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United States National Museum Bulletin 275 SMITHSONIAN INSTITUTIO N MUSEUM O F NATURAL HISTORY UNITED STATES NATIONAL MUSEUM BULLETIN 275 The Rodents of Libya Taxonomy, Ecology and Zoogeographical Relationships GARY L. RANCK Curator, Mammal Identification Service Division of Mammals, U.S. National Museum SMITHSONIAN INSTITUTION PRESS WASHINGTON, D.C. 1968 Publications of the United States National Museum The scientific publications of the United States National Museum include two series, Proceedings of the United States National Museum and United States National Museum Bulletin. In these series are published original articles and monographs dealing with the collections and work of the Museum and setting forth newly acquired facts in the field of anthropology, biology, geology, history, and technology. Copies of each publication are distributed .to libraries and scientific organizations and to specialists and others interested in the various subjects. The Proceedings, begun in 1878, are intended for the publication, in separate form, of shorter papers. These are gathered in volumes, octavo in size, with the publication date of each paper recorded in the table of contents of the volume. In the Bulletin series, the first of which was issued in 1875, appear longer, separate publications consisting of monographs (occasionally in several parts) and volumes in which are collected works on related subjects. Bulletins are either octavo or quarto in size, depending on the needs of the presentation. Since 1902, papers relating to the botanical collections of the Museum have been published in the Bulletin series under the heading Contributions from the United States National Herbarium. This work forms number 275 of the Bulletin series. Frank A. Taylor Director, United States National Museum U.S. GOVERNMENT PRINTING OFFICE WASHINGTON : 1968 For sale by the Superintendent of Documents, U.S. Government Printing Office Washington, D.C. 20402 - Price $1.50 (paper cover) Contents Page Acknowledgments 1 Introduction 3 Physical description of Libya 6 General features 6 The coastal plain 7 The coastal escarpment 7 Pre-Saharan or steppe 7 Hamadas 8 Sand seas 9 Serirs 9 Fesh-fesh 10 Depressions 10 Mountainous areas of the interior 10 The Cyrenaican Plateau 11 The Tripolitanian Gebel 11 Geology 12 Tripolitania and Cyrenaica 12 Fezzan 13 Volcanic activity 14 Climate 14 Libyan interior 14 Arid or desert type 14 Semiarid or steppe type 15 Mediterranean littoral 16 Coastal uplands 17 Additional meteorological data 17 Phytogeography 18 General features of the Saharan flora 18 Phytogeographic characteristics 19 Systematic composition 19 Endemism 20 Adaptation to the desert environment 20 The vegetative cover of Libya 21 Coastal plain 21 Cyrenaican Plateau 22 Tripolitanian Gebel 23 Saharan steppe 23 Sand seas, hamadas, and serirs 23 Depressions or sebchets 24 Oases 24 The grass cover of Libya 25 Mammalian Faunal Areas 26 Mediterranean Faunal Area 27 Saharan Steppe Faunal Area 30 Saharan Desert Faunal Area 30 VI CONTENTS Page Speciation and geographical variation 34 Geographic variation in response to selective factors of the habitat . 35 Modes of speciation 36 Sympatric speciation 37 Geographic speciation 39 Clinal variation 39 Geographic isolates 40 Zoogeographical considerations 44 Effects of the Pleistocene on the dispersal of Saharan and Libyan rodents 44 Zoogeographical relationships of the rodents of the Sahara and South- west Asia 53 Conclusions 61 Systematic treatment 62 Plan of treatment 62 Gazetteer of principal collecting localities 67 Family Cricetidae 69 Subfamily Microtinae 69 Genus Microtus Schrank 69 Microtias mustersi Hinton 69 Subfamily Gerbillinae 73 Genus Gerbillus Desmarest 73 Gerbillus aureus aureus Setzer 87 Gerbillus aureus favillus Setzer 89 Gerbillus aureus nalulensis, new subspecies 90 Gerbillus eatoni eatoni Thomas 95 Gerbillus eatoni inflatus, new subspecies 97 Gerbillus eatoni versicolor, new subspecies 98 Gerbillus gerbillus aeruginosus, new subspecies 103 Gerbillus gerbillus discolor, new subspecies 106 Gerbillus gerbillus gerbillus (Olivier) 109 Gerbillus gerbillus latastei Thomas and Trouessart 110 Gerbillus gerbillus psammophilous, .new subspecies 112 Gerbillus pyramidum hamadensis, new subspecies IIS Gerbillus pyramidum tarabuli Thomas 121 Gerbillus amoenus vivax (Thomas) 125 Gerbillus campestris brunnescens, new subspecies 133 Gerbillus campestris dodsoni (Thomas) 136 Gerbillus campestris haymani Setzer 139 Gerbillus campestris patrizii (de Beaux) 141 Gerbillus campestris wassifi Setzer 143 Gerbillus grobbeni Klaptocz 146 Gerbillus henleyi henleyi (de Winton) 147 Gerbillus kaiseri Setzer 149 Genus Pachyuromys Lataste 153 Pachyuromys duprasi natronensis de Winton 153 Genus Merioncs Illiger 157 Meriones caudatus amplus, new