MAMMALS OF THE EASTERN MEDITERRANEAN REGIONj THEIR ECOLOOf, SYSTSMATICS AND ZOOGEOGRAFHICAL RELATIONSHIPS Sana Isa Atallah, B,S., M,S. American University of Beirut, Beirut, Lebanon, 1963 American University of Beirut, Beirut, Lebanon, 1965 A Dissertation Submitted In Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy at The University of Connecticut 1969 Copyright by SANA ISA ATALLAH 1969 APPROVAL PAGE Doctor of Philosophy Dissertation MAMMALS OF THE EASTERN MEDITERRANEAN REGIONj THEIR ECOLOGY, SYSTEMATICS AND ZOOGEOGRAPHICAL RELATIONSHIPS Sana Isa Atallah, B.S., M.S. Major Adviser \a^. V_a $ -g~tcr o The University of Connecticut 196? ii ACKNOWLEDGMENTS The c<- apletj.cn of this work would have not been possible without the constant assistance and advice of my major advisor. Dr. Ralph M. Wetzel, I am also greatly indebted to Dr. and Mrs. Robert E. Lewis, Iowa State University, Ames, Iowa, previously at the Dept, of Biology, American University of Beirut, Lebanon, for their very kind assistance, direction and advice while in Lebanon during the years 1963-1966, and Dr. David L, Harrison, Sevenoaks, Kent, England, for his help in the field and in identifying and comparing many specimens with material in his personal collection and at the British Museum (Natural History) collections, I am also very grateful to Drs, Ralph M. Wetzel, James A, Slater and George A, Clark at the University of Connecticut and Dr. Homy W, Sotzcr at the Smithsonian Institution for their useful suggestions and cidtical reading of the mar.uyeript. Thanks are also duo to my parents, Mr. Jacob Qumaioh, Miss Jean Uridgwood, Mr. Charles Bridgeood, Mr. Richard Biddgwcod, Mr. George Kassis, Mr, Hanna Qumaioh, Mr. George Khuri and Dr. J. Morton Boyd fur their valuable assistance in the field; Mrs. Stephana Schaefer, Miss Susan Kramer and Dr. David Harrison for providing the text figures; and Miss Kristina Hosko for her help in assembling Appendix X, Finally, I wish to exbend my sincere thanks to all my friends and colleagues for their constant snoouragment and assistance. iii TABLE OF CONTENTS PAGE Aclcno'. ledgmenta ;........... lit Lis b ff tables............t •«vi List lit’ illustrations.................... .......vii Intrci’uction................................................................ -1 Tho E? stem Mediterranean Region.................................................. 12 A. The People and the.Land............ 13 B. Physical Geography........................ 17 C. Climate.......................... 30 D. General Ecology and Fhytogeography..................37 Materials and Methods..................................................................... U? Species Accounts.........................................................................................53 Order Insectivora.5^ Order Chiroptcra............................ 88 Order Lagcmorpha,...^x..,).........1^42 Order Rodentia...c...... Order Carnivora..................... «2/;0 Order Hyracoidea.............«2U5 Order Artiodactyla..2lt6 Domesticated aiid Introduced Mammals.,.................2149 iv PAGE Zoog^ .raphlcal Summary........................................... 2$3 Bibli;. jraphy................... .2^9 Adder* ait to Bibliography.........................................................280 Appen.. Lx I - List of Measurements................281 LIST OF TABLES TABLE PAGE 1. Checklist of the mammals of tho Eastern 7 Mediterranean Region. 2. Comparison of monthly rainfall and mean 31 Kiaximurrt and minimum temperatures of six localities. ■vi LIST OF ILLUSTRATIONS IGUR2 PAGE 1. Tli.) Eactai-n Mediterranean Regionj collection 2 and other important localities cited in tlvi text. 2. Schematic representation of regions and sub- 18 regions pertaining to the physical geography of the Eastern Mediterranean Region. 3. Biocli;.iatic zones of tho Eastern Mediterranean 35 Region. It. Ecological regions of tho Eastern Mediterranean 39 Region, 5. The glans penis of A. Crocidnra russula. SA 60 1153 from Afka, Lebanon, and B, Erinaceus europaeus, SA 1063 from near Damascus, Syria. 6, Head of Nycatalus noctula lebanoticus 129 (HARR. 26,33^0) collected at' Natural Bridge^, Faraya, Lebanon... 