524-ASM Swede GS3526-ASM Saami GS3541-ASM Hungarian GS35179-ASM Cossack GS1687-ASM Ashkenazi Jew, Unknown GS13755-ASM Ukrainian, North GS2417-ASM USA, Unspecified GS1433-ASM Mordvin GS35174-ASM Ukrainian, East GS35176-ASM Ukrainian, East GS16338-ASM Mixed GS35178-ASM Ukrainian, West GS15183-ASM USA, Unspecified GS14566-ASM Mixed GRC Kabardin, North GS16192-ASM Tajik GS16312-ASM Ashkenazi Jew, Unknown Figure S1. Haplogroup R1a phylogeny as reconstructed by BEAST with estimated coalescence ages of the nodes.
3 3282 GRC Arab, Israel 54 GS16213-ASM Arab, Israel 3396 GS13745-ASM Armenian 367 GS1618-ASM Saudi Arabian GS16277-ASM Saudi Arabian GRC Druze B GS1966-ASM Ashkenazi Jew, Unknown 494 GS1845-ASM Ashkenazi Jew, Unknown GS16887-ASM Yemenite Jew, Israelite 367 GS16973-ASM Yemenite Jew, Israelite GRC Bedouin 758 GRC Bedouin 586 GRC Afghani Jew, Cohen GS16136-ASM Druze 5466 GRC Ashkenazi Jew, Cohen GRC Ashkenazi Jew, Cohen 1488 GRC Bulgarian Jew, Cohen 5692 B GRC Ashkenazi Jew, Cohen GRC Ashkenazi Jew, Cohen GRC Moroccan Jew, Cohen GRC Ashkenazi Jew, Cohen GS1797-ASM Ashkenazi Jew, Unknown GS15788-ASM Ashkenazi Jew, Unknown 6216 GRC Yemenite Jew, Cohen GS35124-ASM Armenian GS35434-ASM Albanian 6484 GS14474-ASM Jordanian GS35125-ASM Armenian J1b P GRC Egyptian GRC Turkish Jew, Cohen GS35126-ASM Armenian GS1784-ASM Ashkenazi Jew, Unknown GS11735-ASM Mixed GS14368-ASM Kumyk, North Caucasus GRC Assyrian J1 M J1c PF7256 J1a Z M GS1445-ASM Avar, North Caucasus GS14421-ASM Tabasaran, North Caucasus GS9929-ASM USA, Unspecified GS13724-ASM Lezgin, North Caucasus GS13741-ASM Azerbaijani GRC Jerba Jew, Cohen 176 GRC Jerba Jew, Cohen 677 GRC Tunisian Jew, Unknown GS35115-ASM Kabardin, North Caucasus GS16177-ASM Kyrgyz GS13747-ASM Iranian GRC Ashkenazi Jew, Cohen GS15895-ASM USA, Unspecified J M M GS16445-ASM Unknown GS1682-ASM Kapu, India GS14997-ASM Ashkenazi Jew, Unknown GS1789-ASM Ashkenazi Jew, Unknown GS1692-ASM Ashkenazi Jew, Unknown 1197 GS1635-ASM Ashkenazi Jew, Unknown 1267 GS15875-ASM Tatar J2a M GS1677-ASM Ashkenazi Jew, Unknown GS1611-ASM Ashkenazi Jew, Unknown GS35685-ASM Georgian GS1431-ASM Mixed GS13751-ASM Assyrian GS1441-ASM Abkhazian J2 M B GRC Cochin Jew, Israelite 49 GS16138-ASM Cochin Jew, Israelite GRC Ashkenazi Jew, Cohen GS15174-ASM Mixed M GS35147-ASM Albanian GS16373-ASM Mixed J2b M M GRC Ashkenazi Jew, Cohen GRC Ashkenazi Jew, Cohen GS1434-ASM Mixed GS16882-ASM Mumbai Jew, Israelite GRC Palestinian GS13746-ASM Armenian I GS16181-ASM Jordanian GS13754-ASM Croat GS1614-ASM Belarusian Figure S2. Haplogroup J phylogeny as reconstructed by BEAST with estimated coalescence ages of the nodes.
