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Full text of "The contemporary land mammals of Egypt (including Sinai)"

ILL . / 

AT URBAK ,. ,,!AMPAIGf 
BIOLOGY 

JUL 1 9 1982 



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UNIVERSITY OF ILLINOIS LIBRARY AT URBANA-CHAMPAIGN 



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MAR 2 i 1931 



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5- "lELDIANA ii»iom wsiMY s«iivo 

Zoology AUG2 5 1980 

Published by Field Museum of Natural History UDJIAIIi 

New Series, No. 5 



THE CONTEMPORARY LAND MAMMALS 
OF EGYPT (INCLUDING SINAI) 

DALE J. OSBORN 
IBRAHIM HELMY 



OCT 71S80 



August 15. 1980 
Publication 1309 



FIELDIANA 
Zoology 

Published by Field Museum of Natural History 



New Series, No. 5 



THE CONTEMPORARY LAND MAMMALS 
OF EGYPT (INCLUDING SINAI) 

DALE J. OSBORN 

Research Associate 

Field Museum of Natural History 

and 

Mammalogist 

Department of Medical Zoology 

U.S. Naval Medical Research Unit No. 3 

American Embassy 

Cairo, Arab Republic of Egypt 

IBRAHIM HELMY 

Field Research Supervisor 

Department of Medical Zoology 

U.S. Naval Medical Research Unit No. 3 

American Embassy 

Cairo, Arab Republic of Egypt 



From Research Project MR041.09.01-0152, Naval Medical Research and Develop- 
ment Command, National Naval Medical Center, Bethesda, Maryland. The opinions 
and assertions contained herein are the private ones of the authors and are not to be 
construed as official or as reflecting the views of the Department of the Navy or of 
the naval service at large. 

Dr. Osborn's present address is Ladova 7, 12800 Praha 2, Czechoslovakia, S.R. 
Dr. Osborn's employment at NAMRU-3 was supported by Office of Naval Research 
grant Nonr 4414(00)NR 107-806. 

This publication was supported in part by NIH Grant #LM02765 from The Na- 
tional Library of Medicine. 

August 15, 1980 
Publication 1309 



Library of Congress Catalog Number: 79-51549 

US ISSN: 0015-0754 

PRINTED IN THE UNITED STATES OF AMERICA 



mo. s- 7 



TABLE OF CONTENTS 

List of Illustrations x 

List of Tables xv 

Preface xviii 

Introduction 1 

Review of Literature 1 

Materials and Methods 3 

Study Area 6 

TbPOGRAPHY 9 

Sinai Peninsula 9 

Eastern Desert 10 

Nile Valley and Delta 14 

Western Mediterranean Coastal Desert 16 

Western Desert 18 

Desert Features 24 

Rami 24 

Serir 24 

Hamada 25 

Balata 25 

Nafash 26 

Sebakha 26 

The Wadi 27 

Climatic Factors 30 

Temperature 30 

Wind 33 

RainfaU 33 

Dew 36 

Water and Desert Life 37 

Vegetation 39 

Sinai Peninsula 40 

Red Sea Coastal Desert 41 

Eastern Desert 42 

Gebel Elba 45 

Nile Valley and Delta 45 

Western Mediterranean Coastal Desert 46 

Western Desert 48 

Zoogeography 51 

Special Adaptations of Desert Mammals 53 

Synopsesof Representatives in Ordersof Recent Egyptian Land Mammals. . 55 

Order Insectivora 57 

Key to Famihes of Egyptian Insectivores 57 



iv 

Family 1. Erinaceidae 57 

Key to Egyptian Genera of Erinaceidae 57 

Genus Hemiechinus Fitzinger, 1866 T 57 

Hemiechinus auritus (Gmelin, 1770) 57 

Key to Egyptian Subspecies of Hemiechinus auritus 62 

Hemiechinus auritus aegyptius (Fischer, 1829) 62 

Hemiechinus auritus libycus (Ehrenberg, 1833) 63 

Genus Paraechinus Trouessart, 1879 64 

Key to Egyptian Species of Paraechinus 64 

Paraechinus deserti (Loche, 1858) 64 

Paraechinus deserti deserti (Loche, 1858) 65 

Paraechinus aethiopicus (Ehrenberg, 1833) 68 

Paraechinus aethiopicus aethiopicus (Ehrenberg, 1833) 69 

Paraechinus dorsalis (Anderson and De Winton, 1901) 70 

Paraechinus dorsalis dorsalis (Anderson and De Winton, 1901) 70 

Family 2. Soricidae 71 

Key to Egyptian Genera of Soricidae 71 

Genus Crocidura Wagler, 1832 71 

Key to Egyptian Species of Crocidura 71 

Crocidura flavescens (I. Geoffroy St.-Hilaire, 1827) 73 

Crocidura flavescens deltae Heim de Balsac and Barloy. 1966 73 

Crocidura floweri Dollman, 1916 76 

Crocidura nana Dobson, 1890 77 

Crocidura suaveolens (Pallas, 1811) 78 

Egyptian Subspecies of Crocidura suaveolens 79 

Crocidura suaveolens portali (Thomas, 1920) 79 

Crocidura suaveolens matruhensis Setzer, 1960 80 

Genus Suncus Ehrenberg, 1833 80 

Key to Egyptian Species of Suncus 80 

Suncus murinus Linnaeus, 1766 81 

Suncus murinus sacer (Ehrenberg, 1833) 81 

Suncus etruscus Savi, 1822 82 

Order Lagomorpha 84 

Family Leporidae 84 

Genus Lepus Linnaeus, 1758 84 

Lepus capensis Linnaeus, 1758 84 

Key to Egyptian Subspecies of Lepus capensis 90 

Lepus capensis sinaiticus (Ehrenberg. 1833) 90 

Lepus capensis aegyptius (Demarest, 1822) 91 

Lepus capensis isabelUnus (Cretzschmar, 1826) 92 

Lepus capensis rothschildi (De Winton, 1902) 92 

Order Rodentia 94 

Key to Egyptian Families of Rodentia 94 

Family 1. Cricetidae (Subfamily Gerbillinae) 95 

Keys to Egyptian Genera of Gerbillinae (External Characters and Cranial 

and Dental Characters) 95 

Genus Gerbillus Desmarest, 1804 96 

Key to Egyptian Species of Gerbillus 96 

Gerbillus pyramidum I. Geoffroy St. Hilaire, 1825 96 

Key to Egyptian Subspecies of Gerbillus pyramidum Ill 



Gerbilbis pyramidum pyramidum I. Geoffroy St. Hilaire, 1825 Ill 

Gerbillus pyramidum floweri (Thomas, 1919) 113 

Gerbillus pyramidum gedeedus ssp. nov. Osborn and Helmy 114 

Gerbillus pyramidum elbaensis Setzer, 1958 116 

Gerbillus perpallidus Setzer, 1958 117 

Gerbillus andersoni De Winton, 1902 119 

Key to Egyptian Subspecies of Gerbillus andersoni 127 

Gerbillus andersoni andersoni De Winton, 1902 128 

Gerbillus andersoni inflatus (Ranck, 1968) 128 

Gerbillus andersoni bonhotei (Thomas, 1919) 129 

Gerbillus gerbillus (Olivier, 1801) 130 

Key to Egyptian Subspecies of Gerbillus gerbillus 136 

Gerbillus gerbillus asyutensis Setzer, 1960 136 

Gerbillus gerbillus sudanensis Setzer, 1956 138 

Gerbillus gerbillus gerbillus (Olivier, 1801) 139 

Genus Dipodillus Lataste, 1881 140 

Keys to Egyptian Species of Dipodillus (External Characters and Cranial 
Characters) 141 

Dipodillus campestris (Levaillant, 1857) 141 

Egyptian Subspecies of Dipodillus campestris 152 

Dipodillus campestris wassifi (Setzer, 1958) 152 

Dipodillus campestris haymani (Setzer, 1958) 153 

Dipodillus campestris patrizii (de Beaux, 1932) 154 

Dipodillus campestris venustus (Sundevall, 1843) 154 

Dipodillus dasyurus (Wagner, 1842) 155 

Dipodillus dasyurus dasyurus (Wagner, 1842) 155 

Dipodillus mackilligini Thomas, 1904 159 

Dipodillus simoni (Lataste, 1881) 161 

Dipodillus simoni kaiseri (Setzer, 1958) 161 

Dipodillus amoenus De Winton, 1902 167 

Dipodillus amoenus amoenus De Winton, 1902 168 

Dipodillus henleyi De Winton, 1903 174 

Key to Egyptian Subspecies oi Dipodillus henleyi 179 

Dipodillus henleyi henleyi De Winton, 1903 179 

Dipodillus henleyi mariae (Bonhote, 1909) 180 

Genus Sekeetamys Ellerman, 1947 181 

Sekeetamys calurus (Thomas, 1892) 181 

Key to Egyptian Subspecies of Sekeetamys calurus 188 

Sekeetamys calurus calurus (Thomas, 1892) 188 

Sekeetamys calurus makrami (Setzer, 1961) 189 

Gerais Meriones Illiger, 1811 190 

Key to Egyptian Species of Meriones 190 

Meriones crassus Sundevall, 1842 191 

Key to Egyptian Subspecies of Meriones crassus 201 

Meriones crassus crassus Sundevall, 1842 201 

Meriones crassus perpallidus Setzer, 1961 202 

Meriones crassus paUidus Bonhote, 1912 203 

Meriones sacramenti Thomas, 1922 204 

Meriones libycus (Lichtenstein, 1823) 207 

Meriones libycus libycus (Lichtenstein, 1823) 208 



VI 



Meriones shawi Rozet, 1833 214 

Meriones shawi isis (Thomas, 1919) 215 

Meriones tristrami Thomas. 1892 -. 218 

Meriones tristrami tristrami Thomas, 1892 218 

Genus Pachyuromys Lataste, 1880 220 

Pachyuromys duprasi Lataste, 1880 220 

Pachyuromys duprasi natronensis De Winton, 1903 221 

Genus Psammomys Cretzschmar, 1828 226 

Psammomys obesus Cretzschmar, 1828 227 

Key to Egyptian Subspecies of Psammomys obesus 242 

Psammomys obesus obesus Cretzschmar, 1828 242 

Psammomys obesus nicolli Thomas, 1908 243 

Psammomys obesus terraesanctae Thomas, 1902 244 

Family 2. Spalacidae 245 

Genus Spalax Guldenstaedt, 1770 245 

Spalax ehrenbergi Nehring, 1898 246 

Spalax ehrenbergi aegyptiacus (Nehring, 1898) 246 

Family 3. Muridae 253 

Key to Egyptian Genera of Muridae 254 

Genus Arvicanthis Lesson, 1842 254 

Arvicanthis niloticus (Desmarest, 1822) 254 

Arvicanthis niloticus niloticus (Desmarest, 1822) 255 

Genus Rattus Fischer. 1803 263 

Key to Egyptian Species of Rattus 264 

Rattus rattus (Linnaeus, 1758) 264 

Rattus norvegicus (Berkenhout, 1769) 269 

Genus Mus Linnaeus, 1758 273 

Mus musculus Linnaeus, 1758 274 

Mus musculus praetextus (Brants, 1827) 274 

Genus Acomys I. Geoffroy St. Hilaire, 1838 285 

Key to Egyptian Species of Acomys 285 

Acomys russatus (Wagner, 1840) 286 

Key to Egyptian Subspecies of Acomys russatus 292 

Acomys russatus russatus (Wagner, 1840) 292 

Acomys russatus aegyptiacus Bonhote, 1912 293 

Acomys cahirinus (Desmarest, 1819) 293 

Key to Egyptian Subspecies of Acomys cahirinus 303 

Acomys cahirinus cahirinus (Desmarest. 1819) 303 

Acomys cahirinus viator (Thomas. 1902) 305 

Acomys cahirinus hunteri (De Winton. 1901) 305 

Acomys cahirinus dimidiatus (Cretzschmar. 1826) 306 

Acomys cahirinus megalodus (Setzer. 1959) 307 

Acomys cahirinus helmyi ssp. nov. Osborn 308 

Genus Nesokia Gray, 1842 309 

Nesokia indica (Gray and Hardwicke. 1832) 309 

Nesokia indica suilla (Thomas, 1907) 309 

Family 4. Muscardinidae 315 

Genus Eliomys Wagner. 1840 315 

EUomys quercinus (Linnaeus, 1766) 316 

Key to Egyptian Subspecies of Eliomys quercinus 321 



Eliomys quercinus melanurus (Wagner, 1840) 321 

Eliomys quercinus cyrenaicus (Festa, 1921) 322 

Family 5. Dipodidae 322 

Key to Egyptian Genera of Dipodidae 323 

Genus Allactaga Cuvier, 1836 323 

Allactaga tetradactyla (Lichtenstein, 1823) 327 

Genus Jaculus Erxleben, 1777 333 

Key to Egyptian Species of Jaculus 333 

Jaculus orientalis Erxleben, 1777 334 

Jaculus orientalis orientalis Erxleben, 1777 334 

Jaculus jaculus (Linnaeus, 1758) 339 

Key to Egyptian Subspecies of Jaculus jaculus 352 

Jaculus jaculus schlueteri (Nehring, 1901) 352 

Jaculus jaculus flavillus Setzer, 1955 353 

Jaculus jaculus butleri Thomas, 1922 355 

Jaculus jaculus jaculus (Linnaeus, 1758) 356 

Family 6. Hystricidae 357 

Genus Hystrix Linnaeus, 1758 357 

Hystrix cristata Linnaeus, 1758 357 

Order Carnivora 359 

Key to Egyptian Families of Carnivora (Cranial and Dental Characters and 

External Characters) 359 

Family 1. Canidae 359 

Key to Egyptian Genera of Canidae 360 

Genus Canis Linnaeus, 1758 360 

Canis aureus Linnaeus, 1758 360 

Canis aureus lupaster (Hemprich and Ehrenberg, 1833) 361 

Genus Vulpes Oken, 1816 371 

Key to Egyptian Species of Vulpes 371 

Vulpes vulpes (Linnaeus, 1758) 371 

Vulpes vulpes aegyptiaca (Sonnini, 1816) 372 

Vulpes rueppelli (Schinz, 1825) 379 

Vulpes rueppelli rueppelli (Schinz, 1825) 380 

Genus Fennecus Desmarest, 1804 387 

Fennecus zerda (Zimmermann, 1780) 387 

Family 2. Mustelidae 395 

Key to Egyptian Genera of Mustelidae 395 

Genus Poecilictis Thomas and Hinton, 1920 395 

Poecilictis libyca (Hemprich and Ehrenberg, 1833) 395 

Poecilictis libyca libyca (Hemprich and Ehrenberg, 1833) 396 

Genus Ictonyx Kaup, 1835 403 

Ictonyx striatus (Perry, 1810) 404 

Ictonyx striatus erythreae De Winton, 1898 404 

Genus Mustela Linnaeus, 1758 405 

Mustela nivalis Linnaeus, 1766 406 

Mustela nivalis subpalmata (Hemprich and Ehrenberg, 1833) 406 

Family 3. Viverridae 410 

Key to Egyptian Genera of Viverridae 410 

Genus Genetta Oken, 1816 410 

Genetta genetta (Linnaeus, 1758) 410 



viii 

Genetta genetta senegalensis (Fischer, 1829) 411 

Genus Herpestes Illiger, 1811 415 

Herpestes ichneumon (Linnaeus, 1758) \- 416 

Herpestes ichneumon ichneumon (Linnaeus, 1758) 416 

Family 4. Hyaenidae 422 

Key to Egyptian Genera of Hyaenidae 422 

Genus Hyaena Brisson, 1762 422 

Hyaena hyaena (Linnaeus, 1758) 423 

Hyaena hyaena dubbah (Meyer, 1793) 423 

Genus Proteles \. Geoffroy St.-Hilaire, 1824 432 

Proteles cristatus (Sparrmann, 1783) 432 

Proteles cristatus palUdior Cabrera, 1910 432 

Family 5. Felidae 434 

Key to Egyptian Genera of Felidae 434 

Genus Felis Linnaeus, 1758 434 

Key to Egyptian Species of Genus Felis 435 

Felis chaus GUldenstaedt, 1776 435 

FeUs chaus nilotica De Winton, 1898 435 

FeUs sylvestris Schreber, 1777 440 

Felis sylvestris libyca (Forster, 1780) 443 

FeUs sylvestris tristrami (Pocock, 1944) 443 

Felis margarita Loche, 1858 444 

Felis margarita margarita Loche, 1858 444 

Genus Caracal Gray, 1843 447 

Caracal caracal (Schreber, 1776) 447 

Caracal caracal schmitzi (Matschie, 1912) 448 

Genus Panthera Oken, 1816 451 

Panthera pardus (Linnaeus, 1758) 451 

Panthera pardus jarvisi Pocock, 1932 454 

Panthera pardus pardus (Linnaeus, 1758) 455 

Genus Acinonyx Brookes, 1828 455 

Acinonyx jubatus (Schreber, 1776) 455 

Order Hyracx)idea 460 

Family Procaviidae 460 

Genus Procavia Storr, 1780 460 

Procavia capensis (Pallas, 1766) 460 

Key to Egyptian Subspecies of Procavia capensis 468 

Procavia capensis syriaca (Schreber, 1784) 468 

Procavia capensis ruficeps (Hemprich and Ehrenberg, 1832) 468 

Order Perissodactyla 470 

Family Equidae 470 

Genus Equus Linnaeus, 1758 470 

Equus asinus Linnaeus, 1758 470 

Equus asinus africanus (Fitzinger, 1857) 470 

Order Artiodactyla 475 

Family 1. Suidae 475 

Genus Sus Linnaeus, 1758 475 

Sus scrofa Linnaeus, 1758 475 

Family 2. Hippopotamidae 477 

Genus Hippopotamus Linnaeus, 1758 477 



Hippopotamus amphibius Linnaeus, 1758 477 

Family 3. Bovidae 479 

Key to Egyptian Genera of Bovidae 479 

Genus Oryx Blainville. 1816 480 

Oryx dammah (Cretzschmar, 1826) 480 

Genus Addax Rafinesque, 1815 482 

Addax nasomaculatus (Blainville, 1816) 482 

Genus Alcelaphus Blainville, 1816 484 

Alcelaphus buselaphus (Pallas, 1766) 484 

Genus Gazella Blainville, 1816 486 

Key to Egyptian Species of Gazella 486 

Gazella leptoceros (F. Cuvier, 1842) 487 

Gazella leptoceros leptoceros (F. Cuvier, 1842) 487 

Gazella dorcas (Linnaeus, 1758) 501 

Key to Egyptian Subspecies of Gazella dorcas 507 

Gazella dorcas littoralis (Blaine, 1913) 508 

Gazella dorcas dorcas (Linnaeus, 1758) 509 

Gazella dorcas saudiya (Carruthers and Schwarz, 1935) 512 

Gazella gazella (Pallas, 1766) 513 

Gazella gazella arabica (Lichtenstein, 1827) 513 

Genus Capra Linnaeus, 1758 514 

Capra ibex Linnaeus, 1758 515 

Capra ibex nubiana (F. Cuvier, 1825) 515 

Genus Ammotragus Blyth, 1840 521 

Ammotragus lervia (Pallas, 1777) 521 

Ammotragus lervia omatus (I. Geoffroy St. Hilaire, 1827) 521 

Appendix 1. Explanation of abbreviations 526 

Appendix 2. The auditory bulla 528 

Appendix 3. Gerbillinae tooth terminology 530 

Appendix 4. The governorates of Egypt 532 

Appendix 5. Gazetteer of localities mentioned in the text 533 

Appendix 6. Definitions of terms used in the text 553 

References 554 



LIST OF ILLUSTRATIONS 

1. Geography of Egypt 6 

2. Rainfall map of Egypt 7 

3. Eastern Desert. Red Sea Hills 11 

4. Eastern Desert. Wadi Garawi in Helwan Plateau 13 

5. Eastern Desert. Wadi Gindali south of Cairo-Suez Road 15 

6. Nile Valley near Giza 16 

7. Western Mediterranean Coastal Desert near Burg el Arab 17 

8. Western Mediterranean Coastal Desert 4.8 km. E of Abu Mena 18 

9. Western Desert. Serir (pebble desert) and sand sheet 19 

10. Western Desert. Meandering sand sheet in area between Wadi el Natroun 

and Bir Victoria 20 

11. Western Mediterranean Coastal Desert. Limestone cliffs near Salum 21 

12. Western Desert. Wadi Labaq in northeastern part of Qattara Depression. . 22 

13. Western Desert. Wadi Muwellih 25 

14. Eastern Desert. Wadi Hof 28 

15. Eastern Desert. Wadi el Qreiya at km. 77 on Qena-Safaga road 29 

16. Eastern Desert. Bir Qiseib in Wadi Qiseib 31 

17. Western Desert. El Maghra 34 

18. Western Desert. Acacia raddiana grove 120 km. S of El Maghra 40 

19. Western Mediterranean Coastal Desert 34 km. S of Bahig 47 

20. Western Mediterranean Coastal Desert near area in Figure 19 47 

21 . Western Desert. Bahariya Oasis, Bir Wigaba 49 

22. Western Desert. Bahrein 50 

23. Collection localities of Hemiechinus auritus aegyptius and H. a. lybicus. . . 58 

24. Skull of Hemiechinus auritus 60 

25. Collection localities of Paraechinus deserti deserti, P. aethiopicus 

aethiopicus, and P. dorsalis dorsalis 65 

26. Ventral view of Paraechinus aethiopicus aethiopicus 66 

27. Skull of Paraechinus deserti deserti 67 

28. Collection localities of Crocidura flavescens deltae, C. floweri, C. nana, C. 

suaveolens portali, and C s. matruhensis 72 

29. Skull of Crocidura flavescens deltae 74 

30. Collection localities of Lepus capensis sinaiticus, L. c. aegyptius, L. c. 

isabellinus, L. c. rothschildi, and sight records 85 

31. Skull of Lepus capensis 87 

32. Collection localities of Gerbillus pyramidum pyramidum, G. p. floweri, 

G. p. gedeedus, G. p. elbaensis, and G. perpallidus 97 

33. Palatal ridges of Gerbillus gerbillus, G. andersoni, G. pyramidum, 

Dipodillus campestris, D. simoni, D. amoenus, Sekeetamys calurus, 

Meriones crassus, and Psammomys obesus 99 



34. Palms and soles of DipodiUus campestris, D. dasyurus, D. simoni, D. 

amoenus, D. henleyU Gerbillus gerbillus, Sekeetamys calurus, 

Meriones crassus, and Psammomys obesus 101 

35. Skull of adult Gerbillus pyramidum floweri 102 

36. Mastoid bulla variations in Gerbillus pyramidum and G. perpallidus 103 

37. Posterior margins of nasals and interparietal outlines of Gerbillus 

pyramidum pyramidum, G. p. floweri, and G. perpalUdus 104 

38. Crown views of right upper and left lower molars of mature and 

immature of species of DipodiUus and Gerbillus 105 

39. Collection localities of Gerbillus andersoni andersoni, G. a. inflatus, 

and G. a. bonhotei 120 

40. Comparison of auditory bullae in lateral and dorsal view and skull shape in 

Gerbillus andersoni inflatus, G. a. andersoni, and G. gerbillus gerbillus. . . 122 

41. Posterior margins of nasals and incisive and posterior palatal foramina 

of Gerbillus andersoni and G. gerbillus 123 

42. Scatter diagram of incisive foramina length versus occipitonasal length 

in Gerbillus andersoni and G. gerbillus 127 

43. Collection localities of Gerbillus gerbillus gerbillus, G. g. asyutensis, 

and G. g. sudanensis 131 

44. Cadaver of Gerbillus gerbillus 132 

45. Collection localities of DipodiUus campestris wassifi, D. c. haymani, 

D. c. patrizii, D. c. venustus, D. dasyurus dasyurus, andD. mackilUgini. . 142 

46. Live specimen of DipodiUus campestris wassifi 144 

47. Auditory bullae in lateral view and outlines of skulls in dorsal view of 

species of DipodiUus 145 

48. Western Mediterranean Coastal Desert. Ras el Hekma 166 

49. Scatter diagram of tail length versus head and body length in Eastern 

Desert and Sinai Peninsula samples oi DipodiUus dasyurus 167 

50. Collection localities of DipodiUus simoni kaiseri and D. amoenus amoenus. 168 

51. Skull of DipodiUus simoni kaiseri 169 

52. Crown view of upper first molars of DipodiUus henleyi, D. simoni, and 

D. amoenus 171 

53. Cadaver of DipodiUus amoenus amoenus 172 

54. Collection localities of DipodiUus henleyi henleyi and D. h. mariae 175 

55. Cadaver of DipodiUus henleyi mariae 176 

56. Collection localities of Sekeetamys calurus calurus and 5. c. makrami 

and sight records 182 

57. Live specimen of Sekeetamys calurus makrami 183 

58. Skull of Sekeetamys calurus makrami 184 

59. Crown views of right upper and left lower molars of mature and 

immature Sekeetamys calurus, Meriones crassus, Psammomys obesus, 
and Pachyuromys duprasi 187 

60. Auditory bullae in lateral and dorsal views, interparietal shapes, and 

exposure of infraorbital foramina in lateral view in species of Meriones. . 192 

61. Skull of Meriones sacramenti 193 

62. Collection localities of Meriones crassus crassus, M. c. perpaUidus, 

and M. c. paUidus 194 

63. Live specimen of Meriones crassus 195 

64. Western Desert. About 7 km. SE of km. 208 on Cairo-Bahariya road 199 



xfi 



65. Collection localities of Meriones sacramenti, M. libycua libycua, M. shawi 

isis, and M. tristrami < 205 

66. Collection localities of Pachyuromya duprasi natronensis 222 

67. Live specimen of Pachyummys duprasi natronensis 223 

68. Skull of Pachyuromys duprasi natronensis 225 

69. Collection localities of Psammomys obesus obesus, P. o. nicolli, and 

P. o. terraesanctae 228 

70. Cadaver of Psammomys obesus obesus 229 

71. Diagrams of middorsal hairs from subspecies of Psammomys obesus 231 

72. Skull of Psammomys obesus 232 

73. Lateral view of right upper molars of adult and immature Psammomys 

obesus 233 

74. Anterior and posterior position of posterior margin of nasals in 

subspecies of Psammomys obesus 236 

76. Burrow and toilet of Psammomys obesus 238 

76. Collection localities of Spalax ehrenbergi aegyptiacus and sight record 247 

77. Live specimen of Spalax ehrenbergi aegyptiacus 248 

78. Skull of Spalax ehrenbergi aegyptiacus 249 

79. Crown views of right upper and left lower molars of Egyptian Muridae 

and Spalax ehrenbergi 250 

80. Collection localities of Arvicanthis niloticus niloticus 256 

81. Dorsal views of Arvicanthis niloticus niloticus, Rattus rattus, R. 

norvegicus, and Nesokia indica 257 

82. Skull of Arvicanthis niloticus niloticus 258 

83. Collection localities of Rattus rattus and R. norvegicus 265 

84. Skull of Rattus rattus 266 

85. Skull of Rattus norvegicus 271 

86. Collection localities of Mus musculus praetextus 275 

87. Skull of Af us musculus praetextus 277 

88. Collection localities of Acomys russatus russatus andi4. r. aegyptiacus. . . . 287 

89. Live specimen of Acomys russatus 288 

90. Collection localities of Acomys cahirinus cahirinus, A. c. megalodus, 

A. c. dimidiatus, A. c. hunteri, A. c. helmyi, and A. c. viator 294 

91. Skull of Acomys cahirinus 296 

92. Shells of snails {Eremica desertorum) gnawed by a captive Acomys 

cahirinus 301 

93. Collection localities of Nesokia indica suilla and sites of Paleolithic 

remains 310 

94. Skull of Nesokia indica suilla 312 

95. Collection localities of Eliomys quercinus melanurus andE. q. cyrenaicus. .317 

96. Live specimen of Eliomys quercinus cyrenaicus 318 

97. Skull of Eliomys quercinus 319 

98. Crown views of right upper and left lower molars of mature Allactaga 

tetradactyla, Jaculus orientalis, and J. jaculus 323 

99. Collection localities of Allactaga tetradactyla and Jaculus orientalis 

orientalis 328 

100. Live specimen of Allactaga tetradactyla 329 

101. Skull of Allactaga tetradactyla 330 

102. Posterior margins of nasals of Allactaga tetradactyla, Jaculus orientalis, 

and J. jaculus 331 



103. Skull of Jaculus orientalis orientalis 335 

104. Collection localities of Jaculus jaculus jaculus, J. j. flaviUus, J. j. schlueteri, 

J. j. butleri, and Hystrix cristata 340 

105. Live specimen of Jaculus jaculus jaculus 341 

106. Lower jaw and posteriors of lower jaws in ventral view of Jaculus jaculus. . 343 

107. Frequency diagrams of ear length and hind foot length of subspecies of 

Jaculus jaculus 347 

108. Western Desert 3 km. SSE of Camel Pass Dune. Solitary specimen of 

Comulaca monocantha, an important browse plant of Gazella sp. and 

a food source for Jaculus jaculus 349 

109. Collection localities of Canis aureus lupaster and sight records 362 

1 10. Skull of Canis aureus lupaster 363 

111. Left mandibular teeth of jackal and dog 368 

112. Comparison of nasomaxillary contact and shape of posterior margins of 

nasals in Vulpes uulpes and V. rueppelli 372 

113. Collection localities of Vulpes vulpes aegyptiaca 373 

114. Skull of Vulpes vulpes aegyptiaca 375 

115. Collection localities of Vulpes rueppelli rueppelli and sight record 380 

1 16. Live specimen of Vulpes rueppelli rueppelli 381 

117. Skull of Vulpes rueppelli rueppelli 383 

118. Collection localities of Fennecus zerda and sight record 388 

119. Young Fennecus zerda 389 

120. Skull of Fennecus zerda 391 

121. Collection localities of Poecilictis libyca libyca and records of tracks, 

Ictonyx striatus erythreae, Mustela nivalis subpalmata and sight 

records, and Genetta genetta senegalensis 396 

122. Mustelidae. Dorsal and ventral views of Mustela nivalis subpalmata; 

two dorsal views and a ventral view of Poecilictis libyca libyca; dorsal 
and ventral views of Ictonyx striatus erythreae 397 

123. Skull of Poecilictis libyca libyca 398 

124. Skull of Mustela nivalis subpalmata 407 

125. Museum specimen of Genetta genetta senegalensis 412 

126. Skull of Genetta genetta senegalensis 413 

127. Collection localities of Herpestes ichneumon ichneumon and sight record. . 417 

128. Cadavers of Herpestes ichneumon ichneumon 418 

129. Skull of Herpestes ichneumon ichneumon 419 

130. Collection localities of Hyaena hyaena dubbah, sight records and tracks; 

and Proteles cristata 424 

131. Cadaver of Hyaena hyaena dubbah 425 

132. Skull of Hyaena hyaena dubbah 426 

133. Collection localities of Felis chaus nilotica, F. sylvestris libyca, F. s. 

tristrami, and F. margarita 436 

134. Skull of Felis chaus nilotica 438 

135. Skull of FeUs sylvestris libyca 442 

136. Skull of Felis margarita 446 

137. Collection localities of Caracal caracal schmitzi 448 

138. Skull of Caracal caracal schmitzi 449 

139. Collection localities of Panthera pardus pardus and P. p. jarvisi 452 

140. Skull of Panthera pardus 453 

141. Collection localities, sight records, tracks, and verbal reports of Acinonyx 



jubatus 456 

142. Skull of Acinonyx jubatus 458 

143. Collection localities of Procavia capensis syriaca, P. c. nificeps, and 

sight records 461 

144. Skull of Procavia capensis 462 

145. Comparison of position of anterior end of malar relative to lacrimal in 

Procavia capensis ruficeps and P. c. syriacus 463 

1 46. Habitat of Procavia capensis in Wadi Nagib 466 

147. Localities of observations of Equus asinus africanus in 1800's and 

from 1950 to date; paleolithic and prehistoric remains 471 

148. Previous distribution of Sus scrofa 476 

149. Previous distribution of Hippopotamus amphibius 478 

150. Previous distribution of Oryx damah and recent sight record 481 

151. Previous distribution olAddax nasomaculatus 483 

152. Previous distribution of Alcelaphus buselaphus 485 

153. Collection localities of Gazella leptoceros leptoceros; sight records; 

skulls and horns 488 

154. Skull of Gazella leptoceros leptoceros 492 

155. Lateral views of anterior part of skulls of Gazella leptoceros and 

G. dorcas 494. 495 

156. Nasal bones, interparietals, and lower second and third molars of Gazella 

leptoceros and G. dorcas 496 

157. Frontal views of horns of Gazella dorcas and G. leptoceros 498 

158. Cross-sections of bases of left horn cores of adult males and females of 

Gazella dorcas and G. leptoceros 499 

159. Acacia ehrenbergiana browsed by Gazella dorcas 506 

160. Collection localities of Gazella dorcas dorcas, G. d. littoraUs, G. d. saudiya, 

and G. gazella arabica: sight records and skulls and horns 508 

161. SkuU of Gazella dorcas 510 

162. Collection localities of Capra ibex nubiana, undated and prior to 1932 

and 1948 to date; horns or skulls: and sight records 516 

163. SkuU of Capra ibex nubiana 517 

164. Collection localities of Ammotragus lervia omatus, undated and prior 

to 1932, 1932-1948, and 1949 to date; and horns and skulls; and sight 
records 1932 to date 522 

165. Auditory bullae of Dipodillus campestris 529 

166. Terminology used in describing molars of Gerbillinae 531 

167. Governorates of Egypt 532 



LIST OF TABLES 

1 . Monthly means (and ranges) in temperature in Egypt 32 

2. Means (and ranges) of measurements of adult Hemiechinus auritus 61 

3. Means (and ranges) of measurements and weight of Paraechinus deserti, 

P. aethiopicus, and P. dorsalis 61 

4. Means (and ranges) of measurements and ratios of species of Crocidura 75 

5. Means (and ranges) of measurements and ratios of adult Lepus capensis 88 

6. Means (and ranges) of measurements, ratios, and weight of adult Gerbillus 

pyramidum and G. perpallidus 106 

7. Characters of Gerbillus pyramidum and G. perpallidus 108 

8. Means (and ranges) of measurements, ratios, and weight of adult Gerbillus 

andersoni 110 

9. Comparison of inflation in mastoid chambers of bulla in Gerbillus andersoni. 124 

10. Means (and ranges) of measurements, ratios, and weight of adult Gerbillus 

gerbillus 124 

11. Comparison of characters of Gerbillus andersoni and G. gerbillus 125 

12. Means (and ranges) of measurements, ratios, and weight of adult Dipodillus 

campestris 147 

13. Means (and ranges) of measurements, ratios, and weight of adult Dipodillus 

dasyurus and D. mackiUigini 148 

14. Characters of species of Dipodillus 149 

15. Means (and ranges) of measurements, ratios, and weight of adult Dipodillus 

simoni 164 

16. Means (and ranges) of measurements, ratios, and weight of adult Dipodillus 

a. amoenus 165 

17. Means (and ranges) of measurements, ratios, and weight of adult Dipodillus 

henleyi 178 

18. Means (and ranges) of measurements, ratios, and weight of adult 

Sekeetamys calurus 187 

19. Means (and ranges) of measurements, ratios, and weight of adult Meriones 

crassus 197 

20. Color and cranial variations in Meriones crassus 197 

21. Means (and ranges) of head and body and occipitonasal lengths of adult 

Meriones crassus from Sinai Peninsula and northern to southern localities 
in Eastern and Western Deserts 198 

22. Means (and ranges) of measurements, ratios, and weight of adults of 

species of Meriones 209 

23. Characters of species of Meriones 210 

24. Means (and ranges) of measurements, ratios, and weight of adult 

Pachyuromys duprasi natronensis 224 

25. Means (and ranges) of measurements, ratios, and weight of adult 

Psammomys obesus 234 



XV 



26. Color patterns of Psammomys obesus 235 

27. Variation in position of posterior margins of nasals in Psammomys obesus. . 235 

28. Numbers of burrow systems and occupation: numbers and distribution 

of sexes within age classes of Psammomys obesus 240 

29. Number and per cent of males and fenuiles in three age classes of museum 

samples of Psammomys obesus 242 

30. Means (and ranges) of measurements and weight of adult Spalax ehrenbergi 

aegyptiacus 25 1 

31. Means (and ranges) of measurements, ratios, and weight of adult Rattus 

rattus, R. norvegicus, and Arvicanthis niloticus 259 

32. Characters of Rattus rattus, R. norvegicus, and Arvicanthis niloticus 260 

33. Color phases of Mus musculus praetextus 276 

34. Means (and ranges) of measurements, ratios, and weight of adult Mus 

musculus praetextus 278 

35. Means (and ranges) and ratios of head and body length and tail length 

of Mus musculus praetextus 279 

36. Means (and ranges) or measurements, ratios, and weight of adult Acomys 

russatus 289 

37. Means (and ranges) of measurements, ratios, and weight of adult Acomys 

cahirinus 290 

38. Means (and ranges) of measurements, ratios, and weight of adult Nesokia 

indica suilla 311 

39. Means (and ranges) of measurements, ratios, and weight of adult Eliomys 

quercinus 321 

40. Characters of Egyptian jerboas 324 

41. Means (and ranges) of measurements, ratios, and weight of adult AUactaga 

tetradactyla and Jaculus orientalis 332 

42. Color and cranial variations in Jaculus jaculus 342 

43. Means (and ranges) of measurements, ratios, and weight of adult Jaculus 

jaculus 345 

44. Means (and ranges) of measurements, ratios, and weight of adult Canis 

aureus lupaster 364 

45. Means (and ranges) of measurements, ratios, and weight of adult male and 

female Vulpes vulpes nilotica 366 

46. Means (and ranges) of measurements, ratios, and weight of adult male and 

female Vulpes r rueppelli 367 

47. Means (and ranges) of measurements, ratios, and weight of adult 

Fennecus zerda 390 

48. Means (and ranges) of measurements and ratios of adult Poecilictis L 

libyca and adult male and female Ictonyx striatus erythreae 400 

49. Variation in median lumbar stripe in samples of Poecilictis libyca 403 

50. Means (and ranges) of measurements and ratios of adult male and female 

Mustela nivalis subpalmata 408 

51. Means (and ranges) of measurements and ratios of Genetta genetta and 

Herpestes ichneumon 415 

52. Means (and ranges) of measurements, ratios, and weight of adult Hyaena 

h. dubbah 427 

53. Means (and ranges) of measurements, ratios, and weight of Felis sylvestris 

and F. chaus 437 

54. Means (and ranges) of measurements of Procavia capensis 464 



55. Hind hoof length (and ranges) in subadult and adult gazelles from hard 

and sandy desert 489 

56. Cranial and horn characters of Gazella leptoceros, G. dorcas, and 

0. gazella 490 

57. Means (and ranges) of measurements and weight of adult male and female 

Gazella dorcas and G. leptoceros 493 

58. Measurements of two specimens of Capra ibex nubiana 518 



PREFACE 

A volume of this kind is never the work or the inspiration of its 
authors alone. It is a co-operative effort involving persons of many 
professions and personal interests. 

Dr. Harry Hoogstraal. Head, Medical Zoology Department, 
United States Naval Medical Research Unit Number Three 
(NAMRU-3), laid the foundation in 1950 by beginning a research 
program on the ecologic factors of arthropod-borne diseases. The 
necessity for accurate identification of mammalian host species and 
knowledge of ecologic factors and geographical distribution was of 
immediate importance in this project (Hoogstraal, 1960). His goal, a 
modern treatise on the mammals of Egypt with workable keys, ac- 
curate descriptions, and fundamental biological information, has 
been our goal as well. 

For the loan of study materials, we are indebted to Dr. G. B. 
Corbet, British Museum (Natural History), London; Dr. Jean Dorst, 
Museum National d'Histoire Naturelle, Paris; Dr. H. Felten, Natur- 
Museum und Forschungs Institute Senckenberg, Frankfurt; Dr. 
Kamal Wassif, Ain Shams University, Cairo; Dr. A. M. Monaiery, 
Giza Zoological Gardens, Giza; Drs. Joseph Curtis Moore and Luis 
de la Torre, Field Museum of Natural History, Chicago; Dr. Oliver 
Pearson, Museum of Vertebrate Zoology, Berkeley; and Dr. Henry 
W. Setzer, United States National Museum, Washington, D.C. 

For identifying plants, we are indebted to late Dr. Vivi Tackholm. 
Cairo University; snails, late Dr. William J. Clench. Museum of 
Vertebrate Zoology. Cambridge; and arthropod remains from 
stomach samples, late Mr. Anastase Alfieri. Entomological In- 
stitute. Cairo. 

For figures other than those of the co-author (1. H.). we appreciate 
the help of Mr. Samuel H. Grove of Field Museum and Miss Marian 
Moon of Cairo. 

Almost every employee of NAMRU-3 was involved in some way 



with the collecting phase of our work, albeit administrative, order- 
ing and issuing supplies, maintaining vehicles, and constructing or 
repairing equipment. 

Our greatest debt of appreciation is to the devoted persons of the 
Medical Zoology Department who over the years spent countless 
hours in the field amassing the bulk of the collection which made 
this publication possible — Drs. Harry Hoogstraal and Makram N. 
Kaiser, and Messrs. Sobhy Gaber, Sayed Metwally, Hassan 
Touhamy, Ibrahim Khedr, and Kasim Hadad. 

Mr. Abd el Aziz Salah, Manager, Administrative Liaison, was in- 
dispensible in his ability to secure travel permits and special 
privileges from Egyptian government agencies. 

We also thank the representatives of the Ministry of Public 
Health who accompanied us on field trips and assumed responsibili- 
ty for our welfare. 

Representatives of the Department of Antiquities, Department of 
Locust Control, and the Governors of El Wadi El Gedeed and 
Matruh Governorates were especially helpful in securing guides, 
rest houses, and collecting permits. 

Field assistants recruited from various localities participated in 
the collecting effort. These men dug rodents and foxes from bur- 
rows, set out and retrieved traps, and chased jerboas with nets. 

Lastly, we are grateful to the professional guides who led us 
without map or compass into the most remote regions of the desert. 

The NAMRU-3 collections reported herein are deposited in the 
Smithsonian Institution and in Field Museum of Natural History. 



INTRODUCTION 

Review of Literature.— Egyptian mammals were first recorded in 
rock engravings and cave paintings mainly in the areas of Gebel 
Uweinat and the southern Eastern Desert, especially around Gebel 
Elba. Elephant, giraffe, hartebeest, oryx, addax, wild cattle, hip- 
popotamus, and lion, which are no longer part of the fauna, and ibex, 
gazelle, Barbary sheep, wild ass, leopard, and other cats were known 
to prehistoric man in Egypt (Newbold, 1928; Sandford and Arkell, 
1933, 1939; Winkler, 1938; Dunbar, 1941). Pre-dynastic men made 
the wheel trap with pointed pieces of markh wood (Leptadenia 
pyrotechnica) and captured wild ass, ibex, gazelle (Winkler, 1938), 
and probably other species. This same device, made with the spines 
of date palms, was used by the ancient Egyptians and is still used 
by Bedouins to capture gazelle (Harding- King, 1925; Khairat, 1954; 
Osborn, 1968b). Extensive stone walls were constructed by 
Paleolithic men which enabled them to drive game into small 
enclosures for slaughter (Reed et al., 1967), and these same traps 
were doubtlessly utilized by the Pharaohs in their "hunts." The 
walls still stand in Nubia (Giegengack, 1968). 

Lion, oryx, addax, hartebeest, hippopotamus, gazelle, ibex, and 
Barbary sheep appear in paintings and reliefs on the walls of 
Pharaonic tombs and temples. Animal motifs of the V and VI 
Dynasties at Saqqara are particularly well preserved. In addition to 
these, there are carvings of shrew, hedgehog, mongoose, otter, 
hyena, hare, fox, wild cats, striped weasel, porcupine, and aardvark 
(Paton, 1925; Keimer, 1942, 1944, 1949, 1955; Brunner, 1969). 
Numerous species were worshipped and mummified (Gaillard and 
Daressy, 1905; Morrison-Scott, 1952). Two shrews, Crocidura 
olivieri (C. flavescens) and Sorex religiosus (C. nana), were each 
described from mummified specimens (Geoffroy St.-Hilaire, 1827). 

The Pharaohs hunted and tamed most of the larger animals, in- 
cluding hyenas and lions, and also kept mongooses as pets (Wilkin- 
son, 1878; Newbold, 1928; Khairat, 1954). During the Greek occupa- 



2 FIELDIANA: ZOOLOGY 

tion from the time of Alexander (332 B.C.) until the Arab invasion of 
639 A. I)., gazelle were plentiful, the Nile abounded with hip- 
popotamus, and lions were hunted near Alexandria (Lindsay, 1965). 
Herodotus (fifth century B.C.) can be credited with the first accounts 
of Egyptian mammals, and some correlation seemed to have existed 
between the known fauna and Herodotus' compilations (Keimer. 
1955). Natural history written by Pliny the Elder in the first cen- 
tury A.I), was supposedly influenced by the Egyptians. His state- 
ment that mice were spontaneously generated from Nile mud after 
each annual flood was reported by Dawson (1924) to have been cur- 
rently believed in Egypt. 

Alpini (1735) wrote the first book on Egyptian mammals, Forskal 
(1775) followed with descriptions of wild and domesticated animals, 
and Bruce (1790) listed a few Egyptian species in his "Travels." A 
large number of publications on natural history and the mammalian 
fauna (E. Geoffroy St.-Hilaire, 1818; I. Geoffroy St.-Hilaire, 1827; 
Audouin. 1829; E. Geoffroy St.-Hilaire and Audouin, 1829) were 
based on the collections of the French zoologist, Etienne Geoffroy 
Saint-Hilaire, who accompanied Napoleon's abortive military ex- 
pedition to Egypt during 1798 and 1799 (Pagan, 1975) and remained 
there until 1801. Both he and his son. Isidore (previously cited 
herein in reference to a shrew) described numerous additions to the 
vertebrate fauna. Isidore completed his father's work on the reptiles 
of Egypt, wrote his biography (Geoffroy St.-Hilaire. 1847). and 
published on the fishes of the Nile and the Red Sea. Further notes 
on this era of Egyptian natural history are in Sherborn (1897). The 
great explorer. Riippell (1826. 1829. 1842), added many new forms 
to the list of Egyptian mammals. 