7« Postero-lateral view of skull A. Meriones 169 libycus B. Meriones crassus C. Morionos tristrami D, Sekeetamys calurus. vii INTRODUCTION The E. •item Mediterranean Region, aa understood here (Fig.l), represents the smallest natural unit that can be satisfactorily delineated from tho Middle East ( » Near East), and in essence it correspond:; to the left horn of the so called "Fertile Crescent" and the in' ervening Syrian Desert. The Fertile Crescent includes many centeis of ancient civilizations and extends from the Tigris and Euphrates River Valleys on the east, westward along the southern slopes of the Taurus Mountains and southwards along the land bordering the eastern Mediterranean to the Gulf of Aqaba on the Red Sea (Aramco Handbook, 1968). The Eastern Mediterranean has been of great zoogeographical importance since the Miocene, when this land first emerged from under tho Sea of Tethys* Its importance rests on its location^ along the dispersal routes between the two major Old World land mas­ ses: Eurasia and Africa. Consequently, one finds among tho endemic Palaearctic fauna of this region mammals characteristic of the Oriental and Ethiopian Zoogeographical Regions. The objectives of this vork are to provide,as much as possible, information on the ecology, distribution, systematics and zoogeo­ graphical relationships of mammals presently occurring along the eastern Mediterranean. Data presented is based on published records, unpublished museum records and original observations accumulated through extensive field work in tho area during tho years 1963-1966 and the summer of 1968, 1 2 EASTERN MEDITERRANEAN REGION 0 ZS SO 15 iqo Km. s| y„„ • i Jw...* ■'■---' Haita W H^TAMYK P ',* •N**Wi},?«»1 -r • K»AI . TtbAvivLviv w 0*«O v* *.***» Yafo0 I • *****«*«■•« t*M U • ZlOH i *t.waugnT*«»M f ’ilHOVcFT ^ XUU|«HlH '&tY«kA«%Y / /'• &MI| t OMvtUU vSAh«A e.lM, .W.OftT«AU« . StOHi, , „ VAlA/*_/ #T»LMil-. Giyr%ti.W * kmKm Y»**r% *”®***S"t8A “E»kY»8 \*. »Tat»u» \ SyR^N ShouQAa . EL.-S»fS • HA«g • t.u ma«* • EL-Ou^t'ftft LiuAtV-# _ S\K Fig. 1. The Eastern Mediterranean Region; collection sites and other important localities cited in the text. Elevations are in m. 3 Ninety -Tour species and one hundred subspecies of terrestrial, wild mamma}* are recognized from the Eastern Mediterranean Region (Table l). These figures include only wild mammals occurring at the present time} hoviever thirteen additional species of domes­ ticated anti introduced mammals are briefly mentioned and their impact on the environment discussed, A brief historical review of the principal publications dealing with the mammals of the Eastern Mediterranean Region is given below in chronological order, Pi-e 1600 Conspicuous mammals such as the wild ass, coney, ibex, hyaena, wild boar, leopard, cheetah, etc. are continuously referred to in the Bible and in the works of early travelers and pilgrims (Aharoni, 1938$ Bodenheimer, I960). Later descriptions of many mammals appeared in the writings of Arab scientists such as Ad- Damiri (bom 13h9 A.D.) and Al-Qazwini(born 1281-2 A.D.)j tho former in his two volume book "Hayat El-Hayawan *» The Life of Animals" and the latter in two contributions "Huzhat Ul-Qulub ** Hearts' Delight" and "Aja'eb Ul-Makhloukat - The Wonders of the Living", The earliest systematic collections were those of E. Hasselquist and E. Geoffroy St. Hilaire who traveled into southern Palestine from Sinai and North Africa in the mid and late eighteenth century. 18CO-1900 The foundations of our present knowledge of mammals in the Eastern Mediterranean Region were laid in this period. Many German and English travelers obtained specimens that were later described a by VJagnor, brants, 0. Thomas and A. Nehring. Canon 'f. B. Tristram made four trips to Palestine that re­ sulted in h_s major contribution "Fauna and Flora of Western Palestine" published in 188U, Hovever, he devoted less than 30 pages to his discussion of 113 species, including fossil forms, of domesticated and wild mammals» This work is now outdated and not available except in largo libraries, also many of the species were incorrectly identified. Other scientists of importance at this time were H. C. Hart, T. von Heug.lin, E. Rueppoll and C. Cretscchmar; also valuable collections were made by Baron von Schubert and the botanist Kotschy, 1900-1925 Several collections were made during this period especially through the endeavors of British Army Staff such as S, S. Flower, R. I, Pocock and other collectors such as D. Carruthers, H. Schliiter, P, A. Buxton, E. Schmitz, S. Merrill and I.Aharoni. These collectors provided European museums with indispensable material, and also published in accounts of their travels valuable information on the habits and habitats of many species of mammals. From these collec­ tions many new species were described by museum personnel at London and Berlin, of these 0. Thomas, A. Nehring, P. Matschie and F. Muller are especially worthy of mention. I. C. Phillips assisted by W. M, Mann traversed the country south of Mt. Hermon (March-June, 19lll) and collected over 150 small mammals some of which, wore later described by G. M. Allen (1915) at Harvard. 5 192S-19SO Many p-iblications appeared in this period, but only a few are noted here. Tho mu roids of Palestine were described by B. Aharoni (1932) and those of Asia Minor by G, Neuhauser (1936).