4 GS1846-ASM Ashkenazi Jew, Unknown GS1959-ASM Ashkenazi Jew, Unknown 1272 GS1523-ASM Mixed 1341 GS253-DNA_H1_2_37-ASM Unknown GS15786-ASM Ashkenazi Jew, Unknown GS1799-ASM Ashkenazi Jew, Unknown Z838 GS15783-ASM Ashkenazi Jew, Unknown GS185-ASM Ashkenazi Jew, Unknown GRC Libyian Jew, Unknown 1258 GRC Ashkenazi Jew, Cohen GRC Arab, Israel 55 GS1626-ASM Arab, Israel GRC Bedouin GRC Ashkenazi Jew, Israelite GS14561-ASM Mixed 6188 GRC Palestinian M GRC Ashkenazi Jew, Israelite GRC Yemenite Jew, Cohen GRC Palestinian GRC Iraqi Jew, Israelite M GRC Turkish Jew, Unknown GRC Ashkenazi Jew, Unknown GRC Yemenite Jew, Unknown PF GS1961-ASM Ashkenazi Jew, Unknown 1125 GS253-DNA_F1_2_37-ASM Mixed GS16214-ASM Uzbeki Jew, Israelite 2168 GS16217-ASM Arab, Israel GRC Ashkenazi Jew, Israelite E2b CTS PF378 B GRC Italian Jew, Israelite GS1851-ASM Ashkenazi Jew, Unknown GS1579-ASM Ashkenazi Jew, Unknown GRC Ashkenazi Jew, Israelite 13 GS1679-ASM Ashkenazi Jew, Unknown E2b c Z GS319-DNA_D2_11_37-ASM Sandawe, Tanzania GS319-DNA_E2_11_37-ASM Sandawe, Tanzania GRC Moroccan Jew, Unknown E2c L M GRC Turkish Jew, Unknown GRC Tunisian Jew, Unknown GRC Ashkenazi Jew, Levite GRC Moroccan Jew, Unknown 3523 GS15892-ASM USA, Unspecified 3881 GS3521-ASM Albanian GS1726-ASM Estonian GRC Ashkenazi Jew, Unknown GS35227-ASM Australian GS15153-ASM USA, Unspecified E GRC Druze 857 M GRC Druze GS14562-ASM Mixed 4833 GS16441-ASM Mixed NA ASM Italian 4884 GS15172-ASM Mixed GS13763-ASM Mishari Tatar GRC Turkish Jew, Unknown GRC Ashkenazi Jew, Israelite 1362 B GS1848-ASM Ashkenazi Jew, Unknown GS17449-ASM Ashkenazi Jew, Unknown 199 GRC Ashkenazi Jew, Unknown E2a M NA ASM Maasai 144 NA ASM Maasai GRC Bedouin GRC Palestinian GS1578-ASM Mixed GS16179-ASM Iranian GS11811-ASM USA, Unspecified GS319-DNA_D1_11_37-ASM Bakola, Pygmy GS319-DNA_F1_11_37-ASM Hadza GS319-DNA_B1_11_37-ASM Baka, Pygmy NA ASM Luhya, Kenya NA ASM Yoruban GS35245-ASM Congo, Pygmy NA ASM African American GS319-DNA_C2_11_37-ASM Sandawe, Tanzania 3952 NA ASM Luhya, Kenya 4748 NA ASM Luhya, Kenya M GS319-DNA_E1_11_37-ASM Bedzan, Pygmy NA ASM African American NA ASM African American NA ASM Yoruban 6937 GS253-DNA_B2_2_37-ASM Unknown GS17267-ASM Yoruban NA19239-L ASM Yoruban E4 M GS17275-ASM Yoruban 126 GS17277-ASM Yoruban GS1724-ASM Yoruban GS16315-ASM Ashkenazi Jew, Unknown D GS3537-ASM Tamang, India NA ASM Japanese Figure S3. Haplogroup E phylogeny as reconstructed by BEAST with estimated coalescence ages of the nodes.