Anderson's (1898) introduction in Volume I of Zoology of Egypt is 
a source of considerable historical data compiled from the writings 
of early explorers. Publications of the nineteenth century usually 
mentioned the observable mammals, such as leopard, hyena, jackal, 
fox. gazelle, ibex. Barbary sheep, wild donkey, hippopotamus, 
hyrax, and hare; and some folklore was often included (Sonnini, 
1807; Russell, 1831; Wilkinson. 1832; Adams, 1870; Klunzinger, 
1878; Floyer. 1887. 1893). 

Hart (1891) published the first annotated list of the mammals of 
Sinai, but the first modern treatment of the Egyptian mammal 
fauna was Anderson's (1902) monumental work. Numerous publica- 
tions followed which dealt with local collections and descriptions of 
new forms, and these were summarized by Flower (1932). In the 



OSBORN & HELMY: MAMMALS OF EGYPT 3 

same year, Innes (1932), compiled a resume of the rodents of Egypt. 
Information on the larger mammals continued to appear in 
geological survey reports and miscellaneous books and papers (Bar- 
ron and Hume, 1902; Hume, 1906. 1921; Barron, 1907a; Stuart, 
1910; Murray, 1912). An interesting list appears in Budge's (1926), 
The Dwellers on the Nile. 

Wassif (1944 et seq.) began a series of papers on bats, rodents, and 
local faunal lists. Setzer (1952 et seq.) started a trend in more strict- 
ly taxonomic treatments of the mammals. Negumi (1952) compiled 
a semi-popular account (in Arabic). Sandborn and Hoogstraal (1955) 
brought information on bats up to date, and Hoogstraal (1962, 
1963, 1964) reviewed most of the work that had been done since 
Flower (1932). Papers continue to be published on this fauna (Bauer, 
1963; Osborn, 1968a; Hoogstraal et al., 1968; Lay and Nadler, 1969, 
1972; Wassif et al., 1969; Osborn and Krombein, 1969; Missone, 
1969, 1970; Pelikan et al., 1971; DeBlase, 1972; Gaisler et al., 1972, 
1973; Haim and Tchernov, 1974; Lay et al., 1975). Numerous 
references to Egyptian forms are in the works of Ranck (1968), Har- 
rison (1964, 1968, 1972), and Meester and Setzer (1971). 

The first recorded observation of decline in Egypt's fauna was by 
Floyer (1893). In reference to the paucity of animal life in the 
Eastern Desert, he (Floyer, 1893, p. 41 1) remarked, "The horse, cow, 
ostrich and wild donkey have disappeared from this country, not, 
however, by a diminished rainfall, but perhaps expelled by the 
camel." Concern over the slaughter of Egypt's game mammals and 
threatened species of wildlife has been expressed by too few in- 
dividuals (Dumreicher, 1931; Russell, 1949ab, 1951; Monaiery, 
1955; Hoogstraal, 1964; Hoogstraal et al., 1968). Prince Kemal el 
Din, a renowned trophy hunter, established an ibex sanctuary in 
Wadi Rishrash in 1900 which was maintained for approximately 40 
years (Halton, 1935; Talbot, 1960). According to Dumreicher (1931), 
good, strict game laws had been introduced by the government, and 
a short period of enforcement resulted in an apparent increase in 
hare, gazelle, and cheetah. In 1953, new game laws were established, 
and hope was expressed for the development of more reserves 
(Monaiery, 1955). Game laws, however, are not enforced, and 
animals are killed in all seasons (Hartert, 1913; Wellard, 1965, 
p. 28). 

Materials and Methods. — ¥'\e\A parties from NAMRU-3 have ex- 
plored all of Egypt except for some parts of Sinai. Collections were 



4 FIELDIANA: ZOOLOGY 

made during all seasons at numerous localities. Areas in which the 
most extensive collections were obtained are northern Red Sea 
Hills. Nile Delta. Western Mediterranean Coastal Desert, and El 
Maghra. 

Various means of obtaining sp)ecimens were used: trapping, dig- 
ging out of burrows, and shooting or netting at night under a 
spotlight. Our rodent live trap is shown in Figure 108. Bait was 
usually f)eanut butter and bread, but vegetables, dates, and other 
fruits were sometimes used. Snap traps were not used, because dead 
specimens become damaged by ants (Pelikan et al., 1971) and lose 
ectoparasites, particularly fleas. Digging rodents and small car- 
nivores out of burrows, although laborious and time consuming, has 
its rewards— specimens have most of their ectoparasites, nests can 
be examined if present, and commensals may be found. There is 
always an extra bit of biological knowledge to be gained from a 
burrow. 

Commercial live traps were used for capturing carnivores. Sardine 
bait was used successfully for most species. Steel traps were 
resorted to when other techniques failed to get specimens. Hunting 
at night with a light was an efficient way to obtain cats, hyena, red 
fox. and jackals. Jerboas were often captured at night with nets. 
Various other species of rodents and hedgehogs were captured by 
hand under a spotlight. Small dipodils were pulled from the protec- 
tion of spiny bushes with long forceps. Methods of collection are fur- 
ther discussed under individual species. 

Ectoparasites were removed from sp)ecimens as soon as i>ossible 
after collection and were sent to various specialists. 

Standard techniques were followed in preparation of specimens 
(Corbet. 1966; Harrison. 1972; DeBlase and Martin. 1974). Unless 
otherwise indicated, tables of measurements and weights are of 
adult males and females combined. In most rodents, average weight 
of males is slightly greater than that of females of the same age 
category. Explanation of abbreviations for measurements and 
methods of taking same are given in Appendix 1. In systematic 
work on the Gerbillinae. useful characters were variations in 
chamber sizes of the auditory bulla (Appendix 2. fig. 165). swellings 
of the lip of the external auditory meatus, and shajjes of associated 
bones, such as basisphenoid, basioccipital. and interparietal and 
presence or absence of the accessory tympanum. Terminology used 
in describing rodent molars is given in Appendix 3, Figure 166. 



OSBORN&HELMY: MAMMALS OF EGYPT 5 

Synonymy in the taxonomic sections is, for the most part, that of 
Allen (1939), Ellerman (1941, 1948. 1949), and Ellerman and Mor- 
rison-Scott (1951). Recent controversies and taxonomic changes are 
listed and explained in the text. 

Plant names have been revised to agree with Tackholm's (1974) 
new edition of Student's Flora of Egypt. 

Localities of collections are listed by Governorate, District, or 
other political subdivision (in capitals) followed by the name of a 
town or place. Localities in countries other than Egypt are preceded 
by the name of the country (e.g., Sudan. K ASS ALA: Wadi Onib). 
Names of small villages are sometimes preceded by the name of a 
township or oasis. 

Maps showing Governorates of Egypt are in Appendix 4, Figure 
167. All locahties mentioned in the text are listed in the Gazeteer in 
Appendix 5. Definitions of various local terms used in the text are in 
Appendix 6. 



STUDY AREA 

Egypt (fig. 1) occupies a 1,000,000-km.^ (386,000-mile^) area of 
the northeastern corner of Africa north of parallel 22° N lat. and 
east of parallel 25° E long. About 3.6 per cent of the country is con- 
sidered habitable, the rest is desert. High, naked mountains, bold 
plateaus, sterile, stony plains, fields of sinuous dunes, lush oases. 



,. 2 5* 2 6* 2 7* 28* 2 9* 30* 31° 3 2* 3 3° 34* 35* 36* 37* 




FiG.l. Geography of Egypt. Stippled areas are sand dunes. Black areas are moun- 
tains above 1,200 m. 



OSBORN&HELMY: MAMMALS OF EGYPT 7 

and the green Nile Valley make it a land of striking geographical 
contrasts. 

Egypt is a land of sunshine and intense, dry heat. Were it not for 
the Nile River, which receives no water-bearing tributaries in its 
lower 2,750 km., cultivable land would be limited to the deltas of a 
few wadis, narrow strips bordering the Mediterranean coast, and a 
few oases in the Western Desert. The Nile Valley, deserts west and 
east of the Nile, and the Sinai Peninsula are practically rainless (fig. 
2). 

Egypt is bounded on the north by the Mediterranean Sea and on 
the east by a 200-km. Israeli frontier, the Gulf of Aqaba, and the 
Red Sea. Western and southern political boundaries separate Egypt 
from Libya and Sudan, respectively. The Sudan Government Ad- 
ministration Area (SUDAN ADMIN.) boundary in the southeast 



25* 26* 27* 28* 29* 30* 31* 32* 33* 34* 35* 36* 37* 




25* 26* 27* 28* 29* 30* 31* 32* 33* 34* 35* 36* 37* 



I.HELMY1971 

Fig. 2. Rainfall map of Egypt. Isohyets are in millimeters. 



8 FIELDIANA: ZOOLOGY 

(fig. 1) zigzags between Gebel Muqsim and Bir Shalatein. 
separating the Ababda tribe, which is under Egyptian authority, 
from the Bisharin who are Sudanese (Ball. 1912; Henderson, 1965). 

Certain geopolitical terms of Pharaonic origin still in use today 
are Lower Egypt, referring to the Nile "Delta" (a term invented by 
the Greeks), and Upper Egypt (including El Faiyum) from the apex 
of the Delta south to the first cataract at Aswan. South of Aswan is 
Nubia (fig. 1), a once-settled area, which as a result of the Aswan 
High Dam, has been inundated by Lake Nasser. 

The major geographical provinces of Egypt are: Sinai Peninsula; 
Isthmus of Suez; Eastern Desert (Arabian Desert), including the 
Red Sea Hills (Northern Etbai Range), the Gebel Elba region within 
the Sudan Administrative Area, and the Nubian Desert (the 
southern section, which includes the last two, is also known as 
Bisharin Desert); Nile Vedley and Delta; Western Desert (Libyan 
Desert) and oases; and Western Mediterranean Coastal Desert (fig. 
1). Political subdivisions of Egypt (Governorates) are in Appendix 
4. 

One of the most readable accounts of Egyptian terrain is in 
Anderson's (1898) Zoology of Egypt which, incidentally, included 
Suakin, Berber, and Dongola provinces of Sudan as part of Egypt. 
Additional sources dealing with all parts of the country are Hume 
(1921), Gautier (1935), Jarvis (1937), Ball (1939), Murray (1951), 
Mitwally (1954), Said (1962), and Abu Al-Izz (1971). The majority 
of references concern specific localities or sections. 



TOPOGRAPHY 

Sinai Peninsula.— The Sinai Peninsula (fig. 1) is a triangular area 
of desert 61,000 km.^ (23,200 miles^) linking Africa with Asia and 
usually considered as part of Asia. 

Steep mountains of igneous and metamorphic rock skirted on the 
north and west with remnants of Cretaceous or Nubian sandstone 
form the southern third of the peninsula or Sinai proper. Prominent 
peaks of this region are Gebels Serbal (2,070 m.), Umm Shomer 
(2,586 m.). Musa (2,228 m.), and Catherine (2,637 m.), the highest 
mountain in Egypt. Rocky wadis drain these mountains eastward, 
abruptly into the Gulf of Aqaba, and westward, gradually over the 
broad, sandy plain of Qaa into the Gulf of Suez. Wadis Isla, Hebron, 
Feiran, and Sidri are the most extensive of the western drainage 
systems. 

A high plateau of Cretaceous, Eocene, and Miocene limestones 
dominates central and part of northern Sinai. Gebel Egma (1,620 
m.), a plateau itself, forms the southern extremity of northward dip- 
ping Badiet el Tih (Desert of the Wanderings) Plateau which 
averages between 700- and 900-m. elevation. The 500-m. high 
sandstone cliffs of Gebel Dhulal (1,570 m.) form the southern scarp 
of El Tih. Tributaries of northward flowing Wadi el Arish drain 
most of the surface of El Tih. Incidentally, the outwash plain of this 
great wadi, prior to the Israeli occupation, was the only site of 
agriculture in Sinai. Northern limits of the plateau are marked by a 
series of prominent mesas, ranging from 370- to 1,094-m. elevation, 
between which extend dunes and plains of the northern desert. 
These sand and gravel plains and belts of northwest-southeast 
oriented dunes, 80 to 100 m. in height, cover a triangular area from 
the Isthmus of Suez eastward to Rafa. This area, together with the 
plateau to the south, is called El Tih or, in combination with the 
region north of Wadi Tumilat, Isthmic Desert (Hassib, 1951; 
Tackholm, 1974). 

The Mediterranean coastal desert of Sinai, or Nile-Sinai Mediter- 



10 FIELDIANA: ZOOLOGY 

ranean Littoral (Meigs, 1966), is not as distinctive a region as is the 
Western Mediterranean Coastal Desert discussed below. In con- 
trast, the former desert receives less rainfall, is more sparsely 
vegetated (Zohary, 1944), and is low, featureless, and slowly sub- 
siding. Two shallow saline lakes, Sebakha el Bardawil and Birket 
Serbonis (Sea of Reeds) enclosed by offshore bars, occupy the cen- 
tral sector of the coast. 

Further notes on Sinai are found in Holland (1868), Hart (1891), 
Barron (1907a), Hume (1906, 1921, 1925), Kassas (1952, 1955), 
Zohary (1954), Haim and Tchernov (1974), and general references to 

Egypt- 

The Isthmus of Suez, a low, sandy, and scantily vegetated neck of 
land connecting Egypt with Sinai, contains Lake Timsah and Great 
and Little Bitter Lakes which are connected and traversed by the 
Suez Canal. The isthmus lies in the northwest branch of the Great 
Rift Valley and has, in times past, fluctuated in elevation and 
separated the adjoining land masses. Thus, a partial barrier exists 
here to the interchange of African and Asian faunas. 

Eastern Desert— Comparable geologically with the Sinai Penin- 
sula, the Eastern Desert also has mountains of igneous and 
metamorphic rock skirted on the north and west with remnants of 
sandstone, northwestward-dipping limestone plateaus in the north 
and northwest, and a northern plains section. 

The Red Sea Hills, or Northern Etbai Range, occupy nearly one- 
third of the area of the Eastern Desert. These mountains of great 
topographical complexity, with no single dominating ridge, parallel 
the Red Sea and southern part of the Gulf of Suez from Gebel 
Shellal (1,409 m.) north to Gebel Umm Tenassib (1,110 m.). 
Disintegrated by the rock splitting forces of heat and cold and 
eroded by water, these ancient peaks have become buried in their 
own rubble (fig. 3). In the Sudan Government Administrative Area, 
11 peaks rise above 1,200 m. elevation and four above 1,500 m. The 
Prominent Gebel Elba is 1,437 m. above sea level. Thirteen peaks 
north of the Sudan Administrative Area are above 1,200 m. eleva- 
tion, and eight are higher than 1,500 m. Gebel Shayeb, the second 
highest mountain in Egypt, rises 2,187 m. above sea level. 

East of the mountains, broad gravel plains, averaging 15 to 20 
km. in width, slope down to the sea and are occasionally intersected 
by narrow series of low hills. Wadis draining into the sea are 
relatively short, straight, and often deep compared with those 



OSBORN & HELMY: MAMMALS OF EGYPT 



11 




Fig. 3. Eastern Desert. Red Sea Hills. View eastward from summit of Gebel Abu 
Kharif. Gebel Ras el Barud (1,439 m.) in background. 

draining westward into the Nile Valley. Exceptions are the long, 
winding, and complicated Wadis Diib and Hodein ending at the 
southern and northern ends of El Wadah coastal plain (fig. 1). 

The Nubi'an Desert west of the Gebel Elba area, south and 
southeast of Aswan, is a country of sandy plains, low granite hills, 
and sandstone cones and buttes. This desert extends eastward into 
the Red Sea Hills and a short distance west of the Nile Valley into 
the bordering limestone scarps and sandstone pinnacles. The two 
largest drainage systems crossing the Nubian Desert are Wadi 
Dihmit, with affluents in the Red Sea Hills, and Wadi Allaqi, with 
eastern tributaries in the Elba Mountains and a great northward 
flowing branch, Wadi Gabgaba, originating in Sudan. A source book 
of extensive information on Nubia is Butzer and Hansen (1968). Por- 
tions of Nubia west of the Nile were described by Butzer (1965). 

North and east of Aswan on the Nile side of the Red Sea Hills suc- 
cessive belts of sandstone form a rough, broken terrain known as 
the Ababda Plateau. Adjacent to the Nile are two broad plains: 
Kom Ombo from 30 km. north of Aswan to Gebel Silsila and 



12 FIELDIANA: ZOOLOGY 

Lakeitah between Luxor and Qena. Four major drainage systems 
between Aswan and Qena are those of Wadis Kharit. Shait. Abad, 
and Zeidun running out of the Red Sea HHls and crossing the 
plateau. North of the Red Sea Hills are two vast tablelands, with 
eastern faces of sandstone on the Gulf of Suez overlain to the north 
and west by limestones: Gebel Galala el Qibliya or South Galala 
Plateau (1,464 m.) and Galala el Bahariya or North Galala Plateau 
(1.247 m.). These tablelands are separated by Wadi Araba which is 
30 km. at its widest, penetrates halfway from the Gulf of Suez to the 
Nile, and was a bay in Upper Mesozoic (Said, 1962). 

Incidentally, there is no geological evidence to support Ranck's 
(1968, p. 52) theory that tectonic disturbances during Pliocene- 
Quaternary might have so altered the course of the Nile that it flow- 
ed into the Red Sea via Wadi Araba or Wadi Ghuweiba further 
north. 

West and southwest of the Galalas lies another series of plateaus 
which border the Nile Valley from Qena north to Wadi el Digla and 
dominate two-thirds of the northern part of the Eastern Desert (fig. 
1). Cliffs of limestone also intermittently occur west of the Nile be- 
tween Luxor and Gebel Deshesha, at the level of el Faiyum. 

Southernmost of the aforementioned eastern plateaus is Maaza 
Plateau (Hume, 1921), with elevations of over 500 m. This plateau is 
drained toward the Nile by steep walled Wadis Qasab, Umm Dud. 
and Asyuti and its tributary, Wadi Habeeb. Wadi Gurdi and Wadi 
Umm Omeyid are two of the larger systems draining this plateau 
eastward into Wadi Qena. Plateaus to the north and .their major 
wadi systems are El Saff, drained by Wadis Tarfa and Rishrash, and 
Helwan Plateau, drained by Wadis Garawi (figs. 1, 4), Rished, Hof, 
and El Digla. The last separates Helwan Plateau, with its highest 
point at Gebel Hof (317 m.), from Gebel Mokattam (205 m.). The 
tops of these tablelands are severely eroded to form large areas of 
barren rock fragments or hamada. According to Hume (1925), serir 
plains have also developed on the higher summits. 

Wadi Qena (fig. 1), mentioned earlier, has its northernmost 
tributary west of Gebel Gharib in the Red Sea Hills and flows 
southwards about 320 km. to Qena, receiving tributaries from a 
vast part of the western slopes of these mountains and eastern parts 
of the plateaus. Important tributaries from the Red Sea Hills are 
Wadi el Atrash, Wadi Fatira, Wadi Merkh, and Wadi Hammad. The 
main part of Wadi Qena is a broad, sterile plain of sand. clay, and 
stone. 




ho 



6 



o 



c d 



a 



s «» 

^ § 

•^ 



13 



14 FIELDIANA: ZOOLOGY 

The northern limits of the plateau country are marked by a chain 
of low mesas between the Nile Valley and Suez. Between these 
outlying mesas and the main plateaus is a rubble-covered stretch of 
hillocks and hollows of Oligocene Nile fluviatile. This formation also 
occupies a large area west of the Nile (Hume. 1921; Davis, 1953; 
Said, 1962). The presence of soft Oligocene beds is indicated by the 
numerous burrows of rodents as opposed to compact Miocene rocks 
where burrows are lacking (Barron, 1907b). The Wadi el Gafra-Wadi 
Iseili-Wadi Gindali system (figs. 1, 5) drains the central and nor- 
thern parts of the Eastern Desert plateaus, cutting across the 
fluviatile and between the outlying mesas to debauch in a broad, 
sandy area on the eastern margin of the Delta. Wadi el Baharri 
drains northeastward from the plateau into the northern end of the 
Gulf of Suez. Wadi Ghuweiba, with tributaries cutting over 70 km. 
inland from the Gulf of Suez, separates Gebel Ataqa (871 m.) and 
Akheider Ridge (367 m.) from the main plateau with an outwash 
plain 25 km. wide. A limited agriculture is practiced by Bedouins in 
this area. 

The northern section of the Eastern Desert consists of low, 
featureless hills and plains of serir interspersed with sandy wadis 
and depressions (Tortonese, 1948) lying mainly south of Wadi 
Tumilat, an ancient course of the Nile which is now a cultivated area 
along the Cairo-Ismailia canal. Similar terrain occurs west of the 
Nile Delta (see under Western Desert). 

The Eastern Desert as a whole has a system of external drainage. 
Water is the most obvious agent of erosion and has carved the vast 
and intricately complex wadi systems. Wind erosion and deposition 
are negligible and, other than local sand shadows and drifts, has 
created only three areas of true dunes — El Khanka near Cairo, the 
deltaic plain of Wadi el Laqeita. and a range in the southwestern end 
of El Waddah. the coastal plain north of Gebel Elba. 

Nile Valley and Delta. — For about 300 km. from Wadi Haifa to 
Aswan, prior to inundation by Lake Nasser (fig. 1), the Nile Valley 
in Nubia extended between cliffs of sandstone and granite, intermit- 
tent outwash plains and narrow benches covered with acacias, 
groves of date palms, and gardens. From Aswan to Cairo, a distance 
of about 800 km., the Nile Valley stretches like an elongated oasis 
from 1 to 23 km. wide, separating Egypt into two quite different 
halves. 

The Nile Valley is bordered by cliffs of sandstone from Aswan 



OSBORN & HELMY: MAMMALS OF EGYPT 



15 




I Fig. 5. Eastern Desert. Wadi Gindali south of Cairo-Suez Road. Dominant shrubs 

I are Anabasis articulata and Hammada elegans. 



North to Esna and by limestone from Esna to Cairo. North of 
Asyut, the western cHffs are considerably lower than those to the 
east. The river runs closest to the eastern cliff boundaries and, in the 
great bend at Qena, nearly washes the feet of 300 m. walls. In wider 
stretches of the Valley, ancient terraces bearing traces of prehistoric 
man rise high above the present alluvial level. 

Near Cairo, the river divides into two branches which flow into 
the Mediterranean at Rosetta (Rashid) and Damietta (Dumyat). The 
Delta is about 166 km. long and 250 km. wide and is intensively 
cultivated, being likened to a vast market garden by Russell 
(1949a). The basin system of cultivation and irrigation prevails 
throughout, and the entire Valley and Delta are interlaced with ir- 
rigation and drainage canals (fig. 6). Islands of sand and saline soil 
occur here and there. Ard el Barari (The Barrens) is an area of poorly 
drained, salty wasteland (Hassan, 1953) of marsh and swamp along 
the inner borders of the Delta Lakes (Manzalah, Burullus, Idku, and 
Maryut). Barrier beaches of sand occur on the outer shores of these 
lagoons. Near Baltim and on the northern edge of Lake Burullus are 



16 



FIELDIANA: ZOOLOGY 




Fig. 6. Nile Valley near Giza. 

fields of anchored crescentric dunes (Drar, 1955; Meigs, 1966) be- 
tween which are date palm groves and gardens. 

Western Mediterranean Coastal Desert.— The Western Mediterra- 
nean Coastal Desert (fig. 1) is a distinct northern part of the 
Western Desert (discussed below). This desert extends from Alexan- 
dria westward about 600 km. to Salum and varies in width from 15 
to 30 km. in the eastern and central sections to a few kilometers in 
the west, south of the cliffs at Salum. 

Various names applied to this region are: Marmarica (Hassib, 
1951). Mareotis District (Kassas, 1955), Western Mediterranean 
Coastal Region (Tackholm, 1956). and Qattara Littoral (Meigs, 
1966). 

This coastal desert differs from the Sinai Littoral in the fact that 
it is calcareous rather than siliceous, has a higher rainfall, and has 
the richest flora in Egypt other than that of the Gebel Elba area 
(Tadros, 1953). At various intervals west of Alexandria, dunes of 
white oolitic sand form the coastline. Usually paralleling the sandy 



OSBORN & HELMY: MAMMALS OF EGYPT 



17 




Fig. 7. Western Mediterranean Coastal Desert near Burg el Arab. Coastal sand 
dune, salt marsh, and limestone ridge. Vegetation: foreground, Juncus sp. and 
Limonium delicatulum; sand dune at right, Ficus sp. planted by Bedouins; salt 
marsh, a variety consisting mainly of Halocnemon strobilaceum, Arthrocnemon 
glaucum, and Limoniastrum monopetalum. Suaeda fruticosa occurs on limestone 
slopes in the background. 

coast is a series of two valleys containing salt marshes alternating 
with limestone ridges (fig. 7) (Shata, 1955). The latter were probably 
offshore bars in the Pleistocene (Said, 1962). At Ras el Hekma, 
Ageeba, and Salum, sheer cliffs border the sea. Inland of the cliffs, 
ridges, and salt marshes lies a relatively flat strip of sand and clay 
soils (fig. 8) interspersed with hamada. 

A few relatively short wadis drain the annual runoff from the 
coastal desert. During heavy rains, they become torrents carrying 
large quantities of soil into the sea. 

Figs are cultivated in a semi-wild state on the coastal dunes (fig. 
7); and dates, along the margins of salt marshes and, together with 
olives, in inland valleys. Barley, the chief crop of the semi-nomadic 
Bedouins, is grown on the more level clay and loam soils (fig. 8), 
where earlier, the Romans also practiced dry-farming. Sandy soils, 
particularly of the Sidi Barrani area, are also the site of fig trees 
and, in addition, produce annual crops of watermelons. 

With the introduction of irrigation water via canals from the Nile 



18 



FIELDIANA: ZOOLOGY 




Fio 8. Western Mediterranean Coastal Desert 4.8 km. E of Abu Mena. Soil is 
clay. Vegetation: Lycium sp. in foreground and Anabasis articulata in background. 
This area was previously Bedouin barley fields. Note burrow of Meriones shawi isis 
under Lycium. Rod is divided into 10-cm. units. 

Delta, the coastal area as far west as El Hammam is rapidly being 
changed. As a result, the Nile rat (Arvicanthis niloticus) and the 
mongoose {Herpestes ichneumon) have become part of the fauna. 

Western Desert. — In contrast with Sinai and the Eastern Desert, 
the Western Desert lacks extensive areas of high relief. Gebel 
Uweinat (1.892 m.). with its highest point in Sudan, is an isolated 
sandstone and igneous massive where the borders of Egypt, Libya, 
and Sudan intersect (Osborn and Krombein, 1969). Gilf el Kebir 
(The Great Cliff) of Nubian sandstone. 100 km. N of Gebel Uweinat. 
rises 1 .000 m. above sea level at its southern margin. The northern 
limestone plateaus are no more than 500 m. in elevation. 

The landscape of the Western Desert is wind-dominated (fig. 9). 
Erosion by water is not as obvious as it is in the Eastern Desert and 
Sinai, except for the long, winding wadis of Gebel Uweinat and Gilf 
el Kebir. High plateaus around oases have been dissected to about 
the same extent as plateaus in the Eastern Desert (Murray, 1950); 
otherwise, wind has removed much of the relief. 

The northeastern part of the Western Desert, which is traversed 




19 



20 



FIELDIANA: ZOOLOGY 





^'^ 



4/^:^ A 






Fk;. 10. Western Desert. Meandering sand sheet in area between Wadi el Natroun 
and Bir Victoria. Vegetation: Artemisia monosperma and Pituranthos tortuosus. 

by the Cairo- Alexandria desert road, is a monotony of low, rolling 
hills and plains of Pleistocene and recent material interspersed with 
small sand sheets and narrow, sinuous sandy depressions or 
meanders of sand (fig. 10) sporadically interrupted by low ridges 
capped with weathered Pliocene limestone (Shata, 1955). Similar 
low hills and sandy depressions were described by Tortonese (1948) 
in the Eastern Desert south of Wadi Tumilat. 

El Daffa (Marmarican or Libyan Plateau) in the northwest (fig. 1). 
is a vast pebble-covered plain with a scattering of mud pans and 
large areas of hamada. This is the Miocene limestone that forms 
cliffs along the Mediterranean coast (fig. 11), 200 m. promontories 
on the north of Qattara Depression (fig. 12), and scarps bordering 
Siwa Oasis and smaller depressions west of Siwa. A narrow tongue 
of this formation protrudes eastward south of Wadi el Natroun and 
the aforementioned rolling plains to the Nile Valley (Shata, 1955), 
ending in prominent Gebel Ghigiga (197 m.) near Abu Rawash. 
South of the broken limestone hills is the previously mentioned 



OSBORN & HELMY: MAMMALS OF EGYPT 



21 




Fui 11. Western Mediterranean Coastal Desert. Limestone cliffs near Salum. 
Vegetation: Rhus tripartita, Limoniastrum monopetalum, Limonium pruinosum, 
Pancratium maritimum, Moricandia nitens. Asparagus stipularis, Noaea 
mucronata, Pituranthos tortuosus, Rumex cyprius. and Euphorbia dendroides. 

broad area of Oligocene Nile fluviatile that extends some 200 km. 
SW of Cairo. 

Deep depressions or oases in the Western Desert are a result of in- 
itial erosive action of water on folds and fractures in the limestone 
formations during pre- and Pleistocene pluvial periods. During 
subsequent dry phases of Pleistocene and Recent Periods, wind ac- 
tion completed the excavations to ground water level. Smaller 
depressions, such as Wadi el Natroun and Wadi el Farigh, where 
soft sediments were exposed, are considered to have been excavated 
entirely by wind (Abu Al-Izz, 1971. p. 197). Opinions on the forma- 
tion of these depressions vary slightly (Hume, 1925; Ball, 1927; 
Caton-Thompson and Gardner, 1932; Mitwally, 1953b). Most 
depressions have spectacular northern escarpments, but are open to 
the south and rise gradually to the level of the desert floor. Excep- 
tions are El F'aiyum and Bahariya Oasis which are completely sur- 
rounded by cliffs. Within the latter are isolated cones and buttes. 
Depressions east from Giarabub through Siwa Oasis and Qattara 
Depression follow the Eocene-Miocene boundary. Other oases are 
in Paleocene and Eocene deposits. Depression floors are close to or 



22 



FIELDIANA: ZOOLOGY 




Fk; 12. Western Desert. Wadi Labaq in noil lucisiern part of Qattai a Dtpn s^>l<)n. 
Minqar Abu Dweis in distant background. Shrubs in background are Hammada 
elegans. Small, cushiony plants are Convolvulus lanatus which, if unmolested, grow 
to a height of 50 to 60 cm. The area is browsed continuously by gazelles and 
periodically by camels. 

below sea level. The lowest point (-137 m.) is in Qattara. Springs, 
artesian wells, and cultivable land have permitted conversion of 
parts of some hollows into habitable oases (e.g., Siwa, Qara. 
Farafara, Bahariya, Dakhla. and Kharga Oases; and Wadi el 
Natroun and El Faiyum). The last borders Birket Qarun (-45 m.), 
the remains of prehistoric Lake Moeris. and has been supplied with 
Nile River water for 3.000 years by the 435 km. Canal of Joseph or 
Bahr Yusuf (Whitehouse. 1887). Wadi el Rayan (-64 m.) and Wadi 
Muwellih ( + 25 m.) to the west and southwest of El Faiyum are 
undeveloped depressions with springs of potable water. Between 
these hollows and Bahariya Oasis is VA Bahr (The Sea), a belt of con- 
fused or broken country consisting of cones and table hills. 

Lakes within depressions are usually salty, due to high rate of 
evaporation, and partly surrounded by sebakha (salt pan) and/or 
marsh (fig. 13). 

Winds of excavation are also winds of deposition, and material 
from the depressions is distributed over the Western Desert in the 
form of dunes and sheets of sand. Conflicting north and northwest 



OSBORN&HELMY: MAMMALS OF EGYPT 23 

winds formed the parallel ranges of longitudinal seif dunes over- 
lying the serir or pebble desert between Qattara Depression and 
Camel Pass Dune about 100 km. W of Cairo (figs. 1, 9). Ghard el 
Moharik, a dune range 6 to 8 km. wide and 450 km. long, runs from 
north and east of Bahariya Oasis southward into Kharga Oasis. Ir- 
regular accumulations of dunes occur southwest of El Faiyum and 
in the depressions of Gharak, Wadi el Rayan, and Wadi Muwellih. 
North of Giza Pyramids and between El Bahnassa and Mallawi, 
sand encroaches on the Nile Valley. The Great Sand Sea or Libyan 
Erg, a vast waterless and almost impassable area of high com- 
plicated dunes, some of which are over 100 m. high (Murray, 1967), 
lies west of Farafara Oasis and stretches about 800 km. south of the 
Siwa-Giarabub depressions where it inundates the lower levels of 
the northward-dipping sandstone of Gilf el Kebir. 

South from the latitude of Kharga Oasis, north winds continually 
deposit sand in shallow depressions and also form ranges of 
crescent-shaped or barchan dunes which move slowly over the 
plains. Within Kharga Oasis, marching dunes temporarily inundate 
fields and villages. Sand dunes of the Western Desert were first 
described in detail by Beadnell (1910) and have received con- 
siderable attention since then by various desert explorers (Kemal el 
Din, 1928; Bagnold, 1931, 1933, 1935, 1936, 1942; Mason, 1936; and 
Jarvis, 1937). 



DESERT FEATURES 

Bedouins have more than 20 names for the various land forms in 
the desert (Dumreicher, 1931). Many terms refer to local 
phenomena, such as batikh, or watermelon-shaped rocks east of 
Kharga Oasis, and kharafish, grooved and ridged limestone south of 
the dune lines. Surfaces most frequently traversed will be described. 

Rami— Rami or sand occupies far less total area than does bare 
rock in the Egyptian deserts (Mitwally, 1954; Said, 1962). Sand, 
unless moving as over the surface of a dune, supports most of the 
vegetation in the Western Desert (figs. 9, 10). According to Davis 
(1953, p. 160), "Sand drift enables areas to support vegetation that 
would otherwise be quite bare, the sand acting as a mulch and 
preventing drying out of the substratum." Sand is a substrate to 
which hairy-footed mammals such as Meriones sp., Gerbillus sp., 
Jaculus jaculus, Fennecus zerda, and Felis margarita have become 
adapted. 

Serir.—Serir or pebble desert (fig. 9), a pavement of small to large 
stones or even cobbles, is the main surface formation in the Western 
Desert and northern plains of the Eastern Desert and Sinai. 
Temperature extremes cause fracturing of solid materials from 
which all the finer dust and sand has been swept away, leaving a 
stone veneer overlying hard packed sand (Hume, 1921; Harding- 
King, 1925; Tortonese, 1948; Kassas. 1952; Mitwally, 1953). Some 
authors (Zohary, 1944; Ranck, 1968) classify serir as hamada, A 
mature serir consists of angular, faceted pebbles polished smooth 
by wind and sand. Serir plains remain barren in spite of rain, 
because plants cannot become established. Travel on serir is easy 
and rapid, and wheel tracks and camel tracks remain visible for 
many years on this surface. 

Incidentally, the jerboa Uaculus jaculus) burrows under serir. 
Wind carries away the excavated sand, making its burrows hard to 
find. 



24 



OSBORN & HELMY: MAMMALS OF EGYPT 



25 




Fig. 13. Western Desert. Wadi Muwellih. Sebakha (salt crust) in foreground. 
Behind this is a strip of Juncus rigidus in shallow, salty water. Tamarix nilotica 
covers the sand mound in the background. 

Hamada. —Hamada or rocky desert occurs on limestone and sand- 
stone tablelands. Hamada is rock surface directly covered with a 
layer of rock fragments. Formation is by weathering and deflation 
(removal by wind of small particles). Vegetation occurs where rain- 
fall is sufficient and soil has formed in basins or cracks. Intact or 
jointed rock surfaces on plateaus and granite mountains and barren 
mountain slopes with loose rock are comparable to hamada. 

Balata.—Balata or mud pans are smooth, flat deposits of silt in 
shallow depressions (Mitwally, 1953a). They are common on the Li- 
byan Plateau and vary in area from a few square meters to several 
acres (Omer-Cooper, 1947). Balata support plants such as Capparis 



26 FIELDIANA: ZOOLOGY 

deserti (= C spinosa). Acacia raddiana, Zygophyllum coccineum, 
and Anastatica hierochuntica. 

Nafash.—Nafash is powdery limestone or gypsum with a covering 
of pebbles or limestone fragments or, in some areas, coin-like num- 
milites (e.g., Bahrein and an area between El Faiyum and Wadi 
Muwellih). Nafash is rarely vegetated and can be most difficult to 
traverse. 

SebakhxL—Sebakha or salt pan (fig. 13) is a mixture of sand, salt, 
and varying amounts of water upon which a solid or semi-soUd crust 
forms due to evaporation. The latter condition is usually im- 
passable. Sebakha may overlie solid ground or salty sludge. The 
crust is usually brownish and may be hard or soft, smooth or rough. 
Depressions and oases have extensive areas of sebakha that are sup- 
plied by underground water. It may also border salty lakes and low- 
lying seacoasts. According to Ball (1933), the area of sebakha in 
Qattara Depression is about 5,800 km.^ Sandy hillocks supporting 
date palm (Phoenix dactylifera), Tamarix sp., Nitraria retusa, and 
Juncus sp. occur within sebakha. 



THE WADI 

Wadi (Karkur in Gebel Uweinat, Khor in Nubia) refers to a gully, 
canyon, valley, or any dry stream bed that conveys water at ir- 
regular intervals. A wadi is the result of periodic erosion and deposi- 
tion by water. Some closed-in depressions are erroneously called 
wadis (e.g., Wadi el Rayan, Wadi Muwellih, and Wadi el Natroun). 
The mature wadi "has a main channel which is wide, deep, and well 
defined" (Kassas, 1952), side terraces and alluvial deposits, and is 
connected with numerous small to large tributaries or runnels. 
Young and immature wadis lack deposits and terraces. 

In igneous mountains, runnels may be shallow but precipitous 
and covered by massive blocks and boulders. These runnels debauch 
into drainage lines at the feet of the mountains (fig. 3). In plateaus, 
shallow runnels may traverse the surfaces and cut the slope or cliff 
of the plateau edge or the wall of the mature wadi (fig. 4). Waterfall- 
like cliffs and shelf-like ledges also develop (fig. 14) in wadis of 
plateau country. In limestone plateaus, wadis are generally narrow 
and canyon-like (fig. 4). In Nubian sandstone, running water pro- 
duces broad, open wadis (Abu Al-Izz, 1971). 

The wadi bed is at lower levels than the surrounding terrain and 
receives soil and water via complexly branched tributaries from an 
extensive drainage area (Kassas, 1953). Its water supply is thus 
many times the recorded rainfall (Hassib, 1951). As little as 1 cm. of 
rain is enough to cause flooding (Davis, 1953). Because of the torren- 
tial nature of this water supply, main channels within mountains or 
plateaus are generally devoid of plants. Vegetation is usually 
estabhshed on terraces bordering the channel or on islands within 
broad channels (fig. 15). On plains, where the rate of stream flow is 
negligible, plants may be established in the wadi bed or sometimes 
on rehcts of old terraces (fig. 5). Rodent burrows are usually found in 
these structures. 

Most deposits in wadis are alluvial, but wind-blown sand may ac- 
cumulate within the mountains or plateaus, usually in the 

27 




Fig 14. Eastern Desert. Wadi Hof. Dry waterfall and ledges in Miocene 
limestone. 



28 



OSBORN & HELMY: MAMMALS OF EGYPT 



29 




^»i*- 

^ 



4- 




l"u. 15. l>a.sLci;i iXociL. Vvuu, ci ic^ic.j, u ui kill. 77 on QenaSataga road. Hills are 
granitic. Vegetation: Citrullus colocynthis, prostrate plant in foreground; Aerva 
javenica, low. white shrub; and Leptadenia pyrotechnica, tall, woody shrub. 

downstream parts. Transported materials (silt, sand, gravel), if deep 
enough, usually support some vegetation (Kassas, 1966). 

Flash floods disrupt the wadi ecosystem both by building up and 
removing soil and by destroying vegetation and the habitats of 
animals, if not the animals themselves. Erratic floods sometimes 
uproot trees, move large boulders and masses of soil; totally chang- 
ing long sections of wadis (Kemal el Din, 1928; Kassas and Imam, 
1954; Hoogstraal et al., 1967; Kassas and Girgis, 1970). Flood 
waters or seyal may or may not reach the mouth of a wadi, depen- 
ding upon the amount and locality of rainfall. 



CLIMATIC FACTORS 

The Pleistocene is considered to have been a time of heavy rainfall 
in Egypt, during which the desert wadis were running streams 
(Sandford and Arkell, 1939). Pleistocene terraces carved by recent 
rains are common in the Eastern Desert (fig. 16), and "mud 
sphinxes" or wind-carved remnants of ancient lake sediments occur 
in some parts of the Western Desert (e.g., Kharga Oasis). 

The present, arid, Mediterranean climate of Egypt, except for a 
slight reduction in rainfall from 1910-1940 and increases in moisture 
about 2,350 B.C. and 5,500 B.C., dates back to about 8,000 B.C. 
(Butzer and Twidale, 1966). Contemporary changes that have been 
wrought on vegetation, with consequent reduction or disappearance 
of animal species, are not due to climatic change, as proposed by 
Russell (1949a), but are due solely to actions of man and livestock 
(Hassib, 1951; Long, 1955; Pearse, 1955). 

In Egypt, rains can be expected from November to April, with 
maximums in December and January (table 1), the coolest months 
of the year. Annual precipitation varies from zero or a trace to a 
series of cloudbursts. Years of low rainfall are usually followed by 
several seasons of higher rainfall (El Danasori, 1957). 

The Egyptian deserts are among the most arid regions in the 
world and have the highest air temperatures and lowest rainfall and 
relative humidity of most African and Asian deserts (Migahid and 
Abd el Rahman, 1953a). In combination with the seasonality, ir- 
regularity, and meagerness of rainfall (fig. 2), low atmospheric 
moisture, and high temperatures (table 1), winds are strong and 
dessicating. These factors, except for wind, become more pro- 
nounced with increasing distance from the Mediterranean coast in 
Sinai and Egypt (Kassas, 1952; Migahid et al., 1955), westward into 
the middle of the Sahara (Perret, 1935), and eastward into the Mid- 
dle Eastern and Southwest Asian Deserts (Zohary, 1954). 

Temperature.— Prevailing high temperatures (table 1) in combina- 
tion with low rainfall increases aridity and imposes additional hard- 

30 



OSBORN & HELMY: MAMMALS OF EGYPT 



31 




Fk; 16. Eastern Desert. Bir Qiseib in Wadi Qiseib. Note Pleistocene mud flows. 
Vegetation: date palms (Phoenix dactylifera), reeds {Phragmites australis), Imperata 
cylindrica, ,/uncus rigidus, Tamarix nilotica, Lycium arabicum, and Nitraria retusa. 

ships on desert life. Areas of low relief, plateaus, and south-facing 
chffs and slopes are the surfaces most exposed to solar radiation 
and evaporation and the least likely to support life. North-facing 
slopes of otherwise barren mountains may support a lichen flora 
simply because of subtle microclimatic differences (Abd el Rahman 
and Batanouy, 1966). 

According to Hefny (1953), Egyptian deserts are mostly 
temperate (i.e., mean temperature of coldest month is below 18° C). 
The Red Sea coast south of Quseir is considered Tropical Desert, 
since it has a hot desert climate (i.e., mean temperature of coldest 



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OSBORN&HELMY: MAMMALS OF EGYPT 33 

month is above 18° C). In contrast with intense daily heat in sum- 
mer, desert nights are often very chilly, and cloudy winter days are 
penetratingly cold (table 1). 

In order to escape dessication, small desert animals either retire 
into sand, under stones, or in burrows during the day and become 
active at night. Studies of burrow microclimates of Egyptian 
rodents by Williams (1954), Briscoe (1956), and Yunker and Guirgis 
(1969) will be discussed under appropriate species. Larger mammals 
and men seek the shade of trees or cliffs during the hottest part of 
the day. Gazelles in treeless areas scratch hollows in the mounds 
beneath shrubs in order to obtain shelter from the sun. 

Vyind— Prevailing winds over most of Egypt are from the north- 
west. In the Western Desert south of Kharga Oasis, winds are nor- 
therly. In the southeastern coastal and mountain section (Gebel 
Elba) south of Ras Banas, prevailing winds come from the 
southeast (Abu Al-Izz, 1971). 

From March to May, strong, hot winds known as Khamsin (Fifty- 
Day Winds) or Pentecost Winds veering from south to northwest, 
produce severe sand and dust storms. These winds tend to scatter 
sand, but do not affect the stability of dunes (Ball, 1927). Duration 
is usually two to three days, sometimes recurring every two to three 
days (Hume, 1925). After a rain, such winds are extremely desic- 
cative. Khamsin are common to the greater part of Egypt, but are 
most severe in the non-mountainous areas, particularly the coastal 
deserts. 

Wind velocity is greater on the coast than inland, especially in 
winter (Migahid et al., 1955) and, along with salt spray, is a major 
limiting factor suppressing development of vegetation on exposed 
ridges and cliffs near the seacoast (fig. 11). 

Strong winds dry the upper soil layer, destroy seedlings, shorten 
leaf and flowering periods by increasing transpiration, and suppress 
animal activity (Abd el Rahman and Hadidy, 1959; Abd el Rahman 
and Batanouy, 1959; Migahid and Ayyad, 1959a). Wind and sand 
produce abrasive action that is damaging to plants. Wind-deposited 
materials sometimes cover plants (Montasir, 1954), although 
species such as Calligonum comosum, Nitraria retusa, Zygophyllum 
album, Stipagrostis vulnerans (= Aristida vulnerans) and Tamarix 
sp. (figs. 9, 17), together with wind-blown sand, form hillocks which, 
incidentally, are burrow sites for animals such as rodents and foxes. 