subspecies 165 Meriones caudatus caudatus Thomas 169 Mi r tones caudatus confalonierii de Beaux 172 CONTENTS VII Page Family Cricetidae— Continued Subfamily Gerbillinae—Continued Genus Meriones Illiger—Continued Meriones caudatus luridus, new subspecies 173 Meriones crassus tripolius Thomas 176 Meriones libycus auratus, new subspecies 183 Meriones libycus azizi Setzer 185 Genus Psammomys Cretzschmar 187 Psammomys obesus obesus Cretzschmar 190 Psammomys obesus tripolitanus Thomas 192 Psammomys vexillaris vexillaris Thomas 192 Family Spalacidae 194 Genus Spalax Giildenstaedt 194 Spalax ehrenbergi aegyptiacus Nehring 194 Family Muridae 199 Genus Rattus Fischer 199 Rattus norvegicus (Berkenhout) 199 Rattus rattus (Linnaeus) 200 Genus Mas Linnaeus 203 Mus musculus Linnaeus 203 Genus Acomys Geoffroy 208 Acomys cahirinus viator Thomas 208 Family Gliridae 212 Genus Eliomys Wagner 212 Eliomys quercinus cyrenaicus Festa 214 Eliomys quercinus denticulatus, new subspecies 216 Eliomys quercinus tunetae Thomas 219 Family Dipodidae 220 Genus Jaculus Erxleben 220 Jaculus deserti favilhis Setzer 224 Jaculus deserti fuscipes, new subspecies 226 Jaculus deserti rarus, new subspecies 228 Jaculus deserti vastus, new subspecies 230 Jaculus jaculus arenaceous, new subspecies 234 Jaculus jaculus collinsi, new subspecies 236 Jaculus jaculus cufrensis, new subspecies 238 Jaculus jaculus tripolilanicus, new subspecies 240 Jaculus jaculus whitchurchi, new subspecies 242 Jaculus oricntalis orientalis Erxleben 244 Genus Allactaga Cuvier 248 Allactaga letradactyla (Lichtenstein) 248 Family Hystricidae 251 Genus Hystrix Linnaeus 251 Hystrix cristata Linnaeus 251 Family Ctenodactylidae 252 Genus Clenodactylus Gray 252 Ctenodactylus gundi gundi (Rothman) 254 Ctenodactylus gundi vali Thomas 259 Literature cited 260 Plates 265 Acknowledgments For the collecting of specimens and for aid in some aspects of the field work, I am indebted to James H. Shaw and Henry W. Setzer. My sincere appreciation is extended to the following staff members and technical personnel of the United States National Museum, Smithsonian Institution, and of the Bird and Mammal Laboratories, United States Fish and Wildlife Service, for their assistance in various ways: Terry K. Cederstrom, Helen S. Gaylord, Arthur M. Greenhall, Theodore A. Heist, David H. Johnson, Burrowes T. Lovinggood, Mary W. Mann, Richard H. Manville, John H. Miles, Jr., John L. Paradiso, and Lyman B. Smith. I would like to acknowledge the assistance of Donald J. Ortner, Museum Technician, Division of Physical Anthropology, for informa- tion regarding statistical computations; and Jack Scott and other members of the Photographic Laboratory, Museum of Natural History, for the reproduction of the maps and charts which appear in this publication. Many additional staff members and technical personnel from other divisions of the United States National Museum have also given freely of their advice and services. For the acquisition of various items of field equipment and assist- ance in all of the field work in Libya, I thank the following officers of the 64th Engineering Battalion, Base Topographic Survey, Wheelus Air Force Base, Tripoli, Libya: Lt. Col. Thomas Whitchurch, Major Guy Brunnacci, Captain William McMullen, and Chief Warrant Officer Lynn Walters. In addition, I would like to acknowledge the assistance of the many other officers and enlisted men of the 64th Engineering Battalion, who aided in the field work in Libya. I desire to express my appreciation to the following individuals for their contributions enabling me to operate more efficiently in Libya: Dr. Mohammed Ayub, Director of Antiquities for the Fezzan, Libya; Randy Bowmar, Director of United States Overseas Mission, Fezzan District, Libya; Aiad Ali Elwinsi, guide and interpreter; James R. Jones and Gordon Chase Tibbits, hydrologists of the United States Geological Survey and assigned to the United States Overseas Mis- sion, Libya; Howard Meadows, Agricultural Advisor for the Fezzan, United States Overseas Mission, Libya. For permission to study specimens in their collections and for the loan of comparative materials, I thank Dr. G. C. Corbett and Mr. G. W. Hayman of the Mammal Room, British Museum (Natural 1 2 U.S. NATIONAL MUSEUM BULLETIN 275 History), London; and Dr. Francis Petter, Curator of Mammals, Museum National d'Histoire Naturelle, Paris. I am particularly indebted to Dr. Henry W. Setzer, Associate Curator, Division of Mammals, United States National Museum, Smithsonian Institution, for bis efforts in arranging for my participa- tion in the Libyan project, his untiring criticism of the manuscript, and for his continual encouragement and support of this research. To Professor Stephen D. Durrant, Department of Zoology and Entomology, University of Utah, and to
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