5 NA ASM Mexican GS1186-ASM USA, Unspecified GS1642-ASM European, Northwest 551 GS1548-ASM Ashkenazi Jew, Unknown GS16395-ASM European, Northwest GS16372-ASM Polish GS14413-ASM Kabardin, North Caucasus GS1629-ASM Iraqi Jew, Israelite GS17128-ASM Punjabi, Pakistan GS16463-ASM European, Northwest GRC Italian 1226 GS1696-ASM Ashkenazi Jew, Unknown 1441 GS14475-ASM Iranian G2a P GS16215-ASM Uzbeki Jew, Israelite GS13739-ASM North Ossetian, North Caucasus 3532 GS14414-ASM North Ossetian, North Caucasus 548 GS1449-ASM Abkhazian 6439 GS16899-ASM Moldavian G2 P287_eq 227 GS14366-ASM Lezgin, North Caucasus GRC Ashkenazi Jew, Israelite GS1844-ASM Ashkenazi Jew, Unknown 146 GRC Ashkenazi Jew, Israelite G2b M377_eq 5757 G2b2 BY764 GRC Ashkenazi Jew, Israelite 122 GS1638-ASM Ashkenazi Jew, Unknown 882 GS1952-ASM Ashkenazi Jew, Unknown J G1 M285 GS1714-ASM Punjabi, Pakistan GRC Yemenite Jew, Unknown GRC Assyrian GS35685-ASM Georgian Figure S4. Haplogroup G phylogeny as reconstructed by BEAST with estimated coalescence ages of the nodes.
6 GS16951-ASM Colla, Andean GS16942-ASM Andean, Argentina GS2274-ASM Colla, Andean GS1694-ASM Andean, Argentina GS1642-ASM Peruvian GS19989-ASM Colla, Andean GS1777-ASM Andean, Argentina NA ASM Mexican Q1a M GS16945-ASM Wichi, Andean GS16946-ASM Wichi, Andean GS1996-ASM Colla, Andean 1784 GS19997-ASM Eskimo 2773 GS13717-ASM Ket, Siberia M Q1c L GS13716-ASM Selkup, Siberia GS15393-ASM Ket, Siberia GS353-ASM Uzbeki L Q1d B GS15872-ASM Croat GS3538-ASM Nepalese GRC Yemenite Jew, Levite GS2267-ASM Koryak, Siberia GS2272-ASM Koryak, Siberia 1341 GS34185-ASM Koryak, Siberia GS3497-ASM Koryak, Siberia Q M Q2 F GS16667-ASM Peruvian GS19986-ASM Murut, Borneo GS16436-ASM Punjabi, Pakistan 126 GS1723-ASM Punjabi, Pakistan GS35118-ASM Turkmen 318 GS35119-ASM Turkmen GRC Ashkenazi Jew, Israelite 3496 Q3 M378 B GS1842-ASM Ashkenazi Jew, Unknown GRC Ashkenazi Jew, Israelite GS1843-ASM Ashkenazi Jew, Unknown 142 GS1632-ASM Ashkenazi Jew, Unknown GRC Moroccan Jew, Cohen GS15887-ASM Mixed R GRC1622 Ashkenazi Jew, Levite GS15787-ASM Ashkenazi Jew, Unknown Figure S5. Haplogroup Q phylogeny as reconstructed by BEAST with estimated coalescence ages of the nodes.