Rainfall.— The Western Mediterranean Coastal Desert from 



34 



FIELDIANA: ZOOLOGY 




Fig. 17. Western Desert. El Maghra. Hillocks with Nitraria retusa anu iamanx 
tetragyna interspersed with sand and ./uncus subulatus. Background is salt marsh. 

Salum to the Nile Delta receives an average winter rainfall of 70 to 
200 mm. each year. Alexandria receives an average of 188 mm. per 
year. The amount of rainfall decreases to about 80 mm. at Port Said 
(Hefny, 1953) and to about half that of Alexandria at El Arish in 
Sinai (Migahid et al., 1955). Ali (1952) suggested that west to east 
differences in precipitation were caused by local orientation of the 
coastlines to direction of moisture-bearing northwesterly winds 
(fig. 2). 

Inland, the amount of rainfall decreases sharply. Cairo, 170 km. 
from the sea, has an annual average of 34 mm.; Helwan, 200 km. in- 
land, 31 mm,; El Faiyum, 230 km. inland, 16 mm.; and Siwa, 390 
km. inland, less than 1 mm. 

Omer-Cooper (1947) remarked on the snail belt in El Daffa be- 
tween 31 ° 00 ' and 30 ° 30 ' N lat. and noted that, south of it, the coun- 
try was barren and almost devoid of plants. Note also that this belt 
is slightly beyond the southernmost cistern that was dug in this 
area, indicating how far south effective rainfall penetrates. 

Kassas (1955) reported the average annual minimum and maxi- 
mum rainfall for Alexandria during a 50-year period to have been 59 
and 387 mm., respectively. From 1900-1947, these figures were 83 



OSBORN&HELMY: MAMMALS OF EGYPT 35 

and 287 mm., respectively. For Helwan, minimum and maximum 
from 1904-1947 were 2 and 25 mm., respectively; Siwa, and 44 
mm. for the same period. South of 29° N lat., the average annual 
rainfall is to less than 25 mm. Dakhla Oasis receives an annual in- 
crement of 1 mm. or less of rain. The southern two-thirds of the 
Western Desert is practically rainless (table 1, fig. 2, and Perret, 
1935). 

Missone (1970) thought that no rain had fallen on Gilf el Kebir for 
30 years. Nevertheless, there are green trees in the wadis {Maerua 
crassifolia. Acacia sp., and Balanites aegyptiaca). The exception in 
this sterile desert area is Gebel Uweinat which receives orographic 
rain in some part every 7 to 10 years (Peel, 1939; Williams and Hall, 
1965) and supports a flora in the wadis simileir to that in wadis of 
the Red Sea Hills (Osborn and Krombein, 1969) and some species 
high up on the cliffs (Shaw, 1931). The only natural sources of per- 
manent water in the Western Desert are in the pot holes and springs 
of Gebel Uweinat and springs of oases. In the latter, numerous arte- 
sian wells have been bored for land reclamation. 

The northern part of the Eastern Desert south to the latitudes of 
Cairo and Suez receives an average annual rainfall of between 27 
and 30 mm. The Umestone and sandstone plateaus further south to 
about 29 ° N lat. receive less precipitation, but annual rains can be 
expected. Cloudbursts are not infrequent. Natural water sources are 
Bir Qiseib, Bir Abu Sanduq, Bir Zafarana, and several springs along 
the cUffs bordering Wadi Araba. 

Between 29° and 26° N lat., there is seldom any Mediterranean 
climatic influence. Rainfall is scanty and variable and usually comes 
in irregular cloudbursts at intervals of 10 years or more. 

Asyut receives an average of about 7 mm. of rain per year; Minya, 
2 mm.; Hurghada on the Red Sea coast, about 3 mm. (Kassas and 
Zahran, 1962; Kassas and Girgis, 1965). Higher mountains, pro- 
bably those of 1,500-m. elevation or higher, receive orographic rains 
(Kassas, 1953). Permanent pools of water occur near the summits of 
Gebel Shayeb and Gebel Abu Harba. A few other natural sources of 
water occur in the area, such as Bir Qattar, Bir Sheitun, Bir 
Mellaha, and Bir Ambagi. The last two are brackish. A few pot holes 
(e.g., Qalt Umm Disi) and numerous dug wells occur in this section 
of desert, as indicated on topographical maps. Some wells have been 
in use since Pharaonic times. 

The middle part of the Eastern Desert from 26° N lat. to 24° N 
lat. receives erratic rainfall in the form of cloudbursts. The mean an- 



36 FIELDIANA: ZOOLOGY 

nual precipitation at Qena is 4 mm.; Quseir, 5 mm.; and Aswan, 
3 mm. 

The southern part or Nubian Desert from Aswan south to Wadi 
Haifa is part of the rainless midland of the Sahara and transitional 
between tropical desert to the south and east and the temperate or 
Mediterranean desert to the north. The rare incidents of rain in this 
area may occur in winter (Mediterranean affinity) or in summer 
(tropical affinity). Devastating floods have been recorded in Wadi 
AUaqi in 1830 (Ball, 1912) and 1902. High-water marks of the latter 
are still visible in Wadi Umm Karayiet (Osborn, 1968a). Another 
flood reached the mouth of Wadi Or in 1962 (Giegengack, 1968). The 
Gebel Elba region, east of the dry Nubian Desert, lies within the hot 
desert (tropical desert) of Hefny (1953) which extends along the Red 
Sea coastal area from Quseir southward. Southeasterly winds 
predominate along the Red Sea coast and eastern slopes of the 
mountains as far north as Ras Banas. 

The average rainfall in the Gebel Elba region is 25 mm. per year; 
during 1904-1912, it varied from 5 to 91 mm. Orographic rainfall 
may reach 400 mm. F*recipitation generally falls in winter, but occa- 
sional summer showers occur, although the northern limits of mon- 
soon rains is 18° to 19° N lat. 

Bir Abraq, north of Gebel Elba, is a well-known source of fresh 
water. There are few springs in the Gebel Elba region, and in the dry 
season, naturally occurring water is available from multitudinous 
pot holes iQulut) in steep and sheltered canyons. 

Inland Sinai is practically rainless, although it has a greater rain- 
fall than much of the Eastern Desert (fig. 2, table 1). Higher moun- 
tains in the south receive orographic precipitation, possibly amoun- 
ting to 300 mm. annually (Zohary, 1944). Snow falls occasionally 
above 1,000-m. elevation and may remain for several months. One 
meter of snow was measured at St. Catherine Monastery in 
February 1937 (Drar, 1955). This area, Feiran Oasis, and a few more 
localities, such as Ain el Gedeirat, Ain el Furtaga, Ain el Senned, 
Ain Abu Nateigina, and Ain Sudr, are natural, continuous sources 
of fresh water in Sinai. Water at Hammam Musa and Ayun Musa is 
brackish, although the presence of date palms (Phoenix dactylifera) 
indicates a "fresh water layer among the underground water 
layers" (Zahran, 1969, p. 244). 

Dew. — Fog and dew are common to all parts of the desert (Russell, 
1949a). Dense fogs were reported to have occurred during October 



OSBORN&HELMY: MAMMALS OF EGYPT 37 

and November in the northern part of the Eastern Desert (Tor- 
tonese, 1948). Dew occurs in the mountains of the Eastern Desert 
(Ball, 1912) and in the spring in the Gebel Elba area (Hoogstraal et 
al., 1957a), In the Western Desert, heavy dew was recorded during 
July 2 to 5, 1935, at Qara, and thick fog and dew occurred in late 
July at 30° 26 ' N lat., 26° 27 ' E long., and 29° 57 ' N lat., 25° 56 ' E 
long. Water dripped from cars, formed puddles, and saturated the 
soil to a depth of one-fourth inch (Omer-Cooper, 1947). We ex- 
perienced comparable fog at Bir Shaqqa on August 20, 1964. North- 
west of Cairo near Abu Rawash, early morning mists and occa- 
sionally fogs were common throughout the year, except during 
April, May, and June, and indicated high nocturnal relative humidi- 
ty. Fog usually disappeared before 8 or 9 A.M. and generally oc- 
curred in the lower cultivated areas and semi-desert; rarely in the 
desert (Yunker and Guirgis, 1969). 

A considerable amount of dew is precipitated on the Western 
Mediterranean Coastal Desert during the rainless part of the year 
and is reported to be of significance to shallow rooted plants 
(Migahid and Ayyad, 1959a). During the rainless season, some 
perennials produce ephemeral rootlets close to the soil surface 
(Kassas, 1953). There is a possibility that the greater amount of dew 
in the mountains and coastal desert keeps certain perennials alive. 
Dew is known to be absorbed by the leaves of some desert plants 
(Waisel, 1958), and it is undoubtedly the water source of algae and 
lichens which grow on pebbles and which are a food source of desert 
beetles. The Bedouin belief that the beetles live upon pebbles (Mac- 
Dougal, 1913) is therefore a truism. 

Dew is thought to be a source of water for some birds (Edney, 
1966), and it may also be used by small desert mammals such as 
Jaculus jaculus (Bagnold, 1954). The importance of fog and dew in 
temporarily relieving the physiological pressures of evaporation in 
desert animals and plants appears to be a subject worthy of con- 
sideration. Tortonese (1948, p. 156) noted that dew had a 
"remarkable effect on the vegetation in the first hours of the morn- 
ing," but was unable to clarify the biological importance of dew in 
the desert. 

Water and Desert Life.— Water is singularly the most important 
factor for desert life. Desert plants have evolved numerous adapta- 
tions for conserving water and reducing transpiration (Migahid and 
Abd el Rahman, 1953abc; Zohary, 1954, 1962). 



38 FIELDIANA: ZOOLOGY 

"Scarcity and irregularity of water supplies in the desert make 
the ability of a desert animal to resist drought a matter of life or 
death" (Shehata and Kawashti, 1966, p. 181). Desert animals reduce 
water loss and escape from heat by burrowing underground, seeking 
shade, or being active only at night. Water must either be extracted 
from animal or plant tissues, metabolized from dry food, or obtained 
directly from natural sources. Those animals that must drink can 
usually endure long waterless periods and/or travel long distances 
to obtain water (Adolph et al., 1947). A large amount of literature is 
available on water regulation in desert animals (Schmidt-Nielsen, 
1964; Schmidt-Nielsen and Schmidt-Nielsen, 1949 et seq.). Sum- 
maries are available in Harrison (1964) and Vaughan (1972). Further 
remarks on water are given under individual mammal species. 



VEGETATION 

In the greater part of the Sinai Peninsula, Eastern Desert, and 
Western Desert, perennial vegetation is confined mainly in channels 
and catchments of the drainage systems. These £ire "run-off" 
deserts, in contrast with "rain deserts" which receive enough rain- 
fall to maintain a continuous vegetative cover (Zohary, 1962) as in 
the Sinai Coastal Desert, Western Mediterranean Coastal Desert, 
and the Gebel Elba region. 

Plateaus and mountain slopes are the most exposed to wind, sun, 
and evaporation and are the least vegetated. Exceptions are the 
higher peaks of Sinai and some mountain tops of the Red Sea Hills 
south of Gebel Gharib which are above 1,500-m. elevation and 
receive additional orographic rainfall (Kassas and Zahran, 1965, 
1971). Wadi beds receive a lower wind velocity, collect and contain 
run-off water (Abd el Rahman and Batanouy, 1966), have more 
favorable substrates, and therefore, have the greatest concentra- 
tions of plant life (fig. 4). Likewise, sand sheets and soils of depres- 
sions, if sufficiently deep, support vegetation in otherwise barren 
country (figs. 9, 10, 18). 

Desert vegetation has been classified into three basic subdivi- 
sions: accidental, ephemeral, and perennial. These subdivisions are 
associated with three water sources: accidental, ephemeral, and 
perennial (sub-surface storage) (Kassas, 1966; Kassas and Girgis, 
1970). An occasional or accidental rainstorm "will bring up patches 
of vegetation from seed in the most unlikely parts of the . . . desert" 
(Shaw, 1931, p. 535). Sporadic rainfall, however, limits species 
numbers in both plants and animals and has produced in both, in- 
cluding man, shifting and tenuous populations. 

Ephemerals, although they may be germinated accidentally, are 
more abundant where annual rainfall is a certainty (e.g.. Western 
Mediterranean Coastal Desert, northern part of the Eastern Desert 
and Sinai, and Gebel Elba region). Perennials are found wherever 
there is the meagerest amount of moisture, providing there are 

39 



40 



FIELDI ANA: ZOOLOGY 













[•"k, 18. Western Desert, /icaaa raddiana grove 120 km. S of El Maghra. 

suitable soils in rock crevices, basins, or wadis where water can ac- 
cumulate. Acacia trees are known to grow in catchment basins 
where there is only 1 mm. of effective rain every 15 years (Bagnold, 
1954). 

In general, areas with the deepest soil will have the largest 
number of perennials, regardless of the amount of precipitation. 
Shallow soils support ephemerals or accidentals only. Soils deep 
enough to have a permanently wet soil layer underlying a dessicated 
upper layer in the dry season form a suitable habitat for perennials. 
Climax vegetation in such situations is woody shrubs and trees 
(Kassas, 1952). There are, however, exceptions to every rule. Soils, 
no matter how deep, if covered with a layer of desert armor or serir, 
especially if the stones are cemented together with capillary salts, 
are usually absolutely barren. 

Sinai Peninsula.— The littoral zone or coast of Sinai studied by 
Zohary (1935, 1944) and Kassas (1955) is 10 to 15 km. wide and con- 
sists of dunes alternating with sand plains and serir. 

Typical perennial coastal dune plants of 50 to 100 per cent fre- 



OSBORN&HELMY: MAMMALS OF EGYPT 41 

quency are: Ammophila arenaria, Polygonum equiseti forme, Lotus 
creticus, and Moltkiopsis ciliata {=Moltkea callosa). Plants 
predominating on the sand plains are: Artemisia monosperma, 
Haplophyllum tuberculatum (=H. longi folium), Panicum turgidum, 
Pituranthos tortuosus, Urginea maritima, and Cynodon dactylon. 
Inland dunes and slope-piled dunes on mountains are vegetated 
with Panicum turgidum, Stipagrostis scoparia {=Aristida 
scoparia), Thymelaea hirsuta. Convolvulus lanatus, and Noaea 
mucronata. Lygos raetam (=Retama raetam) also occurs on these 
dunes. Dominant plants on serir, where enough moisture ac- 
cumulates, are Anabasis articulata or Hammada elegens (=Halox- 
ylon salicomicum). These two species, incidentally, extend from sea 
level to 1,000-m. elevation. Haim and Tchernov (1974) listed 
Anabasis articulata, Gymocarpus decandrum, and Zygophyllum 
dumosum as the characteristic plants of northern and central Sinai 
without consideration of site variations. They did not consult the 
works of Kassas and Zohary. Vegetation in plateaus and mountain 
areas is confined chiefly to the wadis, although higher peaks in 
southern Sinai are vegetated because of orographic moisture. 

Haim and Tchernov (1974) listed Artemisia inculta (= A. herba- 
alba), Hammada elegans (H. articulata in their text), and Anabasis 
articulata as dominants on the southern El Tih-Gebel Egma area, 
which, in their opinion, is a transition zone. On the higher moun- 
tains above 1,500 m., Artemisia inculta was noted as being the 
dominant plant. With the exception of a few shrubs, the vegetation 
of Sinai mountains and wadis is essentially the same as that in com- 
parable situations in the Eastern Desert described below. Further 
coverage of the vegetation of Sinai is by Zohary (1935, 1944, 1954) 
who considered the Peninsula on the whole to be in Saharo-Sindian 
territory on the basis of the large number of plant species belonging 
to this element. 

Saline areas around brackish springs and wells (e.g., Ayun Musa, 
Wadi Sudr, Bir Hassana, and Bir Kussaima) are distinguished by 
mound-forming Tamarix nilotica (= T. mannifera), Nitraria retusa, 
and Zygophyllum album, together with Alhagi mannifera (= A. 
maurorum) and clusters of date palms (Zohary, 1944). The same 
plants occur at similar sites in Egypt. 

Red Sea Coastal Vegetation.— At Ain Sukhna, a warm water 
spring at the foot of Khashm el Galala, there are dense stands of 
Juncus rigidus (= J. arabicus), Tammarix nilotica, Phragmites 
australis (= P. communis), scattered date palms, and various salt 



42 FIELDIANA: ZOOLOGY 

marsh species (Tackholm, 1956). Salt marsh communities form an 
intermittent fringe along the Mediterranean coasts, the Sinai and 
Egyptian coasts of the Gulf of Suez, and the coast of the Red Sea. In 
the wet saline or tidal zone and littoral sebakha, which is separated 
from the shoreline by sandbars, common species are Halocnemon 
strobilaceum, Arthrocnemon glaucum, HalopepUs perfoliata, 
Limonium pruinosum, Juncus rigidus, and Cressa cretica. The drier 
inland zone is dominated by mound-forming Tamarix nilotica, 
Nitraria retusa, Zygophyllum album, and in a few localities, 
Aeluropus massauensis {= A. brevifolius). South of Mersa el Alem, 
Suaeda monoica replaces N. retusa. Along the Gulf of Suez and Red 
Sea coasts, mounds and small dunes stabilized by these plants are 
havens for Meriones crassus and Gerbillus gerbillus. Gazella dorcas 
browses A^. retusa and other shrubs, and Vulpes rueppelli hunts 
throughout these areas. 

Mangrove [Avicennia marina) swamps occur on the southern tip 
of Sinai at Ras Muhammed and at various localities along the Red 
Sea coast of Egypt from 22 km. N of Hurghada south to Marsa 
Halaib (Zahran, 1965, 1969). 

Eastern Desert.— The open or serir desert south of Wadi Tumilat 
between the margin of the Delta and the Isthmus of Suez contains 
sand sheets plus a few broad, shallow wadis which originate in the 
limestone plateau. Although an annual rainfall is predictable in this 
area, the amount varies considerably (Kassas and Imam, 1959). 
There is usually enough rain to germinate a few species of 
ephemerals such as Mesembryanthemum forsskalei, Malva par- 
viflora, Erodium pulverulentum, and Melilotus sp. A typical domi- 
nant, Zilla spinosa, forms thickets in deep sand and shelters other 
species (Tortonese, 1948). Large drainage systems, such as Wadi 
Gafra, support stands of Lygos raetam and Panicum turgidum in 
sandy areas. 

In sections of wadis devoid of sand, plant communities are 
dominated by Hammada elegans (fig. 5). This species also occurs on 
slopes of cobble hills, Oligocene gravels, and the Khanka sand 
dunes. Relict specimens of Acacia raddiana occur in some of the 
wadis. More thorough analyses of plant communities of this area 
are in Davis (1953), Kassas (1953), and Kassas and Imam (1959). 

Vegetation in the area adjacent to Cairo and suburbs is suppres- 
sed due to grazing and cutting. The bitter Zygophyllum coccineum 
is dominant in Wadi Liblab and Cassia italica (= C obovata) is com- 
mon, but overgrazed. Further south, Z. coccineum is replaced by 



OSBORN&HELMY: MAMMALS OF EGYPT 43 

Anabasis setifera and other species (Kassas and Imam, 1954). The 
well-vegetated Wadi Garawi is shown in Figure 4. 

On the northernmost plateaus where pockets develop to form rain 
pools, soil accumulates, and Zygophyllum decumbens, Fagonia 
tristis, Limonium pruinosum, Reaumuria hirtella, and Stachys 
aegyptiaca have become established. Vegetation of most wadis in 
this region is continuous and extends to the sea in the east and to 
the Nile Valley in the west. A few ephemerals occur as far south as 
Wadi Araba. Common shrubs in the wadis east and west are Lycium 
shawii (= L. arabicum), Atriplex halimus, and Zilla spinosa. Lygos 
raetam, Tamarix nilotica, and Acacia raddiana are rare in the north- 
west due to cutting and browsing. Important reports dealing with 
the plants of this area are Davis (1953), Kassas and Imam (1954), 
Tackholm (1956), Kassas and Zahran (1962), and Kassas and Girgis 
(1964). 

South of Wadi Araba to Ras Banas, there is a barren coastal plain 
that widens to 15 or 20 km. inland from the thin band of coastal salt 
marsh vegetation discussed above. 

Few wadis, except Wadi Abu Haad, in which there is an Acacia 
raddiana climax, are vegetated on the downstream or coastal sec- 
tion. Around brackish Bir Mellaha, there are clusters of date palms 
phoenix dactylifera), Tamarix nilotica, and mats of Imperata cylin- 
drica and Juncus rigidus. The last, together with T. nilotica^ choke 
the salty stream bed of Wadi Mellaha. Bir Ambagi, another salty 
spring, produces a short stream with dense growths of T. nilotica 
and J. rigidus. 

Vegetation is otherwise limited to the shelter of the mountains 
and foothills (Kassas and Zahran, 1965). On the Nile side of the 
mountains, vegetation is also limited to upstream sections of wadis 
and to deltas along the Nile Valley which receive lateral seepage of 
water. Floods in these wadis rarely reach the Nile Valley. Flood 
waters can more easily traverse the shorter and steeper sloping 
wadis debauching into the Red Sea. 

Three subdivisions of the Eastern Desert recognized by Kassas 
and Girgis (1970) were: (A) Northern, 30° to 26° N lat.; (B) Middle, 
26° to 24° N lat.; and (C) Southern or Nubian Desert south of 24° N 
lat. 

Eighteen communities were described from these areas. Domi- 
nant plants and their areas of distribution are as follows: Lep- 
tadenia pyrotechnica (fig. 15) and Acacia raddiana occur on sand 



44 FIELDIANA: ZOOLOGY 

and gravel deposits of large tributaries throughout the Eastern 
Desert. Acacia ehrenbergiana occurs south of 27° N lat. Acax:ia tor- 
tilis is found in the eastern part of the Nubian Desert. Zygophyllum 
coccineum and Calligonum comosum are in the northern part only. 
Aerva javanica {=A. persica) (fig. 15) is found in the northern and 
southern parts. Zilla spinosa and Crotalaria aegyptiaca are northern 
and middle in distribution. Cassia italica is central, and C. senna, 
central and southern. Francoeuria crispa (=Pulicaria crispa) is 
widespread in the middle section and extends into the northern part 
as well. Salsola baryosma is central and southern in distribution. In- 
digofera argentea is southern. Salvadora persica is found in the 
eastern parts only of the central and southern sections. Extensive 
stands of Tamarix sp. occur in the deltaic parts of Wadi Araba, 
Wadi Qena, Wadi Zeidun, Wadi el Asyuti, and in Wadi Haimur 
(Kassas and Zahran, 1965; Kassas and Girgis, 1970, 1972). 

In the rocky talus and boulder-strewn gorges at the bases of 
mountains higher than 1,300- to 1,500-m. elevation, Moringa 
peregrina, the yassar tree, lives in small groves (Kassas and Zahran, 
1971). According to Kassas and Zahran (1965), the demand for the 
valuable Ben oil produced in the seeds have saved this tree from 
being cut for fuel. Rhus tripartita {=R. oxycantha) and Pistacia 
khinjuk occur on some of the higher mountain tops which receive 
orographic rain, but not on dry slopes or in wadis (Tregenza, 1958). 
Kassas and Zahran (1971) have compiled estimates of abundance of 
selected species in different habitats within coastal mountain 
groups— Shayeb, Nugrus, Samiuki, and Elba. Unfortunately, these 
higher mountains have not been explored for their zoogeographical 
affinities. 

Wadi Qena which flows southward between the Maaza limestone 
plateau and northern Red Sea Hills deserves mention. The main 
part of this great southward-flowing wadi is a broad, sterile plain of 
clay, sand, and stone with a few islands of acacia and thin bands of 
shrub along the bordering terraces (Barron and Hume, 1902). 
Kassas and Girgis (1972) showed that the dominant shrubs in Wadi 
Qena are Zilla spinosa, Calligonum comosum, Crotalaria aegyptiaca, 
Zygophyllum coccineum. Acacia ehrenbergiana, Leptadenia 
pyrotechnica, Tamarix aphylla, and T. nilotica. The last form large 
hillocks; particularly in the broader channels and deltaic parts of 
large wadis. Wadis el Asyuti, Zeidun, and el Laqeita were found to 
have approximately the same vegetation as Wadi Qena. 

On the eastern slope of the Red Sea Hills, southward from Ras 



OSBORN&HELMY: MAMMALS OF EGYPT 45 

Banas, the variety and frequency of plants increases noticeably and 
is continuous in the wadis from the mountains to the sea. In years of 
plentiful rainfall, plains and foothills are covered with vegetation, a 
very different sight indeed from the barren mountain slopes further 
north and west. 

Gebel Elba.— The Gebel Elba region in the Sudan Government 
Administrative Area has the richest flora in Egypt (Fahmy, 1936), 
with a type of tropical forest (Tadros, 1953) mostly of Ethiopian 
species. Vegetation of wadis draining seawards in this area is 
luxurious in comparison with those draining toward the Nile (Ball, 
1912). Rainfall is variable, yet may come in winter and in summer 
(monsoon affinity). Fahmy (1936), in reference to G. Schweinfurth, 
mentioned 40 days of continuous rain in the spring of 1864, during 
the season when no rain falls in the rest of Egypt. Kassas (1966) con- 
siders the area to be a "mist oasis" similar in many respects to 
Erkwit, Sudan (Kassas, 1956). The two areas share a number of 
tropical plant species, one of the most obvious of which is Dracaena 
ombet. 

Acacia etbaica, A. mellifera, A. nubica, and A. tortilis occur in the 
wadis, and A. laeta, A. raddiana, and A. tortilis, on the plains. The 
beautiful Delonix elata (=Poinciana elata) grows in rocky, in- 
accessible wadis. Several ornamental flowering shrubs of this area 
are Abutilon fruticosum, Hibiscus micranthus, and Pavonia triloba. 
In the big acacia forests, trees are entwined with lianas such as Coc- 
culus pendulus, Ochradenus baccatus, and Ephedra ciliata (=E. 
foliata). Hassib (1951) pubUshed on the vertical distribution of plant 
species on Gebel Elba by numbers. Kassas and Zahran (1971) 
distinguished four main altitudinal zones on the north- and east- 
facing slopes of Elba which varied somewhat in height due to com- 
bination of altitude, degree of slope, and other factors. These zones 
are (1) a base zone of Euphorbia cuneata scrub, (2) a zone of E. 
nubica scrub, (3) a zone of Acacia etbaica scrub, and (4) a top and 
wettest zone with patches of Dracaena ombet, Euclea schimperi, 
Dodonaea viscosa, Jasminum sp., Rhus abyssinica, R. tripartita, 
and a variety of ferns, mosses, and liverworts. Vegetation on south- 
facing slopes is confined mainly to runnels and is dominated by 
Commiphora opobalsamum at upper levels and Acacia tortilis scrub 
at the mountain base. 

Nile Valley and Delta.— According to El Hadidi and Ghabbour 
(1968, p. 394), the Nile Valley was "among the least explored 
phytogeographical regions in Egypt." In an intensively cultivated 



46 FIELDIANA: ZOOLOGY 

area such as this (fig. 6), the weed population consists of a great 
number of species (Boulos 1966b, 1967). Cultivated plants, especial- 
ly date palms, dominate the scene. Roads and canals are lined with 
Eucalyptus camaldulensis and Casuarina stricta. Dense growths of 
halfa grasses {Desmostachya bipinnata and Imperata cylindrica) 
occur along the canal banks. Marshy areas are dominated by Juncus 
rigidus and Phragmites australis. Alhagi mannifera occurs around 
the sandy edges of such areas. 

In the Nile Delta, few patches of wild land remain. Sandy islands 
support coarse grasses; low, poorly drained areas usually have a salt 
marsh type of vegetation. The ecotone between Nile Valley and 
Delta and the true desert east and west is very narrow and occurs as 
intermittent stands of Panicum turgidum, Stipagrostis plumosa 
{=Aristida plumosa), S. pungens {=A. pungens), Calligonum com- 
osum, and other less abundant species (Thomas, 1921) on sand 
sheets and as Anabasis articulata and Hammada elegans in sand- 
filled cracks of rocky areas (Davis, 1953). 

Western Mediterranean Coastal Desert— Four phytogeographical 
zones have been defined by Kassas (1955) and others for this littoral 
semi-desert, as follows: 

1. Littor£d oolitic sand dunes (Euphorbia paralias. Pancratium 
maritimum, Ammophila arenaria. Ononis vaginalis, etc., and 
cultivated figs). 

2. Sub-littoral and inland oolitic limestone ridges 3 km. apart 
{Globularia arabica. Thymus capitatus, Helianthemum lippii, 
Asphodelus microcarpus, lichen growth, etc.). 

3. Salt marsh between the two rocky ridges [Halocnemon strobi- 
laceum, Limoniastrum monopetalum, Arthrocnemon glaucum, etc.). 

4. Inland plains (barley fields, olive groves, Thymelaea hirsuta, 
Anabasis articulata, and a variety of annuals including Papaver 
dubium. Chrysanthemum coronarium, and Carthamus tenuis). 

These zones are covered with a continuous vegetation (figs. 7, 8, 
19), except as noted below, which changes to discontinuous patches 
inland on sand sheets (figs. 9, 20). The latter represents the transi- 
tion between coastal desert and the interior barren desert, which has 
only isolated patches of vegetation or solitary plants (figs. 9, 12, 17, 
18). This transitional zone appears to be comparable with what 
Ranck (1968) called the Saharan steppe in Libya. 

Grazing and cutting for fuel have completely removed the vegeta- 
tion from extensive areas around coastal towns and villages and 




Fig. 19. Western Mediterranean Coastal Desert 34 km. S of Bahig. Vegetation is 
primarily Thymelaea hirsuta and Anabasis articulata. Soil is sand and clay. 




Fig. 20. Western Mediterranean Coastal Desert. Artemisia monosperma and 
Panicum turgidum on sand sheet near area in Figure 19. 



47 



48 FIELDIANA: ZOOLOGY 

affected it elsewhere. In areas devoted to agriculture, native plants 
have been reduced to a few species. 

Numerous studies have been made of the coastal desert vegeta- 
tion, particularly on Ras el Hekma (Tadros, 1953, 1956; Montasir, 
1954: Long, 1955; Pearse, 1955; Migahid et al., 1955; Tadros and 
Berlanta, 1958ab; Migahid and Ayyad, 1959abc; Tadros and 
Sharkawi, 1960ab). 

Western Desert— The northeastern plains of the Western Desert 
between Wadi el Natroun and the Nile Delta receive 50 to 100 mm. 
of rain per year. Shallow catchment basins of Bir Victoria and a 
nearby Acacia raddiana grove in this area support the following 
plants: Carduncellus mareoticus (=Carthamus mareoticus), 
Panicum turgidum, Alhagi mannifera, Artemisia monosperma, Zilla 
spinosa, Pituranthos tortuosus, and Hyoscyamus muticus. Similar 
associations occur on sand sheets and meanders of sand (fig. 10). 

To the west of Qattara Depression, El Daffa is a sterile plateau 
with scattered mud pans (balata) that supjwrt a few species of 
plants: Capparis deserti, Acacia raddiana, Zygophyllum coccineum, 
and Anastatica hierochuntica. 

Sand dunes in the Western Desert are rarely vegetated, except for 
occasional clumps of Stipagrostis scoparia, S. vulnerans, and Cor- 
nulaca monacantha. Sand sheets in the dune areas are usually 
vegetated (fig. 9). 

Groves of Acacia raddiana occur in shallow basins scattered 
throughout the north central section (fig. 18). Omer-Cooper (1947) 
mentioned groves in the western part of Siwa Depression. There tire 
other small stands between Siwa and El Bahrein, at Talh el 
Fawakhir north of Qara, near Minqar Abu Dweiss, west of El 
Maghra about 16 km., and a few kilometers north of El Maghra at 
Hatiyet el Maghra. To the east, isolated groves occur at 28° 55 ' N 
lat., 29° 31 ' E long, and at Hatiyet el Sunt (28° 26 ' N lat., 29° 42 ' E 
long.). The largest groves, which are now almost completely dead 
(personal observation of D. Osborn, April, 1977), are near the 
southeastern end of the Depression in the vicinity of 29° 37 ' N lat., 
27° 32 ' E long, and are marked on topographic maps as "numerous 
groves of trees in shallow depressions." These trees survive in an 
area that receives an annual rainfall of to 5 mm., with larger in- 
crements about every 10 years. In such basins where sand or mud 
have collected, other plants may occur, such as Francoueria crispa, 
Astragalus trigonus, Conyza lini folia (—Erigeron crispus). 



OSBORN & HELMY: MAMMALS OF EGYPT 



49 







Fig. 21. Western Desert. Bahariya Oasis, Bir \\ igaha. \ egetaLion: Typhu dom- 
ingensis in foreground and smaller marsh plants such as, /uncus rigidus, Epilobium 
hirsutum, Centaurium spicatum, and C. pulchellum; dense stand of Panicum repens; 
and Salix subserrata. 

Hyoscyamus muticus, Stipagrostis plumosa, Fagonia arabica, Mon- 
sonia nivea, Capparis deserti, Zygophyllum coccineum, and 
Anastatica hierochuntica. The southern one-half of the Western 
Desert, except for Gebel Uweinat, the oases, and an elongate hollow 
south of Kharga Oasis, is almost barren (Ascherson, 1874; Mac- 
Dougal, 1913; Shaw, 1931, 1934; Shaw and Hutchinson, 1931). 
Often, the only plant to be seen over vast areas is Stipagrostis 
plumosa on sand sheets and in shallow, sandy runnels. 

In oases, great numbers of plants grow around non-saline springs 
and cultivated areas (fig. 21). Lakes within oases are usually salty, 
partly surrounded by sebakha and/or marsh (figs. 13, 22) with 
Phragmites australis, Typha domingensis, Juncus sp., and Phoenix 
dactylifera. Outside the damp, salty zone there are usually hillocks 
of sand covered with Nitraria retusa (fig. 17), Tamarix sp., or 
Zygophyllum album. Patches of Alhagi mannifera and stands of 
Panicum. turgidum, Imperata cylindrical and Desmostachya bi- 
pinnata occur on the deeper sand sheets. Trees (Maerua crassifolia. 
Acacia sp., and Balanites aegyptiaca) in the wadis of Gilf el Kebir 




(N «J 



C 



50 



OSBORN&HELMY: MAMMALS OF EGYPT 51 

north of Gebel Uweinat were mentioned previously (Missone, 1970). 
Gebel Uweinat in the southwestern corner of Egypt (fig. 1) has a 
flora of at least 55 species which is comparable with that of the Red 
Sea Hills (Shaw, 1931; Osborn and Krombein, 1969). Common 
species in the wadis of this area are Acacia ehrenbergiana, A. rad- 
diana,and Panicum turgidum. 

In the huge, elongated hollows of El Wadi el Gedeed between 
Kharga Oasis and the Sudanese border, islands of vegetation mark 
the presence of water, oftentimes brackish, a meter or so below the 
surface. There are clusters of dom {Hyphaene thebaica) and date 
palms [Phoenix dactylifera); areas of hummock-forming grasses 
(Stipagrostis vulnerans, Sporobolus spicatus, and Cynodon dac- 
tylon) and mound-forming Cornulaca monacantha, Tamarix amplex- 
icaulis, Acacia ehrenbergiana, and Ziziphus spina-christi; some 
extensive tracts covered with Alhagi mannifera, Imperata cylin- 
drica, and Phragmites australis; and occasional solitary specimens 
of Capparis decidua. Cattails, Typha domingensis (=T. australis), 
were found in a shallow well at Bir Qiseiba. 

A large, solitary specimen of Salvadora persica, visible for 20 km., 

is noted on topographic sheets at 23° 09' N lat., 29° 44' E long. 

' Tamarisc and acacia mounds, together with patches of grass 

(Stipagrostis sp.), occur in the areas of Bir Terfawi and Bir Safsaf 

which are west of the area under discussion (Beadnell, 1931). 

On the Nile side of this area, in Nubia, several small oases occur 
(Kurkur, Dunqul, and Dineigil) which have high underground water 

I supplies. Vegetation in Dunqul Oasis and Wadi Dunqul consists of 
Imperata cylindrical Salsola baryosma, Tamarix amplexicaulis, T. 
aphylla, and Stipagrostis vulnerans. In Dineigil, there are patches 

1^ of Alhagi mannifera, Juncus rigidus, and Imperata cylindrica 
(Zahran, 1966). Kurkur Oasis and Wadi Kurkur (Reed, 1964) sup- 
port these and many additional plants, including dom and date 
palms. Acacia ehrenbergiana, A. raddiana (Boulos, 1966a), and the 
rare Medemia argun (Boulos, 1968). 

Zoogeography.— Tine to striking taxonomic similarities of floras, 
the almost continuous deserts extending from the Sahara through 
the Middle East to Sind in Northwestern India became known as 
the North African -Indian Desert Province (Muschler, 1912) or the 
Saharo-Sindian Phytogeographic Region (Shaw, 1931; Zohary, 
1935, 1944, 1954). Likewise, the mammal fauna that is adapted to 
these deserts has been referred to as Saharo-Sindian (Harrison, 
1964; Ranck, 1968). From a more strictly zoogeographical point of 



62 FIELDIANA: ZOOLOGY 

view, the area should be called the Saharo-Sindian Sub -Region of 
the Palearctic. 

Saharo-Sindian species of mammals which are widely distributed 
in North African and Southwest Asian deserts are Vulpes rueppelli, 
Fennecus zerda, Felis margarita, Gazella dorcas, Jaculus jaculus, 
Meriones libycus, M. crassus, and Psammomys obesus. The latter is 
the more restricted ecologically. 

Most desert species are indigenous to either North African or 
Southwest Asian deserts. Penetration east and west has been 
restricted either by the Isthmus of Suez or the Nile Valley and 
Delta. Most of the faunal interchange between these deserts and 
subsequent isolation, at least in rodents, took place during the Up- 
per Pleistocene (Zahavi and Wahram, 1957; Tchernov, 1968). 

Species endemic to Egypt are Crocidura floweri in the Nile Valley, 
Dipodillus mackilligini in the southern Eastern Desert, and Ger- 
billus perpallidus in the northern Western Desert. The ranges of a 
few North African species extend through Egypt into Sinai or as far 
east as southern Israel or Jordan, e.g., Jaculus orientalis, Ger billus 
gerbillus, G. pyramidum, and G. andersoni. Dipodillus henleyi is 
known from northern Yemen (Bahmanyar and Lay, 1975). Two 
species, closely related to G. andersoni and G. gerbillus, and which 
occur in similar habitats in Israel and Southwest Asia, are G. allen- 
byi and G. cheesmani, respectively. Note that J. orientalis belongs 
to the Dipodidae, a family of Asian origin. North African species 
that range into the Eastern Desert of Egypt are Paraechinus 
aethiopicus, Ammotragus lervia, and Dipodillus amoenus. North 
African species in Egypt which are not known to occur east of the 
Nile Valley and Delta are Paraechinus deserti, Gazella leptoceros, 
Poecilictis libyca, Dipodillus simoni, D. campestris, and Meriones 
shawl. In addition, Pachyuromys duprasi and Allactaga tetradac- 
tyla have limited distributions in the northern limits of the Sahara. 
Allactaga euphratica, a species similar to A. tetradactyla, is widely 
distributed in Southwest Asia. 

Saharo-Sindian species of limited distribution that occur in 
Egypt and/or Sinai are Meriones sacramenti in Sinai, Israel, and 
Jordan; Sekeetamys calurus in the Eastern Desert, Sinai, 
southeastern Israel, and Jordan; and Acomys russatus in the 
Eastern Desert, Sinai, Israel, and parts of western Saudi Arabia. 

Species of Southwest Asian origin which have penetrated into 
North Africa are: Spalax ehrenbergi, which occurs as relict popula- 



OSBORN&HELMY: MAMMALS OF EGYPT 53 

tions in the Western Mediterranean Coastal Desert of Egypt and 
the C5a'enaican Plateau and Coastal Plain of Libya, and Dipodillus 
dasyurus, which occurs in Sinai and the northern part of the 
Eastern Desert. Paraechinus dorsalis is known from Sinai. 

Although the Saharo-Sindian Deserts are a barrier between 
Ethiopian and Palearctic faunas, numerous wide-ranging species of 
Oriental, Palearctic, or Ethiopian origin contain desert-adapted 
subspecies in this Sub-Region. Those that occur in Egypt are 
Hemiechinus auritus, Canis aureus, Vulpes vulpes, Hyaena hyaena. 
Caracal caracal, Felis sylvestris, F. chaus, Panthera pardus, Genetta 
genetta, Procavia capensis, Lepus capensis, Eliomys quercinus, 
Acomys cahirinus, and Hystrix indica. Some species in this 
category, such as Herpestes ichneuman and Mustela nivalis, are 
restricted to river valleys or other areas where water is available. 

A few species of known Ethiopian origin have penetrated Egypt 
via the Nile Valley: Crocidura flavescens, C. nana, and Arvicanthus 
niloticus. Two desert-adapted species of Ethiopian origin, Ictonyx 
striatus and Proteles cristatus, have been collected in southeastern 
Egypt. 

A few relicts of Palearctic origin are Crocidura suaveolens in Sinai 
and the Western Mediterranean Coastal Desert and Suncus 
etruscus in the Nile Delta. Nesokia indica, of Oriental or Indian 
origin, occurs sporadically in Egypt and the Arabian Peninsula. 
Suncus murinus, another Oriental form and a commensal of man, is 
known from the port city of Suez. Its distribution in the Arabian 
Peninsula (Harrison, 1964) indicates transportation by man. Mus 
musculus, Rattus rattus, and R. norvegicus no doubt arrived in 
Egypt and other parts of the Saharo-Sindian Region via man. 

Special Adaptations of Desert Mammals.— The enlarged tym- 
panic and mastoid chambers of the middle ear of certain desert 
rodents, foxes, cats, and weasels, and probably the swollen 
pterygoids in hedgehogs, increase their ability to hear low- 
frequency sounds in warm dry air, which has poor sound-carrying 
quahties (Legouix et al., 1954; Wisner et al., 1954; Vaughan, 1972; 
Harrison, 1972), or vibrations in densely packed sand. Lay's (1972) 
data show a high correlation between specialization in audio- 
sensitivity and environmental aridity. Beecher (1969) suggested 
that, in addition to promoting better hearing, air trapped in the 
sinuses of the middle ear provided an accessory motion sense in 
saltatorial forms. 



54 FIELDIANA: ZOOLOGY 

Soles of the feet of sand-dwelling mammals are usually covered 
with hair. This characteristic is thought to be advantageous for 
movement on sand. Hairy-footed rodents never penetrate the soft, 
salty, clinging soils of salt marshes, as do the bare-footed dipodils; 
nor do they climb rock walls and leap from boulder to boulder, like 
the latter and the agile spiny mice. However, bare-footed dipodils 
and gerbils with partially furred soles (e.g., Dipodillus campestris, 
D. henleyi, D. amoenus, Meriones sp., and Psammomys obesus) are 
commonly found on sandy substrates. 

The paleness of desert mammals appears not to be primarily pro- 
tective, but a result of the physiological effect of dry heat on the 
development of pigments (Bodenheimer, 1935). 

The physical, behavioral, and physiological adaptations of desert 
mammals which enable them to avoid dehydration and the ability of 
some rodents to metabolize water are well known (Schmidt-Nielsen 
and Schmidt-Nielsen 1949 et seq.; Cloudsley-Thompson, 1954; 
Cloudsley-Thompson and Chadwick, 1964; Schmidt-Nielsen, K., 
1964; Harrison, 1964; Mountfort, 1965; Hills, 1966; McGinnies et 
al., 1968; Vaughan, 1972; Prakash and Ghosh, 1975). Further 
discussion herein on the subject are in sections on climate, dew, 
water and desert life, and under individual mammal species. 



SYNOPSES OF REPRESENTATIVES 
IN ORDERS OF RECENT EGYPTIAN LAND MAMMALS 

Order I. I nsectivora.— Hedgehogs and Shrews. Pelage spinous or 
soft. Muzzle forming flexible snout beyond mouth. Feet plan- 
tigrade, palm and sole naked, toes usually five. Tympanic bone 
annular, not fused to skull. First upper and lower incisors pro- 
truding, canines unmodified, molars tuberculosectorial (crushing- 
piercing). Two families in Egypt, p. 57. 

Order II. Chiroptera.— Bats and Flying Mammals. Membranes 
extending between elongated fingers, fore limbs and body, legs and 
tail. The most recent treatments of this group in Egypt are Sanborn 
and Hoogstraal (1955), Hoogstraal (1962), and Zein el Din and Hafez 
(1959). Additional references are Wassif (1946b, 1948, 1949, 1950, 
1951, 1953b, 1960ab, 1962, 1971), Wassif and Madkour (1963, 
1969a, 1969b, 1972abc), Madkour (1961), De Blase (1972), and 
Gaisler et al. (1972, 1973). 

Order III. Lagomorpha.— Hares. Pelage very soft. Ear extremely 
long, lower margin of external opening above level of crown of head. 
Hind limb considerably longer than fore limb in Egyptian forms. 
Tail short. Feet digitigrade, four functional toes, one vestigial. 
Claws, palm, and sole concealed by fur. Skull rodent-like, except for 
fenestrated facial portion of maxilla. Incisive foramina extremely 
long, bony palate very short, postorbital process broad and 
triangular in outline. Incisors continuously growing; two upper 
pairs and one lower. Canines absent, long diastema between incisors 
and premolars. Check teeth six above, five below; occlusal surfaces 
elliptical. One family and one genus in Egypt, p. 84. 

Order IV. Rodentia.— Mice, Rats, Jerboas, Jirds, Gerbils, etc. 
Pelage usually soft, spinous in two genera (Acomys and Hystrix). 
Feet plantigrade to unguiculate; toes usually five, number reduced 
in Dipodidae; palm and sole naked or haired. One pair only of upper 
and lower continuously growing, chisel-shaped incisors. Canines 
absent. Long diastema between incisors and molariform teeth. 

