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Geographical Structure of the Y-Chromosomal Genetic Landscape of the Levant: a Coastal-Inland Contrast

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Geographical Structure of the Y-Chromosomal Genetic Landscape of the Levant: a Coastal-Inland Contrast RC24853 (W0909-003) September 1, 2009 Other IBM Research Report Geographical Structure of the Y-chromosomal Genetic Landscape of the Levant: A Coastal-inland Contrast Mirvat El-Sibai1, Daniel E. Platt2, Marc Haber1, Yali Xue3, Sonia C. Youhanna1, R. Spencer Wells4, Hassan Izaabel5, May F. Sanyoura1, Haidar Harmanani1, Maziar Ashrafian Bonab6, Jaafar Behbehani7, Fuad Hashwa1, Chris Tyler-Smith3, Pierre A Zalloua1,8, Genographic Consortium9 1The Lebanese American University, Chouran, Beirut 1102 2801, Lebanon 2IBM Research Division, Thomas J. Watson Research Ctr., P.O. Box 218, Yorktown Heights, NY 10598, USA 3The Wellcome Trust Sanger Institute, Wellcom Trust Genome Campus, Hinxton, Cambs. CB10 1SA UK 4The Genographic Project, National Geographic Society, Washington, D.C. 20036, USA 5Laboratoire de Biologie Cellulaire & Génétique Moléculaire (LBCGM), Université IBNZOHR, Agadir, Maroc 6University of Portsmouth, School of Biological Sciences, King Henry I Street, Portsmouth PO1 2DY, UK 7Dept. of Community Medicine and Behavioural Sciences, Kuwait University, P.O. Box 24923, Safat, Kuwait 8Harvard School of Public Health, Boston, MA 02215, USA 9Consortium members are listed after Acknowledgements Research Division Almaden - Austin - Beijing - Cambridge - Haifa - India - T. J. Watson - Tokyo - Zurich LIMITED DISTRIBUTION NOTICE: This report has been submitted for publication outside of IBM and will probably be copyrighted if accepted for publication. It has been issued as a Research Report for early dissemination of its contents. In view of the transfer of copyright to the outside publisher, its distribution outside of IBM prior to publication should be limited to peer communications and specific requests. After outside publication, requests should be filled only by reprints or legally obtained copies of the article (e.g. , payment of royalties). Copies may be requested from IBM T. J. Watson Research Center , P. O. Box 218, Yorktown Heights, NY 10598 USA (email: [email protected]). Some reports are available on the internet at http://domino.watson.ibm.com/library/CyberDig.nsf/home . 1 2 3 Geographical Structure of the Y-chromosomal Genetic 4 5 Landscape of the Levant: A coastal-inland contrast 6 1 2 1 3 1 4 7 Mirvat El-Sibai , Daniel E. Platt , Marc Haber ,YaliXue , Sonia C. Youhanna , R. Spencer Wells , 8 Hassan Izaabel5, May F. Sanyoura1, Haidar Harmanani1, Maziar Ashrafian Bonab6, Jaafar Behbehani7, 9 Fuad Hashwa1, Chris Tyler-Smith3, Pierre A. Zalloua1,8,∗ and The Genographic Consortium9 10 1The Lebanese American University, Chouran, Beirut 1102 2801, Lebanon 11 2Bioinformatics and Pattern Discovery, IBM T. J. Watson Research Centre, Yorktown Hgts, NY 10598, USA 12 3The Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, Cambs. CB10 1SA, UK 13 4The Genographic Project, National Geographic Society, Washington, DC 20036, USA; 14 5Laboratoire de Biologie Cellulaire & Gen´ etique´ Moleculaire´ (LBCGM), Universite´ IBNZOHR, Agadir, Maroc 15 6Biological Sciences; University of Portsmouth, School of Biological Sciences, King Henry I Street, Portsmouth PO1 2DY, U.K. 16 7Department of Community Medicine and Behavioural Sciences, Faculty of Medicine, Kuwait University, PO Box 24923, Safat, Kuwait 17 8Harvard School of Public Health, Boston, MA 02215, USA. 18 9Consortium members are listed after Acknowledgements 19 20 21 22 SUMMARY 23 24 We have examined the male-specific phylogeography of the Levant and its surroundings by analyzing Y-chromosomal 25 haplogroup distributions using 5,874 samples (885 new) from 23 countries. The diversity within some of these haplogroups 26 was also examined. The Levantine populations showed clustering in SNP and STR analyses when considered against 27 a broad Middle-East and North African background. However, we also found a coastal-inland, east-west pattern of 28 diversity and frequency distribution in several haplogroups within the small region of the Levant. Since estimates of 29 effective population size are similar in the two regions, this strong pattern is likely to have arisen mainly from differential 30 migrations, with different lineages introduced from the east and west. 31 32 33 Keywords: Y chromosome, Y-SNP, Y-STR, Levant 34 35 36 Introduction Within recent millennia, the Levant has been a frequent 37 focus of conquest by a variety of states and imperial pow- The Levant lies in the eastern Mediterranean region, south 38 ers (Issawi, 1988). It was home to some of the oldest cities of the mountains of Cilicia (South Turkey) and north of the 39 (Jericho, Byblos and Damascus), a path of some of the old- Sinai Peninsula. Throughout human prehistory and history, 40 est and later most important trade routes, and a cradle to this territory has been a key area, due to its geographical 41 three of the World’s Great Religions (Hitti, 1957). Signa- location linking three continents: Europe, Asia and Africa. 