7 GRC Ashkenazi Jew, Israelite GS1621-ASM Georgian Jew, Israelite GS16212-ASM Arab, Israel 8738 GRC Ashkenazi Jew, Unknown 819 T1a GRC Ashkenazi Jew, Unknown L162_eq GRC Turkish Jew, Unknown 1951 T1a M GRC Moroccan Jew, Israelite GRC Palestinian LT B GRC Iranian Jew, Unknown GS16135-ASM Druze L1b M GS1639-ASM Ashkenazi Jew, Unknown GS1375-ASM Tatar Q 3253 GRC Moroccan Jew, Cohen NA ASM Mexican Figure S6. Haplogroup LT phylogeny as reconstructed by BEAST with estimated coalescence ages of the nodes.
8 GS16891-ASM German B2_eq PS_GS16891-ASM German GRC Ashkenazi Jew, Levite GS15711-ASM Puerto Rican HG ASM Puerto Rican GS15233-ASM Portugese GS1637-ASM Ashkenazi Jew, Unknown GS1645-ASM European, Northwest GS1647-ASM European, Northwest GS16375-ASM USA, Unspecified GS19959-ASM Colla, Andean GS19988-ASM Colla, Andean GS1571-ASM USA, Unspecified GS11814-ASM Mixed GS15229-ASM French GS1615-ASM Mixed GS1796-ASM Ashkenazi Jew, Unknown GS15874-ASM Mordvin GS11737-ASM USA, Unspecified GS15186-ASM Unknown NA12886-L ASM CEPH pedigree GS15223-ASM USA, Unspecified R1b1 GS16139-ASM Ashkenazi Jew, Unknown GRC Ashkenazi Jew, Levite L48 GS15787-ASM Ashkenazi Jew, Unknown GS11738-ASM USA, Unspecified GS16333-ASM USA, Unspecified GRC Tunisian Jew, Israelite GRC Druze NA ASM European, Northwest GS14563-ASM USA, Unspecified NA ASM Mexican GS1189-ASM USA, Unspecified GS16885-ASM Arab, Israel GS1436-ASM USA, Unspecified GS1636-ASM Ashkenazi Jew, Unknown NA ASM Italian GS14568-ASM Mixed GRC Belmontese Jew, Israelite GS13195-ASM Peruvian GS1181-ASM USA, Unspecified GS14565-ASM USA, Unspecified GRC Italian NA ASM European, Northwest GS35175-ASM Ukrainian, West GS15227-ASM Mixed GS1629-ASM Mixed GS12712-ASM Mixed GS1627-ASM USA, Unspecified GRC Estonian GS2414-ASM USA, Unspecified L11_eq GS1174-ASM Mixed GS699-ASM USA, Unspecified GS15175-ASM Mixed GS179-ASM Ashkenazi Jew, Unknown GS15155-ASM Mixed GS1426-ASM Mixed GS1437-ASM USA, Unspecified M412 GS1187-ASM USA, Unspecified GS992-ASM Mixed GS9926-ASM Mixed GRC Italian Jew, Unknown GS12713-ASM Mixed GS1798-ASM Ashkenazi Jew, Unknown GS1375-ASM Assyrian M269 GS14559-ASM Mixed GS35173-ASM Armenian GS16133-ASM Turkish Jew, Unknown GS16886-ASM Moroccan Jew, Israelite GS14364-ASM Tabasaran, North Caucasus GS1446-ASM Avar, North Caucasus R1b11 Z215_eq GRC15561 Ashkenazi Jew, Unknown GS1632-ASM USA, Unspecified GRC Estonian L754 B38 GS16881-ASM Kurdish Jew, Israelite GRC Georgian Jew, Unknown GRC Yemenite Jew, Unknown R1b R1b13 GS16134-ASM Bashkir M343 M73 GS35234-ASM Mongolian R1b15 B1 R1b14 PF6292 GS16312-ASM Ashkenazi Jew, Unknown GS16192-ASM Tajik R1a GRC1622 Ashkenazi Jew, Levite GS35176-ASM Ukrainian, East Figure S7. Haplogroup R1b phylogeny as reconstructed by BEAST with estimated coalescence ages of the nodes.
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