55 



56 FIELDIANA: ZOOLOGY 

Check teeth three-four, upper and lower; cusp pattern variable, Six 
families in Egypt, p. 94. 

Order V. Carnivora.— Foxes, Jackals, Cats, Hyenas, Weasels, 
Civets, etc. Great diversity of external form. Feet plantigrade to 
digitigrade, toes four or five. Mandible with condyle transversely 
elongate. Three pairs of upper and lower incisors. Canines long, 
slightly recurved, pointed; p^ and m, modified as camassials or 
flesh-cutting teeth. Five families in Egypt, p. 359. 

Order VI. Hyracoidea.— Hyraxes. Size and appearance hare-like 
except for small ear and absence of tail. Feet broad, plantigrade; 
fore foot four-toed, hind foot three-toed; nails flat except for curved 
claw on inner hind toe. One pair of tusk-like continuously growing 
upper incisors separated by a gap, triangular in cross section, tips 
pointed. Two pairs of rooted lower incisors, chisel-shaped with edges 
pectinate. Canines absent. Long diastema between upper incisors 
and premolars. Premolars and molars dissimilar. Check teeth seven, 
crushing-type with low cusps and ridges. One family and one genus 
in Egypt, p. 460. 

Order VII. Perissodactyla.— Asses. Feet unguligrade with single 
functional toe-bearing hoof. Remnants of metacarpals and metatar- 
sals two and four as splint bones. Preorbital part of skull and 
diastema elongate. Upper canine small and in diastema between in- 
cisors and premolars. Lower canine incisiform. Premolars and 
molars alike, hypsodont, crowns squarish, complexly ridged. One 
family and one genus in Egypt, p. 470. 

Order VIII. Artiodactyla.— Pigs, Hippopotamuses, Oryxes, Ad- 
daxes, Hartebeests, Gazelles, Ibex, and Barbary Sheep. Feet 
unguligrade, usually with two functional and two lateral rudimen- 
tary digits bearing hoofs; sometimes with four functional toes. 
Horns present in both sexes in family Bovidae. Upper incisors and 
canines absent in Bovidae. Lower canines usually incisor-like; upper 
canines may be modified. Diastema usually present between lower 
canines and premolars. Premolars simpler than molars. Cheek teeth 
five-seven, bunodont to selenodont, brachydont to hypsodont. 
Three families known to have occurred in Egypt, p. 475. 



ORDER INSECTIVORA 

Key to Families of Egyptian Insectivores 

1. Size large, head and body length average 200 mm. Pelage spinous. Ear large. 

Zygomatic arch complete Family 1. Erinaceidae, p. 57. 

2. Size smaller, head and body length 44-1 18 mm. Pelage soft. Ear small. Zygomatic 

arch incomplete Family 2. Soricidae. p. 71. 

Family 1, Erinaceidae 

Hedgehogs. Large insectivores, head and body length average 
200 mm. Pelage spinous. Ear large. Zygomatic arch complete, para- 
pterygoid plate and fossa present, tympanic bulla partially 
developed, mandible with one condylar articulation. First upper in- 
cisor caniniform, m' and m^ with four subequal cusps. Dental for- 
mula: i, I, ¥. fx 2=34 or 36. 

Key to Egyptian Genera of Erinaceidae 

1. Spines on forehead not parted. Belly white. Pterygoid and bulla normal 

Hemiechinus, p. 57. 

2. Spines on forehead parted. Belly not all white. Pterygoid inflated, cavity com- 

municating with bulla Paraechinus, p. 64. 

Genus Hemiechinus Fitzinger, 1866 

Ears proportionately large. Forehead without bare gap in spines. 
Tips of dorsal spines pale. Face pale brown, belly and feet whitish. 
Pterygoid fossa not invaded by tympanic cavity. First upper in- 
cisors proodont. Dental formula: I, \, |, |x 2=36. 

Hemiechinus auritus (Gmelin, 1770) 

Erinaceus auritus Gmelin, 1770, Nov. Comment. Acad. Sci. Petropoli, 14, p. 519. 

Type locality. -V.S.S.R.: ASTRAKHAN. 

General distribution.— Mongolia, Chinese Turkestan, India, 
Afghanistan, Iran, Iraq, Ukranian S.S.R., Transcaspia, Trans- 
caucasia, southern U.S.S.R., Asian Turkey, Cyprus, northern parts 



57 




68 



OSBORN & HELMY: MAMMALS OF EGYPT 59 

of Arabian Peninsula, Syria, Jordan, Israel, Sinai Peninsula, Egypt, 
Libya, 

Common names.— hong Eared Hedgehog, Qunfid, Abu Ghunfis. 

Distribution of subspecies in Egypt— Figure 23. Hemiechinus 
auritus aegyptius: Northern Sinai Peninsula, northern part of 
Eastern Desert, Nile Delta, Nile Valley south to Samalut, and El 
Faiyum; Hemiechinus auritus libycus: Wadi el Natroun and 
Western Mediterranean Coastal Desert. 

Dm^nos/s.— Small hedgehog with long ears. Face pale brown, 
belly white. Tips of dorsal spines white. Gap in spines of forehead 
lacking. Pterygoid not inflated nor communicating with tympanic 
cavity. First upper incisors proodont. 

Adult head and body length average 181 mm., tail 24 mm., foot 35 
mm., ear 41 mm., skull length 44.7 mm. 

External characters.— Dorsal spines white tipped, with buffy ter- 
minal band, and brownish bases. Face and forehead pale brown. 
Venter white. Feet whitish to pale brown. Ear large, whitish. Gap in 
forehead spines lacking. 

Cranial characters.— Figure 24. Premaxillary and frontal bones 
never in contact. Postpalatal bridge straight. Pterygoid not in- 
flated. Post-glenoid process of squamosal and mastoid process sube- 
qual, not inflated. 

Dental characters.— First upper incisors proodont. Last lower 
premolar lacking medial tubercle, but with posterior accessory cusp 
present in 62 per cent of specimens examined. 

Measurements.— Table 2. Male and female dimensions are sub- 
equal. Flower (1932) gave the weights of two specimens as 0.4 and 
0.5 kg. 

Comparisons.— Hemiechinus auritus is distinguishable from 
other Egyptian hedgehogs by smaller dimensions (tables 2, 3), paler 
color, lack of a gap in forehead spines, proodont first upper incisors, 
and pterygoid inflation nil. 

Variation.— Specimens from the Nile Delta and Valley and El 
Faiyum are generally darker than those from desert localities. A 
specimen from northern Sinai is paler than any from the Eastern 
Desert. Specimens from the Western Mediterranean Coastal Desert 
have the auditory bulla slightly more inflated. An accessory 






Fig. 24. Skull of Hemiechinus auritus. 



60 



OSBORN&HELMY: MAMMALS OF EGYPT 61 

Table 2. — Means (and ranges) of measurements followed by number of specimens 
of adult Hemiechinus auritus. 

H. a. aegyptius H. a. libycus 

HBL 179.1 (156-206) 35 182.6 (136-206) 19 

TL 24.6(18-39)36 21.8(15-26)19 

FL 35.9 (28-39) 37 34.2 (32-38) 19 

EL 41.0(34-45)37 41.0(38-45)19 

CBL 44.6(42.4-46.7)32 45.0(42.2-47.5)19 

ZW 26.5(24.1-29.7)32 25.9(24.7-27.8)18 

POW 11.0(10.2-11.7)33 11.3(10.5-11.8)19 

RW 9.8(9.1-10.7)33 10.0(9.4-10.7)19 

BCW 21.3(20.1-22.8)33 21.2(20.2-22.5)19 

NL 14.8(12.4-16.3)33 15.9(14.1-17.5)18 

PPF 6.2 (5.0-6.8) 33 6.6 (5.5-7.2) 19 

SH 11.6(10.6-12.5)32 11.8(10.8-12.4)19 

Table 3. — Means (and ranges) of measurements and weight of Paraechinus deserti, 
P. aethiopicus, and P. dorsalis. 





P. d. deserti 


P. a. aethiopicus 


P. d. dorsalis 


HBL 


215.0(192-226)5 


196.1 (169-217) 7 


199.0(182-228) 10 


TL 


21.6 (19-29) 5 


19.1 (15-22)7 


20.6 (13-30) 9 


FL 


33.4 (31-35) 5 


32.8 (30-37) 7 


34.4 (32-36) 10 


EL 


50.8 (45-58) 5 


43.6 (41-45) 7 


49.8 (44-55) 7 


Wt 


285. 380 


500 





CBL 


50.2 (49.0-52.2) 3 


47.0(46.3-48.1)3 


49.8 (48.1-53.3) 10 


ZW 


30.4(30.1-30.6)3 


26.8. 29.3 


28.9 (27.9-30.0) 9 


POW 


11.8(11.5-12.1)3 


10.3(10.0-10.5)4 


11.3(10.9-12.1) 10 


BCW 


27.5 (26.9-27.9) 3 


25.6 (24.7-26.8) 3 


24.9 (23.7-26.2) 10 


RW 


11.5(11.2-12.0)3 


10.8(10.3-11.4)4 


10.8(10.1-11.9) 10 


NL 


14.0, 15.7 


16.5(14.4-17.8)4 


17.1 (15.9-18.2)9 


PPF 


6.4 (6.2-6.6) 3 


5.8(5.4-6.1)4 


6.0(5.0-7.1) 10 


SH 


13.5(13.0-14.2)3 


12.0(11.5-12.4)4 


12.4(11.7-13.3) 10 



posterior cusp on pm2 occurred in 28 (70 per cent) of 40 specimens of 
H. a. libycus and in 18 (52 per cent) of 34 H. a. aegyptius. 

Collection.— Dug occasionally from shallow burrows, but usually 
found in buildings, crevices in walls, brick and stone piles, small 
caves (Hoogstraal, 1962), graveyards, and burrows of fat sand rats 
(Psammomys obesus) in salt marsh habitat. 

Habitats.— Gardens, olive groves, cultivated areas, and more 
densely vegetated areas of the Coastal Desert (fig. 7). As noted 
above, this hedgehog is everywhere associated with human ac- 
tivities. It is never found in scantily vegetated desert (Hoogstraal, 



62 FIELDIANA: ZOOLOGY 

1962) and is not a true desert hedgehog (Harrison, 1964) like species 
of Paraechinus. 

Reasons for lack of hedgehogs in the Nile Valley south of Samalut, 
as mentioned by Hoogstraal (1962), remain obscure. 

//a6its.— Nocturnal. According to Flower (1932), it is very noisy 
when fighting and "growls" when alarmed or angry. 

Food.— Said to eat grapes in some parts of the Nile Delta (Wassif, 
1953a), thought to be mainly insectivorous (Harrison, 1964) or om- 
nivorous (Hoogstraal, 1962). Laboratory specimens kill and eat 
adult white mice. 

Reproduction.— Flower (1932) Hsted litters of one and two young 
found in May and five in August. 

Sex ratio.— A museum sample of 90 specimens contained 50 (55.5 
per cent) males and 40 females. 

Remarks.— Tv/o species of hedgehogs, H. auritus and Paraechinus 
aethiopicus, have been distinguished from illustrations in tombs of 
Giza Pyramid area, Abu Sir, and Sakkara (Anderson, 1902; Wassif, 
1954b). 

Key to Egyptian Subspecies 
OF Hemiechinus auritus 

1. Color darker, average tail length longer, bulla smaller, mesopterygoid space 

deeper aegyptius, p. 62. 

2. Color paler, average tail length shorter, bulla larger, mesopterygoid space 

shallower Ubycus, p. 63. 

Hemiechinus auritus aegyptius (Fischer, 1829) 

Hemiechinus aegyptius Fischer. 1829, Synopsis Mammalia, p. 262. 

Type locality.— CAIRO: Cairo area. 

Distribution in Egypt— Figure 23. Northern Sinai Peninsula, nor- 
thern part of Eastern Desert, Nile Delta, Nile Valley south to 
Samalut, and El Faiyum. Listing of southern Sinai by Flower (1932) 
was without documentation. 

External characters.— Dorsal color variable but generally darker 
than in Ubycus. Tail averages longer and postpalatal foramen 
shorter than in the latter (table 2). 

Cranial characters. — Bulla slightly less inflated and 
mesopterygoid space slightly deeper than in Ubycus. 



OSBORN&HELMY: MAMMALS OF EGYPT 63 

Dental characters.— Accessory posterior cusp on pm^ was found 
in 52 per cent of aegyptius and 70 per cent of libycus. 

iJemar^s.— Characters listed by Setzer (1957b) as diagnostic of//, 
a. metwallyi ssp. nov. are considered to be too obscure, and all such 
specimens are placed in subspecies aegyptius. 

Material from Sinai Peninsula is considered tentatively to be 
aegyptius. 

Specimens examined.— Total 48. 

SINAI: El Arish(l). 

CAIRO: Cairo 10 km. E (1), Maadi (1). 

DAQAHLIYA: Mit Ghamr (1). 

DAM I ETTA: Damietta (1). 

SHARQIYA: Tel el Kebir (1), Tel Basta (6). Bilbeis (1), Abu Hammad (1). 

KAFR EL SHEIKH: Baltim (2). 

MINUFIYA: Quweisna (1), Dinshawi (1). 

QALYUBIYA: El Barada (1), Kafr Abu Sir (1). Kafr el Shobak (1), El Marq (1). 

BEHEIRA: Damanhour (3). Hafs (1). 

GIZA: Abu Rawash (3), El Kuraimat (3). Mit Riheina (1), Sakkara (2), Minshat el 
Bakkari (1), El Badr Shein (1), Giza Pyramid area (1), El Mansuriya (1), Cairo- 
Alexandria desert road km. 4 (1), Beni Yusef (4). 

BENI SUEF:BeniSuef (1). 

MINYA: Samalut, Qulugan (1). 

EL FAIYUM: Qasr Rashwan (1). SeUa (1). 

Published records.— Records are from Flower (1932), Wassif 
(1953b), Setzer (1957b), and Bodenheimer (1958). 

SINAI: Gaza. El Arish. 

SHARQIYA: Tel Basta, Tel el Kebir. Abu Hammad. 
QALYUBIYA: Kafr el Shobak, Kafr Abu Sir, El Barada, El Marq. 
KAFR EL SHEIKH: El Burg N of Baltim, Baltim. Biyala. 
MINUFIYA: Quweisna, Dinshawi. 
DAQAHLIYA: Mit Ghamr. 
CAIRO: Cairo 10 km. E. Maadi, Helwan. 

GIZA: Kafr Ammar, Giza Pyramid area, Sakkara, El Kuraimat, Kafret el Gabel, 
El Badrshein, Minshat el Bakkari, Minyet el Sultan, Mit Riheina, Abu Rawash. 
EL FAIYUM: Qasr Rashwan. Kom O Shim, SeUa. 
BENI SUEF: Beni Suef. 

Hemichinus auritus libycus (Ehrenberg, 1833). 

Erinaceus libycus Ehrenberg in Hemprich and Ehrenberg, 1833, Symbolae 
Physical Mamm., Dec. 2, sheet k (footnote). 

Type locality.— Desert near Alexandria. 

Distribution in Egypt.— Figure 23. Wadi el Natroun and Western 
Mediterranean Coastal Desert. 



64 FIELDIANA: ZOOLOGY 

External characters.— 'Dorsal color generally paler than in aegyp- 
tius. Tail averages shorter than in the latter and postpalatal 
foramen longer (table 2). 

Cranial characters.— Bulla slightly more inflated and mesoptery- 
goid space slightly shallower than in aegyptius. 

Dental characters. —See under H. a. aegyptius. 

Specimens examined. —Total 46. 

ALEXANDRIA: Ramleh 40 km. W (1). Amiriya (3). 

BEHEIRA: Kom Hamada (1). Wadi el Natroun (2). 

MATRUH: Burg el Arab (14); El Daba (1); Ras el Hekma (1); Mersa Matruh (6). 1.6 
km. E (4). 60 km. W (1); Ageeba (1); Sidi Barrani (7). 1.6 km. S (1). 55 km. W (1); Salum 
4.8 km. E (2). 

Published records.— Records are from Setzer (1957b) and 
Hoogstraal (1962). 

ALEXANDRIA: Ramleh 40 km. W. 

BEHEIRA: El Birigat 1 km. W. Wadi el Natroun. 

MATRUH: El Amiriya; Bahig; Burg el Arab; El Daba; Ras el Hekma; Mersa 
Matruh, L6 km. E. 60 km. W; Sidi Barrani, 55 km. W, 1.6 km. S; Salum 66 km. E, 50 
km. E. 

Genus Paraechinus Trouessart, 1879 

Forehead spines divided by bare gap. First upper incisor not proo- 
dont. Tympanic bulla large, cavity communicating with pterygoid. 
Dental formula: |, {, f , fx 2=34 or 36. 

Key to Egyptian Species of Paraechinus 

1. Belly dark medially only. Face partly white. Postpalatal bridge straight. 

a . Tips of dorsal spines pale. Premaxilla contacting frontal. Posterior lower 
premolar with small medial cusp or metaconid. (Western Mediterranean 
Coastal Desert) deserti, p. 64. 

b. Tips of dorsal spines dark. Premaxilla not contacting frontal. Posterior lower 
premolar lacking medial cusp. (Southern part of Eastern Desert) 
aethiopicus, p. 68. 

2. Belly black. Face blackish. Tips of dorsal spines dark. Postpalatal bridge 
V-shaped (Sinai Peninsula) dorsalis, p. 70. 

Paraechinus deserti (Loche, 1858) 

Erinaceus deserti Loche, 1858, Cat. Mamm. Oiseaux Algerie. p. 20. 

Type locality .— Algeria: Oases of Beni Mzab, Ouargla, and 
Tuggurt. 

General distribution.— Egypt, Libya, Tunisia, Algeria, and 
Morocco. 



OSBORN&HELMY: MAMMALS OF EGYPT 65 

,. 2 5* 2 6* 2 7* 28* 2 9* 30* 31* 3 2* 3 3' 34* 35* 36* 37* 




Fig. 25. Collection localities oi Paraechinus deserti deserti (circles), P. aethiopicus 
aethiopicus (dots), and P. dorsalis dorsalis (squares). 

Common name.— Desert Hedgehog. 
Subspecies in Egypt — 

Paraechinus deserti deserti (Loche, 1858) 
Type locality.— See under species. 

Distribution in Egypt— Figure 25. Northwestern margin of Nile 
Delta and Western Mediterranean Coastal Desert. 

Diagnosis.— Tips of dorsal spines pale. Face partly white. Large 
brown area on belly. Parapterygoid slightly inflated, fossa deep. 
Posterior lower premolar with small medial cusp. 

Head and body length average 215 mm., tail 22 mm., foot 33 mm., 
ear 50 mm., skull length 50.2 mm. 



66 



FIELDIANA: ZOOLOGY 




Fig. 26. Ventral view of Paraechinus aethiopicus aethiopicus. 



External characters.— Dorsal spines with terminal band white (3 
to 4 mm,), subterminal band blackish (4 to 5 mm.), basal bands 
white (4 to 6 mm.) and grayish (5 to 7 mm.) (Setzer, 1957b). Face 
brownish anteriorly, frontal area whitish. Chin and throat white, 
belly white with brown patches. Ear, legs, and feet brownish. 

Cranial characters.— Figure 27. Nasal processes of frontal and 
premaxilla usually in contact. Postpalatal bridge straight to slight- 
ly wing-shaped. Bulla moderately inflated, parapterygoid slightly 
swollen; cavities in communication. Parapterygoid fossa deep. 

Teeth.— Posterior lower premolar (p.2) with small medial cusp. 






Fig. 27. Skull of Paraechinus deserti deserti. 



67 



68 FIELDIANA: ZOOLOGY 

Measurements.— Table 3. Male and female dimensions appear to 
be subequal. 

Variation. —Considerable variation with age occurs in the swelling 
of bulla and parapterygoid. 

/?emar/js.— Smaller dimensions, together with less inflation of 
parapterygoid and auditory bulla were used as characters by Setzer 
(1957b) in describing P. d. wassifi subsp. nov. Unfortunately, he was 
unaware of the fact that inflation in these structures increases with 
age. Further comparison between specimens of comparable ages in- 
dicated that all specimens previously recorded as P. d. wassifi 
(Setzer, 1957b; Hoogstraal, 1962) are P. d. deserti. 

Paraechinus d. deserti is rare and localized in Egypt (Hoogstraal, 
1962). 

Comparisons.— Paraechinus d. deserti differs from other Egyp- 
tian paraechine hedgehogs in having dorsal spines with whitish tips 
instead of brown, nasal processes of frontal and premaxilla in con- 
tact, less inflated pterygoids, smaller bulla, and a medial cusp on p2. 

Specimens examined. —Total eight. 

BEHEIRA: El Birigat 3 km. W (2). 

MATRUH: El Hammam 20 km. S (1); Qara road E of Mersa Matruh (1); Sidi 
Barrani 33 km. W (1); Salum (1), 66 km. E (1), 6.6 km. E (1). 

Published records.— Records are from Wassif (1954b), Setzer 
(1957b), and Hoogstraal (1962). 

BEHEIRA: El Birigat 3 km. W. 

MATRUH: Sidi Barrani 33 km. W; Salum. 66 km. E. 6.6 km. E. 

//afeitats.— Collected from a burrow in barren sandy-clay hills and 
barren desert west of El Birigat (Wassif, 1953a), vegetated areas of 
the Western Mediterranean Coastal Desert (fig. 7), and a garden in 
Salum. 

//a6i(s.— Nocturnal. Very little is known about this hedgehog. 

Reproduction.— V/assif (1953a) found a lactating female in June. 

Paraechinus aethiopicus (Ehrenberg, 1833) 

Erinaceus aethiopicus Ehrenberg in Hemprich and Ehrenberg, 1833, Symbolae 
Physical Mamm., Dec. 2, sheet k (footnote). 

Type locality. -Sudan. NORTHERN: Dongola Desert. 

General distribution.— Egypt, Sudan, Eritrea, Somalia, Asben, 
Algeria, Morocco, and Mauretania. 



OSBORN & HELMY: MAMMALS OF EGYPT 69 

Common name.— Ethiopian Hedgehog. 

Subspecies in Egypt— 

Paraechinus aethiopicus aethiopicus (Ehrenberg, 1833) 

Type locality. —See under species. 

Distribution in Egypt— Fig\ire 25. Southern part of Eastern 
desert. 

Diagnosis. —Tips of dorsal spines dark. Face partly white. Central 
part of belly dark brown. Feet and legs dark brown. Parapterygoid 
strongly inflated, fossa obliterated. 

Head and body length average 196 mm., tail 19 mm., foot 32 mm., 
ear 44 mm., skull length 47.0 mm. 

External characters.— Figure 26. Dorsal spines blackish-brown 
with light brown tips. Anterior of face, lower lip, chin, large area on 
throat and center of belly, genital region, legs, and outside of ear 
dark brown. Frontal area of head and most of throat and belly 
white. Inside of ear white in some individuals. 

Cranial characters.— Nasal processes of frontal and premaxillary 
widely separated. Postpalatal bridge straight. Bulla enormously in- 
flated, alisphenoid and parapterygoid hollow and inflated. 
Parapterygoid fossa obliterated. 

TeetA. — Posterior lower molar without small medial cusp. 

Measurements.— Table 3. Male and female dimensions are sube- 
qual. 

Comparisons.— Paraechinus aethiopicus differs from other Egyp- 
tian paraechine hedgehogs in having generally smaller average 
dimensions, particularly ear length (table 3). In color, aethiopicus is 
darker than deserti, paler than dorsalis. The postpalatal margin is 
straight in aethiopicus, V-shaped in dorsalis. Comparisons with 
deserti are under the latter. 

Specimens examined.— Total nine. 

REDSEA: Wadi Naam (1). 

SUDAN ADMINISTRATIVE: Wadi Akwamtra (1); Bir Kansisrob (2), 1.6 km. N 
(1). 5 km. N (2); Wadi Kansisrob (1); Wadi AUaqi. about 22° N lat.. 35° E long. (1). 

Published records.— Records are from Setzer (1957b), Hoogstraal 
et al. (1957b), and Hoogstraal (1962). 

RED SEA: Wadi Naam near Bir Abraq. 



70 FIELDIANA: ZOOLOGY 

SUDAN ADMINISTRATIVE: Bir Kansisrob 1.6 to 5 km. N. Wadi Abu SaaU 
(remains), summit of Gebel Elba (remains). 

Habitats.— Collected in burrows under dense stands of vegetation 
or large shrubs on coastal plain and in rocky wadis. 

//a6i(s.— Nocturnal. Very little is known about this species, par- 
ticularly in Egypt. 

Paraechinus dorsalis (Anderson and De Winton, 1901) 

Erinaceus dorsalis Anderson and De Winton, 1901, Ann. Mag. Nat. Hist., (ser. 7), 
7, p. 42. 

Type locality.— Saudi Arabia: Hadramaut. 

General distribution. — Iraq, Saudi Arabia, Lebanon, Jordan, 
Israel, and Sinai Peninsula. 

Common name.— Desert Hedgehog. 

Subspecies in Egypt.— 

Paraechinus dorsalis dorsalis (Anderson and De Winton, 1901) 

Type locality. —See under species. 

Distribution in Egypt.— Figure 25. Sinai Peninsula. 

Diagnosis.— Dorsal spines blackish. Face, belly, and limbs 
blackish. White mark on throat. Postpalatal bridge V-shaped. Para- 
pterygoid inflated, fossa obliterated. 

Head and body length average 199 mm., tail 20 mm., foot 34 mm., 
ear 50 mm., skull length 49.8 mm. 

External characters.— Dorsal spines blackish, flank spines with 
brownish tips. Face and forehead all black. Chin and throat black, 
latter with white band sometimes extending to base of ear. Venter 
and Umbs black. 

Cranial characters.— Nasal processes of premaxilla and frontal 
sometimes in contact. Postpalatal bridge V-shaped. Bulla greatly 
inflated, alisphenoid and parapterygoid hollow and inflated. 
Parapterygoid fossa obHterated by inflation. 

Tee^A. — Posterior lower molar with vestigial medial cusp. Second 
upper premolar reduced or absent. 

Measurements.— Table 3. Male and female dimensions are 
subequal. 

Comparisons.— Paraechinus dorsalis differs from other Egyptian 



OSBORN&HELMY: MAMMALS OF EGYPT 71 

hedgehogs in having color generally darker, white areas smaller, 
postpalatal margin V-shaped, bulla more inflated. Pterygoids are, 
incidentally, slightly less inflated than those in P. aethiopicus. 

Specimens examined.— TotaX 10. 

SINAI: Wadi el Sheikh (2), St. Catherine Monastery area (5). Wadi Raha (2). 
Feiran Oasis 1.6 km. N (1). 

Published records.— Records are from Wassif and Hoogstraal 
(1954), Setzer (1957b), and Hoogstraal (1962). 

SINAI: St. Catherine Monastery and vicinity, Wadi Raha, Feiran Oasis 1.6 km. 

N. 

Habitats.— Occurs from lowland to high elevations in Sinai Penin- 
sula and in gardens near St. Catherine Monastery. 

//a6its.— Nocturnal. Information on this species is meager. 

Family 2. Soricidae 

Shrews. Small to large insectivores, head and body length averag- 
ing 44 to 118 mm. Pelage soft. Ear small. Zygomatic arch in- 
complete, parapterygoid plate and fossa absent, tympanic bulla ab- 
sent, mandible with two condylar articulations. First upper incisor 
two cusped, m^ and m^ with cusps of unequal height. Dental for- 
mula: i, 5, T. fx 2=28 or 30. 

Key to Egyptian Genera of Soricidae 

1. Upper unicuspids 3-3 Crocidura, p. 71. 

2. Upper unicuspids usually 4-4 Suncus, p. 80. 

Genus Crocidura Wagler, 1832 

White toothed shrews. Fur short, dense. Ear scarcely protruding 
beyond fur. Tail with conspicuous and scattered bristle hairs. Teeth 
unpigmented. Anterior upper incisor with main cusp long, slender, 
and hooked downward; basal lobe less than one-half height of 
anterior lobe. Anterior lower incisor without lobes on cutting edge, 
proodont. Upper unicuspid teeth three. Dental formula: 2. 6. 1. 
|x2=28. 

Key to Egyptian Species of Crocidura 

1. Larger shrews. Head and body length 70 mm. or more, condyloincisive length 15 
mm. or more. 

a. Bristle hairs on proximal two-thirds of tail. Venter dark gray. Hind foot 18 mm. 
or more flavescens, p. 73. 

b. Bristle hairs along entire length of tail. Venter whitish. Hind foot less than 13 
mm suaveolens, p. 78. 




72 



OSBORN&HELMY: MAMMALS OF EGYPT 73 

2. Smaller shrews. Head and body length 72 mm. or less, condyloincisive length 20 
mm. or less. 

a. Bristle hairs of tail inconspicuous, sparse, Umited to proximal one-half. Dorsum 
brownish. Cranium convex. Hind foot length 12 mm. or more. . . . floweri, p. 76. 

b. Bristle hairs of tail conspicuous, numerous. Dorsum grayish. Cranium flat, 
very small. Hind foot length 10 mm. or less nana, p. 77. 

Crocidura flavescens (I. Geoffroy St.-Hilaire, 1827) 

Sorex flavescens I. Geoffroy St.-Hilaire, 1827. Diet. Class. Hist. Nat., p. 324. 
Crocidura olivieri Lesson, 1827, Man. Mamm., p. 127. 

Type locality.— South Africa, "Le Cafr^rie et le pays des Hotten- 
tots" (I. Geoffroy St.-Hilaire, 1827), Eastern CAPE PROVINCE: 
King Williams Town (Roberts, 1951). 

General distribution.— Egypt, Sudan, Ethiopia, and the rest of 
Africa south into South Africa; west Africa north to Sierra Leone. 

Common names:— Giant Musk Shrew, Far, Ersa. 

Subspecies in Egypt — 

Crocidura flavescens deltae Heim de Balsac and Barloy, 1966 

Crocidura flavescens deltae Heim de Balsac and Barloy, 1966, Mammalia, 30, p. 
631. 

Type locality.— Egypt. No exact locality. 

Distribution in Egypt— Figure 28. Nile Delta and Valley as far 
south as Dahshur and in El Faiyum. 

Diagnosis.— Largest of Egyptian species of Crocidura. Dorsum 
dark brown, venter grayish. Tail slightly darker than dorsum, 
unicolored; bristles on proximal two-thirds. 

Third upper unicuspid larger in transverse section than second. 

Adult head and body length average 118 mm.; tail 71 mm., 60 per 
cent of head and body length; foot 20 mm.; ear 12 mm.; condyloin- 
cisive length 28 mm. 

External characters.— Dorsum uniform dark brown with silvery 
sheen shading on lower side to dark gray venter. Some individuals 
with brownish throat patch. Feet grayish to brown. Tail dark 
brown, unicolored; bristles on proximal two-thirds. Scent gland on 
side with whitish hairs. Ear large, almost naked, with pronounced 
ventral fold. 

Cranial characters.— Figure 29. Cranium normal, much larger and 
heavier than in other Egyptian shrews. 

Teeth.— Third upper unicuspid larger in transverse section than 





E 

U 




Fu; 29. Skull of Cmcidura flavescens deltae. 



74 



OSBORN&HELMY: MAMMALS OF EGYPT 75 

Table 4. — Means (and ranges) of measurements and ratios of species of Crocidura. 

C. floweri C. flavescens C. nana 

HBL (57-71)* 117.8(106-135)27 53.8(48-62)8 

TL (55-58)* 71.2(57-84)27 34.8(28-40)8 

TL/HBL% 75* 60.6 (45.6-69.0) 27 65.2 (61.0-75.0) 8 

FL (12-13.5)* 20.4(18-22)27 9.4(8.5-10.0)8 

EL 8* 12.8(11-14)25 7.0(5.5-9.0)7 

CIL 18.4(18.0-19.2)4 27.9(26.0-29.6)26 15.3(14.4-16.1)8 

RW 5.4 (5.2-5.6) 4 8.8 (8.3-9.3) 28 4.5 (4.3-4.8) 13 

BCW 8.0(7.8-8.4)4 11.9(11.2-12.4)26 6.8(6.6-7.1)12 

POW 3.8 (3.7-4.0) 4 5.2 (5.1-5.5) 24 3.2 (3.0-3.5) 13 

TRL 7.8(7.6-8.1)4 12.4(11.3-13.2)28 6.2(5.5-6.7)12 

PL 7.4(7.3-7.5)4 12.0(10.8-12.5)18 6.4(5.8-7.2)9 

SH 4.2(4.1-4.3)4 6.6(6.0-7.5)24 3.4(3.2-3.7)6 
*DaU from Flower (1932). 

second. Basal cusp of anterior upper incisor very small. Setzer 
(1957a) discovered a supernumerary molarifom tooth in a specimen 
from Abu Ghalib. 

Measurements.— Table 4. 

Comparisons.— Crocidura flavescens is the largest of Egyptian 
shrews. The subspecies deltae averages longer in head and body 
length and shorter in tail length than most other African races. 

Remarks.— YAlervciSin et al. (1953) tentatively united all the larger 
forms of southern and eastern Africa under C. flavescens and includ- 
ed olivieri as an outlying race. Setzer (1957b) doubted the correct- 
ness of these assignments. Heim de Balsac and Barloy (1966) 
substantiated the conclusions of EUerman et al., but proposed the 
Egyptian form be called C. /. deltae, as olivieri was inappropriate 
due to having been applied to a diversity of mummified specimens 
of unknown origin. 

Specimens examined.— TotsA. 104. 

DAMIETTA: Damietta (3). 

KAFR EL SHEIKH: Baltim (1). 1 km. S (1). 

MINUFIYA: Mit Faris (1). 

GIZA: Mena House area (6). Tanash (6), El Baragil (4). Abu Rawash (18). Abu 
Ghalib (9), Nahya (11). Wardan (8), Kirdasa (4). Giza Pyramid area (1). Minshat el 
Bakkari (1). Kafr Hakim (1). El Mansuriya (10), Manshiyet Radwan (1). Talbia (1), 
Saft el Laban (3). El Qatta (1), Birqash (1). El Badrshein (1). Dahshur (2). 

EL FAI YUM: Minshat Beni Osman (3). Kom O Shim (2). Royal Shooting Club (4). 

Published records.— Records are from Anderson (1902), Bonhote 
(1909), Flower (1932), Setzer (1952, 1957b). 



76 FIELDIANA: ZOOLOGY 

GIZA: Mena House area (6), Tanash (6). El Baragil (4), Abu Rawash (18), Abu 
Ghalib (9), Nahya (11), Wardan (8), Kirdasa (4), Giza Pyramid area (1), Minshat el 
Bakkari (1), Kafr Hakim (1). El Mansuriya (10), Manshiyet Radwan (1). Talbia (1). 
Saft el Laban (3), El QatU (1). Birqash (1). El Badrshein (1). Dahshur (2). 

EL FAI YUM: Minshat Beni Osman (3). Kom O Shim (2), Royal Shooting Club (4). 

Habitats. — ''Richiy weeded canal margins in Nile Delta and 
Valley" (Hoogstraal, 1962, p. 150); dry wells in summer. A specimen 
from Zagazig "fell from the talons of an eagle owl shot at by Dr. 
Walter Innes" (Anderson, 1902, p. 167). Many have been collected 
in cultivated fields (fig. 6). 

Nests.— Nests are balls of grass and always moist (Hoogstraal, 
1962). 

Food— Nests sometimes contain insect remains and broken snail 
shells (Hoogstraal, 1962). Captive shrews kill and eat adult albino 
Mus musculus. 

Crocidura floweri Dollman, 1915 

Crocidura floweri, Dollman, 1915. Ann. Mag. Nat. Hist., (ser. 8), 15, p. 515 (Actual 
description in 1916 ibid., (ser. 8). 17, p. 192). 

Type locality.— Egypt. GIZA: Giza. 

Distribution.— Figure 28. Nile Delta and El Faiyum. 

Common name.— Flower's shrew. 

Diagnosis. -Small shrew. Dorsum pale cinnamon brown. Venter 
whitish. Tail bicolored, bristles scattered along proximal one-half. 
Skull convex. Second and third unicuspids subequal; third overlap- 
ping second. Head and body length ranges from 57 to 71 mm.; tail 55 
to 58 mm., 75 per cent or more of head and body length; foot 12 to 
13.5 mm., ear 8 mm.; and condyloincisive length 17.1 to 19.2 mm. 

External characters.— Dorsum and side light cinnamon brown, 
flank and belly whitish. Tips of belly hairs whitish, bases gray. Feet 
dirty whitish. Tail bicolored, color of back above, whitish below; 
thinly haired; bristle hairs grayish white, inconspicuous, scattered, 
and confined to proximal one-half. 

Cranial characters. —Skull normal except slightly more convex 
than in other Egyptian shrews. 

Teeth.— Secorxd and third unicuspids subequal, third overlapping 
second. According to Heim de Balsac and Mein (1971, p. 238), the 
molars have a "protometacone union, that is of a metaloph and 
equally that of a metaconule." 



OSBORN&HELMY: MAMMALS OF EGYPT 77 

Measurements.— Table 4. 

Comparisons.— Crocidura floweri is distinguishable from other 
species of Egyptian Crocidura by lack of bristles on distal one-half 
of tail, and second and third unicuspids subequal. 

iJemarAjs.— Practically nothing is known about this shrew. It is 
rare (Setzer, 1957b) and possibly extinct (Hoogstraal, 1962). 
Specimens have been collected in fields. One was from the stomach 
of a cattle egret (Flower, 1932). 

Specimens examined. — Total five. 

GIZA: Giza (Type. 3). 
EL FAIYUM: Faiyum (1). 

Published records.— Records are from Flower (1932) and Heim de 
Balsac and Mein (1971). 

KAFR EL SHEIKH: Baltim. 

GIZA: Giza. 

QENA: Thebes (mummified). 

Crocidura nana Dobson, 1890 

Crocidura nana Dobson, 1890, Ann. Mag. Nat. Hist., (ser. 6), 5, p. 225. 
Sore jc religiosus I. Geoffroy St.-Hilaire, 1827, Mem. Mus. Hist. Nat., Paris, 15, p. 
128. 

Type locality.— Somalia: Dollo. 

General distribution.— Egypt and Eastern Africa from Sudan 
south to Rhodesia. 

Common name.— Dwarf Shrew. 

Distribution in Egypt— Figure 28. Nile Delta and possibly Nile 
Valley. 

Diagnosis.— Size very small. Dorsum grayish with tinge of brown. 
Venter light gray. Tail bicolored, bristles numerous. 

Head and body length average 54 mm.; tail 35 mm., 64 per cent of 
head and body length; foot 10 mm.; ear 7 mm.; condyloincisive 
length 15.4 mm. 

External characters.— Dorsum dull gray with brownish tinge. 
Flank paler. Venter light gray due to hairs with whitish tips and 
gray bases. Chin and throat paler than belly. Feet whitish, almost 
hairless. Tail gray above, whitish below; bristle hairs numerous, 
whitish. 



78 FIELDIANA: ZOOLOGY 

Cranial characters.—SkuW very small, delicate and "flat." 

Measurements.— Table 4. 

Comparisons.— Crocidura nana is distinguishable from other 
species of Egyptian Crocidura on the bases of smaller size, paler 
color, and flatness of skull. 

jRemar^s.— According to Heim de Balsac and Mein (1971, p. 243), 
C nana is synonymous with C religiosa, and the latter name should 
be suppressed because it "did not designate any exact species." 
Crocidura religiosa (I. Geoffroy St.-Hilaire, 1827) was described 
from mummified material. 

Specimens examined.— Total 18. 

QALYUBIYA: Qalyub (1). 

GIZA: Kafr Hakim (1|, Nahya (2). Minshat el Bakkari (1), Abu Rawash (7). Giza (2). 

CAIRO: Cairo (4). 

Published records.— Records are from Flower (1932), Setzer 
(1957b), and Heim de Balsac and Mein (1971). 

GIZA: Abu Rawash, Kafr Hakim, Nahya. 
QENA: Thebes (mummified). 

i?emar/2s. — Found "under stones, bricks, and clumps of earth in 
moist cultivated fields" (Hoogstraal, 1962, p. 150), in canal banks, 
in dry wells, and under piles of grass, cotton, and corn stalks. A nest 
of cotton bolls and small sticks was noted on a label. 

Crocidura suaveolens (Pallas, 1811) 

Sorex suaveolens Pallas, 1811, Zoographia Rosso Asiatica, 1, pi. 9, fig. 2, p. 139. 

Type locality. —Southern Russia. Crimea: Khersomes. 

General distribution. — Korea, Japan, Mongolia, Siberia, China, 
Central and Southern U.S.S.R., Europe, Iran, Iraq, Russian 
Turkestan, Armenian S.S.R., Asian Turkey, Lebanon, Israel, 
Egypt, Algeria, and Morocco. 

Common name.— hesser White Toothed Shrew. 

Distribution of subspecies in Egypt.— Figure 28. Crocidura 
suaveolens portali: Sinai Peninsula; Crocidura suaveolens 
matruhensis: Western Mediterranean Coastal Desert. 

Diagnosis. SrtvaW shrew, tail about one-half length of head and 
body. Dorsum dull brownish gray. Venter and feet whitish. Tail 
bicolored, bristles scattered along entire length. 



OSBORN&HELMY: MAMMALS OF EGYPT 79 

Skull lacking distinctive features. Second upper unicuspid smaller 
than third. 

Head and body length ranges from 55 to 72 mm., tail 25 to 40 
mm., 38 to 60 per cent of head and body length; foot 10 to 12.5 mm.; 
condyloincisive length 16.0 to 17.6 mm. 

External characters.— Dorsum and side dull brownish gray. Hairs 
gray with minute brownish tips. Venter and feet whitish. Tail in- 
distinctly bicolored, grayish or brownish above, paler below. 

Cranial characters.— No distinctive features. 

Teeth.— Large upper premolar with prominent cutting blade and 
paracone well developed. Second upper unicuspid smaller in crown 
area than third. 

Measurements. —See under diagnosis and subspecies. 

Comparisons.— Crocidura suaveolens differs from C. nana in 
larger size. From other Egyptian shrews, it differs in having bristles 
extending along the entire length of tail and the second upper 
unicuspid smaller than third. 

Collection.— Crocidura suaveolens has been found in two 
locahties: inside El Arbaein Monastery near St. Catherine 
Monastery, Sinai Peninsula; and in burrows of fat sand rat (Psam- 
momys obesus) in coastal salt marsh near Mersa Matruh (Wassif 
and Hoogstraal, 1953; Setzer, 1957b, 1960b; Hoogstraal, 1962). 

Habits.— This species appears to be adapted to semidesert condi- 
tions. It is rare and little known (Harrison, 1964). 

Egyptian Subspecies of Crocidura suaveolens 

Crocidura suaveolens portali (Thomas, 1920) 

Crocidura portali Thomas, 1920, Ann. Mag. Nat. Hist., (ser. 9), 5, p. 119. 

Type locality.— Israel: Ramleh. 

Distribution in Egypt— Figure 28. Sinai Peninsula. 

Comparison.— Crocidura s. portali differs from C. s. matruhensis 
in larger body size and longer tail (Setzer, 1960b). 

Collection.— Trapped in old El Arbaein Monastery near St. 
Catherine Monastery, Sinai. 

Measurements.— Measurements from Wassif and Hoogstral 
(1954) of a female specimen are: head and body length 56 mm.; tail 



80 FIELDIANA: ZOOLOGY 

37 mm., about 66 per cent of head and body; foot 10.5 mm.; ear 7 
mm.; occipitonasal length 16.5 mm. 

Specimen examined.— Total one. 

SINAI: El Arbaein Monastery 9 km. W of St. Catherine Monastery (1). 

Published records.— Above-mentioned single specimen is referred 
to in Wassif and Hoogstraal (1953), Setzer (1957b), Hoogstraal 
(1962), and Harrison (1964). 

Crocidura suaveolens matruhensis Setzer, 1960 

Crocidura suaveolens matruhensis Setzer, 1960, J. Egypt. Publ. Health Assn., 35, 
p. 2. 

Type locality.— MATRIJH: Mersa Matruh 4.8 km. W. 

Distribution in Egypt— Known only from type locality 4.8 and 
1.6 km. W of Mersa Matruh (fig. 28). 

Comparisons.— Crocidura s. matruhensis differs from C. s. portali 
of Sinai in smaller body size and shorter tail and from C. s. whitakeri 
of Morocco in proportionately shorter tail, 45 per cent of head and 
body in whitakeri and 38 per cent in matruhensis (Setzer, 1960b). 

Collection.— ''Taken in burrows of fat sand-rat, Psammomys 
obesus, in damp, saline depressions just behind the sea" (Setzer, 
1960b, p. 3). 

Measurements.— Measurements of type, an adult male, are: head 
and body length 63 mm.; tail 25 mm., 38 per cent of head and body; 
foot 11 mm.; ear 9.5 nmi.; condyloincisive length 17.1 mm. (Setzer, 
1960b). 

Specimen examined.— Total one. 

MATRUH: Mersa Matruh 4.8 km. W (1). 

Published records.— Records of above-mentioned specimen and 
another 1.6 km. W of Mersa Matruh are in Setzer (1960b) and 
Hoogstraal (1962). 

Genus Suncus Ehrenberg, 1833 

White-toothed shrews of the genus Suncus are distinguishable 
from Crocidura by presence of small fourth upper unicuspid. Body 
size is minute or very large. Dental formula: I, 5, f, fx 2=30. 

Kky to Eoyitian Spfxiks of Suncus 

1. Size large, head and body length 108-135 mm. Tail thick at base, tapering: not 
bicolored murinus, p. 81. 



OSBORN&HELMY: MAMMALS OF EGYPT 81 

2. Size small, head and body length 40-54 mm. Tail slender, bicolored 
etruscus, p. 82. 

Suncus murinus Linnaeus, 1766 

Suncus murinus Linnaeus, 1766, Syst. Nat., 12th ed., p. 74. 

Type locality.— Java. 