42 tures of the genetic legacy of the Phoenician expansion and It was on one of the early out-of-Africa migration routes 43 the Diaspora (Zalloua et al., 2008a; Aubet, 1993), and traces (Bar-Yosef, 1992; Tchernov, 1994; Luis et al., 2004; Stringer 44 of the Muslim expansion and the Crusaders (Zalloua et al., et al., 1989; Lahr & Foley, 1998), is believed to the be the first 45 2008b; Lamb, 1930) have been identified. However, there recipient of migration waves from East Africa seeking milder 46 remain a number of known historical events, ranging from climatic conditions after the Last Glacial Maximum (LGM) 47 the Byzantine expansion, the Ottoman expansion, periods and was the corridor for Neolithic migrations from the Fertile 48 of Egyptian, Babylonian, Persian, Hittite (Hitti, 1957) and Crescent to Europe and North Africa (Cavalli-Sforza, 1997). 49 other subjugations, as well as the likelihood of many un- 50 known prehistoric and historic events, whose genetic impact 51 ∗ is yet to be revealed. During the Bronze Age, the land south- Corresponding author: Dr. Pierre Zalloua, The Lebanese Amer- west of the Fertile Crescent was called the land of Canaan, 52 ican University, Chouran, Beirut 1102 2801, Lebanon. Tel: 53 + 961-1-784408 Ext. 2855; Fax: + 961-9-546090; E-mail: part of which became Phoenician territory after 1200 BCE 54 [email protected] (Issawi, 1988). This land included most of the coastal territory C 1 M. El-Sibai et al. 2 of the eastern Mediterranean countries. These countries Materials and Methods 3 shared a common culture and history (Hourani, 1946) 4 but some population expansions and migrations have af- Subjects and Comparative Datasets 5 fected Lebanon and Syria in different ways (Harris, 2003). 6 Such movements include the Aramean rule in Syria A total of 885 new samples from five populations (Syria, Jor- dan, Iran, Egypt and Kuwait) were collected and analyzed for 7 (Harris, 2003), the Sea Peoples disruption to some coastal this study. In addition, samples and genotyping data from 951 8 cities north and south of modern day Lebanon (in the Phoeni- Lebanese, 200 Syrian and 101 Palestinian men (Akka) were al- 9 cian era) (Harden, 1971) and the Ottoman occupation (Akarli, ready available (Zalloua et al., 2008a; 2008b). All the participants 10 1993). had three generations of paternal ancestry in their country of 11 Modern Jewish populations have a special phylogeograph- birth. Each provided detailed information on their geographical 12 ical status within the Levant for several reasons. Even by the origin (Table S1) and gave informed consent for this study, which 13 Roman era, there was a significant Diaspora, as attested to by was approved by the IRB committee of the Lebanese Ameri- 14 Strabo, Philo, Josephus, and Cicero (Stern, 2007). The Dias- can University. The 1879 Levantine samples (Lebanese, Syrians, 15 pora fragmented into branches with distinct and well-studied Jordanians and Palestinians) were classified into eight regions. 16 genetic signatures, such as those of Sephardim (Adams et al., These regions mainly reflect geographical subdivisions within 17 2008) and Ashkenazi (Behar et al., 2003; Nebel et al., 2005) modern day Lebanon and Syria (Table S3a and Fig. 1). The Lebanese samples, collected from 20 different cities (Table S3b), 18 and incorporated significant levels of male admixture (Adams were classified into the Lebanese coast (LC) which contains 19 et al., 2008). Current Jewish populations in the Levant derive coastal Lebanese cities including Beirut, Tyre, Sidon, Jounieh 20 largely from a complex pattern of resettlement from multi- and Tripoli, and Lebanese inland (LI) which contains samples ∼ 21 ple sources within the last 50 years and may not represent from Bekaa and Nabatiyeh. The Palestinian samples (PS), were 22 the pre-Diaspora distribution (Baron, 2007). Jewish genetics collected from men currently residing in Lebanon but who orig- 23 have been studied more than those of many other groups inated from Akka. Finally, Syrian samples were from 17 different 24 (Carmeli, 2004). The current study therefore seeks to fo- Syrian cities (Table S3b) and were divided into coast (SC) and 25 cus on a geographical genetic profile through the Levant and four inland regions: inland north (SIN), centre (SIC), south (SIS) 26 surrounding regions among the relatively less well character- and east (SIE) (Table S3a and Fig. 1). 27 ized populations, highlighting the product of other migra- For comparative data on haplogroup frequencies, we used pub- 28 tory and population processes that influenced the Levantine lished sources (Table S2). In addition to the previously
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