General distribution.— Nev/ Guinea, Java, Sumatra, Borneo, 
Celebes, Philippines, Japan, Taiwan, Southeast China, Indochina, 
Burma, Malay States, Bali, Ceylon, and India; and seaports in Iran, 
Iraq, Oman, Aden, Yemen, Saudi Arabia, Egypt (Suez), Sudan 
(Suakin), and possibly Ethiopia. 

Common name.— House Shrew. 

Subspecies in Egypt — 

Suncus murinus sacer (Ehrenberg, 1833) 

Suncus sacer Ehrenberg, 1833, in Hemprich and Ehrenberg, Symbolae Physicae 
Mamm., Dec. 2, folio k. 

Type locality.— SUEZ: Suez. 

Distribution in Egypt— A single specimen was captured in a 
house in Suez. 

Diagnosis.— Yery large shrew. Tail about one-half length of head 
and body. Dorsum brown, venter grayish, tail color of back with 
silvery bristles along entire length. 

Skull massive in comparison with other shrews and strongly 
ridged. 

Head and body length average 118 mm.; tail 70 mm., 58 per cent 
of head and body length; foot 20 mm.; ear 14 mm.; skull length 32,2 
mm. (data from Harrison, 1964). 

External characters.— Large, rat-sized shrew. Tail thick at base, 
tapering; slightly more than one-half length of head and body. Dor- 
sum grayish brown, hairs brown-tipped with gray base. Venter 
grayish white. Line of demarcation between side and beUy in- 
distinct. Feet whitish. 

Cranial characters. —Skull massive compared with that of other 
shrews and heavily ridged. 

Teeth.— Chief characteristics are large size and presence of four 
instead of three upper unicuspids as in Crocidura. . 

Comparisons.— Suncus murinus is distinguishable from all other 



82 FIELDIANA: ZOOLOGY 

Egyptian shrews by its much larger size, tail being shorter than one- 
half head and body length and thick at base. 

Remarks. —Setzer (1952, p. 345) thought that individual 
specimens of this shrew from North Africa were "merely fortuitous 
travelers come ashore from some trading vessel." He omitted 
discussion of the species (Setzer, 1957b, p. 2), because it is "ap- 
parently only infrequently imported from the Indian region and is 
not a member of the native Egyptian mammal fauna." We are in- 
clined to agree with these conclusions and so does Harrison (1964) 
with regard to S. murinus in the Arabian Peninsula. 

Further notes on characters of S. m. sacer are in Harrison (1964). 
Specimens examined,— None from Egypt. One from Saudi Arabia. 

Co//ec^ion.— Easily trapped with a variety of baits (Sanborn and 
Hoogstraal, 1953). 

Habitats. — Houses and buildings in this area; probably not occur- 
ring in nature. 

//a6i(s.— Commensal, at least in the Arabian Peninsula, and ac- 
tive at any time of day or night (Sanborn and Hoogstraal, 1953; 
Harrison, 1964). 

Suncus etruscus Savi, 1822 

Suncus etruscus Savi, 1822, Nuovo Giorn de Letterati, Pisa, 1, p. 60. 
Type locality. — Italy: Pisa. 

General distribution.— Spain, France, Corsica, Sicily, Sardinia, 
Italy, Yugoslavia, Hungary, Albania, Greece, Crete, Turkey, 
Rhodes, Tiflis, Azerbaijan S.S.R., Turkmen S.S.R., Uzbek, Iran, 
Irak, Israel, Aden, and Egypt (Harrison, 1964; Spitzenberger, 
1970). 

Common names.— Savi 's Dwarf Shrew, Pygmy White Toothed 
Shrew. 

Distribution in Egypt— Nile Delta. 

Diagnosis.— Very tiny shrew. Tail about two-thirds of head and 
body length. Dorsum grayish to light brown, venter whitish. Tail 
bicolored with bristles along entire length. Skull minute, flattened, 
and fragile. 

Head and body length average 46 mm.; tail 26 mm., 64 per cent of 
head and body length; foot 7 mm.; skull length 13.0 mm. (data from 
Harrison, 1964). 



OSBORN&HELMY: MAMMALS OF EGYPT 83 

External characters.— One of the smallest shrews. Resembles a 
dwarf Crocidura suaveolens. Tail thin, not tapering, and about two- 
thirds of head and body length; bicolored, brown above, whitish 
below. Dorsum light brown to grayish, and hairs with brown tips 
and gray bases. Venter whitish, hairs with white tips and gray 
bases. Line of demarcation between side and belly indistinct. Feet 
whitish. 

Cranial characters.— S)sm\\ very small and fragile, lacking ridges, 
and dorsoventrally flattened relative to width. 

Teeth.— Chiei characteristic is the presence of four instead of 
three upper unicuspids as in Crocidura. 

Comparisons.— Suncus etruscus is distinguishable from most 
Egyptian shrews by its small size. It is distinguishable from 
Crocidura nana by having four instead of three upper unicuspids. 

/2emar/js.— Further notes on characters of S. etruscus are in 
Harrison (1964). 

Specimens examined.— None from Egypt. One from Israel. 

Published records.— Heim de Balsac and Lamotte (1957) 
discovered a single specimen from the Nile Delta in the Paris 
Museum. 

Habitats.— Knov/n to live in houses in the Mediterranean region, 
also found in gardens, under stones, and in old walls (Corbet, 1966). 



ORDER LAGOMORPHA 

Family Leporidae 

Genus Lepus Linnaeus, 1758 

Hares. Ear long, prominent. Hind foot long, slender; sole densely 
haired. Tail short, black above, white below. Pelage soft, woolly, 
yellowish to buffy gray. 

Skull elongate, convex dorsally. Palate very short, posterior 
margin opposite PM^. Mesopterygoid space wider than length of 
palatine bridge. Incisive foramen very long, broadened posteriorly. 
Palatal foramen minute. Postorbital process broad and triangular 
with distinct anterior and posterior projections. Two pairs of con- 
tinuously growing upper incisors. Upper tooth rows further apart 
than lower. Dental formula: f, 5, i, |x 2=28. 

Lepus capensis Linnaeus, 1758 

Lepus capensis Linnaeus, 1758, Syst. Nat., 10th ed., p. 58. 

Type locality.— Union of South Africa: Cape of Good Hope. 

General distribution.— Britain; Europe eastward through 
U.S.S.R. into Siberia, Mongolia, and China; Afghanistan, Iran, 
Iraq, Syria, Turkey, Lebanon, Israel, Jordan, Sinai Peninsula, and 
Egypt; Sudan west to Rio de Oro and south into eastern and 
southern Africa. 

Common names.— Hare, Arnab. 

Distribution of subspecies in Egypt.— Figure 30. Lepus capensis 
sinaiticus: Sinai Peninsula; Lepus capensis aegyptius: northern two- 
thirds of Eastern Desert; Lepus capensis isabellinus: southern one- 
third of Eastern Desert; Lepus capensis rothschildi: Western 
Desert. 

Diagnosis.— ¥,ar and hind foot long, tail relatively short. Pelage 
soft, yellowish or buffy gray; tail black above, white below. Skull 
elongate, arched in profile; palate very short; incisive foramen very 



84 



OSBORN & HELMY: MAMMALS OF EGYPT 



85 



,. 2 5* 2 6* 2 7* 28* 2 9* 30* 31* 3 2* 3 3* 3 



4 35 36 37 




(do 
(SI. 



30. Collection localities of Lepus capensis sinaiticus (circlesi, L. c. aegyptius 
L. c. isabellinus (solid squares), L. c. rothschildi (hexagons), and sight records 



long; maxilla and posterior parts of cranium fenestrated. Upper in- 
cisors two pairs, continuously growing; diastema very long; cheek- 
teeth hypsodont, six above, five below. Occlusal surfaces are ellip- 
tical. Adult head and body length average 400 mm.; tail 76 mm., 19 
per cent of head and body length; foot 105 mm; ear 116 mm. 

External characters. — Pelage soft, woolly, yellowish to buffy 
gray. Dorsal hairs with blackish tips, subterminal bands yellowish 
to buffy, and basal bands grading from dark gray to pale gray or 
whitish. Narrow stripe on side and flank of buff-tipped hairs with 
white bases. Venter hairs are pure white except for buff-tipped hair 
of groin. Throat buffy. Circumorbital area whitish. Nape buffy to 
rufous. Ear covered with short hair; outer side brownish, tip 
blackish, base color of nape; inner side whitish, margin cream 



86 FIELDIANA: ZOOLOGY 

colored, with long hair on inner margin; anterior tip sometimes 
blackish or brownish. Tail black above, white below. Legs and feet 
with outer surface buffy, inner white. Hair of palm and sole 
brownish, long, nearly concealing claws. 

Cranial characters.— Figure 31. Skull elongate, convex dorsally. 
Nasals long, broadened posteriorly, and separated by a deep, 
U-shaped frontal extension. Postorbital processes are prominent 
and have anterior and posterior extensions. Supraoccipital sloping 
caudad; external occipital protuberance posterior to level of 
occipital condyle and auditory bulla and bearing superior nuchal 
crest. There is a deep groove between the occipital protuberance and 
petromastoid. The interparietal is sometimes absent. Exoccipital is 
wide and flaring. Paroccipital process long, adnate to bulla. Lateral 
part of maxilla, posterior part of parietal, interparietal, supra- 
occipital, temporal process of parietal, temporal, alisphenoid, parts 
of petromastoid and parapterygoid are fenestrated. Auditory bulla 
conspicuously inflated. External auditory meatus tube-like, extend- 
ing dorsocaudad. Malar long, deep, thin, and with prominent 
posterior projection. Diastema very long. Incisive foramen also 
long, broadened posteriorly. Palatine foramen minute. Palatal 
bridge shorter than width of mesopterygoid space, posterior margin 
opposite pm^. Parapterygoid process long, hook-shaped. Angle and 
condyloid process of lower jaw broad, thin; coronoid process 
vestigial. 

Teeth. — Figyire 31. Two pairs of continuously growing upper in- 
cisors. Anterior pair with groove nearer medial than lateral border. 
Cheekteeth hypsodont. Pm' and pm, with reentrant angles on 
anterior surface. Pm' with one deep and two shallow reentrant 
angles, remaining upper cheekteeth with one lingual reentrant 
angle. Crowns elliptical in outline, divided by single transverse 
lamina. 

Measurements.— Table 5. Male and female measurements sub- 
equal. Means and ranges of condyloincisive length (in millimeters) 
of sixteen adult males and eight adult females are 78.8 (75.3 to 82.3) 
and 76.8 (74.9 to 78.9), respectively. 

Age determination. — Adults have all cheekteeth slightly worn, 
basioccipital-basisphenoid suture closed. 

Variation.— Lepus c. rothschildi specimens from the desert edge of 
Giza Governorate are more reddish than those from elsewhere in 
Egypt (Setzer, 1958b). White forehead spots are common in 





Fig. 31. Skull of Lepus capensis. 



87 



»o ■V 



«« 



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OSBORN&HELMY: MAMMALS OF EGYPT 89 

rothschildi, but occur in other Egyptian subspecies. Color in general 
vewies from slightly more grayish Sinai specimens through 
yellowish and pale brownish gray in Eastern Desert specimens to 
brownish and reddish gray in the Western Desert. Black anterior 
ear tip predominates in subspecies sinaiticus and aegyptius, 
appears on intergrades between aegyptius and isabellinus, but is an 
occasional individual variation in Western Desert rothschildi. The 
latter averages smaller in most dimensions than other Egyptian 
subspecies (table 5). Other variations in measurements are dis- 
cussed under comparisons of subspecies. 

Comparisons.— Lepus capensis differs externally from all other 
Egyptian mammals by the combination of long ear, long hind foot, 
and short tail which is black above, white below. Cranially, L. capen- 
sis differs in the marked fenestration of maxilla and posterior parts 
of skull, short palate, two pairs of upper incisors, and upper tooth 
row wider than lower. 

The Western Desert subspecies, L. c. rothschildi, differs from 
other Egyptian subspecies by slightly more brownish to reddish 
color of dorsum, lack of black anterior border on ear tip, and smaller 
average dimensions (table 5). Lepus c. sinaiticus is distinguishable 
from other subspecies by slightly more grayish color, due to length 
of black terminal bands on dorsum hairs, and greater average ear 
length. 

Lepus c. aegyptius in comparison with sinaiticus and isabellinus 
appears to have a higher frequency of specimens with anterior 
border of ear tip black, but shows intergradation with those two 
subspecies in this character. Lepus c. aegyptius averages in- 
termediate in measurements between sinaiticus and isabellinus. 
Lepus c. isabellinus differs little in color from aegyptius, but has a 
considerably longer tail (table 5). 

Remarks. —Setzer (1958b) declined applying a subspecific name to 
1 2 specimens from the Western Desert, chiefly from the desert area 
adjacent to the Nile Delta. We consider them to be rothschildi on 
geographic grounds. Due to the fact that this area represents a 
habitat limit, different phenotypic expressions are to be expected. 

Collection. —Shooting from a vehicle is the most efficient means of 
collecting desert hares. Coni-bear traps set at 1 -meter intervals on 
both sides of a net have taken hares in areas impassable for vehicles. 
Bedouins near Khatatba "trap large numbers in weir-type nets set 
in the desert and baited with clover" (Hoogstraal, 1963, p. 5). 



90 FIELDIANA: ZOOLOGY 

//a6i(afs.— Vegetated desert and wadi mouths along the Red Sea, 
Gulf of Suez, and Western Mediterranean ^coast. In the area of Bir 
Abraq, hares were found hiding in rock crevices (Hoogstraal et al., 
1957ab). In the Western Mediterranean Coastal Desert (fig. 8), hares 
spend the day in clumps of Lycium sp. At Zeitun, Siwa Oasis, hares 
were frightened from clumps of Tamarix sp. in sebakha. South of 
Qena hares were seen in stands of Zygophyllum coccineum. Near Bir 
Zafarana in the Eastern Desert, one was flushed from a patch of 
Juncus sp. Wassif (1953b) reported hares from stands of Panicum 
turgidum in wadis in the Risan Eineiza area of north Sinai 
Peninsula. 

Food.— Little is known of plants eaten by hares in Egypt. The 
variety is doubtlessly great. We have observed that Zygophyllum 
coccineum was nibbled by hares and were told by a Bedouin that 
they eat fallen acacia blossoms. Gnawed bark of yassar trees 
{Moringa peregrina) was noted by Tregenza (1958). 

Reproduction.— Records of reproduction are from three females 
with two, two, and three fetuses from Wadi Ibib in March. 

Sex ratio.— A sample of 71 museum specimens of L. capensis con- 
tained 38 (54 per cent) males and 33 females. 

Key to Egyptian Subspecies of Lepus capensis 

1. Ear tip usually black anteriorly. 

a. Ear length more than 120 mm., average 126 mm. (Sinai Peninsula) 
sinaiticus, p. 90. 

b. Ear length 120 mm. or less, average 115 mm. (Northern two-thirds of Eastern 
Desert) aegyptius, p. 91. 

2. Ear tip usually not black anteriorly. 

a. Tail long, average about 90 mm. (Southern one-third of Eastern Desert) 
isabellinus, p. 92. 

b. Tail shorter, average about 70 mm. (Western Desert) rothschildi, p. 92. 

Lepus capensis sinaiticus (Ehrenberg, 1833) 

Lepus sinaiticus Ehrenberg, 1833. in Hemprich and Ehrenberg. Symbolae 
Physicae. Mamm., Dec. 2, pi. 14. fig. 1. 

Type locality. —SIN Ah Gebel Musa. 

Distribution in Egypt.— Figure 30. Sinai Peninsula. 

External characters.— See species description. Dorsum grayish. 
Anterior border of ear tip usually blackish. 

Cranial characters.— See species description and Figure 31. 

Measurements.— Table 5. 



OSBORN&HELMY: MAMMALS OF EGYPT 91 

Comparisons.— Slightly more grayish than other subspecies. Ear 
length averaging longer than in other Egyptian subspecies. 

Specimens examined.— Total six. 

SINAI: El Quseima (1), Wadi Raha (1). Feiran Oasis (1), Wadi Ain el Gefeef (1), 
Wadi el Sheikh (1), St. Catherine Monastery (1). 

Published records. — Records are from Murray (1912), Flower 
(1932), Wassif (1953b), Setzer (1958b). 

SINAI: Mount Sinai (Typie); Ayun Musa E of; Wadi Feiran. 16 km. W Feiran 
Oasis; El Quseima; Wadi el Sheikh, near St. Catherine Monastery; St. Catherine 
Monastery. Wadi Raha. 

SINAI: Risan Eineza area south of El Arish (sight record), southwest Sinai (sight 
record). 

Lepus capensis aegyptius (Demarest, 1822) 

Lepus aegyptius Demarest, 1822, Encyclop. Methodique, Mammalogie, suppL, p. 
350. 

Type locality.— QEN A: between Luxor and Kamak. 

Distribution in Egypt.— Figure 30. Northern two-thirds of 
Eastern Desert. 

External characters. — See species description. Dorsum 
yellowish to brownish gray. Anterior border of ear tip blackish. 

Cranial characters.— See species description and Figure 31. 

Measurements.— Table 5. 

Comparisons.— Lepus c. aegyptius differs from other Egyptian 
subspecies in having a larger proportion of individuals with 
anterior border of ear tip black, from sinaiticus in more yellowish 
or brownish dorsum and shorter ear, from isabellinus in having a 
shorter tail, and from rothschildi in paler color and generally 
larger dimensions. 

Specimens examined.— Total five. 

SUEZ: Wadi Katamiya mouth (1), Wadi Gindali (1), Wadi Iseili (1). 
RED SEA: Wadi Abu Qarayia (1). Ezzeit (1). 

Sight records of I. Helmy and D. Osborn.— 

CAIRO: Wadi Garawi. 

SUEZ: Wadi Qiseib. Cairo-Suez road km. 77. El Dar el Beida. 

RED SEA: Bir Zafarana. 

QEN A: Qena 30 km. S. 

Published sight records. — Records are from Bedan (1928) and 
Tregenza (1955, 1958). 

RED SEA: Wadi Habeeb, Wadi Umm Sidri, Wadi Shawak. 



92 FIELDIANA: ZOOLOGY 

Lepus capensis isabellinus (Cretzchmar, 1826) 

Lepus isabellinus Cretzschmar. 1826. in Riippell. Atlas zu der Reise im nordliche 
Afrika von Ruppell. Saugeth.. pi. 20, p. 52. 

Type locality.— SxiAan. NORTHERN: Ambukol. deserts south of. 

Distribution in Egypt — Figure 30. Southern one-third of Eastern 
Desert. 

External characters. —See species description. Dorsum yellowish 
to brownish gray. Ear tip usually not black on anterior border. 

Cranial characters. — See species description and Figure 31. 

Measurements.— Table 5, 

Comparisons.— Lepus c. isabellinus is about the same color as L. c. 
aegyptius, less grayish than L. c. sinaiticus, and markedly paler 
than L. c. rothschildi. Coloration is rather uniform in isabellinus 
except for black inner tip of ear in about one-third of specimens 
examined. Tail length is longer in isabellinus than in other Egyptian 
subspecies. Ear length is less in isabellinus than in sinaiticus. In 
comparison with rothschildi, isabellinus averages larger in most 
dimensions, except head and body length and postorbital width. 

Specimens examined.— Total 14. 

RED SEA: Bir Abraq (1), Wadi Naam (1). 
ASWAN: Near Aswan (1). Gebel Ain (1). 

SUDAN ADMINISTRATIVE: Wadi Ibib (2|, Wadi Darawena (2). Wadi Adeib 3.2 
km. N of Bir Kansisrob (2). Wadi Adeib (1), Wadi Kansisrob (2). 
Sudan. NORTHERN: Wadi Haifa (1). 

Sight record of I. Helmy.— 

ASWAN: Qustul East. 

Published records. — Records are from Hoogstraal et al. (1957b) 
and Setzer (1956, 1958b). 

RED SEA: Bir Abraq. Wadi Naam. 

ASWAN: Naikhala. 

SUDAN ADMINISTRATIVE: Wadi Adeib; 3.2 km. N of Bir Kansisrob; 4.8 km. 

N of Bir Kansisrob. 6.4 km. N of Bir Kansisrob; Wadi Darawena. 

Lepus capensis rothschildi (De Winton, 1902) 
Lepus rothschildi De Winton, 1902. Nov. Zool.. 9. p. 444. 

Type locality.— GIZ A: Giza. 

Distribution in Egypt — Figyire 30. Western Desert. 

External characters. — See species description. Dorsum 



OSBORN&HELMY: MAMMALS OF EGYPT 93 

brownish to reddish gray. Ear tip usually entirely yellowish white 
on anterior border. 

Cranial characters.— See species description and Figure 31. 

Measurements.— Table 5. Lepus c. rothschildi averages smaller 
in most dimensions than all other Egyptian subspecies. 

Variation. Setzer (1958b) noted reddish coloration in 
specimens from near Abu Rawash. 

Comparisons. — In comparison with other Egyptian subspecies, 
rothschildi is generally darker and more brownish, lacks black on 
anterior margin of ear tip, and averages smaller in most dimen- 
sions. 

Remarks.— Three specimens from Wadi Nassim, west side of 
Nile Valley, were considered "virtual topotypes of L. c. 
aegyptius" (Setzer, 1958b, p. 147; Hoogstraal, 1963). Note that 
the type locality of L. c. aegyptius has been fixed as between 
Luxor and Karnak on the eastern side of the Nile Valley. On a 
geographical basis, these specimens belong to rothschildi. Their 
chief similarity to aegyptius is the brownish anterior border of ear 
tip. 

Specimens examined.— Total 69. 

BEHEIRA: El Khatatba (6); Kom Hamada (3); Zaghig (6); Bir Victoria (7), 8 km. S 
(4); Wadi el Natroun (1). 

GIZA: Wardan (3), Beni Salami (2), Giza (1). 

QENA: Isna. Wadi Nassim (3). 

MATRUH: Bahig (1), 8 km. S (2). 16 km. S (3). 18 km. S (1), 20 km. S (2); Abu Mena 
(2); Burg el Arab (1); El Hammam 10 km. S (2). 17 km. SW (2); Sidi Barrani (1), 4.8 
km. S (1). 32 km. W (2); Salum 6.4 km. E (1); Bir Wair area (2): Bir Shaqqa 4.8 km. N 
(1); Marsaba (1); El Ferinat E of El Maghra (1); El Malfa 1 10 km. W of Siwa (2); Siwa 
Oasis, Abu Shuruf (2); El Zeitun (2). 

ASWAN: Near Aswan West (1). 

Sight records of I. Helmy and D. Osborn.— 

ASWAN: Aswan West, Nag Farqanda West. 

MATRUH: El Maghra; Qaret el Mashruka W of; Siwa Oasis. El Zeitun. 

Published records.— Records are from Anderson (1902), De 
Winton (1902b. 1903), Kasim (1912), and Setzer (1958b). 

GIZA: Wardan, Abu Rawash 1 km. S. 
EL FAIYUM: Gattah, Birket Qarun. 

BEHEIRA: Bir Victoria; Bir Victoria 8 km. S; Wadi el Natroun; Wadi el Natroun, 
Zaghig: between El Khatatba and Wadi el Natroun. 

MATRUH: Burg el Arab, Wadi el Ghazal S of Sidi Barrani (sight record). 



ORDER RODENTIA 

Kky r) Egyptian Families of Rodkntia 

1. Pelage soft, harsh, or spinous. Head and body length not exceeding 260 mm. 
Nasofrontal region normal. Angular process of lower jaw arising ventral to 
alveoli. 

a. External ear and tail present. Eyes normal. Supraoccipital normal, except in 
genus Nesokia. Median sagittal ridge absent. Interparietal present. Molar pat- 
terns variable, never S-shaped. 

i. Hind limbs elongate, tibia and fibula fused. Functional hind toes three. Tail 
length averages 150 per cent of head and body. Tail tip feathered black and 
white. Infraorbital foramen greatly enlarged, with upper root of zygomatic 
process posterior to lower. Maxillary plate minute. Check teeth "3-, enamel 

patterns E- and Z-shaped Family 5. Dipodidae, p. 322. 

ii. Hind limbs normal. Functional hind toes five. Tail length averages less than 
150 per cent of head and body. Tail tip not feathered black and white. In- 
fraorbital foramen not greatly enlarged. Upper root of zygomatic process 
anterior to lower. Maxillary plate normal, except in Muscardinidae. Cheek 
teeth §.4, crowns transversely ridged, tuberculate, prismatic, or laminate. 
(a)Tail bushy nearly to base. Black facial markings present. Skull smooth. 
Tympanic bulla with three septa. Cheek teeth 4, crowns concave. 

transversely ridged Family 4. Muscardinidae, p. 315. 

(b)Tail not bushy, except in genus Sekeetamys. Black facial markings ab- 
sent. Skull ridged. Tympanic bulla with no more than two septa. Cheek 
teeth 5, crowns not as above. 

(1) Pelage soft. Tail hair concealing annulations. Tail tip usually tufted. 
Supraorbital ridge present, tempoparietal ridge usually absent. In- 
cisive and palatal foramina large. Bulla conspicuously inflated. Molars 
with tubercles or prisms in two longitudinal rows, laminate in genus 

Pachyuromys Family 1. Cricetidae, p. 95. 

(21 Pelage soft, harsh, or spinous. Tail hair not concealing annulations. Tail 
never tufted. Supraorbital and tempoparietal ridges present, except in 
genus Mus. Incisive foramen large, except in genus Nesokia: palatal 
foramen minute. Bulla not conspicuously inflated. Molars with cusps in 

two longitudinal rows, laminate in genus Nesokia 

Family 3. Muridae. p. 253. 

b. External ear absent. Tail not visible externally. Eyes small, covered with hairy 
skin. Supraoccipital large, sloping forward to level of zygomatic process of tem- 
poral. Median sagittal ridge present. Interparietal absent. Enamel pattern of 
molars S-shaped Family 2. Spalacidae, p. 245. 

2. Pelage of dorsum and tail of long, round, hollow quills. Head and body length 

94 



OSBORN&HELMY: MAMMALS OF EGYPT 95 

average 600 mm. Nasofrontal region inflated. Angular process of mandible aris- 
ing lateral to alveoli. Crowns of cheek teeth flat, complexly folded 

Family 6. Hystricidae, p. 357. 

Family 1. Cricetidae 
(Subfamily Gerbillinae) 

Small to relatively large rodents. Head and body length average 
66 to 120 mm. Fur soft. Supraorbital spots usually prominent. Tail 
annulations concealed by hair, apical brush usually present. 
Supraorbital ridge present, tempoparietal ridge usually not con- 
spicuously developed; infraorbital foramen relatively small and in- 
cisive and palatine foramina long. Tympanic and mastoid bullae 
conspicuously inflated. Upper incisor with anterior surface grooved, 
except in genus Psammomys. Cheek teeth tuberculate (tubercles in 
two longitudinal rows), laminate, or prismatic. Dental formula: \, 5, 
5,3X2=16. 

Keys to Egyptian Genera of Gerbillinae 
External Characters 

1. Tail slender, usually longer than head and body length (95 per cent or more). 

a. Sole partly or completely haired, with a single lobed subdigital pad and a 
subhallucal tubercle Ifig. 34). 

i. Sole partly haired. Belly hairs usually with gray bases Meriones, p. 190. 

ii. Sole completely haired. Belly hairs never with gray bases. . Gerbillus, p. 96. 

b. Sole naked, with six tubercles (three subdigital, one subhallucal, and two plan- 
tar) (fig. 34). 

i. Tail bushy Sekeetamys, p. 181. 

ii. Tail not bushy Dipodillus, p. 140. 

2. Tail thick, always shorter than head and body length (90 per cent or less). 

a. Tail normal, with black tip. Belly hairs yellowish Psammomys, p. 226. 

b. Tail clavate (club-shaped), without black tip. Belly hairs white 

Pachyuromys, p. 220. 

Cranial and Dental Characters 
1. Anterior surface of upper incisor grooved. 

a. Molars tuberculate in immatures, laminate in adults. Supraoccipital swollen, 
posterior margin beyond level of occipital condyles. 

i. First libial and lingual cusps of m' opposite. Mastoid bulla never inflated 

posterior to level of supraoccipital Gerbillus, p. 96. 

ii. First libial and lingual cusps of m' alternate, at least in immatures. Mastoid 
bulla sometimes inflated posterior to level of supraoccipital. 
(a) Mastoid bulla always inflated beyond level of supraoccipital. Superior 

wall of parapterygoid perforated Sekeetamys, p. 181. 

(b)Mastoid bulla inflated beyond level of supraoccipital in some species. 
Superior wall of parapterygoid fossa not perforated Dipodillus, p. 140. 

b. Molars laminate or prismatic in all ages. Supraoccipital not swollen, posterior 
margin at or slightly beyond level of occipital condyles. 



96 FIELDIANA: ZOOLOGY 

i. Supraoccipital slightly constricted by swelling of bullae. Palatine foramen 

narrow, inconspicuous Meriones, p. 190. 

ii. Supraoccipital narrowly constricted by swelling of bullae. Palatine foramen 

broad, conspicuous Pachyuromys, p. 220. 

2. Anterior surface of upper incisor smooth Psammomys, p. 226. 

Genus Gerbillus Desmarest, 1804 

Orangish to brownish rodents of varying size. Tail longer than 
head and body in all species. Tail brush variable. Palm and sole 
haired. Hand with large palmar pad bearing appendix. Foot with 
single, lobed subdigital pad and subhallucal tubercle, no plantar 
tubercles (fig. 34). 

Brain case usually inflated and supraoccipital swollen beyond 
level of occipital condyle. Cranial ridges developed in Icirger species, 
supraorbital ridges in all species. Lip of auditory meatus never 
modified or swollen. Accessory tympanum present. Some variation 
among species in size of tympanic bulla and chambers of mastoid 
bulla. Subarcuate fossa small, never separating anterior and lateral 
superior p)osterior mastoid chambers as in some species of 
Dipodillus (figs. 36, 47). 

Upper incisors with single groove on anterior surface. Molars 
tuberculate, becoming laminate with wear. First labial and lingual 
cusps of m' opposite (fig. 38). 

Key to Egyptian Species of Gerbillus 

1. Larger species, hind foot length 30-36 mm., occipitonasal length 30-38 mm. 

a. Dorsum dark to pale. Posterior margin of nasals broadly to narrowly truncate. 
Interparietal usually deep and narrow (table 7. fig. 37 pyramidum, p. 96. 

b. Dorsum very pale. Posterior portion of nasals tapering and narrow, "bottle- 
shaped." Interparietal shallow and broad (table 7. fig. 37) . .perpaUidus, p. 117. 

2. Smaller species, hind foot length 25-32 mm., occipitonasal length 25 to 31 mm. 

a. Dorsum dark, brownish orange. Ear and sole pigmented. Posterior margin of 
nasals truncate (table 11. fig. 41) andersoni, p. 1 19. 

b. Dorsum orangish. Ear and sole not pigmented. Posterior margin of nasals 
round or pointed (table 11. fig. 41) gerbillus, p. 130. 

Gerbillus pyramidum I. Geoffroy St. Hilaire, 1825 

Gerbillus pyramidum I. Geoffroy St. Hilaire. 1825. Diet. Class. Hist. Nat.. Vol. 
VII. p. 321. 

Type locality.— Egypt. GIZA: Giza Pyramids area. 

General distribution. — Israel, Sinai Peninsula, Egypt, Sudan, 
Libya, Tunisia, Algeria, Morocco, Northern Chad, Niger, and 
Mauritania. 

Common names.— Greater Gerbil, Demsy. 



OSBORN & HELMY: MAMMALS OF EGYPT 



97 



2 6* 




Fig, 32. Collection localities of Gerbillus pyramidum pyramidum (dots), G. p. 
floweri (solid squares), G. p. gedeedus (hexagons), G. p. elbaensis (open squares), and 
G. perpallidus (circles). 



Distribution of subspecies in Egypt— Figure 32. Gerbillus 
pyramidum floweri: northern Sinai and northern part of Eastern 
Desert; Gerbillus pyramidum pyramidum: Nile Delta and Valley, 
Wadi el Natroun, El Faiyum, Wadi Muwellih, and Siwa Oasis; Ger- 
billus pyramidum gedeedus ssp. nov.: Bahariya, Kharga, and 
Dakhla Oases and probably Farafara Oasis; Gerbillus pyramidum 
elbaensis: southeastern part of Eastern Desert. 

Diagnosis.— Large gerbil with orangish to tawny and brownish 
upper parts. Dorsal stripe dark and usually distinct from sides. Tail 
long; brush brownish, usually conspicuous. Ears pigmented in 
nominate subspecies. Eye prominent. Skull large, heavily ridged. 
Nasals broadly to narrowly truncate posteriorly. Bulla extending or 
not extending posteriorly slightly beyond paroccipital. 



98 FIELDIANA: ZOOLOGY 

Largest of Egyptian species of Gerbillus. Adult head and body 
length average 120 mm.; tail 158 mm., 134 per cent of head and 
body length; hind foot 36 mm.; ear 17 mifi.; occipitonasal length 
33.4 mm.; weight 60 gm. 

External characters. — Upper parts varying through orangish cin- 
namon, cinnamon, and tawny to brownish. Dorsal stripe, if distinct, 
broad and dark. All hairs of dorsum and side, except for narrow ven- 
trolateral strip, with gray bases. Widths of agouti bands and 
brownish tips of dorsal hairs variable. Hairs of underparts and feet 
white to base. Upper surface of tail either as dorsum (with brownish 
tipped hairs) or side (lacking brownish tipped hairs). Tail brush one- 
third or more of tail length, brownish or fuscous. Under surface of 
tail either entirely white or with buffy base. Rump patch of white 
hairs, hairs with white bands, or lacking. Mystacial and circum- 
orbital areas pale in desert populations, darker and less conspicuous 
in Nile Delta and Valley populations. 

Palatal ridges.— Figure 33. Palatal ridges of two or three 
specimens of species of Gerbillinae were examined, with the excep- 
tion of Gerbillus perpallidus, Dipodillus mackilligini, Meriones 
sacramenti, and M. tristrami. They appear to be of taxonomic value 
in this group. Eisentraut (1969) recognized the usefulness of palatal 
ridges in the Muridae, but noted that extra ridges and abnormalities 
may occur. 

In 0. pyramidum, the first diastemal or antemolar ridge is broad- 
ly U-shaped, the second is transverse. The first to fourth intermolar 
ridges reach the midline and are recurved; the fourth is slenderest; 
the fifth is thickest, reaches midline, but is not recurved. 

Glans penis and 6acu/um.— Slight differences in shape of the 
penis among species of the family Gerbillinae were illustrated by 
Wassif et al. (1969, p. 84). In Gerbillus and Dipodillus, the surface of 
the penis is covered with minute spines in small, circular pockets. 
The baculum consists of a basal plate distinguished from the bony 
shaft, except in G. andersoni, with "three separate cartilaginous 
digitigrade processes." 

Feet. — Palm and sole haired, pads and tubercles comparable with 
G. gerbillus (fig. 34). 

Cranial characters.— Figures 35-37. Skull largest and most heavi- 
ly developed of Egyptian species of Gerbillus. Supraorbital ridge 
thick and prominent, parietal ridge variable. Zygomatic plate large, 
anterior margin usually reaching level of premaxillary-maxillary 











r?\5^^f*"^j '""-^^^tjy 




r^y^ 



^i^ 



G.GERBILLUS 



G.ANDERSONI 




G. PYRAMIDUM 



D.CAMPeSTRIS 



0. SIMONI 




D.AMOENUS 




SCALURUS 



M.CRASSUS 



P. OBESUS 



Fig. 33. Palatal ridges of Gerbillus gerbillus, G. andersoni, G. pyramidum, 
Dipodillus campestris, D. simoni, D. amoenus, Sekeetamys calurus, Meriones 
cmssus, and Psammomys obesus. Not drawn to same scale. 



99 



100 FIELDIANA: ZOOLOGY 

suture. Posterior margin of nasals broadly to narrowly truncate. In- 
terparietal shape varies between subspecies, as shown in Figure 37. 
Anterior surface of tympanic bulla reaching level of posterior 
margin of foramen ovale. Posterior surface of mastoid bulla ex- 
ceeding or not exceeding level of paroccipital process. Partition be- 
tween anterior mastoid chamber and posterior superior mastoid 
chamber at level of or slightly behind level of posterior margin of 
hamular (suprameatal) process of temporal (fig. 36). 

Teeth.— Figure 38. Upper incisor grooved. First labial and lingual 
cusps of m' opposite. Molars tuberculate in immatures, becoming 
fused into laminae with rounded margins. Confluency of cusps 
begins between anterior and first lingual cusps in m', anterior and 
first labial cusps in m,, and is completed earliest in m,. M and mj 
become laminated prior to confluency of cusps of m\ mj, but 
anteroposterior union is limited. M^ small, with three transient 
cusps. 

Measurements.— Table 6. Male dimensions average slightly larger 
than female. Means (and ranges) of occipitonasal length (in 
millimeters) of 21 adult males and 13 adult females, respectively, 
are 35.9 (34.6 to 38.1) and 34.8 (33.2 to 37.3). 

Age determmation.— Individuals are considered adult when 
anterior and first lingual cusps of m' become confluent or almost 
confluent and/or the basioccipital-basisphenoid suture closes. 

Variation.— The greatest amount of variation is between G. p. 
pyramidum of the Nile Valley and Delta and G. p. floweri of the 
northern Eastern Desert and Sinai Peninsula. 

Intergradation between G. p. pyramidum and G. p. floweri ex- 
tends a few kilometers east of the Nile Delta where floweri becomes 
considerably paler than pyramidum; ear tip loses pigmentation; tail 
becomes completely bicolored; upper surface of tail loses blackish 
heiirs; tail brush becomes smaller and paler; rump patch increases in 
size and whiteness; mystacial, suborbital, postorbital, and 
postauricular areas become lighter or white; white of underside ex- 
tends onto shoulders and over whole of limbs; and dorsal stripe 
becomes less distinct or lacking. The anterior mastoid chamber is 
slightly more expanded in desert populations (fig. 36), posterior 
margin of nasals narrower, interparietal less deep (fig. 37), and 
means of most dimensions smaller. 

Differences between G. p. pyramidum and other subspecies are 



OSBORN & HELMY: MAMMALS OF EGYPT 



101 




G.GERBrauS S.CAIURUS M.CRASSUS P.OBESUS 

Fk; 34. Palms and soles of Dipodillus campestris, I), dasyurus. I), simoni, I), 
amoenus, D. henleyi. Gerbillus gerbillus. Sekeetamys calurus. Meriones crassus, and 
Psammomys obesus. Not drawn to same scale. 

considerably less striking. Tail brush is larger in G. p. gedeedus ssp. 
nov. (table 7), dorsal stripe indistinct, skull more heavily ridged, and 
bulla inflation slightly greater. 

Gerbillus p. elbaensis is slightly paler than G. p. pyramidum, tail 
proportionally longer (table 6), posterior margin of nasals narrower, 
and anterior mastoid chamber of bulla more swollen in about 50 per 
cent of the samples. 

The statement of Innes (1932, p. 22) that ear length is longer than 
one-half hind foot length in G. pyramidum is true only for samples 
from the Nile Delta. 

Comparisons.— Gerbillus pyramidum differs from related species. 




Fk; 35. Skull of adult Gerbillus pyramidum floueri. 



102 



OSBORN & HELMY: MAMMALS OF EGYPT 



103 





B 



Fui 36. Mastoid bulla variations in Gerbillus pyramidum and G. perpallidus. (A) 
Anterior position of partition (at arrow) between anterior mastoid (shaded) and 
superior posterior lateral mastoid chambers as in G. pyramidum pyramidum. (B) 
Posterior position of partition (at arrow) as in desert subspecies of G. pyramidum 
and in G. perpallidus. 

G. perpallidus, in much darker color, presence of a broad dorsal 
stripe, white rump patch smaller or absent, nasals truncate 
posteriorly instead of tapering narrowly, narrower and deeper inter- 
parietal, thicker and longer supraorbital ridge, less inflated mastoid 
bulla, and significantly larger dimensions. Characters of larger ger- 
bils are summarized in Table 7, and shown in Figures 36, 37. 

From G. gerbillus and G. andersoni, G. pyramidum differs in 
much larger dimensions (tables 6, 8, 10), relatively smaller bulla, 







<0 





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104 



OSBORN & HELMY: MAMMALS OF EGYPT 



105 






t 

4^ 



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D.CAMPESTRIS 






0. AMOBNUS 




D.DASYURUS 







^ 



D.HENIEY/ 



11 



DS/MON/ 







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(^ 



^ ^ 

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G.GERBIUUS 



t 



D.MACKILUGINI 




G.PYHAMIDUM 



Fig. 38. Crown views of right upper (U) and left lower (L) molars of mature (A) and 
immature (I) of species oi Dipodillus and Gerbillus. Mature only of D. mackilligini 
shown. 

longer and more prominent tail brush; from G. gerbillus, in darker 
coloration and presence of dorsal stripe. 

Wassif et al. (1969) reported the diploid chromosome numbers of 
G. pyramidum, G. andersoni, and G. gerbillus to be 38, 40, and 42 
(43 in males), respectively. Lay et al. (1975) found that G. per- 
pallidus exhibited a 2N=40. Zahavi and Wahrman (1957) listed two 
IsraeU forms of G. pyramidum, from the Negev and Coastal plain, 
as having diploid numbers of 66 and 52. 



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106 



OSBORN&HELMY: MAMMALS OF EGYPT 107 

Collection.— Dug from burrows in sand and trapped alive near 
burrows and where tracks are obvious. 

Habitats.— Sinai Peninsula: Gerbillus p. floweri was collected in 
palm groves and near cultivation (Hoogstraal, 1963) and in sandy 
dunes of the northern section (Wassif, 1953b), but not in dunes of 
southern Sinai (Haim and Tchernov, 1974). The last authors gave 
the southern limit at Wadi Wardan. 

Eastern Desert: Gerbillus p. floweri was collected in Wadi el Gafra 
in sand beneath Lygos raetum. Hoogstraal (1963, p. 12) listed it 
from "palm groves and near cultivation" in Suez and "from around 
old army barracks" in Qalyubiya Governorate. Gerbillus p. elbaen- 
sis is from "grassy valleys in Gebel Elba," sandy coastal plain 
(Hoogstraal, 1963, p. 11), and lower reaches of Wadi Ibib beneath 
close stands of Panicum turgidum. 

Nile Delta: Sandy areas near cultivation, palm groves, and sandy 
canal banks supporting half a grasses (Desmostachya bipinnata and 
Imperata cylindrica); desert edge areas in stands of Panicum 
turgidum and other bunch grasses; and barren sand and gravel 
around tents and houses south of Giza Pyramids. 

Nile Valley: East and west banks in situations described above 
and inside larger tributary wadis, such as Wadi Asyuti, where 
vegetation is scattered Acacia raddiana, Leptadenia pyrotechnica, 
and Zilla spinosa; dry cracked mud beside the Nile; and palm groves 
with patches of halfa grass. 

Wadi el Natroun: Sand sheets vegetated with Panicum turgidum 
and small shrubs. 

El Faiyum: Under Tamarix sp. in sandy areas at desert edge. 

Wadi Muwellih: Sand mounds supporting Nitraria retusa and 
Desmostachya bipinnata. 

Oases: Sandy areas with cover of halfa grasses and other vegeta- 
tion in palm groves; sand sheets supporting camel thorn {Alhagi 
mannifera) and bunch grasses (Sporobolus spicatus and 
Stipagrostis vulnerans); heavily vegetated areas around walls, 
springs, and leaking aqueducts (fig. 21); beneath Acacia sp. and 
Lagonychium farctum; under date palms and Tamarix sp.; and in 
drifted sand in uninhabited buildings. 

ficAafior.— Nocturnal. Extremely nervous and highly sensitive to 
slightest noise and movement. Difficult to handle and bites readily. 



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109 



no FIELDIANA: ZOOLOGY 



Table 8. — Means (and ranges) of measurements, ratios, and weight of adult Ger- 
billus andersoni. 





G. a. inflatus 


G. a. andersoni 


G. a. bonhotei 


HBL 


96.0(83-112)38 


92.2(90-115)26 


96.2 (95-98) 4 


TL 


128.0(118-140131 


124.0(110-150)26 


120.5(117-125)4 


TL/HBL% 


134.3(118.2-159.0)32 


125.4(115.5-139.6)25 


125.2(119.4-131.6)4 


FL 


28.4 (25-30) 38 


29.4 (27-32) 27 


27.8(26-31)4 


EL 


15.2(15-18)39 


16.0(15-19)25 


15.5(14-16)4 


Wt 


28.6(15.9-38.4)31 


30.0 (27.7-33.2) 9 





ONL 


28.4 (27.5-30.3) 32 


29.2(27.7-31.0)26 


28.9 (28.3-30.0) 5 


ZW 


16.0(14.8-16.6)27 


16.4 (14.9-17.6) 17 


15.8, 16.0 


lOW 


6.0(5.1-6.2)34 


6.0 (5.4-6.0) 28 


5.6 (5.4-5.8) 5 


BCW 


14.0(13.4-14.4)31 


14.0(13.0-15.6)25 


14.0(13.7-14.3)5 


NL 


11.2(10.4-12.0)32 


12.0(10.5-14.0)27 


10.9(10.5-11.3)5 


IFL 


5.2 (4.6-5.6) 35 


5.0 (4.8-5.6) 27 


5.1 (4.8-5.4) 5 


AL 


4.2 (4.0-4.4) 34 


4.0 (3.6-4.8) 28 


4.6 (4.2-5.4) 5 


RW 


4.0 (3.6-4.4) 34 


4.0 (3.6-4.5) 28 


3.8,4.1,4.2 


BL 


9.0 (8.4-9.5) 31 


8.5(7.8-9.1)27 


9.1 (8.9-9.3) 5 


SH 


12.2(11.7-12.8)27 


12.0(11.1-12.5)22 


12.0(11.9-12.2)4 



Burrows.— The similarity between G. pyramidum and G. gerbillus 
burrows was noted by Yunker and Guirgis (1969) (see p. 135). 

Food.— See G. gerbillus. In northern Sinai, Wassif (1953b) found 
camel dung and seeds of Citrullus (=Colocynthis) in burrows of G. 
p. floweri and assumed they were part of the animal's food ration. 

Reproduction.— Data available only from January through March 
on three males with testes descended and two females, one with 
three fetal scars, and one with five fetuses. 

Sex ratio. — In a sample of 74 museum specimens, there were 38 
(51 per cent) males and 36 females. 

Associates.— Gerbillus pyramidum shares habitats with G. ger- 
billus, G. andersoni, G. perpallidus, Arvicanthis niloticus, Rattus 
rattus, Meriones libycus, and M. crassus. Commensal inhabitants of 
burrows are Usted by Yunker and Guirgis (1969) and discussed 
under G. gerbillus. 

Remarks. — Happold (1967a) considered G. pyramidum to be bet- 
ter adapted to desert environment of the Sudan than Jaculus 
jaculus, owing to its ability to breed during a longer period of the 
year and lose less weight when fed on whole barley for four weeks at 
30 °C. (Schmidt-Nielsen, 1964, p. 182). Nevertheless, G. pyramidum 
is never found in isolated, barren desert situations where J. jaculus 
survives. 



OSBORN&HELMY: MAMMALS OF EGYPT 111 

Key to Egyptian Subspecies of Gerbillus pyramidum 

1. Dorsum dark to pale. Tail with conspicuous black brush extending one-third or 
more of tail length. Nasals broadly truncate on posterior margin. 

a. Dorsum darker, stripe distinct. Brush extending up to one-third of tail length. 

Underside of tail base buffy. (Nile Delta, Nile Valley, El Faiyum) 

pyramidum, p. 111. 

b. Dorsum paler, stripe indistinct. Brush extending more than one-third of tail 
length. Underside of tail base white. (Western Desert oases), .gedeedus, p. 114. 

2. Dorsum pale, dorsal stripe indistinct to lacking. Tail with inconspicuous, fuscous, 
or grayish brush extending less than one-third of tail length. Nasals usually nar- 
rowly truncate posteriorly. 

a. Interparietal shallow, broad. Partition between mastoid bulla chambers in 

posterior position. (Sinai Peninsula and northern part of Eastern Desert) 

floweri, p. 1 13. 

b. Interparietal deep, narrow. Partition between mastoid bulla chambers in 
posterior position in 50 per cent of sample. (Southeastern part of Eastern 
Desert) elbaensis, p. 116. 

Gerbillus pyramidum pyramidum I. Geoff roy St. Hilaire, 1825 

Type locality.— Egypt. GIZA: Giza Pyramids area. 

Distribution in Egypt— Figure 32. Nile Delta and Nile Valley. 

External characters.— Upper parts tawny to brownish, dorsal 
stripe broad, dark brown. Side tawny to pale brownish. Some in- 
dividuals almost completely brown on back and side. Venter and 
feet white. Broad pigmented band extending from mystacial area 
below ear and continuous with color of sides. Circumorbital area 
pigmented. Postauricular spot white. Rump patch small or lacking. 
Upper surface of tail with blackish tipped hairs to base. Tail brush 
prominent. Under surface of tail white with buffy base. Ear 
pigmented. 

Cranial characters.— Figure 35. Skull large, angular, prominently 
ridged. Posterior margin of nasals broadly truncate. Partition be- 
tween anterior and posterior superior mastoid chambers in anterior 
position (fig. 36). Posterior margin of mastoid bulla not inflated 
beyond level of paroccipital. Interparietal deep and narrow (fig. 37). 

Measurements.— Table 6. Except for tail length, dimensions of G. 
p. pyramidum differ slightly from those of other subspecies. Ear 
length is obviously longer in Nile Delta samples only. 

Vanation.— Samples of G. p. pyramidum from dark soils are 
darker and have a less clearly defined dorsal stripe than those from 
pallid soils. Ear length decreases from north to south in Nile Delta 
and Nile Valley populations. Tail length increases slightly from 



112 FIELDI ANA: ZOOLOGY 

north to south along the Nile Valley. Albinistic specimens have been 
collected from near Abu Rawash. 

Comparisons.— The subspecies pyramidum can be distinguished 
from others by darker color, more clearly defined dorsal stripes, 
blackish hairs along entire length of tail, and buffy underside of tail 
base. Ears are pigmented in pyramidum and gedeedus, but not in 
other subspecies (table 7). 

From the desert-inhabiting subspecies, floweri and elbaensis, 
pyramidum can be disting^shed by less inflated bulla (fig. 36), 
deeper and narrower interparietal (fig. 37), partition between 
mastoid chambers in anterior position, and broadly truncate 
posterior nasal margin (fig. 37). From gedeedus, pyramidum differs 
in having average tail length 10 per cent shorter and having a less 
conspicuous tail brush. From elbaensis, pyramidum differs in 
having average tail length about 20 per cent shorter, but more con- 
spicuous brush. 

Remarks.— Two specimens (skulls only) of G. pyramidum were ob- 
tained in 1974 from beneath date palms at El Zeitun and El Maragi 
which are in the Siwa depression. They could have arrived acciden- 
tally via camel pack or they could be relicts of a formerly continuous 
distribution. This species is found in western Libya and far to the 
east in Egypt (fig. 32). All we can suggest at this time is the old 
cliche: "Further investigation is obviously necessary." 

Specimens examined.— Total 193. 

ALEXANDRIA: Near Alexandria (3). 

BEHEIRA: El Khatatba (3). El Birigat 2 km. W (1). Kom Hamada (1). Wadi el 
Natroun (2). Ezbet Beni Salami (2). 

GIZA: Giza (8); Abu GhaUb (13); Abu Rawash (19). 3 km. N (1); Gebel el Ghigiga 
(5); Giza Pyramids area (1), 3.6 km. W (1), 1 km. S (1); Cairo-Alexandria desert road 
km. 10 (2); Kafr Hakim (2). 

CAIRO: Cairo (4). 

MATRUH: El Maragi (1). El Zeitun (1). 

EL FAIYUM: Faiyum (3). 4.8 km. N (2); Kom O Shim (2): Kom Ashmun (2); 
Shooting Club (4); Ezbet Ayub Ali (2); Sinnurus (3); Fanus (2); Minshat el Amir 
(Mohamed Ali Pasha) (2); Wadi Muwellih. Bir Dakaar (2). 

ASYUT: Asyut 4.8 km. SW (1). Beni Adi (24). Durunka (2). Wadi Asyuti (2). 

QENA: Luxor (1). 4.8 km. N (1); Wadi Nassim (11). 

ASWAN: Aswan (2). 1.6 km. SE (1). 16 km. N (1): West Aswan (6); Kom Ombo (19), 
El Kagug Cave (2); Muneiha (3); El Biyara (10); Kom Ombo Temple 0.8 km. E (1): 
West Armina (4); Adindan (2); El Dirr. Amada (4); Abu Simbil (1). 

Published records. — Records are from Anderson (1902), Flower 
(1932), Setzer (1952, 1958d), and Bauer (1963). 



OSBORN&HELMY: MAMMALS OF EGYPT 113 

QALYUBIYA: Kafr Abu Sir. 

SHARQIYA: Tel el Kebir. 

KAFR EL SHEIKH: Baltim, El Burg. 

BEHEIRA: Wadi el Natroun; Bir Hooker: Kafr Dawud; El Khatatba: Kom 
Hamada, Talha Station. 

GIZA: Abu Rawash. 3 km. N, 1.6 km. N; Gebel Abu Rawash 4.8 km. NE. 1.6 km. 
NE: Giza Pyramids 3.2 km. W: Abu Ghalib: El Aiyat: Mit Riheina: Atfih; El 
Mansuriya: Zawyet Abu Mussalam: Birqash: Mena suburbs: Kafr Hakim: El Lisht 
Pyramid: Cairo- Alexandria desert road km. 10. 

CAIRO: Abassia Fever Hospital. 

EL FAIYUM: Faiyum. El Nassariya, Ezbet Ayub Ali, Idwa. Ezbet el Asfar, 
Fanus, Minshat Tantawi, Ezbet Abu Zeid, Kom O Shim 3.6 km. NE. 

MINYA: Tel el Amarna. 

QENA: Luxor, Wadi Nassim. 

ASWAN: Aswan: West Faras 4 km. S; El Dirr, Amada: Abu Simbil. 

Sudan. NORTHERN: Wadi Haifa. 

Gerbillus pyramidum floweri (Thomas, 1919) 

Gerbillus floweri Thomas. 1919, Ann. Mag. Nat. Hist., (ser. 9), 3, p. 559. 

Type locality. -Egypt. SINAI: Wadi Hareidin 22.4 km. S of El 
Arish. 

Distribution in Egypt.— Figure 32. Northern parts of Sinai Penin- 
sula and Eastern Desert. 

External characters.— Pale cinnamon to tawny above with in- 
distinct dorsal stripe. Mystacial area with white hairs only, area 
below eye very pale to whitish. Preorbital, postorbital, and post- 
auricular areas and rump patch white and prominent. Ear usually 
not pigmented. Tail usually without brownish or blackish hairs dor- 
sally, brush relatively inconspicuous, fuscous. 

Cranial characters.— Figures 35, 37. Skull with pronounced 
supraorbital ridge, posterior margin of nasals narrowly truncate, 
anterior margin of zygomatic plate usually reaching level of pre- 
maxillary-maxillary suture, posterior margin of mastoid bulla 
usually not inflated beyond level of paroccipital. Partition between 
anterior and posterior superior mastoid chambers usually in the 
posterior position (fig. 36). Interparietal a little less deep than in 
pyramidum (fig. 37). 

Measurements.— Table 6. Slightly smaller than G. p. pyramidum 
in some measurements. Ear length appears to be the only 
significantly smaller measurement. 

Vana^/on.— Samples from eastern edge of the Nile Delta and Wadi 
el Gafra have a more distinct dorsal stripe than those from vicinities 



114 FIELDIANA: ZOOLOGY 

of Suez Canal and northern Sinai. Other variations are described 
under the species. 

Comparisons.— The subspecies flowed is distinguishable from 
pyramidum in decidedly paler color, less distinct dorsal stripe and 
tail brush, narrower posterior margin of nasals, and posterior posi- 
tion of partition between anterior and posterior superior lateral 
mastoid chambers (fig. 36). Intergradation in color, nasal shape, and 
bulla conformation exist between G. pyramidum floweri and G. p. 
pyramidum. From gedeedus, floweri differs in the same details, and 
tail length averages 10 per cent shorter. Color markings in floweri 
differ slightly from those of elbaensis, but body size is larger, tail 
length shorter, posterior nasal margin narrower, interparietal 
shallower, and mastoid partition is posterior in position (table 7). 
Intergradation in characters between floweri and elbaensis is evi- 
dent. 

Gerbillus p. floweri and G. perpallidus are strikingly similar in 
color, and bulla shape and, in some individuals, posterior margin of 
nasals. 

Remarks.— VLBirci and Tchemov (1974) considered the Sinai form 
to be G. p. negev without recourse to substantiative data. 

Specimens examined.— Total 81. 

SINAI: El Arish (32|. Wadi Hareidin (Type). El Has el Ahmar (2). El Quseima (1). 
Ain Sudr (1), Ayun Musa (1). 

ISMAILIA: Abu Sultan (1), Lake Timsah west side (1). El Ballah (1). 
SHARQIYA: Bilbeis (6). 
QALYUBIYA: Kafr Abu Sir (6). 
SUEZ: Wadi el Gafra (28). el Kubri (1). 

Published records.— Records are from Flower (1932), Wassif 
(1954c), Setzer (1958d). and Haim and Tchemov (1974). 

SINAI: El Arish, Wadi Hareidin, El Quseima, Wadi Wardan near Has el Sudr 
(Sidr). 

ISMAILIA: Abu Sultan. 

QALYUBIYA: Kafr Abu Sir. Mazaret el Gebel el Asfar. El Khanka. 

Gerbillus pyramidum gedeedus ssp. nov. Osborn and Helmy 

Type.— Adult male, skin and skull, Field Museum of Natural 
History number 106213; original number 17230 in H. Hoogstraal 
catalog. Collected December 6, 1966, by Ibrahim Helmy. 

Type locality-Egypt. EL WADI EL GEDEED: Dakhla Oasis, 
El Mawhoub. 



OSBORN&HELMY: MAMMALS OF EGYPT 115 

External characters.— Tawny to light brown dorsally. Most 
without distinct dorsal stripe. Usually no black hairs on upper sur- 
face of tail except for long, distinctive black brush. 

Circumorbital markings indistinct. Postauricular patch white, 
prominent. White rump patch small when present. Belly, inside of 
legs, and underside of tail white. Ear pigmented. 

Cranial characters. —SkuW similar to G. p. pyramidum, but more 
massive. Posterior margin of nasals broadly truncate. Partition be- 
tween anterior and posterior superior mastoid chambers in anterior 
position (fig. 36). Interparietal as in pyramidum (fig. 37). 

Measurements.— Tables 6, 7. Type head and body length 121 
mm.; tail 180 mm., 148 per cent of head and body length; hind foot 
37 mm.; ear 17 mm.; occipitonasal length 35.9 mm.; weight 54.2 gm. 
Ear length is less than one-half hind foot length. 

Compansons.— Subspecies gedeedus differs from pyramidum in 
paler color; lack of or less distinct dorsal stripe; completely white 
underside of tail; lack of blackish hairs on upper side of tail; larger, 
more conspicuous tail brush; longer tail; slightly more inflated 
mastoid bulla; and slightly more angular and strongly ridged skull. 
Measurements other than tail length are about equal in the two 
subspecies (tables 6, 7). 

Gerbillus p. gedeedus cannot be confused with G. p. floweri, G. p. 
elbaensis, or G. perpallidus because of its larger size, darker color, 
and longer tail brush. The tail is relatively and actually longer than 
in other subspecies except elbaensis. 

Habitat.—See Oases under G. pyramidum and Figure 21. 

Specimens examined.— Total 101. 

GIZA: Bahariya Oasis. Bir Qasr No. 1 (7). Bir Qasr No. 3 (2); El Aguz (9); Bir 
Wigaba (3); Ain Marun (7); Mandisha (3); Bawiti (1); Ain el Qht (7); El Hara (9); Ain el 
Beilda (1); Wadi Ghorabi (2). 

EL WADI EL GEDEED: Dakhla Oasis. El Mawhoub (10. Type). Mut (2); Kharga 
Oasis (1). El Kharga 14 km. E (3), 3 km. S (3). 4 km. S (1); Bulaq (3): Nasser ViUage (3): 
Baris (2); El Gezira (11); El Farag (2); Ain Eede 8 km. E (1); Ginah (7). 

Published records.— Records are from Flower (1932) and Wassif 
(1960ab, as G. p. pyramidum). 

GIZA: Bahariya Oasis, Mandisha. 

EL WADI EL GEDEED: Kharga Oasis; Kharga Oasis, El Mahariq, Baris. 



116 FIELDIANA: ZOOLOGY 

Gerbillus pyramidum elbaensis Setzer, 1958 

Gerbiltus pyramidum elbaensis Setzer, 1959, J. Egypt. Publ. Health Assn., 33, p. 
223. 

Type locality.-Egypt. SUDAN ADMINISTRATIVE AREA: 
Wadi Adeib, 3.2 km. N of Bir Kansisrob. 

Distribution in Egypt— Figure 32. Southeastern part of Eastern 
Desert. 

External characters.— Dorsum pale cinnamon to tawny, dorsal 
stripe inconspicuous to absent. Color and markings very similar to 
G. p. floweri. Tail indistinctly bicolored, upper surface without 
brownish or blackish hairs; brush about one-third of tail length, 
fuscous, less conspicuous than in G. p. pyramidum or G. p. 
gedeedus. 

Cranial characters.— Nasal with posterior margin tap)ering and 
narrowly truncate or rounded in about 70 per cent of specimens ex- 
amined, broadly truncate in 30 per cent. Partition between anterior 
and superior posterior mastoid chambers in posterior position (fig. 
36) in about 50 per cent of specimens examined, posterior margin of 
mastoid bulla posterior to level of paroccipital in about 80 per cent, 
and interparietal deep and narrow, as m pyramidum, in about 70 per 
cent. Interparietal shape similar to that of floweri in about 30 per 
cent. 

Measurements.— Table 6. Gerbillus p. elbaensis averages smaller 
in most dimensions, except tail length, than other subspecies; 
however, the sample is much smaller than others. Anterior palatine 
(incisive) foramen is not as "long and narrow" (table 6) as one would 
assume from the comment of Setzer (1958d, p. 223). 

Comparisons. — Gerbillus p. elbaensis is distinguishable from 
floweri and other subspecies by smaller dimensions, except tail 
length. From pyramidum and gedeedus it differs in lack of a dorsal 
stripe, less conspicuous tail brush, relatively larger bulla, and nar- 
rower posterior nasal margins. 

Similarities between this subspecies, G. p. floweri, and G. per- 
pallidus in color and bulla conformation may be correlated with 
adaptation to desert habitat. 

Specimens examined.— Total 18. 

SUDAN ADMINISTRATIVE: Wadi Kansisrob (8); Bir Kansisrob 4.8 km. N (1); 
Wadi Adeib. 3.2 km. N of Bir Kansisrob (2); Wadi Serimtai, 16 km. N of Halaib (4); 
Halaib 20 km. N (1). Wadi Hodein (1); Wadi Ibib (1). 



OSBORN&HELMY: MAMMALS OF EGYPT 117 

Published records.— Records are from Hoogstraal et al. (1957b) 
and Setzer (1958d). 

SUDAN ADMINISTRATIVE: Halaib 20 km. N; Wadi Serimtai; Bir Kansisrob 
4.8 km. N, 3.2 km. N; Wadi Kansisrob. 

Gerbillus perpallidus Setzer, 1958 

GerbiUus perpallidus Setzer, 1958. J. Egypt. Publ. Health Assn., 33, p. 221. 

Type locality.— Egypt. BE HE IRA: Bir Victoria. 

Common name.— Pallid Gerbil. 

Distribution in Egypt— Figure 32. Western Desert between the 
western part of Nile Delta, Qattara Depression, and Western 
Mediterranean Coastal Desert environs of El Hamman. 

Diagnosis.— Medium size gerbil with pale orangish upper parts 
lacking dorsal stripe. Tail brush relatively inconspicuous, fuscous. 
Ears not pigmented. Skull not strongly developed, supraorbital 
ridges not heavy. Nasals tapering and narrow or "bottle shaped" 
posteriorly. Interparietal shallow and broad. Bulla extending 
posteriorly beyond paroccipital. 

Adult head and body length average 107 mm.; tail 137 mm., 129 
per cent of head and body length; hind foot 34 mm.; ear 16 mm.; 
occipitonasal length 32.3 mm.; weight 36.3 gm. 

External characters.— Dorsal color pale yellowish orange to light 
reddish orange with no dorsal stripe, but dark-tipped hairs on rump. 
Hairs of dorsum and part of side have gray bases. Underparts, feet, 
and underside of tail white. Mystacial and suborbital areas without 
pigmented hairs. Postorbital and postauricular spots and rump 
patch white, conspicuous. Dorsal tail color as back, lacking 
brownish tipped hairs; brush about one-third of tail length, fuscous. 
Ears not pigmented. 

Palatal ridges. — Pattern similar to G. andersoni (fig. 33). 

Glans penis and baculum.— Not observed. 

Cranial characters.— Figures 36, 37. Skull without strongly 
developed supraorbital ridge. Posterior portion of nasals tapering 
narrowly or "bottle shaped." Anterior margin of zygomatic plate 
slightly posterior to or reaching level of premaxillary-maxillary 
suture. Anterior margin of tympanic bulla reaching level of foramen 
ovale or beyond. Posterior margin of mastoid bulla beyond margin 
of paroccipital. Partition between anterior mastoid and superior 



118 FIELDIANA: ZOOLOGY 

posterior mastoid chambers clearly posterior to margin of hamular 
process of temporal. Interparietal shallow ^nd broad. 

Teeth. — Upper incisor grooved. Cusp pattern of molars as in other 
Gerbillus species (fig. 38). 

Feet — Palm and sole haired, pads and tubercles as in G. gerbillus 
(fig. 34). 

Measurements.— Table 6. Males slightly larger than females. Ear 
length averages less than one-half hind foot length. 

Age determination.— Adults are separated from immatures as in 
other species of Gerbillus on bases of tooth wear and suture closure. 

Variation. —Color varies from pale yellowish orange in the Wadi el 
Natroun area to light reddish orange in El Maghra. 

Comparisons.— Gerbillus perpallidus differs from G. gerbillus and 
G. andersoni in larger size and from the last in paler color and lack 
of dorsal stripe. Skins of young G. perpallidus can be distinguished 
from G. g. gerbillus by comparing the size of the white pygal area 
which is comparatively larger and extends further forward on the 
hip of the latter. It also differs in color from G. p. pyramidum, but 
not from all individuals of G. p. floweri. It differs from G. pyra- 
midum subspecies in less strongly ridged skull and greater inflation 
of mastoid chambers, and markedly from other Gerbillus species in 
shape of interparietal and posterior nasal shape (fig. 37). Anterior 
margin of zygomatic plate does not extend over the premaxillary- 
maxillary suture in G. perpallidus as in most G. pyramidum. The 
similarity between G. perpallidus and G. p. floweri is noted under 
the latter form, and characters of larger gerbils are summarized in 
Table 7. The validity of taxon G. perpallidus was tentatively con- 
firmed by chromosomal studies of Lay et al. (1975), who reported a 
2N = 40 and FN = 76 for the latter and a 2N=38 and FN = 76 for G. 
pyramidum from Egypt. 

Collection.— Dug from burrows in sand and trapped alive beside 
open burrows or with lines of traps in habitat. 

Habitats. — Idku: Coastal sand dunes. 

Western Mediterranean Coastal Desert: Coastal dunes of white, 
nummilitic sand (fig. 7); sandy areas in stands of Thymelaea hirsuta 
and Artemisia monosperma in the southern limits of vegetation 
(figs. 19, 20). 

Wadi el Natroun: Lake shore areas of mud and salty sand support- 



OSBORN&HELMY: MAMMALS OF EGYPT 119 

ing Typha sp. and Desmostachya bipinnata; almost barren sand 
slop)es with dry, ephemeral Mesembryanthemum sp.; soft sand 
sheets supporting stands of Panic um turgidum; dunes under exotic 
Prosopis juliflora; sandy slopes near clover fields, but not within the 
fields. 

Bir Victoria: Sand sheets and meanders with dominants of Arte- 
misia monosperma, Panicum turgidum, and Pityranthus tortuosa 
(fig. 10). 

El Maghra: Sand mounds around Nitraria retusa and Zygophyl- 
lum alburn^ barren sand and gravel slopes, and 15 km. W in soft 
sand in scattered Acacia raddiana. 

Reproduction.— One record of five fetuses in April. 

Associates.— Gerbillus gerbillus, G. andersoni, G. pyramidum, 
Jaculus jaculus, Meriones libycus, and M. shawi and, probably, M. 
crassus. 

Specimens examined.— Total 218. 

GIZA: Abu GhaUb (1), Abu Rawash (1). 

EL FAIYUM: Ezbet el Asfar (1). 

ALEXANDRIA: Idku (4). 

EL TAHREER: Cairo-Alexandria desert road km. 102 (3). km. 143 (1). 

BEHEIRA: Bir Victoria (27); wadi el Natroun (45); Wadi el Natroun V, to 1 km. E 
(9). Bir Hooker (5); Gebel Muluk (3). Zaghig (17); El Birigat 2 km. W (4). Kom 
Hamada (1). 

MATRUH: Burg el Arab (2); El Hawa 20 km. S of El Hamman (1); Qasr el Qatagi 
(1); Nakhlat el Barraq (1); El Maghra (72). 15 km. W (5); Bir Nahid (7); Qur el Hilab 
(7). 

Published records.— Records are from De Winton (1903 as G. 
tarabuli) and Setzer (1958d). 

BEHEIRA: Bir Victoria; Wadi el Natroun; Wadi el Natroun, Gebel Muluk. and 
Zaghig. 

Gerbillus andersoni De Winton, 1902 

Gerbillus andersoni De Winton, 1902, Ann. Mag. Nat. Hist., (ser. 7). 9, p. 45. 
Gerbillus eatoni Thomas, 1902, Proc. Zool. Soc., London, 2, pt. 1, p. 6. 
Gerbillus bonhotei Thomas 1919, Ann. Mag. Nat. Hist., (ser. 9). 3, p. 5. 

Type locality.— Egypt. ALEXANDRIA: Alexandria, El Mandara. 

General distribution.— Jordan, Northern Sinai Peninsula, Egypt, 
Libya, and Tunisia. 

Common names.— Anderson s Gerbil, Bayoudi. 




OS 



o 



120 



OSBORN&HELMY: MAMMALS OF EGYPT 121 

Distribution of subspecies in Egypt— Figure 39. Gerbillus under- 
soni bonhotei: Northern Sinai Peninsula; Gerbillus andersoni an- 
dersoni: Nile Delta and El Faiyum; Gerbillus andersoni inflatus: 
Northern Western Desert and Siwa Oasis. 

Diagnosis.— Brownish orange gerbil slightly larger than G. ger- 
billus, with ear and sole pigmented. Tail not bicolored at base; brush 
small, brownish. Whitish supraorbital, postauricular markings, and 
rump patch inconspicuous. Ear length equal to or greater than one- 
half of hind foot length. Bulla large. Incisive foramina relatively 
long, palatine foramina relatively short. Posterior margin of nasals 
truncate. 

Adult head and body length average 97 mm.; tail 126 mm., 130 
per cent of head and body length; foot 28 mm.; ear 15 mm.; occipi to- 
nasal length 28.8 mm.; weight 28 gm. 

External characters.— Dorsum brownish orange, darkest on 
rump. Side clear orange. Color of side extending onto upper foreleg 
and heel. Hairs of dorsum and portion of sides with gray bases. 
Underparts, feet, and distal portion of underside of tail white. 
Underside of tail base buffy. Broad, conspicuous band of dark- 
tipped hairs extending from mystacial area beneath eye to base of 
ear. Whitish postorbital and postauricular areas small, inconspicu- 
ous. White rump patch small. Dorsal tail color as back and with 
blackish hairs; brush fuscous or brownish, not conspicuous, and 
about one-fourth tail length. Ear and sole pigmented. 

Palatal ridges.— Figure 33. Pattern similar to G. gerbillus, but 
intermolar ridges somewhat thicker, with tips of first to third more 
medial and recurving. 

Glans penis and baculum.— These structures as described in 
Wassif et al. (1969) show little difference between species of Ger- 
billinae, except that the bacular shaft in G. andersoni broadens 
gradually into a wide, thickened base. 

Feet.— Palm and sole almost completely haired as in G. gerbillus 
(fig. 34). Sole pigmented. 

Cranial characters.— Figures 40, 41. Skull slightly larger than G. 
gerbillus, supraorbital ridge not well developed, not extending 
anterior to level of posterior plane of lacrimal bone. Posterior margin 
of nasals truncate. Incisive foramina relatively long and palatine 
foramina relatively short. Anterior margin of zygomatic plate never 
reaching level of premaxillary-maxillary suture. Parapterygoid 



122 



FIELDIANA: ZOOLOGY 




^ 






O 




Q.A.INFLATUS G.A.ANDERSONI G.GERBILLUS 

Fio 40. Comparison of auditory bullae in lateral and dorsal view (heavy lines) and 
skull shape in Gerbillus andersoni inflatus, G. a. andersoni, and 0. gerbiUus ger- 
billus. Lateral superior posterior mastoid chambers are shaded. G. a. bonhotei (not 
shown) is identical with G. a inflatus. 

fossa deep and partly closed. Anterior margin of tympanic bulla 
reaching level of anterior margin of foramen ovale. Posterior margin 
of mastoid chambers extending to and sometimes beyond level of 
occipital condyle, but not to level of supraoccipital. Posterior supjer- 
ior mastoid chamber (app. 2, fig. 165) more inflated than inferior 
chamber (fig. 40). Lip of external auditory meatus slightly thickened 
in adults. 

Teeth.— As in other species of Gerbillus (fig. 38). 

Measurements.— Table 8. Male and female dimensions subequal. 
Means (and ranges) of occipitonasal length (in millimeters) of 11 
adult males and 10 adult females, respectively: 29.4 (28.2 to 31.0) 
and 29.0 (27.7 to 29.8). Note that ear length averages half or greater 
than half of hind foot length. 

Age determination.— Adults are determined by tooth wear and 
suture closure as in G. pyramidum. 

Variation.— Color varies slightly from dark G. a. andersoni of the 
Nile Delta area through paler forms of G. a. inflatus inhabiting light- 
colored beach sand near Burg el Arab and pallid soils of Ras el 





^ 




G.ANDERSONI 



G.GERBILLUS 



Fig 41. Posterior margins of nasals (upper) and incisive and posterior palatal 
foramina (lower) of Gerbillus andersoni and G. gerbillus. 



123 



124 



FIELDIANA: ZOOLOGY 



Hekma, to darker individuals of that subspecies from more western 
localities. Specimens from northern Sinai are paler than those from 
the Nile Delta. 

Superior posterior mastoid and anterior mastoid chambers show 
more inflation in inflatus west of the Nile Delta and in bonhotei of 
northeastern Sinai (fig. 40, table 9). 



Table 9. — Comparison of inflation in mastoid chambers of bulla in Gerbillus 
andersonL 







Number 


Number with < 


degi 


Subspecies 


Locality 


examined 


Maximum M 


edii 


G. a. andersoni 


Baltim, Idku, other 
Delta localities 


26 


— 


— 


G. a. inflatus 


Cairo- Alexandria 
desert road km. 164 


3 


3 


— 




Burg el Arab 


16 


10 


3 




Ras el Hekma. 


21 


16 


5 




Mersa Matruh 










Buq Buq 


18 


12 


6 




Salum 


25 


14 


4 


G. a. bonhotei 


Northeastern Sinai 


5 


5 


— 



Minimum 
26 



Table 10. — Means (and ranges) of measurements, ratios, and weight of adult 
Gerbillus gerbillus. 

G. g. gerbillus G. g. asyutensis G. g. sudanensis 

HBL 90.0 (77-104) 164 88.4 (76-97) 108 84.0 (79-93) 30 

TL 124.0(107-137)161 124.0(100-143)104 117.4(91-128)29 

TL/HBL% 138.0(118.0-161.5)159 134.9(125.8-167.1)108 140.0(109.6-151.2)29 

FL 30.2 (28-32) 175 29.4 (26-32) 109 28.0 (25-30) 30 

EL 13.2(12.0-15.0)171 12.9(12.0-14.5)109 12.3(11.0-14.0)30 

Wt 24.2 (15.0-34.7) 112 22.1 (16.0-33.0) 49 19.2 (13.6-24.2) 17 

ONL 28.0 (25.8-30.0) 142 28.0 (25.6-29.5) 99 26.8 (26.4-28.4) 23 

ZW 15.2(13.7-16.2)122 15.1(14.1-16.0)79 14.5(13.9-15.1)17 

lOW 5.6 (5.0-6.4) 159 5.5 (4.9-6.1) 110 5.2 (4.7-5.7) 19 

BCW 13.5(12.9-14.2)156 13.4(12.8-13.9)109 13.0(12.7-13.5)24 

NL 10.4 (9.2-1 1.4) 132 10.3 (9.5-1 1.4) 95 9.9 (9.3-10.7) 23 

IFL 4.4 (3.7-4.9) 156 4.3 (3.8-4.8) 100 4.1 (3.8-4.6) 27 

AL 3.9(3.4-4.6)155 3.7(3.2-4.2)101 3.6(3.2-4.1)27 

RW 3.8 (3.6-4.2) 158 3.8 (3.5-4.3) 109 3.6 (3.4-3.9) 20 

BL 8.5 (7.9-9.3) 154 8.4 (7.4-9.2) 103 8.3 (7.8-8.8) 22 

SH 11.2(10.6-12.0)142 11.2(10.5-11.9)99 10.9(10.5-11.9)20 



OSBORN & HELMY: MAMMALS OF EGYPT 



125 



Dimensions given in Table 8 indicate negligible differences be- 
tween subspecies except in alveolar and bulla lengths. 

Comparisons.— Gerbillus andersoni is distinguishable from G. 
gerbillus by darker coloration, smaller white rump patch, pigmented 
ear and sole, base of tail usually not bicolored, longer ear length 
relative to hind foot length, posterior margin of nasals truncate 
rather than round or pointed, longer incisive foramina, shorter pala- 
tine foramina, shaft of baculum not distinct from base (Wassif et al., 
1969), and other characters listed in Table 11 and shown in Figures 
40-42, From Dipodillus campestris, which it resembles superficially, 
G. andersoni differs in having palm and sole hairy rather than bare, 
smaller tail brush, greater amount of white hair on rump, larger 
auditory bulla, opposite rather than alternate arrangement of first 
lingual and labial cusps of first upper molar, narrower basioccipital, 

Table 11. — Comparison of characters of Gerbillus andersoni and G. gerbillus. 



Character 


G. andersoni 


G. gerbillus 


Dorsum 


darker (brownish orange) 


lighter (orange) 


Ear 


pigmented 


not pigmented 


Sole 


pigmented 


not pigmented 


Mystacial and 
suborbital areas 


with pigmented hairs 


without pigmented hair 


Supraorbital areas 


inconspicuous 


conspicuous 


White rump patch 


half size of gerbillus 


twice size of andersoni 


Tail 


not bicolored at base 
brush small, dark 


bicolored 
brush large, pale 


Posterior margin 
of nasals 


truncate 


round or pointed 


Mastoid bulla 


superior posterior 
chamber more inflated 


inferior posterior 
chamber more inflated 


Incisive (anterior 
palatine) foramina 


longer 


shorter 


Posterior palatine 
foramina 


shorter 


longer 


Supraorbital ridge 


not anterior to posterior 
plane of lacrimals 


anterior to posterior 
plane of lacrimals 


Ear length 


half or slightly more than 
half hind foot length 


less than half hind foot 
length 


Diploid chromosome 
number (Wassif et al., 
1969) 


40 


male 43 
female 42 



126 FIELDIANA: ZOOLOGY 

deeper parapterygoid fossa and other characters in Figures 33, 34, 
40. Tables 8 and 12. 

From G. pyramidum, G. andersoni differs in having relatively 
more inflated bulla, smaller dimensions, less conspicuous dorsal 
stripe and much smaller and less conspicuous tail brush, although 
De Winton (1902a, p. 45) described it as "a miniature of G. 
pyramidum/' From G. perpallidus, G. andersoni differs in having 
darker coloration, smaller white rump patch, sole and ear 
pigmented, posterior margin of nasals truncate, and averaging less 
in all dimensions. Further comparisons can be made by examining 
Figures 37 and 41 and Tables 6 and 8. 

For discussion of chromosomal variation, see under Comparisons 
in G. pyramidum. 

Remarks.— Synonymy of G. andersoni with G. eatoni and G. bon- 
hotei is based on similarity in coloration, dimensions (table 9), and 
other features in common. Relationships between G. andersoni and 
G. bonhotei were mentioned by Thomas (1919b), Flower (1932), 
EUerman (1948), Ellerman and Morrison-Scott (1951), and Wassif 
(1953b). Note under G. g. asyutensis the inclusion of Sinai Peninsula 
specimens that were misnamed G. gerbillus bonhotei by Setzer 
(1958d). Cockrum et al. (1976a) verified the conspecificity oi ander- 
soni and eatoni. 

Collection.— Dug from burrows in sand or captured with live 
traps. 

Habitats. —Sinai Peninsula: Sandy areas in the northeast. 

Nile Valley and Delta: Sandy areas in palm groves. Vegetated, 
cultivated, and noncultivated semidesert. 

Oases (Faiyum, Wadi el Natroun, and Siwa): Vegetated sandy 
areas. 

Western Mediterranean Coastal Desert: Coastal dunes of white, 
nummulitic sand. Dunes adjacent to salt marsh in Atriplex halimus. 
Salt marsh beside Limoniastrum monopetalum (fig. 7). Sandy and 
rocky slopes supporting Lycium sp., Noaea mucronata, Thymelaea 
hirsuta, and Onopordon alexandrinum (a thistle) where it was usu- 
ally trapped beside Lycium bushes (fig. 48). Depressions in sandy 
loam supporting almost pure stands of Artemisia inculta. Sandy 
areas within stands of Thymelaea hirsuta, Anabasis articulata, and 
Artemisia monosperma and further inland in discontinuous patches 
of A. monosperma (figs. 19, 20). 



OSBORN & HELMY: MAMMALS OF EGYPT 



127 



e. AuotHsom % 
a. otKniLus O 



• 9^ 



• • 






>--^|^^»?>V i 






°o^ 



goo o o 
o 



Occipito-nasal Length 

Fig. 42. Scatter diagram of incisive foramina length versus occipitonasal length 
in Gerbillus andersoni and G. gerbillus. Half-circles represent individuals of both 
species at same point. Data from all age groups. 

Ranck (1968) mentioned finding the species (under G. eatoni) in 
areas devoid of sand. It is also found in similar situations in Egypt. 
Gerbillus andersoni appears to be more numerous in the Mediter- 
ranean Coastal Desert vegetation and coastal sands than G. gerbil- 
lus and "does not inhabit more rigorous desert areas" (Hoogstraal, 
1963, p. 10) as does the latter. 

jBe/iafior.— Nocturnal. More docile in captivity than G. gerbillus. 

Burrows.— Not distinguishable from those of G. gerbillus. 

Reproduction.— Scattered data from September through June 
indicate a rather long breeding season. Mean (and range) of litter 
size from 10 females captured during September, October, and June 
is 3.9 (3 to 7). 

Sex ratio.— In a sample of 251 museum specimens, males num- 
bered 127 (50.6 per cent), and females numbered 124. 

Associates.— Gerbillus andersoni lives in association with G. ger- 
billus, G. perpallidus, Meriones shawi, Dipodillus amoenus, D. 
simoni, probably G. pyramidum and Jaculus orientalis, and occa- 
sionally D. campestris and Mus musculus. It may also live in close 
proximity with Psammomys obesus, which burrows beneath 
Lycium bushes (fig. 48). 

Key to Egyptian Subspecies of Gerbillus andersoni 

1. Mastoid chambers not prominently inflated (fig. 40). Dorsum dark. (Nile Delta 
and El Faiyum) andersoni, p. 128. 



128 FIELDIANA: ZOOLOGY 

2. Mastoid chambers prominently inflated (fig. 40). 

a. Dorsum dark. (Western Desert) inflatus, p. 128. 

b. Dorsum paler (Sinai Peninsula) V bonhotei, p. 129. 

Gerbillus andersoni andersoni De Winton, 1902 

Type /oca/ity. -Egypt. ALEXANDRIA: Alexandria, El Mandara. 

Distribution in Egypt.— Figure 39. Nile Delta, south to Helwan; 
El Faiyum. 

External characters.— See species description. Generally darker 
than other subspecies. 

Cranial characters.— Figure 40. Bulla less swollen than in other 
subspecies, particularly mastoid chambers (table 9). 

Measurements.— Table 8. Dimensions about equal to those of 
inflatus and bonhotei, except for longer nasal length, shorter alve- 
olar length, and shorter bulla length in andersoni. 

Variations.— One albinistic specimen was collected from Imbaba, 
Giza Governorate. 

Specimens examined.— Total 89. 

DAM I ETTA: Damietta W of (6). Kafr el Battikh (2). 

KAFR EL SHEIKH: Baltim (19). El Burg (8). Lake Burullus eastern shore (2). 

ALEXANDRIA: DikheUa airfield 0.8 km. W (1); El Mandara (Type); Abu Qir (2); 
El Muntaza 0.8 km. W (1), 0.8 km. E (1): Amiriya (2). 

MUNUFIYA: Quweisna (6). 

BEHEIRA: Rosetta (1). Lake Burullus area (6). Idku (12). Kom Hamada (2). Kafr 
Dawud (2). El Khatatba (3). 

GIZA: Wardan (3); Abu Rawash (2); Cairo- Faiyum road km. 3(1). km. 30 (3). 

CAIRO: Helwan (2). 

EL FAIYUM: Faiyum (1). Kom O Shim (1). 

Published records.— Records are from De Winton, (1902a, 1903), 
Innes (1932), EUerman (1949), and Setzer (1952, 1958d). 

DAMIETTA: Damietta. Kafr el Battikh. 

KAFR EL SHEIKH: Burullus W of Damietta. Baltim. El Burg. 
MINUFIYA: Quweisna. 

ALEXANDRIA: Idku. El Mandara, Dikheila airfield 0.8 km. W. Muntaza 0.8 km. 
E. Amiriya. 
BEHEIRA: Rosetta, Kom Hamada, Kafr Dawud. EI Khatatba. 
GIZA: Wardan. 

Gerbillus andersoni inflatus (Ranck, 1968) 

Gerbillus eatoni inflatus Ranck. 1968. U.S. Nat. Mus. Bull.. No. 275. p. 97. 

Type locality.-Uhya. CYRENAICA: Fort Cappuzo 10 km. SW. 



OSBORN&HELMY: MAMMALS OF EGYPT 129 

Distribution in Egypt— Figure 39. Western Mediterranean 
Coastal Desert west of Nile Delta to northeastern Libya; Wadi el 
Natroun and Siwa Oasis. 

External characters.— Brov/nish orange dorsally; differs little 
from nominate subspecies. Pale individuals occur on light-colored 
coastal sands and pallid soils of Ras el Hekma and sand sheets in 
Wadi el Natroun. 

Cranial characters.— Figure 40. Mastoid chamber of auditory 
bulla more inflated than in andersoni, but about the same as in bon- 
hotel (table 9). 

Measurements.— Table 8. Dimensions about equal to those of the 
nominate subspecies, except for greater bulla and alveolar lengths 
and shorter nasal length. Means (and ranges) of measurements (in 
millimeters) from Ranck (1968) of five adult males from the type 
locality are: total length 226 (218 to 236), tail length 125 (121 to 
131), hind foot length 27.2 (26 to 28), ear length 15.4 (15 to 16), occip- 
itonasal length 30.6 (29.9 to 31.7), zygomatic width 16.1 (15.6 to 
16.4), interorbital width 5.9 (5.5 to 6.5), and nasal length 11.7 (11.4 
to 12.3). 

Specimens examined.— Total 178. 

TAHREER: Cairo-Alexandria desert road km. 90 (1). km. 164 (8). km. 195 (1). 

BEHEIRA: Wadi el Natroun (3). 

MATRUH: Alexandria-Salum road km. 54 (8); Lake Mariut (1); Bahig 33.6 km. S 
(6). 38.4 km. S (1). 51.2 km. S (2); El Alamein (1); Abu Haggag 3.2 km. E (1); Ras el 
Hekma (38); Mersa Matruh (16), 32 km. E (2); Matruh-Qara desert road 18 km. S (3); 
Zawyet el Mithniyan (3); Sidi Barrani (1), 32 km. E (3), 19.2 km. E (2). 19.2 km. S (2), 
4.8 km. S (1), 19.2 km. SW (5), 32 km. SW (1). 52.8 km. W (1); Buq Buq 1 1.2 km. SW 
(17): Salum (5), 24 km. E (1). 22 km. E (4). 19.2 km. E (6). 18.7 km. E (9), 16 km. E (13), 
10 km. E (1), 8 km. SE (1). 9.6 km. S (1), Bir Bosslanga (Bir Wair) (1); Siwa Oasis (3). 

Libya. CYRENAICA: Fort Capuzzo 10 km. SW (5). 

Published records.— Records are from EUerman (1949), Setzer 
(1958d), and Ranck (1968). 

Egypt. TAHREER: Cairo-Alexandria desert road km. 195 (junction of Alexandria 
and Mersa Matruh roads). 

MATRUH: Burg el Arab; Mersa Matruh; Sidi Barrani 19.2 km. SW. 32 km. E, 
52.8 km. W. 

Libya. CYRENAICA: Fort Capuzzo 10 km. SW. 

Gerbillus andersoni bonhotei (Thomas, 1919) 
Type locality.— Egypt. SINAI: Khubra Abu Guzoar. 
Distribution in Egypt.— Figure 39. Northeastern Sinai Peninsula. 



130 FIELDIANA: ZOOLOGY 

External characters. — Palest of three subspecies; being more 
similar in color to G. gerbillus. 

Cranial characters.— The outstanding feature separating bonhotei 
from andersoni is greater inflation of bulla, in which it is comparable 
to inflatus in Figure 40 and Table 9. 

Measurements.— Table 8. Dimensions somewhat smaller in 
general than other subspecies, but sample is smallest of three and 
data may not be too reliable. Bulla length, however, appears to be 
slightly longer in bonhotei than in other subspecies. 

/?emar/js.— Setzer (1958d) and then Hoogstraal (1963) erroneously 
applied G. gerbillus bonhotei to Sinai and Suez specimens of G. ger- 
billus asyutensis. 

Specimens examined.— Total six. 

SINAI: Khubra Abu Guzoar. (Type. 3). Wadi Hareidin (1), Gebel Lehfan (1). 

Published records.— Records are from Thomas (1919b), Flower 
(1932). Wassif (1953c). 

SINAI: Khubra Abu Guzoar, Wadi Hareidin, Gebel Lehfan. 
Gerbillus gerbillus (Olivier, 1801) 

Dipus gerbillus Olivier. 1801, Bull. Sci. Philom. Paris, 2. p. 121. 

Type locality.— Egypt. GIZA: Probably near the pyramids. 

General distribution. — Israel, Sinai Peninsula, Egypt, Libya, 
Sudan, Uganda, parts of Niger, Mauritania, Chad, and Mali. 

Common names.— Lesser Gerbil, Bayoudi. 

Distribution of subspecies in Egypt.— Figure 43. Gerbillus gerbil- 
lus asuytensis: Sinai Peninsula, northern part of Eastern Desert; 
Gerbillus gerbillus sudanensis: Southern part of Eastern Desert; 
Gerbillus gerbillus gerbillus: Nile Delta and Western Desert. 

Diagnosis. —Small yellowish orange gerbil with ear and sole not 
pigmented. Tail bicolored; brush moderately conspicuous, grayish 
to brownish. White supraorbital and postauricular markings and 
white rump patch prominent. Ear length less than one-half hind foot 
length. Skull with large bulla, incisive foramina relatively short, 
palatine foramina relatively long, and posterior margin of nasals 
round or pointed. 

Adult head and body length average 88 mm.; tail 123 mm., 138 



OSBORN & HELMY: MAMMALS OF EGYPT 



131 



25* 26* 27* 28* 2 9* 30* 31* 32* 3 3* 3 4* 35* 36* 37* 




Fig. 43. Collection localities of Gerbillus gerbillus gerbitlus (circles), G. g. asyuten- 
sis (dots), and G. g. sudanensis (squares). 

per cent of head and body length; foot 30 mm.; ear 13 mm.; occipito- 
nasal length 28.0 mm.; weight 23 gm. 

External characters.— Figure 44. Upper parts pale yellowish 
orange to reddish orange. Dorsum slightly darker than side, con- 
trasted further in some specimens by brownish tipped hairs, especi- 
ally on rump. Color of side not extending onto foreleg or further 
than thigh on hind limb. Hairs of dorsum with gray bases, of side 
with white bases. Hair of underparts, feet, and entire ventral sur- 
face of tail white. Mystacial, suborbital, supraorbital, and 
postauricular areas white. White hairs nearly circumorbital in 
palest specimens. White rump patch large, conspicuous. Tail com- 
pletely bicolored; dorsal color as back; brush fairly conspicuous, 



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ft ^ 




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^ 1 ~n 




'" ^^B^ 


• ^ 




./> B 


e • ^E 


^^B 


^^E 


1 

•:sl 






• -fH 


^^^^^^^^^^^v 


^^V^^^^^^^E 


t 




^H ^^^B^ 


' ^n 


K^Ji^^^^^^^l 


^^^Hk ^^^^K'' 


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Fig. 44. Cadaver of Gerbillus gerbillus. 



132 



OSBORN&HELMY: MAMMALS OF EGYPT 133 

I grayish to brownish, and slightly more than one-third of tail length. 
r Ear margin blackish, but ear not pigmented. 

Palatal ridges.— Figure 33. Diastemal ridges broadly U-shaped; 
I first to third intermolar ridges recurved; fourth intermolar small, 
crenulated; fifth directed medially, also crenulated. 

Glans penis and haculum.—See under G. pyramidum and G. 
andersoni. 

Feef.— Figure 34. Palm and sole almost completely haired. Front 
foot with one large, soft postidigital pad bearing proximal accessory 
lobe. Hind foot with large, lobed postdigital pad and indistinct 
posthallucal tubercle. Sole not pigmented. 

Cranial characters.— Figure 40. Skull smallest of Egyptian species 
of Gerbillus. Supraorbital ridge well developed, extending anterior 
to level of posterior plane of lacrymal bone. Bulla more inflated than 
in other species of Gerbillus. Posterior margin of nasals round or 
pointed. Incisive foramina relatively short; palatine foramina rela- 
tively long (fig. 41). Anterior margin of zygomatic plate sloping 
anteriorly and usually reaching level of premaxillary-maxillary 
suture. Parapterygoid fossa deep, partly closed. Anterior margin of 
tympanic bulla reaching level of anterior margin of foramen ovale. 
Posterior margin of mastoid chambers extending beyond occipital 
condyle, but not to level of supraoccipital. Lateral inferior posterior 
mastoid chamber more inflated than lateral superior posterior 
chamber (fig. 40). Lip of external auditory meatus slightly thickened 
in adults. 

Teeth.— Figure 38. See description under G. pyramidum. 

Measurements.— Table 10. Smallest Gerbillus species in Egypt. 
Male and female dimensions subequal. Means (and ranges) of occipi- 
tonasal length (in millimeters) of 10 adult males and 10 adult fe- 
males, respectively, are 27.9 (26.7 to 29.3) and 27.4 (26.7 to 28.2). 
Note that ear length averages less than one-half hind foot length. 

Age determination.— Adults are determined by tooth wear and 
suture closure as in G. pyramidum. 

Variation.— Color varies from pale yellowish orange through 
orangish and reddish orange in Western Desert G. g. gerbillus. Indi- 
viduals from oases and western Nile Valley and Delta localities are 
slightly darker. Reddish orange specimens are from southern Wadi 
el Gedeed (Bir Qiseiba, Bir Kurayim, and Bir el Shab). 



134 FIELDIANA: ZOOLOGY 

Samples from Sinai Peninsula and northern Eastern Desert, 
assigned to G. g. asyutensis, are slightly paler than G. g. gerbillus 
and have brownish hairs on dorsum and rump. Specimens from 
southern Eastern Desert, assigned to G. g. sudanensis, are darker 
than G. g. asyutensis and G. g. gerbillus from west bank of the Nile. 

Size variation shows no definite directional trends among Western 
Desert samples. In the Eastern Desert, there is a decrease in most 
dimensions from north to south between G. g. asyutensis and G. g. 
sudanensis (table 10). 

Co mpansons.— Differences between G. gerbillus and G. andersoni 
are discussed under the latter, listed in Table 11, and shown in Fig- 
ures 40-42. From G. perpallidus and G. pyramidum, G. gerbillus dif- 
fers distinctly in dimensions (tables 7, 10) and shape of posterior 
margin of nasals (figs. 37, 41), and from G. pyramidum, in much 
paler color. From Dipodillus campestris, G. gerbillus can be distin- 
gfuished by paler color, restriction of color in facial region, hair on 
soles, larger auditory bulla, shorter incisive foramina, opposite 
rather than alternate arrangement of first lingual and labial cusps 
of m', and narrower basioccipital. 

Co//ection.— Easily trapped or dug from burrows in sand. 

Habitats. —Sinai Peninsula: Dunes and alluvium in wadis. Limit- 
ed, according to Haim and Tchernov (1974), to elevations below 
1,100 m. and correlated with the distribution oi Hammada elegans. 

Nile Valley and Delta: Sand patches in palm groves and cultivated 
areas. Sandy areas of semidesert supporting bunch grasses (Stipa- 
grostis scoparia and Panicum turgidum), reeds (Phragmites aus- 
tralis), and shrubs such as Heliotropium digynum (Briscoe, 1956; 
Yunker and Guirgis, 1969). 

Eastern Desert: Patches of windblown sand in wadis; sand and 
fine gravel accumulations around trees and shrubs (fig. 15); patches 
of prostrate Citrullus colocynthis (bitter melon), as noted also by 
Bauer (1963), together with the jird, Meriones crassus; and littoral 
areas in sand mounds around Nitraria retusa. Common where sand 
accumulates beside houses and in sand littered with empty tins, 
broken pottery, and remains of reed matting. 

Western Mediterranean Coastal Desert: Coastal dunes of white 
nummilitic sand (fig. 7). Adjacent to and sometimes in salt marshes. 
Sandy areas within the coastal vegetation. Incidentally, G. ander- 



OSBORN & HELMY: MAMMALS OF EGYPT 135 

soni is more numerous than G. gerbillus throughout the Httoral 
desert. 

Western Desert: Wherever there are plants and sand (figs. 9, 10), 
although seldom in clumps of Tamarix sp. Occasionally found in 
barren areas far from vegetation where windblown detritus appar- 
ently provides food. 

Oases: Vegetated, sandy areas similar to habitats described 
above, and debris beneath palm trees. 

In all barren or vegetated desert areas, campsites attract G. ger- 
billus. As G. pyramidum, G. perpallidus, and Meriones crassus, it is 
also attracted to camel dung. 

Be^aL'ior.— Nocturnal. Very nervous, bites when first handled, 
but becomes quite tame with repeated handling. 

Burrows.— Yunker and Guirgis (1969) studied gerbil burrows in 
the desert and semidesert near Cairo. Burrows in desert were in flat 
sandy areas, rarely under plants. Burrows in semidesert were some- 
times dug among roots of plants. Occupied burrows were plugged 
with sand during the warm period of the year (March through 
December) and open during the cooler wet season (January and Feb- 
ruary). A typical burrow reached a depth of 30 to 60 cm. Desert bur- 
rows were deeper (50 to 80 cm.) during the cool period. The main 
passage ranged from about 50 cm. to about 4.5 m. with one to four 
short passages, some ending blindly and containing food caches, 
nest materials, and/or gerbils. 

Burrow microclimate.— Despite extreme temperature fluctua- 
tions in sand and outside air, Yunker and Guirgis (1969) found that 
temperature in burrows varied only a few degrees during a 24-hour 
period. Relative humidity (RH) tended to follow the 24-hour curve of 
readings for outside air, but at a 30 to 60 per cent higher average 
value. On January 24 and 25, outside air temperatures ranged from 
47 to 72 degrees F, surface sand, 46 to 82 degrees F, whereas burrow 
temperatures were 57 degrees ± 7 in semidesert and 56 degrees ± 2 
in desert. On May 23 and 24, outside air RH ranged from 15 to 88 
per cent, whereas burrow RH was 81 to 100 per cent. From January 
to August, semidesert burrow RH averaged 10 per cent higher than 
desert burrow RH. 

Food— Although Bodenheimer (1935) stated that gerbils feed on 
seeds, roots, and insects, no locusts, crickets, butterflies, or beetles 



136 FIELDIANA: ZOOLOGY 

placed in cages with Egyptian G. gerbillus were killed or eaten. 
Food in the desert is mainly seeds, leaves, buds, and fruits. Camel 
dung is torn apart by gerbils in search of seeds and fibers. Schmidt- 
Nielsen (1964) reported that this rodent can live on dry food. The 
physiologic features of water metabolism on this diet was studied 
by Burns (1956). 

Food items and remnants in semidesert burrows consisted of date 
seeds; camel droppings; and seeds, spikes, and husks of cereal 
grains and other plants. Dry seeds were thought to be the staple 
food of desert gerbils by Yunker and Guirgis (1969). 

Associates.— Wthongh. G. gerbillus, G. andersoni, G. perpallidus, 
and G. pyramidum are sometimes collected in the same habitat, 
behavioral and other relationships among them are unknown. Other 
species occurring with G. gerbillus are Jaculus jaculus and, in some 
areas, Dipodillus amoenus, Pachyuromys duprasi, Meriones 
crassus, and M libycus. Burrow inhabitants listed by Yunker and 
Guirgis (1969) include lizards and toads (in semidesert only) and 
about 44 species in 14 orders of arthropods, mostly in semidesert 
burrows. 

Reproduction.— From evidence of swollen, descended testes of 
males and pregnant and lactating females, the breeding period ap- 
pears to be January through May. Mean (and range) of litter size of 
seven females captured during April and May was 4.3 (3 to 6). 

Sex ratio.— In a sample of 247 museum specimens of lesser ger- 
bils, males numbered 138 (56 per cent) and females numbered 109. 

Commensalism.— Readily enters permanent or temporary dwell- 
ings in search of food or shelter (Flower, 1932, p. 416). 

Key to Egyptian Subspecies of Gerbillus gerbillus 

1. Dorsum hairs brownish tipped; pale, yellowish orange. Dimensions about as in 

gerbillus. (Sinai Peninsula and northern part of Eastern Desert) 

asyutensis, p. 136. 

2. Dorsum hairs not brownish tipped. 

a. Dark, clear orange. Size smaller. (Southern part of Eastern Desert) 

sudanensis, p. 138. 

b. Orange to reddish orange. Size larger. (Western Desert) gerbillus, p. 139. 

Gerbillus gerbillus asyutensis Setzer, 1960 

Gerbillus gerbillus asyutensis Setzer, 1960, J. Egypt. I'ubl. Health Assn., 33, No. 
1, p. 3. 

Type locality. -Egypt. ASYUT: Wadi Asyuti. 



OSBORN&HELMY: MAMMALS OF EGYPT 137 

Distribution in Egypt.— Figure 43. Sinai Peninsula and northern 
part of Eastern Desert. 

External characters.— Dorsum pale yellowish orange with con- 
spicuous brownish-tipped hairs, particularly on rump. 

Cranial characters.— See species description. 

Measurements. — Table 10. 

Variation.— There is slight variation in size among samples of G. 
g. asyutensis. Between samples of this subspecies and G. g. 
sudanensis, however, there is a noticeable decrease in the means of 
most measurements. 

Color varies from very pale in Wadi Asyuti (type locality) to 
darker along the Gulf of Suez and Red Sea coast. Samples from 
northwestern Eastern Desert are slightly darker than those from 
Sinai. Scattered individuals from wadis east of the Red Sea Hills are 
as pale as Wadi Asyuti sample described by Setzer (1960b). 

Comparison.— Of 38 G. g. gerbillus specimens from the north- 
eastern part of the Western Desert, three were as pale as Eastern 
Desert G. g. asyutensis. Of 51 specimens of G. g. asyutensis from 
the northern part of the Eastern Desert, six were as dark as Western 
Desert G. g. gerbillus. 

The only difference between G. g. asyutensis and the nominate 
subspecies is slightly paler basic coloration and conspicuous dark- 
tipped hairs on the dorsum, particularly on the rump. Size difference 
between these two subspecies is negligible (table 10). 

Between G. g. asyutensis and G. g. sudanensis, there are more 
noticeable differences in color and size. The latter is darker and 
averages smaller in nearly all dimensions (table 10). 

Remarks.—Setzer's (1959d) application of the trinomen bonhotei 
to G. gerbillus from Sinai was in error. Specimens recognizable as G. 
bonhotei Thomas are synonymous with G. andersoni De Winton. 
The subspecies of G. gerbillus in Sinai is G. g. asyutensis. 

Specimens examined.— Total 236. 

SINAI: El Arish (7). Abu Aweigila (2), El Quseima (7), Abu Zenima (1). Ras Abu 
Rudeis (2). Wadi Sidr (4). Feiran Oasis (1). Ayun Musa (1). 

ISMAILIA: Fayid 4.8 km. NW (13). Ismailia (1). 

SUEZ: Cairo-Suez road km. 29 (1). Maadi 32 km. E (1), Wadi el Gafra (25). Wadi 
Iseili (8). Suez (1), Ain Sukhna (5). Wadi Nakhl (1). Wadi Abu Seyala (2). Wadi Qiseib 
(1). Wadi Dom (1). 



138 FIELDIANA: ZOOLOGY 

RED SEA: Ras Abu el Darag 1 km. N (1); Wadi el Nil (231: Bir Zafarana (6); Ras 
Zafarana (9): Wadi Araba (3); Bir Abu Shaar (5): Wadi Abu Shaar (2): Wadi BaU (3); 
El Ahiah (21: Hurghada (1). 12 km. S (3). 14 km. S (1). 20 km. S (1): Wadi el Qreiya, 
QenaSafaga road km. 77 (4): Wadi Abu Sheeh. Qena-Safaga road km. 80 (3): Wadi 
Umm Seleimat (2): El Kanayis (2): Wadi Umm Huweiut (1): Wadi Abu Quraiya (2): 
Disht el Daba road (2): Safaga 6.4 km. S (6): Abu Kharif mine area (5): Wadi Abu 
ZawU (4): Bir Abu Zawil (3). 6.4 km. W (6): Wadi Fatira (2): Wadi Abu Ziran (3). 

SHARQIYA: Bilbeis (1). 

CAIRO: HeUopoUs 8 km. E (1). 12.8 km. E (9); Wadi Digla (1); Helwan (5). 

ASYUT: Wadi Asyuti (20). 

QENA: Wadi Qena. el Saqiya (1): Luxor (8). 

Published records.— Records are from Anderson (1902), Allen 
(1915), Flower (1932), Wassif (1953b), Wassif and Hoogstraal (1954), 
Setzer (1952, 1958, 1960b), and Hoogstraal (1963). 

SINAI: El Arish. Abu Aweigila, El Quseima, Ras Abu Rudeis, Abu Zenima 8 km. 
N, Feiran Oasis 14 km. W, Tor: Ayun Musa, Wadi Gharandal (Shurandel), Bir el 
Suweir (Suweira). 

SUEZ: Suez, Fani 

CAIRO: Heliopolis 8 km. E. Helwan 2 km. SE. 

ASYUT: Wadi Asyuti. 

Gerbillus gerbillus sudanensis Setzer, 1956 

Gerbillus gerbillus sudanensis Setzer, 1956, J. Egypt Publ. Health Assn., 33, p. 
220. 

Type locality. -Sudan. KASSALA: Port Sudan. 

Distribution in Egypt— Figure 43. Southern part of Eastern 
Desert. 

External characters.— Dorsum dark, clear orangish. 

Cranial characters. —See species description. 

Measurements.— Table 10. Smallest Egyptian subspecies of G. 
gerbillus. 

Comparison.— See subspecies gerbillus and asyutensis. The zone 
of asyutensis and sudanensis intergradation is between Luxor and 
Aswan and extends eastward to the Red Sea. There is no intergra- 
dation with G. g. gerbillus. 

Specimens examined.— Total 63. 

RED SEA: Wadi Hodein (I), Bir Abraq (4). 

SUDAN ADMINISTRATIVE: Wadi Adeib (5): Bir Kansisrob (I), 1.6 km. N (1|; 
Abu Ramad (1). 

ASWAN: Kom Ombo (6). Muneiha (8), Adindan (1), Armina Temple (1), Wadi Or 
(1). QustuI (II. AUaqi 11.2 km. S (8), Wadi AUaqi (1), Wadi Umm Qareiyat (3), Wadi 



OSBORN&HELMY: MAMMALS OF EGYPT 139 

Nagib (5). Wadi Haimur mine area (2), Bir Murra 3.2 km. N (2), Wadi Abusku (4), 
Wadi Quleib (6), Gebel Magal Gabril (1). 

Published records.— Records are from Hoogstraal et al. (1957b), 
Setzer (1958d, 1960b), and Bauer (1963). 

RED SEA: Wadi Hodein. Bir Abraq. 

SUDAN ADMINISTRATIVE: Wadi Adeib. Bir Kansisrob 1.6 km. N. 

ASWAN: Aswan 1.6 km. SE. 

Sudan. NORTHERN: Wadi Haifa, Khor Musa Pasha. 

Gerbillus gerbillus gerbillus (Olivier, 1801) 

Type locality.— Egypt. GIZA: Probably near the pyramids. 

Distribution in Egypt— Figiire 43. Nile Delta and Western 
Desert. 

External characters.— Dorsxim orangish to reddish orange and 
usually lacking brownish tipped hairs. 

Cranial characters.—See species description. 

Measurements.— Table 10. See species description. 

Variation.—See species description. 

Comparison.— Differs from G. g. asyutensis in darker, clearer, and 
more orangish dorsum; and lack of brownish tipped hairs. Dimen- 
sions average slightly larger (table 10). 

Differs from G. g. sudanesis in slightly paler dorsum and averages 
considerably larger in most dimensions (table 10), 

Specimens examined.— Total 500. 

ALEXANDRIA: El Amiriya (1). 

TAHREER: Cairo-Alexandria desert road km. 110, 0.5 km. E (4), km. 143 (3). 

BEHEIRA: Abu el Matamir (1); El Khatatba (2); El Birigat (1); Kafr Dawud (2); 
Kom Hamada (5), Bir Victoria (5); Wadi el Natroun (31), 5 km. W (9). 

MATRUH: Alexandria-Salum road km. 54, 0.5 km. N (1); Bahig (1), 42 km. S (4), 51 
km. S (7); Abu Mena E of (1); Burg el Arab (1); El Hawa 20 km. S of El Hamman (1); 
El Quweirat el Sud (1); Qasr el Qatagi (2); Nakhlat el Barraq (6); El Maghra (27), 120 
km. S (10); Bir Nahid (5); Wadi Labaq (6); Minqar Abu Dweiss area (3); Camel Pass 
Dune area (6); Mersa Matruh 19 km. E (3); Salum 10 km. E (2), 16 km. E (1), 18 km. E 
(2), 19 km. E (5); Sidi Omar (1); French Camp No. 2(1); Qara Oasis (1); Siwa Oasis (2); 
Aghurmi 5 km. E (3); El Maragi (5); Bahrein (6); Ain el Dakrur (10); Ain el Baqar (1). 

GIZA: Cairo-Alexandria desert road km. 10 (1), km. 12 (1), km. 31 (1); Abu Ghalib 
(7): Abu Rawash (6); Abu Sir (4); El Mansuriya (5); Giza Pyramids (3); Mena (1); El 
Qatta (2); Sakkara (2); Cairo-Bahariya Oasis road km. 20 (2); Cairo-Bahariya Oasis 
Track km. 208, acacia grove area 6 km. SE (1); Bahariya Oasis, Bir Qasr No. 1 (40), 
No. 2 (7), No. 3 (13). El Hara (10), Bawiti (3), 14 km. S (6); El Aguz (6); Ain Marun (2); 
Ain el Beilda (8); Wadi Ghorabi (7); Uyun Tab-Limun (9); Hatiyet Tabany (1). 



140 FIELDIANA: ZOOLOGY 

EL FA I YUM: Shooting club (11. Qasr Rashwan (U. Tamiya (1). Wadi Muwellih 
<15). 

MINYA: Beni Mazar (1). Hatiyet el Sunt (2). 

ASYUT: Beni Adi (8|. 

QENA: Dandara 6 to 8 km. S (1). 

ASWAN: Kurkur Oasis (2). Seiyala (2|. Abu Simbel (1|. 

EL WADI EL GEDEED: Farafara Oasis. 4.8 km. N (2); El Khanafis (2); Ain El 
Tinnin (2); Ain Besai (3); Ain el Wadi (2). Qokshira (2); Wadi Hennis (3); Batras 6.4 
km. E (2); El Kharga 60 km. N (2); Kharga Oasis (4); Ganah (2); El Gezira (8); Ezbet 
Muhib (4); Nasser village (3); Ain Eede (11); El Mahariq (2); Ganah (2); Baris (3), 17 
km. S (8): Bir Qiseiba (19); Bir Kurayim (17). 1 km. E (1). 1.5 km. E (1). 5 km. E (2); Bir 
el Shab (3), 6.5 km. ESE (10); Gilf el Kebir (1). 

Sudan. NORTHERN: Gebel Uweinat. Karkur Murr (7). 

Published records. — Records are from Anderson (1902), De 
Winton (1903), Bonhote (1912), Flower (1932), Hayman (1948). 
Setzer (1952, 1958d), Wassif (1960ab), Bauer (1963), and Osborn and 
Krombein (1969). 

BEHEIRA: El Birigat; Zaghig; Bir Victoria; Wadi el Natroun. 

GIZA: El Mansuriya; Ezbet Afifi Pasha; Abu Sir; Imbaba. Abu Ghalib; Abu 
Rawash; Atfih; Giza; Giza Pyramid area; Giza pyramid 8 km. NW; Sakkara; 
Bahariya Oasis. El Heiz. El Bawiti 40 km. S. 

EL FAIYUM: Kom O Shim, Ezbet Aiyub All. Qasr Rashwan. 

ASYUT: Beni Adi. 

MATRUH: El Maghra. Siwa 6.4 km. E. 

EL WADI EL GEDEED: Kharga Oasis, El Kharga. El Mahariq. Baris. 

Sudan. NORTHERN: Faras West 4 km. S; Gebel Uweinat, Karkur Murr. 

Genus Dipodillus Lataste, 1881 

Orangish brown to yellowish brown rodents of varying size. Tail 
longer than head and body, except in D. simoni. Tail brush variable. 
Palm and sole bare. Hand with three subdigital tubercles and two 
palmar pads. Foot with three subdigital, one subhallucal, and two 
plantar tubercles (fig. 34). 

Braincase usually slightly inflated and supraoccipital swollen 
beyond level of occipital condyle. Cranial ridges not strongly 
developed except supraorbitals. Some species have auditory meatus 
lip modified or swollen. Accessory tympanum present or absent. 
Chambers and cavities in mastoid bulla vary with species (fig. 47). 

Upper incisor with single groove on anterior surface. Molars 
tuberculate, becoming laminate with wear in most species. First 
labial and lingual cusps of m' alternate, at least in immatures (fig. 
38). 



OSBORN&HELMY: MAMMALS OF EGYPT 141 

Keys to Egyptian Species of Dipodillus 

External Characters 

1. Tail considerably longer than head and body. 

a. Tail usually with a conspicuous brush. White rump patch inconspicuous or ab- 
sent. 

i. Color orangish brown. (Western Desert) campestris, p. 141. 

ii. Color yellowish brown. 

(a) Paler form without blackish hairs extending to base on upper side of tail 
surface. (Sinai Peninsula and northern Eastern Desert), dasyurus, p. 155. 

(b) Darker form with blackish hairs extending to base on upper side of tail. 
(Southern Eastern Desert) mackilligini, p. 159. 

b. Tail without a conspicuous brush. White rump patch conspicuous. 

i. Ear tip pigmented. Dorsum dark amoenus, p. 167. 

ii. Ear tip not pigmented. Dorsum pale henleyi, p. 174. 

2. Tail less than to slightly longer than head and body and lacking a brush. Whitish 
rump patch absent. (Western Mediterranean Coastal Desert) simoni, p. 161. 

Cranial Characters 

1. Lip of auditory meatus not swollen. 

a. Accessory tympanum absent. 

i. Cavity of subarcuate fossa large and conspicuous (fig. 47). 

(a) Bulla not inflated beyond exoccipital. (Western Desert)campesfns, p. 141. 

(b) Bulla inflated beyond exoccipital. (Sinai Peninsula and northern Eastern 
Desert) dasyurus, p. 155. 

ii. Cavity of subarcuate fossa small (fig. 47). Bulla inflated beyond level of ex- 
occipital. (Southern Eastern Desert) mackilligini, p. 159. 

b. Accessory tympanum present. Cavity of subarcuate fossa large (fig. 47). Bulla 

not inflated beyond exoccipital. Shape of m' distinctive (figs. 38, 52) 

simoni, p. 161. 

2. Lip of auditory meatus swollen. 

a. Swelling is an anterodorsal protuberance (fig. 35). Shape of m', m^ distinctive 
(figs. 38. 52) amoenus, p. 167. 

b. Entire lip swollen (fig. 47). Shape of m' distinctive (figs. 38, 52). henleyi, p. 174. 

Dipodillus campestris (Levaillant, 1857) 
Gerbillus campestris Levaillant. 1857. Atlas Expl. Sci. Alg. Mamm.. pi. V, fig. 2. 

Type /oca/ity.- Algeria. CONSTANTINE: Phillipeville. 

General distribution.— Egypt west of Nile River and Delta, 
northern Sudan, Libya, Tunisia, Algeria, Morocco, and probably 
northern Chad and Niger. 

Common name.— Large North African Dipodil. 

Distribution of subspecies in Egypt.— Figure 45. Dipodillus 
campestris wassifi: Western Mediterranean Coastal Desert; 
Dipodillus campestris haymani: Farafara Oasis, Qattara Depres- 
sion, Siwa Oasis, and depressions west to Giarabub; Dipodillus 



142 



FIELDIANA: ZOOLOGY 




Fic; 45. Collection localities of Dipodillus campestris uassifi (circles). D. c. 
haymani (dots), I), c. patrizii (open squares), D. c. venustus (half circle), D. dasyurus 
dasyurus (solid squares), and 1). mackilligini (triangles). 

campestris patrizii: wadis of Gebel Uweinat and probably Gilf el 
Kebir; Dipodillus campestris venustus: west bank of Nile in Upper 
Egypt. 

Diagnosis.— Orangish brown, slightly larger than lesser gerbil 
(Gerbillus gerbillus). Fur long, soft. Tail long, brush of varying size. 
Ears prominent, pigmented. Supraorbital and postauricular mark- 
ings and whitish rump patch inconspicuous. Skull with bulla 
moderately inflated, lip of external auditory meatus unmodified, ac- 
cessory tympanum absent, parapterygoid fossa shallow and open, 
basioccipital conspicuously broad, incisive foramina elongate, and 
posterior margin of nasals usually divided. 



OSBORN&HELMY: MAMMALS OF EGYPT 143 

Largest of Egyptian species of Dipodillus. Adult head and body 
length average 100 mm.; tail 134 mm., 138 per cent of head and 
body length; foot 26 mm.; ear 17 mm.; occipitonasal length 30.0 
mm.; weight 30.8 gm. 

External characters. — Figure 46. Upper parts orangish to 
brownish. Dorsum with or without coarse agouti pattern ("streaked 
appearance" of Ranck, 1968, p. 141). Color gradually paUng to nar- 
row border of clear orangish on side and foreleg. All hairs of dorsum 
and side, except for narrow ventrolateral strip, with gray bases. 
Width of orangish subterminal bands and brownish tips variable. 
Darkest individuals without streaking on dorsum and rump. Hair of 
underparts and feet white. Conspicuous, broad band of dark-tipped 
hairs extending from mystacial area beneath eye to base of ear. 
Postorbital and postauricular areas of whitish hairs with dark tips 
and inconspicuous. Rump patch inconspicuous, white bands on 
hairs present or absent. Tail distinctly or indistinctly bicolored, up- 
per surface as back, ventral surface whitish to brownish; brush con- 
spicuous or inconspicuous, graying to fuscous or blackish, one-third 
to one-half of tail length. Ear pigmented. 

Palatal ridges.— Figure 33. First diastemal ridge slightly curved, 
second diastemal ridge relatively straight. First, second, and third 
intermolar ridges relatively long and recurved; fourth very small; 
fifth large and slightly recurved. 

Glans penis and baculum.— Compare notes under other species of 
Dipodillus and Gerbillus. 

Feet— Figure 34. Palm and sole hairless. Plantar tubercles 
distinct, but slightly smaller than in D. dasyurus. Two proximal 
plantar tubercles and hallucal tubercle about equidistantly spaced. 
Proximal nongranular part of sole pigmented. 

Cranial characters.— Figure 47. Skull largest, zygomatic arch 
heaviest, supraorbital ridge thickest, basioccipital broadest, and 
auditory bulla least inflated of Egyptian species of Dipodillus. 
Posterior margin of nasals usually bifurcated, otherwise irregular or 
truncate. Incisive foramina relatively long, zygomatic plate large 
and, in adults, covering infroaorbital foramen and posterior part of 
premaxillary-maxillary suture. Parapterygoid fossa open and 
shallow. Anterior margin of tympanic bulla reaching level of middle 
of foramen ovale. Posterior margin of mastoid bulla not extending 
beyond exoccipital. Anterior mastoid chamber filling less than one- 




Fk; 46. Live specimen of IJipodillus campestris wassifi. 



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146 FIELDIANA: ZOOLOGY 

half of suprameatal triangle. Hamular process of temporal 
T-shaped, closing suprameatal triangle posteriorly. External 
auditory meatus lip unmodified. Subarcuate fossa very large, par- 
tially separating anterior mastoid and lateral superior mastoid 
chambers. Medial superior posterior cavity small, not visible in 
lateral view. Accessory tympanum absent, neck and body of 
malleus hidden by enlarged bony anterior and posterior areas of 
tympanum. Figure 165 shows details of auditory bulla of D. 
campestris in posterolateral view. 

Tee ^/i. — Figure 38. Upper incisor grooved; first labial and Ungual 
cusps of m' alternate, at least in immatures. Cusps of m' separate 
and somewhat angular in immatures, becoming confluent and 
rounded in adults. First libial and second lingual folds of m' promi- 
nent and deep. Posterolateral folds of m', m"^ transient. 

Confluency of three anterior cusps of m' begins between first 
labial and first lingual followed by union of anterior and first 
lingual. Anterior cusp of m, unites with first labial about the same 
time or slightly before union of three anterior cusps of m'. Confluen- 
cy of anterior cusps and posterior lamina of m' is followed by union 
of lamina of m^, which precedes completion of confluency in m,, m2. 
M ' has three transient cusps. 

Measurements.— Table 12. Largest species of Dipodillus in 
Egypt. Male and female dimensions subequal. Means (and ranges) 
of occipitonasal length (in millimeters) of 10 adult males and 10 
adult females, respectively, are 30.7 (30.0 to 31.9) and 30.2 (28.9 to 
31.1). 

Age determmat/on. — Individuals are considered adult when 
anterior and first lingual cusps of m' become confluent (fig. 38) 
and/or basioccipital-basisphenoid suture closes. 

Variation.— Co\oT varies within subspecies in correlation with 
substrate. Palest specimens are D. c. wassifi from white limestone 
and pallid soils of Ras el Hekma, although wassifi is generally 
slightly darker than haymani from Qara and Siwa. Specimens of D. 
c. patrizii from Gebel Uweinat are slightly darker than haymani, 
and the darkest race is venustus of northern Sudan and southern 
Egypt. Degree of prominence of tail tuft varies geographically in 
correlation with color density. 

Means of head and body length, hind foot, and occipitonasal 
length show slight clinal decrease west to east in samples from 



Table 12. — Means (and ranges) of measurements, ratios, and weight of adult 
DipodiUus campestris. 



Libya 



Egypt 





D. c. brunnescens 


D. c. wassifi 


D. c. haymani 


HBL 


103.5(99-112)8 


96.2(86-112)59 


105.6 (86-125) 30 


TL 


142.2 (132-150) 7 


133.8(118-153)51 


133.6(118-147)31 


TL/HBL% 


138.0(128.2-151.5)7 


140.0 (120.7-159.3) 51 


127.2 (124.4-145.0) 29 


FL 


28.1 (27-30) 8 


26.6 (25-29) 58 


28.4 (25-30) 32 


EL 


17.6(17-18)8 


16.7 (15-20) 59 


17.5(16-20)32 


Wt 





29.2 (21.3-38.3) 39 


34.1 (26.7-44.1) 10 


ONL 


30.4 (29.9-31.6) 8 


29.6 (27.9-31.9) 51 


31.5(28.8-33.6)31 


ZW 


15.6 (15.2-16.6) 8 


15.3 (14.0-17.1) 27 


16.2(15.3-17.4) 17 


lOW 


5.4(5.1-5.7)8 


5.1 (4.8-5.8) 55 


5.5 (5.2-6.1) 31 


BCW 


13.9 (13.4-14.4) 8 


13.8(12.9-15.0)49 


14.2(13.3-15.1)31 


NL 


12.0(11.6-12.6)8 


11.6(10.7-12.8)49 


12.4(11.3-13.8)28 


IFL 


5.4 (5.2-5.8) 8 


5.4 (4.8-6.1) 54 


5.6(5.1-6.2)31 


AL 


4.2 (3.8-4.5) 8 


4.1 (3.6-4.6) 54 


4.4 (4.1-5.4) 31 


RW 





4.1 (3.9-4.7)49 


4.2 (3.8-4.6) 21 


BL 


7.6 (7.4-7.9) 8 


7.6 (7.2-8.2) 45 


8.1 (7.5-8.7) 31 


SH 


11.6(11.5-12.0)8 


11.4(11.0-12.3)42 


11.7(11.3-12.3)31 




Libya 


Libya-Sudan 






various localities 


(Cufra-Uweinat) 


Sudan 




D. c. dodsoni 


D. c. patrizii 


D. c. venustus 


HBL 


100.8(92-113)8 


88.2 (80-95) 10 


89.8 (80-98) 5 


TL 


130.8(117-138)8 


125.6(111-144)9 


134.0(115-145)5 


TL/HBL% 


130.2(116.8-150.0)8 


141.6(126.1-165.4)9 


149.2 (137.8-157.0) 5 


FL 


26.3 (25-28) 9 


26.2 (25-28) 10 


25.0 (24-26) 5* 


EL 


16.4 (16-18) 9 


15.9(14-17) 10 


13.6(13-15)5 


Wt 





24.0 


26.4 (22.0-32.0) 5 


ONL 


30.4 (29.1-31.5) 10 


29.2 (28.2-31.0) 12 


29.3 (28.6-29.9) 5 


ZW 


15.8(15.3-16.4)8 


15.4(14.4-15.9)9 


14.9(14.4-15.3)5 


lOW 


5.3 (4.9-5.8) 9 


5.0 (4.7-5.3) 12 


5.1 (5.0-5.3)5 


BCW 


13.9(13.2-14.4) 10 


13.5 (12.9-14.4) 12 


13.3(12.9-13.7)5 


NL 


12.0(11.7-12.4) 10 


11.4(11.0-12.3)12 


11.2(10.7-11.4)5 


IFL 


5.4 (5.1-5.6) 10 


5.1 (4.8-5.5) 12 


5.2 (4.9-5.4) 5 


AL 


4.1 (3.8-4.4) 10 


3.9(3.8-4.1) 12 


4.3 (4.1-4.6) 5 


RW 


4.2 (3.9-4.5) 10 


3.9 (3.6-4.2) 12 


4.1 (4.0-4.3) 5 


BL 


8.1 (7.7-8.5) 10 


7.6(7.3-8.1) 11 


7.6 (7.3-7.8) 5 


SH 


11.6(11.2-11.8) 10 


10.9(10.2-11.6) 11 


11.0(10.9-11.3)5 
♦Without claw. 



147 



148 FIELDIANA: ZOOLOGY 

Table 13. — Means (and ranges) of measurements, ratios, and weight of adult 
DipodiUus dasyurus and D. mackilligini. 





Sinai Peninsula 


Eastern Desert 




D. dasyurus 


D. dasyurus 


D. mackilligini 


HBL 


92.9 (80-102) 26 


90.0 (82-94) 18 


77.8 (72-86) 6 


TL 


120.9(109-130)23 


126.8(114-136) 16 


120.2 (99-138) 6 


TL/HBL% 


130.0(118.4-143.6)23 


142.2(128.0-155.6) 15 


154.0(137.5-176.9)5 


FL 


25.6 (25-27) 26 


25.1 (24-27) 19 


24.2 (22-26) 6 


EL 


14.9 (14-16) 26 


14.5(14-16) 19 


12.9(12-14)6 


Wt 





22.8 (16.0-34.9) 15 





ONL 


28.6 (27.9-29.2) 17 


28.3 (27.2-29.5) 18 


26.9 (26.2-27.7) 3 


ZW 


14.9(14.4-15.8)6 


14.8(14.2-15.4) 14 


13.3(13.2-13.5)3 


lOW 


5.0 (4.7-5.3) 22 


5.0 (4.8-5.3) 19 


4.8 (4.7-5.0) 6 


BCW 


13.4(12.6-14.1) 13 


13.2(12.9-13.5)8 


13.2(12.8-13.5)5 


NL 


10.9(10.2-11.8) 18 


10.8(10.3-11.5) 18 


9.8 (9.7-10.6) 4 


IFL 


5.2 (4.8-5.5) 21 


5.0 (4.4-5.8) 18 


4.6 (4.3-4.8) 5 


AL 


4.0 (3.8-4.3) 22 


4.0 (3.6-4.5) 18 


3.8 (3.5-4.0) 4 


RW 


4.0 (3.6-4.4) 26 


4.0 (3.8-4.7) 19 


3.8 (3.7-3.9) 3 


BL 


8.0 (7.4-8.5) 18 


8.0 (7.5-8.6) 19 


8.2 (8.0-8.3) 3 


SH 


11.2(10.8-11.7) 12 


11.2(10.9-11.8) 19 


11.1 (10.9-11.3)2 



Mediterranean coast localities (table 12). For comparison, samples 
of D. c. brunnescens (Ranck, 1968) from the Cyrenaican Plateau and 
adjacent littoral areas of Libya and dodsoni from the interior are in- 
cluded in the table. Size increases southward slightly in Qara and 
Siwa, then decreases through Libyan Desert samples of dodsoni and 
patrizii and in venustus of Sudan. Other measurements, except tail 
length, show similar geographic variation (table 12). 

Inflation of auditory bulla is greater in D. c. haymani than in 
other subspecies. Ear length is smaller in venustus than in other 
subspecies (table 12). 

Comparisons.— DipodiUus campestris differs fromZ). dasyurus in 
more orangish or brownish color, slightly larger external and cranial 
measurements (tables 12, 13); smaller, less swollen auditory bulla 
(fig. 47); larger subarcuate fossa; and larger molars (fig. 38 and 
tables 12, 13). 

Skins of D. campestris may be misidentified as Gerbillus 
andersoni if soles of feet are not examined. Immatures may be con- 
fused with G. henleyi. Otherwise,/), campestris is easily identifiable 
on the basis of cranial characters listed in Table 14 and shown in 
Figure 47. 

Remarks.— The trinomen D. c. venustus (Sundevall, 1843), which 
was listed in Allen's (1939) "Checklist," has no referable type 



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150 



OSBORN&HELMY: MAMMALS OF EGYPT 151 

specimen. However, Happold (1967c, p. 316) applied the name to 
specimens from northern Sudan because they "fit Sundevall's 
description." Later, Petter (Part 6.3, p. 11 in Meester and Setzer, 
1971) listed the subspecies, "(fide Happold 1967)." We believe that 
retention of the name is warranted. 

Collection.— Easily trapped in all habitats and dug from burrows 
in sand. 

Habitats.— Western Mediterranean Coastal Desert: Limestone 
cliffs beside the sea and inland near Salum (fig. 11), Ageeba and Ras 
el Hekma; quarries in limestone ridges near Burg el Arab (fig. 7); 
stone houses, walls, and temple ruins (Abu Mena and Abu Sir); 
vegetated rocky slopes, barren gullies, dunes beneath Nitraria 
retusa, and exotic Acacia saligna and Atriplex sp. on Ras el Hekma; 
shallow depressions with sandy-loam soil and boulder areas near 
Abu Haggag. Reported from sand in fig groves at Sidi Barreuii 
(Hoogstraal, 1963). 

Qattara Depression: Houses, rock outcroppings, date palm 
clusters, and piles of dead branches. 

Farafara Oasis: Date palm clusters. 

Siwa Oasis: Barley fields, palm groves, fallow sandy fields 
(Hoogstraal, 1963), and houses. 

Gebel Uweinat, Karkur Murr: Sand among boulders and beneath 
sandstone blocks (Osborn and Krombein, 1969). 

Considered "most widely distributed of all Libyan rodents" by 
Ranck (1968, p. 133) who found it living in dense growths of grass 
and sedge associated with agriculture; dense stands of sedges and 
other mesophytes including Phragmites in hard, salty clay margins 
of oases lakes; sedge pockets and areas of sand in palm groves of 
oases and outlying zones of Tamarix and Acacia; abundant in cliffs, 
rocky outcroppings and talus, and around thorny bushy perennials 
near the coast, but never in coastal dunes. Ranck (p. 139) stated that 
"Near Ghat the habitat was typical 'hamada' desert without any 
visible plant cover." Happold (1966a, 1967c, p. 317) trapped D. c. 
venustus in granitic Sabaloka Hills and syenitic Gebel Qeili in 
northern Sudan and observed that larger hills "with many crevices 
and cracks, are suitable habitats." 

This species is less restricted to rocky situations than D. dasyurus 
of the Eastern Desert and Sinai. 



152 FIELDIANA: ZOOLOGY 

Behavior.— More nervous and prone to bite than other species of 
Dipodillus. 

Reproduction.— Two females from Salum, captured on March 
26-28, contained six and three embryos, and a third female was lac- 
tating. Two lactating females were trapped on May 9 near Qara. 
One female with six embryos came from Mariut on December 3. 
These data indicate an extensive breeding period of about six 
months, but in coastal desert and inland desert populations, periods 
may not be synchronous. The breeding season of coastal desert 
populations appears to coincide with the winter rainy season 
(November-April), but inland populations may breed later. 
Happold's (1967c, p. 317) data indicate that, in Northern Sudan, the 
September-November breeding period "is at the end of the rains." 

Sex ratio. — In a sample of 128 museum specimens of D. 
campestris, males numbered 66 (52 per cent) and females numbered 
62. 

Commensalism.— Found in mud and stone houses, but probably 
because of similarity to the natural habitat. 

Economic importance.— The tag on a British Museum specimen 
from Siwa (misidentified originally as G. andersoni) reads: "The 
common species frequents houses. The natives consider this mouse 
a great delicacy." 

Associates.— Dipodillus campestris occurs in sand with G. ger- 
billus and G. andersoni; cliffs and rocky areas with ^4 corny s 
cahirinus and Eliomys quercinus; and salt marshes inhabited by 
Psammomys obesus, D. amoenus, D. simoni, and D. henleyi (Wassif , 
1960c). 

Egyptian Subspecies of Dipodillus campestris 
Dipodillus campestris wassifi (Setzer, 1958) 

Gerbillus campestris wassifi Setzer. 1958, J. Egypt. Publ. Health Assn., 33, No. 6, 
p. 209. 

Type locality.-Egypt. MATRUH: Salum, Libyan Plateau 60 ± 
m. 

Distribution in Egypt— Figure 45. Western Mediterranean 
Coastal Desert from Abu Qir west to Libyan border. 

External c/iaracfers. — Individuals vary in dorsal color from 
orangish to brownish, as noted in specimens from Salum by Wassif 



OSBORN&HELMY: MAMMALS OF EGYPT 153 

(1956a, p. 190). Summer pelage is paler and more orangish than 
winter pelage. With exception of the sample from Ras el Hekma, 
wassifi is darker than haymani. The tail is more distinctly bicolored 
and brush slightly less conspicuous than in haymani. 

Cranial characters.— Skull not so strongly developed and supraor- 
bital ridge thinner in wassifi than in haymani. 

Measurements.— Table 12. Size averages smaller than haymani. 

Specimens examined.— Total 90. 

ALEXANDRIA: Abu Qir (1), Mariut (5). 

MATRUH: Burg el Arab (8), Abu Sir (1). Abu Mens (2); Abu Haggag (2); Ras el 
Hekma (25); Mersa Matruh-Qara road 18 km. S (3); Ageeba (7); Sidi Barrani (3); 
Salum (25). 16 km. S (2); Bir Qattar (6). 

Published records.— Records are from Wassif (1956a), Setzer 
(1958d), Hoogstraal (1963), and Ranck (1968). 

Egypt. MATRUH: Salum, Mersa Matruh, Burg el Arab, Sidi Barrani. 
Libya. CYRENAICA: Bardia, 5 km. W. 

Dipodillus campestris haymani (Setzer, 1958) 

Gerbillus campestris haymani Setzer, 1958, J. Egypt. Publ. Health Assn., 33, No. 
6. p. 208. 

Type locality.— Egypt. MATRUH: Siwa Oasis, Siwa. 

Distribution in Egypt— Figure 45. Farafara Oasis, Qattara 
Depression, Siwa Oasis, and probably depressions west to 
Giarabub. 

External characters.— Palest subspecies. Tail less distinctly 
bicolored; brush slightly more conspicuous than in wassifi, but 
much less than in patrizii or venustus. 

Cranial characters.— SlavXl generally larger, more angular and 
strongly developed and supraorbital ridge thicker than in other 
subspecies. 

Measurements.— Tahle 12. Larger in external and most cranial 
measurements than other Egyptian subspecies. A sample from 
Qara is intermediate in some measurements between wassifi from 
Salum and haymani from Siwa. 

Specimens examined.— Total 60. 

MATRUH: Bir Abd el Nabi (1); Qara (14); Siwa Oasis (29), 5 km. N (4); Aghurmi 5 
km. E (4); El Maragi (4); El Zeitun (2). 
EL WADI EL GEDEED: Farafara Oasis, Batras (2). 



164 FIELDIANA: ZOOLOGY 

Published records.— Records are from de Beaux (1928), Hayman 
(1946). Setzer (1958d). and Ranck (1968). 

Egypt. MATRUH: Siwa Oasis. 

Libya. CYRENAICA: Giarabub and neighboring regions, Bahr el Tubal. 

Dipodillus campestris patrizii (de Beaux, 1932) 

Dipoditlus dodsoni patrizii de Beaux, 1932, Ann. Mus. Civ. Stor. Nat. Genova. 55, 
p. 379. 

Type locality.-Uhya. CYRENAICA: Cufra Oasis, El Giof. 

Distribution in Egypt— Figure 45. Probably all canyons of Gebel 
Uweinat and possibly Gilf el Kebir. 

External characters.— Color variable but generally darker with 
more prominent agouti pattern than wassifi, haymani, and dodsoni, 
but paler than venustus. Circumorbital and postauricular areas 
buffy. Tail indistinctly bicolored, brownish below. Tail brush 
distinct, fuscous. 

Cranial characters.— Skull not so strongly developed as in 
haymani. 

Measurements.— Table 12. Smaller generally in external and 
cranial measurements than haymani. Larger than venustus. 

Specimens examined.— Total 12. 

Libya. CYRENAICA: Cufra Oasis, el Hauuari (6). 
Sudan. NORTHERN: Gebel Uweinat, Karkur Murr (6). 

Published records.— Records are from de Beaux (1928), Ranck 
(1968), and Osborn and Krombein (1969). 

Libya. CYRENAICA: Cufra Oasis. El Giof and El Hauuari. 
Sudan. NORTHERN: Gebel Uweinat, Karkur Murr. 

Dipodillus campestris venustus (Sundevall, 1843) 

Meriones venustus Sundevall, 1843. Kongl. Svenska Vet.-Akad. Handl., 
Stockholm, p. 230. 

Type locality.— Sudan. Near White Nile. 

Distribution in Egypt— Figure 45. Single specimen from 
Kalabsha. 

External characters.— Smallest, darkest subspecies with most 
conspicuous tail brush. Dark brownish orange dorsally; circum- 
orbital and postauricular areas buffy; tail brownish above, lighter 
below; brush blackish. 



OSBORN & HELM Y: MAMMALS OF EGYPT 155 

Cranial characters.— About as other subspecies aside from 
haymani. 

Measurements.— Table 12. Most dimensions average smaller than 
other subspecies. Large ratio of tail length to head and body length 
in Happold's (1967c) Sudan sample is possibly due to method of 
measuring. 

Remarks.— A single specimen from near Kalabsha in Upper 
Egypt (Yale University Peabody Museum No. 2647) was trapped 
"in a tiny garden near water edge" (notes of collector, Thomas 
Lovejoy III, 13 January 1963). Attempts to obtain more specimens 
from the same and adjacent areas failed (unpublished field notes). 
The locality is now inundated by Lake Nasser. 

Specimens examined.— Total 13. 

Egypt. ASWAN: Kalabsha, Beit el Wali Temple (1). 

Sudan. KASSALA: Gebel Qeili (9). KHARTOUM: Sabaloka Hills (3). 

Published records.— Records are from Happold (1967c). 

Sudan. KHARTOUM: Sabaloka Hills and Gebel Qeili. 

Dipodillus dasyurus (Wagner, 1842) 
Meriones dasyurus Wagner, 1842, Arch. Nat., 8th year, 1, p. 20. 

Type locality.— West coast of Arabia, exact locality not known. 

General distribution. — Iraq, Arabia, Israel, Yemen, Sinai Penin- 
sula, Eastern Desert of Egypt, and Sudan. 

Common names.— Wagner's Dipodil, Rough-tailed Dipodil, Wadi 
Hof Gerbil. 

Subspecies in Egypt — 

Dipodillus dasyurus dasyurus (Wagner, 1842) 

Distribution in Egypt.— Figure 45. Sinai Peninsula and northern 
part of Eastern Desert. 

Diagnosis.— Yellowish brown about the size of lesser gerbil {Ger- 
billus gerbillus). Fur long, soft. Tail long, brush conspicuous. Ear 
prominent, pigmented. Supraorbital and postauricular markings 
inconspicuous. Whitish rump patch absent. Skull with bulla 
markedly inflated, lip of external auditory meatus unmodified, ac- 
cessory tympanum absent, parapterygoid fossa moderately deep 
and partly closed. 



166 FIELDIANA: ZOOLOGY 

Adult head and body length average 93 mm.; tail 124 mm., 136 
per cent of head and body length; foot 25 mm,; ear 14 mm.; 
occipitonasal length 28.4 mm.; weight 23.6 gm. 

External characters. — Upper parts yellowish brown. Dorsum with 
fine agouti pattern. Color paling to narrow border of clear yellowish 
on side and foreleg. All hairs of dorsum and side with gray bases. 

Dorsal hairs with blackish tip and yellowish subterminal band. 
Hairs of underparts and feet white to base. Tail bicolored, upper sur- 
face as back; ventral surface whitish. Brush about one-half length of 
tail, fuscous. Broad, conspicuous band of dark-tipped hairs extend- 
ing from mystacial area beneath eye to base of ear. Whitish rump 
patch lacking. Ear pigmented. 

Palatal ridges. —Similar to D. campestris in Figure 33, except 
fifth intermolar ridge is directed medially. 

Glans penis and baculum.— Compare notes under other species of 
Dipodillus and Gerbillus. 

Feet— Figure 34. Palm and sole hairless. Tubercles distinct and 
slightly larger than in D. campestris. Two proximal plantar 
tubercles and hallucal tubercle spaced about equidistantly. 

Cranial characters. — Figure 47. Supraorbital ridge moderately 
developed, particularly in comparison withD. campestris. Posterior 
margin of nasals irregular, occasionally rounded or truncate. 
Anterior palatine foramina relatively long. Zygomatic plate not 
reaching premaxillary-maxillary suture nor covering infraorbital 
foramen. Parapterygoid fossa moderately deep and partially closed. 
Basioccipital narrow. Anterior margin of tympanic bulla almost 
reaching anterior edge of foramen ovale and contacting posterior 
shelf of parapterygoid fossa. Posterior margin of bulla extending 
beyond exoccipital, but not beyond supraoccipital. Anterior 
mastoid chamber filling more than one-half of suprameatal triangle 
(fig. 47). Subarcuate fossa fairly large and partially separating 
anterior mastoid and lateral superior posterior mastoid chambers. 
Medial superior posterior cavity small and not visible in lateral 
view. Accessory tympanum absent (fig, 47). Lip of external auditory 
meatus unmodified. Hamular process of temporal usually L-shap)ed 
and closing suprameatal triangle posteriorly. 

Teeth,— Figure 38, Upper incisor grooved; first labial and lingual 
cusps of m' alternate in immatures and adults. Molars with narrow, 



OSBORN&HELMY: MAMMALS OF EGYPT 157 

sharp laminae in immatures similar to simoni, but becoming 
somewhat more rounded with wear. Folds deep and open. First 
labial fold of m' prominent, equal in depth to second lingual fold. 
Posterolateral folds of m\ m^ transient. Confluency of cusps of m', 
m,, m^, and mg follow sequence of £>. campestris (p. 146 and fig. 46). 
M^ has two transient cusps. 

Measurements.— Table 13. Male and female dimensions subequal. 
Means (and ranges) of occipitonasal length (in millimeters) of 16 
adult males and 18 adult females, respectively, are 28.6 (27.9 to 
29.5) and 28.3 (27.2 to 30.0). 

Age determination.— Adults are determined by confluency of 
cusps of m' (fig. 38) and fusion of the basioccipital-basisphenoid 
suture. 

Variation.— There are no apparent differences in color or in brush 
size between Z>. dasyurus samples from Sinai Peninsula and Eastern 
Desert. Means of all measurements are approximately equal except- 
ing length and proportions of tail (table 13), However, the 
uselessness of tail length as a taxon in this species is illustrated in 
the scatter diagram of tail length versus head and body length (fig. 
49). 

Comparisons.— Differs from D. campestris in paler, more 
yellowish color, slightly smaller average external and cranial 
measurements, more prominent tail brush, more inflated chambers 
of auditory bulla, smaller subarcuate fossa, etc. (table 14 and fig. 
47). Externally, D. dasyurus is very similar to D. mackilligini, but 
differs from the latter in having a slightly less conspicuous tail 
brush, paler color, less blackish hair on upper tail surface, larger 
subarcuate fossa which is visible externally, and less inflated bulla 
(fig. 47). Comparisons with other species are summarized in Table 
14 and Figure 47. 

Specimens examined.— Total 83. 

SINAI: El Arish (1); Bir el Maghara (1): Wadi el Sheikh (19); Wadi Raha (5); El 
Raba (6); St. Catherine Monastery area (3), 4.8 km. W (1); Tor (3); Gebel Dhalfa (1). 

SUEZ: Ain Sukhna. 3.5 km. S (1); Wadi Nakhl (1); Wadi Abu Seyala (2); Wadi Bir el 
Abd (3). Wadi Dom (18); Wadi Yesein (1); Gebel Katamiya (4); Wadi Iseili (1). 

CAIRO: Gebel Mokkatam (4), Wadi Hof (2), Wadi Garawi (6). 

Published records.— Records are from Anderson (1902), Allen 
(1915), Flower (1932), Wassif and Hoogstraal (1954), Wassif (1956a), 
and Setzer (1958d). 



168 FIELDIANA: ZOOLOGY 

SINAI: Gebel Dhalfa. Ain Sudr, Tor. Bir el Maghara. El Quseima. Gebel Umm 
Shomer, St. Catherine Monastery area, Suweira, Nuweibah, El Raba, Wadi Raha, 
and Wadi el Sheikh. 

CAIRO: Wadi Hof near Helwan. 

Collection. — Enters live traps readily and can be dug from sandy 
patches in rocky wadis. 

//a6i7a(s.— Eastern Desert: Trapped where first collected by 
Anderson (1902, p. 261) '*. . .under a shelving rock below a cliff in 
Wadi Hoaf, Helwan" (fig. 14); in crevices and shelves in sandstone 
and limestone cliffs; boulder-strewn, vegetated wadis; and beneath 
Nitraria retusa bushes on the sea shore. 

Southern Sinai Peninsula: Burrows in sand patches among rocks, 
rocky situations to high elevations, including gardens of St. 
Catherine Monastery (Wassif and Hoogstraal, 1954; Wassif, 1956a), 
but "absent from the littoral areas (below 500 m.) of the southern 
zone" (Haim and Tchernov, 1974). 

Buxton (1923) listed D. dasyurus as a salt flat dweller among 
Chenopodiaceae and Tamarix sp. in Mesopotamia, and Lewis et al. 
(1965) trapped it beneath thorny Rosaceae shrubs in a playa in 
northern Saudi Arabia. 

Be/iaL'ior.— Details of the behavior of D. dasyurus in captivity 
were reported by Fiedler (1973). 

fiurroi^s.— Tortuous burrows 15 to 20 cm. below the surface of 
sand between rocks, with entrances plugged with sand, were found 
in southern Sinai Peninsula in May (Wassif and Hoogstraal, 1954). 
Buxton (1923) mentioned D. dasyurus among desert mice which 
plug burrows by day to retain a favorable microclimate. 

Food. — In captivity, D. dasyurus ate foliage of Zygophyllum 
coccineum, raw carrots, dry bread, but no insects. According to 
Buxton (1923, p. 127), this species depends upon succulent, 
halophytic Suaeda sp. and insects in salt marshes of Mesopotamia. 

Associates.— Sinai Peninsula: Sekeetamys calurus (Wassif, 
1956a), Acomys cahirinus, A. russatus, and Eliomys quercinus. 

Eastern Desert: Sekeetamys calurus, Acomys cahirinus, and A. 
russatus. 

Northern Saudi Arabia: Trapped with D. nanus beside burrows of 
Meriones libycus (Lewis et al., 1965). 

Reproduction.— Thiee males with testes descended from Wadi 



OSBORN&HELMY: MAMMALS OF EGYPT 159 

Hof and Wadi Garawi near Helwan, February 3 and 4; and one from 
Wadi Dom on the Gulf of Suez, March 11. 

Flower (1932) reported 12 litters born in Giza Zoological Gardens 
during months of October, December, January, February, March, 
April, May, and June, consisting of one litter of 6, one of 5, three of 
4, three of 3, and four of 2 (average, 3.3). 

Sex ratio. — In a sample of 76 museum specimens, males and 
females each numbered 38. 

Dipodillus mackilligini Thomas, 1904 

Dipodillus mackilligini Thomas, 1904, Ann. Mag. Nat. Hist., (ser. 7), 14, p. 158. 

Type locality.-Egypt. SUDAN ADMINISTRATIVE: Wadi 
Allaqi (about 22° N lat.. 35° E long.). 

Common name.— Mackilligin's Dipodil. 

Distribution in Egypt— Figure 45. Southern part of Eastern 
Desert. 

Dm^nosis.— Yellowish brown, slightly smaller than lesser gerbil 
(Gerbillus gerbillus). Fur long, soft. Tail long; brush conspicuous, 
dark. Ear prominent, pigmented. Supraorbital and postauricular 
markings inconspicuous. White rump patch absent. Bulla markedly 
inflated, lip of external auditory meatus unmodified, accessory 
tympanum absent, subarcuate fossa small and not separating 
anterior and lateral superior posterior mastoid chambers. 

Adult head and body length average 78 mm.; tail 120 mm., 154 
per cent of head and body length; foot 24 mm.; ear 13 mm.; 
occipitonasal length 26.9 mm. 

External characters.— Upper parts dark yellowish brown. Dorsum 
with fine agouti pattern. Sides paler with narrow border of clear 
yellowish extending onto fore and hind limbs. All hairs of dorsum 
and sides, except a very narrow margin, with gray bases. Dorsal 
hairs with blackish tips and yellowish subterminal bands. Hairs of 
underparts and feet white to base. Broad, conspicuous band of dark 
tipped hairs extending from mystacial area beneath eye to base of 
ear. Whitish supraorbital and postauricular areas inconspicuous. 
White rump patch absent. Tail indistinctly bicolored; upper surface 
darker than back, with blackish hairs to base. Underside of tail 
whitish to buff. Tail brush about one-half of tjiil length, fuscous. Ear 
pigmented. 



160 FIELDIANA: ZOOLOGY 

Palatal ridges.— Not observed. 

Glans penis and baculum.— Not observed, 

Fee^ — Palm and sole hairless. Tubercles distinct. Proximal tuber- 
cle further from others than in D. dasyurus. Sole not pigmented. 

Cranial characters. — Figure 47. Supraorbital ridge moderately 
developed in comparison with D. campestris. Posterior margin of 
nasals broadly divided or truncate. Incisive foramina relatively 
long. Palatine foramina relatively long, open. Zygomatic plate not 
reaching level of premaxillary-maxillary suture, but covering 
infraorbital foramen. Parapterygoid fossa deep and partly closed. 
Basioccipital narrow. Anterior margin of tympanic bulla level with 
middle of foramen ovale and contacting shelf of parapterygoid 
fossa. Posterior margin of mastoid bulla extending beyond level of 
paroccipital process. Anterior mastoid chamber filling slightly more 
than one-half of the suprameatal triangle. 

Subarcuate fossa deeper than D. amoenus or D. henleyi, but not 
separating anterior mastoid and lateral superior posterior mastoid 
chambers. Medial superior posterior cavity small, not visible in 
lateral view. Accessory tympanum absent (fig. 47). Lip of external 
auditory meatus unmodified. Hamular process of squamosal 
L-shaped, closing suprameatal triangle posteriorly. 

Teeth.— Figure 38. Upper incisor grooved, first labial and Ungual 
cusps of m' alternate, angular, and almost separate in immatures, 
becoming confluent and rounded in adults. Second lingual fold of m' 
deeper and more prominent than first labial fold. Posterolateral 
folds of m\ m^ transient. 

Confluency of cusps of m', m^ m,, and mg appear to follow 
sequence of D. campestris andD. dasyurus (p. 146 and fig. 38). M^ 
has two transient cusps separated by a labial fold. 

Measurements.— Table 13. 

Age determination.— Adults are determined by confluency of 
cusps of m' (fig. 38) and fusion of the basioccipital-basisphenoid 
suture as in other species of Dipodillus. 

Comparisons.— Dipodillus mackilligini can be distinguished from 
other Egyptian species only by a combination of characters (table 
14, fig. 38). Tooth characters of D. mackilligini, combined with 
cranial and external characters, indicate a closer relationship with 



OSBORN & HELMY: MAMMALS OF EGYPT 161 

D. dasyurus, D. campestris, and perhaps D. simoni, than with D. 
amoenus and D. henleyi. 

Ellerman (1941) referred mackilligini to the dasyurus group 
because of its long tail. Later, on the basis of bulla size, it was given 
subspecific status under D. nanus (Ellerman, 1949; Ellerman and 
Morrison-Scott, 1951; Wassif, 1956). Setzer (1959d) considered 
mackilligini a full species distinguishable from nanus by smaller 
bulla, narrower incisive foramina, curved and proportionately 
longer tooth row, and relatively more open parapterygoid fossa. A 
few more obvious differences between the two species are in Table 
14. Note also that, except for size differences in the subarcuate 
fossa, D. mackilligini is very similar to D. dasyurus (table 14, fig. 
47). 

Specimens examined.— TotaX eight. 

ASWAN: Khor Rhama el Bahari (2). 

SUDAN ADMINISTRATIVE: Wadi AUaqi (3), Wadi Kansisrob (3). 

Published records.— Records are from Thomas (1904), Setzer 
(1958d) and Hoogstraal (1963). 

SUDAN ADMINISTRATIVE: Wadi AUaqi, Wadi Kansisrob. 

//a6itats.— East bank of Nile River in Nubia: Grassy plot beside 
water, bush beside water in abandoned village (from notes of collec- 
tor, Christopher O. Maser). 

Wadi Allaqi: Ancient ruins. 

Gebel Elba: "Dug from plugged burrows in a grassy valley at 
about 2500 feet" altitude (Hoogstraal, 1963, p. 14). 

No further information has been recorded on this species. 

Dipodillus simoni (Lataste, 1881) 
Gerbillus simoni Lataste, 1881, Naturaliste, Paris, 1, p. 497. 

Type locality.- Algeria. CONSTANTINE: Wadi Maghra, north of 
Hodna. 

Common name.— Simon's Dipodil. 

General Distribution,— Egypt west of Nile Delta, Libya, Tunisia, 
and Algeria. 

Subspecies in Egypt — 

Dipodillus simoni kaiseri (Setzer, 1958) 
Dipodillus kaiseri Setzer, 1958. J. Egypt. Pub. Health Assn., 33. No. 6, p. 214. 



162 FIELDIANA: ZOOLOGY 

Type locality.— Egypt. MATRUH: Mersa Matruh. 

Distribution in Egypt. — Figure 50. Western Mediterranean 
Coastal Desert west of Nile Delta. 

Diagnosis.— YeWovfish brown mice smaller than lesser gerbils 
{Gerbillus gerbillus); tail about same length as head and body, brush 
lacking; ear pigmented; palm and sole naked. Whitish area around 
eye inconspicuous. Whitish rump patch absent. 

Skull with bulla moderately inflated, lip of external auditory 
meatus unmodified, accessory tympanum present, and posterior 
margin of nasals truncate. 

One of the smaller Egyptian species of Dipodillus. Adult head and 
body length average 80 mm.; tail 86 mm., 106 per cent of head and 
body length; foot 21 mm.; ear 12 mm.; occipitonasal length 25.2 
mm.; weight 17.4 gm. 

External characters. — Upper parts yellowish brown. Dorsal color 
paling gradually to a narrow line of clear yellowish on side. Color of 
side not extending onto foreleg. Dorsum with fine agouti pattern. 
Dorsal hair tips blackish, subterminal bands yellowish, and base 
gray. Hair of side yellowish with gray base, except for narrow 
ventrolateral strip with white base. Tail buffy with scattering of 
black hairs on upper surface. Brush very inconspicuous, long hairs 
on tip of tail only. 

Underparts and upper surfaces of feet white. Band of dark-tipped 
hairs from mystacinal area beneath eye to base of ear, broad and 
conspicuous. Whitish supraorbital area inconspicuous (as opposed 
to conspicuous in D. amoenus). Whitish postauricular patch large. 
Whitish rump patch lacking. Ear pigmented. 

Palatal ridges. — Figure 33. First diastemal ridge slightly curved; 
second diastemal ridge broadly U-shaped, sometimes divided; 
fourth intermolar ridge very small or missing; fifth intermolar not 
recurved. 

Glans penis and baculum.—See under Gerbillus pyramidum, D. 
campestris, and Wassif et al. (1969). 

Feet — Figure 34. Palm and sole hairless. Two plantar tubercles 
and hallucal tubercle spaced about equidistantly. Sole of hind foot 
not pigmented. 

Cranial characters.— Figures 47, 51. Supraorbital ridge moderate- 
ly developed. Posterior margin of nasals truncate. Zygomatic plate 



OSBORN&HELMY: MAMMALS OF EGYPT 163 

sloping posteriorly, therefore not reaching level of premaxillary- 
maxillary suture. Incisive foramina long. Parapterygoid fossa open 
and shallow. Basioccipital relatively broad. Anterior margin of tym- 
panic bulla usually reaching level of middle of foramen ovale. 
Posterior margin of mastoid bulla not or barely surpassing level of 
exoccipital ridge. Anterior mastoid chamber filling about one-half of 
suprameatal triangle. Subarcuate fossa very large and partially 
separating anterior mastoid and lateral superior posterior mastoid 
chambers. Medial superior posterior cavity small and not visible in 
lateral view. Accessory tympanum present. Lip of external auditory 
meatus unmodified. Hamular process of squamosal T- or L-shaped 
closing suprameatal triangle posteriorly. 

Teeth.— Fig[ires 38, 52. Upper incisor grooved; first labial and 
Ungual cusps of m^ alternate in immatures and adults. Margins of 
cusps and laminae angular; folds deep and open. First labial fold in 
m' equal in depth to second lingual. A posterolateral fold is tran- 
sient in m'; lacking in m^. Confluency of anterior cusps of m\ m^ 
occurs at same time or slightly later than in m^ m2. All cusps and 
laminae confluent in immatures. M^ with two transient cusps. Note 
m' is distinctive ofD. simoni. 

Measurements.— Table 15. Male and female dimensions subequal. 
Means (and ranges) of occipitonasal length (in milUmeters) of 10 
adult males and seven adult females, respectively, are 25.3 (23.2 to 
26.7) and 25.0 (23.4 to 25.9). 

Variation.— Data from Algeria, Tunisia, Libya, and Egypt (table 
15) indicate west to east clinal increase in tail length, decrease in 
zygomatic width, and reduced inflation of auditory bulla. Libyan 
specimens are darker than Egyptian, although color of dorsum, 
according to Ranck (1968), varies widely. 

Comparisons.— Dipodillus simoni kaiseri from Egypt is 
distinguishable from the nominate form by slightly longer tail, 
darker color and smaller bulla. Dipodillus simoni can be distin- 
guished from all other Egyptian dipodils by the relatively short, 
almost unicolorous, tail which lacks a brush; individual 
characteristics in palatal ridges (fig. 33) and molars, particularly m' 
(figs. 38, 52); and early confluency of cusps and laminae. Combina- 
tions of characters presented in Table 14 are of further use in 
distinguishing D. simoni from other species. In dimensions, D. 
simoni is more comparable with D. amoenus (tables 15, 16), except 
for shorter tail, shorter foot, longer incisive foramina, and shorter 



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164 



OSBORN&HELMY: MAMMALS OF EGYPT 165 

Table 16. — Means (and ranges) of measurements, ratios, and weight of adult 
DipodiUus a. amoenus. 







Wadi el Natroun, 




El Faiyum 


Bir Victoria 


HBL 


84.0 (70-100) 13 


79.9 (73-86) 18 


TL 


104.6(93-115) 12 


106.4(99-116) 17 


TL/HBL% 


124.6(112.0-147.4) 12 


134.1 (118.8-155.4) 17 


FL 


22.9 (20-24) 12 


22.6 (21-24) 18 


EL 


13.0(12-14) 12 


12.2(11-14) 18 


Wt 





13.2(10.7-17.5) 10 


ONL 


25.8 (23.6-26.7) 10 


25.2 (24.4-26.3) 15 


ZW 


13.9(12.9-14.4)5 


13.4(13.0-14.1) 10 


lOW 


4.6(4.2-4.8) 11 


4.4 (4.2-4.9) 18 


BCW 


12.4(11.8-12.9) 10 


12.2(11.7-12.8) 18 


NL 


9.3 (7.9-10.0) 10 


9.2 (8.6-9.8) 15 


IFL 


4.2 (3.8-4.6) 12 


4.0 (3.7-4.2) 18 


AL 


3.6(3.2-4.1) 12 


3.4 (3.2-3.7) 18 


RW 


3.6 (3.4-3.8) 13 


3.4 (3.3-3.7) 19 


BL 


7.8 (7.2-8.2) 12 


7.7 (7.3-8.2) 19 


SH 


10.2 (9.6-10.5) 9 


10.0 (9.6-10.3) 18 



bulla. DipodiUus simoni is also heavier on the average than D. 
amoenus. 

Remarks.— Greater average tail length of Egyptian and Libyan 
specimens of D. simoni signified for Setzer (1958d) and Ranck (1968) 
a distinct species, kaiseri. However, other measurements in Table 
15 from Egypt, Libya, and Algeria, as well as tooth characters (figs. 
38, 52 and Petter, 1959, fig. 1), support Wassif's (1956a, 1960c) con- 
clusion that simoni and kaiseri are synonymous. Later Wassif et al. 
(1969) recorded the karyotype of Egyptian D. simoni as 2N=60 
with 8 to 10 biarms and FN =68-69. Cockrum et al. (1976a) reported 
identical karyotypes from Tunisian specimens. 

Specimens examined.— Total 39 

ALEXANDRIA: El Amiriya (5). 1.6 km. E (1). 

MATRUH: Lake Mariut (1); Bahig (7). 7 km. S (2); Abu Mena (1). 1.6 km. E (2); 
Burg el Arab (9); El Hammam 15 km. SSW (1): El Daba (2); Ras el Hekma (2); Mersa 
Matruh (3), 4.8 km. E (1); El Qasr (1); Sidi Barrani (1). 

Published records.— Records are from Wassif (1956a, 1960c) and 
Setzer (1958d). 

MATRUH: Lake Mariut, Burg el Arab, El Daba, Mersa Matruh, Sidi Barrani. 



166 



FIELDIANA: ZOOLOGY 




Fig 48. Western Mediterranean Coastal Desert. Ras el Hekma. Rocky and sandy 
slope. Vegetation: Lycium shawii in foreground surrounded by thistle {Onopordon 
alexandrinum). Burrow under Lycium is of Psammomys obesus. 

Collection.— Dug from burrows in salt marshes. Readily enters 
live traps. May be picked up by hand under a spotlight at night. 

//a6/^af.— Littoral salt marshes (fig. 7) (Wassif, 1960c; 
Hoogstraal, 1963) in salty, sandy loam with halophytic vegetation; 
olive groves; barley fields; clay soil in Thymelaea hirsuta and 
Anabasis articulata associations (fig. 8); slopes above salt marshes 
supporting a variety of shrubs, including Lycium shawii and 
Thymelaea hirsuta (fig. 48). 

Occurs also in high plateaus of Algerian Atlas and vegetated 
littoral desert in Tunisia and Libya (Harrison, 1967; Ranck, 1968). 

Be/iauior.— Nocturnal. A very docile mouse, easily handled. When 
approached with a light at night, D. simoni either "freezes" or 
crawls under an available shrub. 

Burrows. — In late July, Wassif (1960c) found unplugged burrows 
near salt marsh vegetation. 



OSBORN&HELMY: MAMMALS OF EGYPT 167 



r 

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Tail Length | 

Fig. 49. Scatter diagram of tail length versus head and body length in Eastern 
Desert and Sinai Peninsula samples of Dipodillus dasyurus. Half-circles represent 
individuals of both samples at same point. Data from all ages. 

Reproduction.— Female with four embryos, October 4 (Harrison, 
1967) from Tunisia. No data from Egypt. 

Associates.— Occurs in coastal salt marshes with D. henleyi, D. 
amoenus, D. campestris (Wassif, 1960c), Psammomys ohesus, Allac- 
taga tetradactyla, and Jaculus orientalis; and in the littoral desert 
with Gerbillus andersoni, G. gerbillus, J. jaculus, J. orientalis, A. 
tetradactyla, Meriones shawi, and possibly P. ohesus. 

Dipodillus amoenus De Winton, 1902 
Dipodillus amoenus De Winton, 1902, Ann. Mag. Nat. Hist., (ser. 7), 9, p. 46. 
Type locality.— Egypt. GIZA. 

General distribution.— Egypt and Libya and possibly Tunisia, 
Algeria, Morocco, and Mauritania. 

Common name.— Charming Dipodil. 

Subspecies in Egypt— 



168 



FIELDIANA: ZOOLOGY 




Fig. 50. Collection localities of Dipodillus simoni kaiseri (dotsi and D. amoenus 
amoenus (circles). 



Dipodillus amoenus amoenus De Winton, 1902 

Distribution in Egypt— Figure 50. Western part of Nile Delta, 
Western Desert to Libyan frontier, and southern part of the Eastern 
Desert. 

Diagnosis. —Size smaller than lesser gerbil (Gerbillus gerbillus). 
Dorsum brownish with contrasting whitish areas above eye, behind 
ear, and above base of tail. Fur relatively short and soft. Tail longer 
than head and body with inconspicuous brush. Ear prominent and 
partly pigmented. 

Incisive foramina relatively long. Bulla inflated posteriorly 
beyond level of supraoccipital. Lip of external auditory meatus with 
prominent anterodorsal swelling. Accessory tympanum present. 




Fig. 51. Skull ol Dipodillus simoni kaiseri. 



169 



170 FIELDIANA: ZOOLOGY 

One of the smaller Egyptian species of Dipodillus. Adult head and 
body length average 81 mm.; tail 106 mm., 130 per cent of head and 
body length; hind foot 23 mm.; ear 12 mm.; occipitonasal length 
25.5 mm.; weight 13 gm. 

External characters.— Figare 53. Dorsum dark yellowish brown. 
Dorsum brown in darker individuals with color gradually paling to 
border of clear yellowish on side. Color of side not extending onto 
forelimb. Dorsal hairs with blackish tips, yellowish subterminal 
bands, gray bases. Side with broad area of hairs with white bases. 
Venter, upper surface of feet, and underside of tail white. Tail 
bicolored; upper surface as back. Brush one-third to one-fourth 
length of tail, inconspicuous, fuscous. Whitish area on rump large, 
conspicuous. Band of dark-tipped hairs from mystacial area beneath 
eye to base of ear broad and conspicuous. White areas above eye and 
behind ear prominent. Tip of ear pigmented. 

Palatal ridges.— Figure 33. Diastemal ridges slightly curved. The 
first to third intermolar ridges long and recurved, fourth intermolar 
vestigial, fifth long and curving anteriorly. 

Glans penis and baculum.— In this and the following species, D. 
henleyU the basal plate is lozenge-shaped (see also under G. 
pyramidum, G. andersoni, and Wassif et al., 1969). 

Feet.— Figure 34. Palm and sole naked. Proximal plantar 
tubercles small, indistinct, and close together. Sole not pigmented. 

Cranial characters.— Figure 47. Supraorbital ridge moderately 
developed. Interparietal deep and rectangular. Posterior margin of 
nasals irregularly truncate to broadly rounded. Incisive foramina 
relatively long. Zygomatic plate projecting forward to level of 
premaxillary-maxillary suture. Parapterygoid fossa deep. Basioc- 
cipital relatively broad. Anterior margin of tympanic bulla extend- 
ing beyond posterior margin of foramen ovale. Posterior margin of 
mastoid bulla extending beyond level of paroccipital and supraoc- 
cipital. Subarcuate fossa small, not dividing anterior mastoid and 
lateral sui)erior posterior mastoid chambers. Medial superior 
posterior cavity large and visible in lateral view. Accessory 
tympanum present. Lip of external auditory meatus distinctive 
with a large anterodorsal swelling nearly touching zygomatic pro- 
cess of temporal. Hamular process of temijoral usually L-shaped 
and closing the suprameatal triangle posteriorly. 

Teef A.— Figures 38, 52. Upper incisor grooved; first labial and 



OSBORN & HELM Y: MAMMALS OF EGYPT 



171 



E 
E 






D.HENLEYI 



D.SIMONI 




D.AMOENUS 

Fig. 52. Crown views of upper first molars of Dipodillus henleyi, D. simoni, and D. 
amoenus illustrating differences in pattern and changes with wear; and of D. 
amoenus, individual variation. 

lingual cusps of m^ alternate, at least in immatures. Margins of 
cusps and laminae rounded. Second lingual fold of m' prominent and 
deeper than first labial fold. Small posterolateral folds on m, m^; 
mj, m2. All cusps of m^ confluent in adults. Transverse lamination in 
m , m, and posterior cusps of m\ mi completed in immatures. 
Stages of confluency of the three anterior cusps of m variable and 
may begin between the anterior cusp and first lingual, as in D. 
henleyi, or between the first labial and first lingual (fig. 52). Stages 
in m, also appear variable, but confluency may be completed slight- 
ly before m\ 

Posterior lamina of mj not united with anterior cusps. Anterior 
and posterior laminae of m^, mg not always confluent. M has two or 
three transient cusps. Enamel pattern of m\ m^ is distinctive of D. 
amoenus. 



172 



FIELDIANA: ZOOLOGY 




Fig. 53. Cadaver of Dipodillus umoenus amoenus. Note distinctive white rump 
patch. 

Measurements.— Table 16. Male and female dimensions subequal. 
Means (and ranges) of occipitonasal length (in millimeters) of 19 
adult males and nine adult females, respectively, are 25.6 (23.6 to 
26.7) and 25.2 (24.4 to 26.6). 

Age determination.— Adults have part of all cusps of m' confluent 
(figs. 38, 52) and cranial sutures closed. 

Variation.— Means of measurements within D. amoenus vary only 
slightly from north to south in Egypt. Samples are too small for 
further analyses (table 16). Ranck (1968) reported decreased "size" 
and reduction in inflation of auditory bulla from west to east in 
Libya, 

Comparisons.— The nominate subspecies from Egypt is reported 
to be darker than Dipodillus amoenus vivax from Libya and with 



OSBORN&HELMY: MAMMALS OF EGYPT 173 

skull less strongly developed, bulla smaller, tail shorter with brush 
less conspicuous (Ranck, 1968). 

Relationships between D. amoenus and D. nanus or D. dasyurus 
were assumed by several authors (EUerman and Morrison-Scott, 
1951; Petter, 1961; Harrison, 1967) without evidence. The most 
closely related species recognized in this work are nanus and 
henleyi. These species and amoenus have similar color pattern, 
including whitish rump patch; anterior lip of auditory meatus com- 
pletely or partially inflated (fig. 47); a small subarcuate fossa; large 
medial superior posterior cavity, which is visible in lateral view; and 
patterns of occlusal surfaces of molariform teeth similar, particular- 
ly in amoenus and henleyi (figs. 38, 52). 

Dipodillus amoenus and D. simoni appear simileir superficially, 
but the latter lacks a whitish rump patch, and the white marking 
above the eye is inconspicuous; the tail of simoni is shorter, with 
almost no brush; tooth characters of simoni are distinctive (fig. 52); 
lip of the auditory meatus is unmodified; and in simoni, the 
averages of most measurements are smaller except for incisive 
foramina length (tables 15, 16). Further comparisons are sum- 
marized in Table 14. 

Remarks.— A specimen reported to be D. mackilligini from near 
Cairo by Bonhote (1909, p. 792) isD. amoenus (B. M. 9.7.1.43). 

Specimens examined.— Tota\ 102. 

ALEXANDRIA: El Amiriya (8). 

TAHREER: Cairo-Alexandria desert road km. 102 (1). 

ASWAN: Wadi AUaqi (2). Wadi Quleib (2). 

BEHEIRA: Hafs (1); Bir Victoria (3); Wadi el Natroun (18); Wadi el Natroun. 
Zaghig (1): El Beida (1), 5 km. S (1), 4.8 km. E (1). 

MATRUH: Burg el Arab (1). Salum 17.6 km. SE (1). El Maghra (4). Camel Pass 
Dune area (6). 

GIZA: Hawamdiya (2); El Aiyat. Mit Riheina (2); Giza Pyramids (4). 

EL FAIYUM: El Auberge (1); Minshat Tantawi (3); Minshat el Amir (5); Lake 
Qarun (2): Abu Gandir (1); Kom O Shim (6), 1.6 km. NE (1); Tahreer Forest (2); 
Shooting Club (12); Seila (1). 

EL WADI EL GEDEED: Dakhla Oasis. Gharb El Mawhoub (3): Bir el Nokta (1); 
Mut 3.2 km. S (1). 10 km. N (3). 

RED SEA: Wadi Naam (1). 

Published records.— Records are from De Winton (1903), Flower 
(1932), Setzer (1952, 1958d), and Wassif (1956a). 

BEHEIRA: Hafs; Bir Victoria; Wadi el Natroun. 
MATRUH: Burg el Arab, El Alamein. 

EL FAIYUM: Kom O Shim. Kom O Shim 1.6 km. NE; Lake Qarun; El Auberge; 
Sinnuris; Seila; Minshat Tantawi; Shooting Club. 



174 FIELDIANA: ZOOLOGY 

Collection.— Dug from burrows in salt marshes and hard desert 
soil and sand. Readily enters live traps. May be picked up by hand 
under a spotlight at night. 

//a6t tats.— Southeastern Desert: Burrows found in sand under 
Zilla spinosa in Wadi Naam (Hoogstraal et al., 1957b). 

Northeastern Delta (Hafs, Amiriya): Found in burrows of Psam- 
momys obesus in dark, saline soil and salt marsh. Burrows found in 
dikes in a soil reclamation area. 

Wadi el Natroun: Clover fields, salty areas, canal banks support- 
ing halfa grasses {Desmostachya bipinnata and Imperata cylin- 
drica). 

El Faiyum: Under Tamarix sp. at edge of cultivated fields and in 
cultivated ground. 

Camel Pass Dune area: Tents of an oil company camp on barren, 
hard, pebble desert like that between sand sheets in Figure 9. 

Kharga Oasis: Under Tamarix sp. and beside stands of 
Hyoscyamus muticus in wasteland. 

Mediterranean Coastal Desert: Salt marsh near the coast and 
sandy depressions inland (fig. 7). 

El Maghra: In sand under dead fronds and other debris of wild 
date palms. 

Dipodillus amoenus was reported from salt marsh and semidesert 
areas of Egypt by Wassif (1956a) and Hoogstraal (1963). According 
to Ranck (1968), it is the common rodent of oases in Libya, occur- 
ring under dead fronds of date palms and in cultivated land. 

Burrows.— Simple and extending about 25 cm. below ground sur- 
face (Wassif, 1956a). 

Commensa/ism.— Occasionally found in tents (see above). 

i4ssocmtes.— Occurs with Psammomys obesus and other rodents 
inhabiting salt marshes (see under D. simoni) and with Gerbillus 
gerbillus or G. andersoni in sandy areas. 

Dipodillus henleyi De Winton, 1903 

Dipodillus henleyi De Winton. 1903. Nov. Zool.. 10. p. 284. 

Type locality.— Egypt. BEHEIRA: Wadi el Natroun, Zaghig. 

General distribution.— Jordan, Yemen, Sinai Peninsula, Egypt, 
Libya, Algeria, and probably Tunisia. 



OSBORN & HELMY: MAMMALS OF EGYPT 



175 



2 5* 2 6* 2 7* 28* 2 9* 30* 31* 32* 33* 34* 35* 36* 37 




Fig. 54. Collection localities of Dipodillus henleyi henleyi (circles) andD. h. mariae 
<dots). 

Common names.— Henley's Gerbil, Pigmy Dipodil. 

Distribution of subspecies in Egypt— Figure 54. Dipodillus 
henleyi henleyi: northern part of Western Desert; Dipodillus henleyi 
mariae: Sinai Peninsula and Eastern Desert. 

Diagnosis. —Small, buffy brown with contrasting white areas 
above eye, behind ear, and on rump; tail longer than head and body, 
with inconspicuous terminal brush; ear not prominent nor 
pigmented. Incisive foramina relatively short. Bulla inflated 
posteriorly slightly beyond level of exoccipital. Entire anterior lip of 
external auditory meatus inflated. Accessory tympanum present. 

Adult head and body length average 66 mm.; tail 88 mm., 134 per 
cent of head and body length; hind foot 20 mm.; ear 10 mm.; 
occipitonasal length 22.0 mm.; weight 10 gm. 



FIELDIANA: ZOOLOGY 




Fig. 55. Cadaver of Dipodiltus henleyi mariae. 

External characters.— Figure 55. Dorsum buffy brown and darker 
than sides due to brownish tipped hairs. Dorsal and side hairs with 
gray base, except for narrow ventrolateral strip with white base. 
Side buffy, color not extending onto forelimb. Underparts, foreleg, 
and feet white. Band of dark-tipped hairs extending from mystacial 
area beneath eye to base of ear, width variable. Supraorbital and 
postauricular areas and rump patch white, prominent. Tail 
bicolored; upper surface as back, ventral white. Tail brush 
inconspicuous, about one-fourth or less of tail length, fuscous. Ear 
not pigmented. 

Palatal nd^es.— Similar toD. amoenus in Figure 33, except fourth 
and fifth intermolar ridges are crenulated. 

Glans penis and baculum.—See under D. amoenus. 

Feet— Figure 34. Palm and sole hairless. Tubercles distinct. Prox- 
imal plantar tubercles close together. Sole not pigmented. 

Cranial characters. — Figure 47. Skull small, fragile. Supraorbital 
ridges thin. Posterior margin of nasals irregularly truncate. Inter- 



OSBORN&HELMY: MAMMALS OF EGYPT 177 

parietal proportionately deeper than in other species and distinc- 
tive. Anterior palatine foramen relatively short. Zygomatic plate 
with nearly vertical anterior margin reaching or almost reaching 
level of premaxillary-maxillary suture. Parapterygoid fossa 
moderately deep. Basioccipital slender. Anterior margin of 
tympanic bulla reaching level of anterior edge of foramen ovale. 
Posterior margin of mastoid bulla inflated almost to level of 
supraoccipital. Entire lip of the external auditory meatus swollen, a 
distinctive species character (fig. 47). Accessory tympanum present. 
Subarcuate fossa small and not separating anterior mastoid and 
lateral superior posterior mastoid chamber. Medial superior interior 
posterior mastoid cavity large and visible in lateral view (fig. 47). 
Squamosal T-shaped and closing suprameatal triangle posteriorly. 

TeetA.— Figures 38, 52. Upper incisor grooved; first labial and 
lingual cusps of m^ alternate; sometimes appearing opposite in worn 
teeth. Molars tuberculate in immatures, becoming fused with wear 
into laminae with rounded margins. A deep first labial fold in m^ 
separates the combined anterior and lingual cusps from the first 
labial in immatures and adults. Union between first lingual and first 
labial cusps occurs in old individuals after union of first labial and 
posterior lamina. Second lingual fold large and deep. Posterolateral 
folds of m\ m^ transient. Anterior and first lingual cusps of mi 
usually united and confluency between first labial and lingual 
follows pattern of m\ Confluency of posterior lamina of mj with 
anterior cusps and union of laminae of mg occurs in teeth showing 
considerable wear. 

Note that size and pattern of molars, particularly m\ are distinc- 
tive of D. henleyi (fig. 52). 

Measurements.— Table 17. Smallest Egyptian species of 
Dipodillus. Male and female dimensions subequal. Means (and 
ranges) of occipitonasal length (in millimeters) of eight adult males 
and eight adult females, respectively, are 22.1 (21.4 to 22.8) and 22.0 
(22.9 to 23.0). 

Variation.— Color of specimens from the Eastern Desert is slight- 
ly paler than Western Desert samples, but with dark brownish hairs 
on the posterior dorsum more prominent. White hairs are more 
extensive in D. h. mariae. The rump patch is more conspicuous and 
tail about 10 per cent longer in D. h. henleyi. 

Compansons.— Relationship between Z). henleyi, D. amoenus, and 
D. nanus are evident in the presence of a prominent white rump 



178 



FIELDIANA: ZOOLOGY 



Tablk 17. — Means (and ranges) of measurements, ratios, and weight of adult 
Dipodillus henleyi. 





Western Desert 


Eastern Desert 




D. h. henleyi 


D. h. mariae 


HBL 


66.4 (62-75) 15 


67.6 (62-75) 9 


TL 


85.4 (72-95) 15 


94.4 (89-99) 8 


TL/HBL% 


128.8(101.4-146.8) 15 


139.8 (128.0-159.6) 8 


FL 


19.2(19-20) 15 


20.4(19.5-21.0)9 


EL 


9.4 (9-10) 15 


9.6 (9.0-10.5) 9 


Wt 


9.8(8.0-11.4)9 


7.2. 10.0 


ONL 


22.0 (20.9-23.0) 14 


22.0(21.0-22.6)7 


ZW 


12.3(11.9-12.8)8 


12.2(11.8-12.5)4 


lOW 


3.8 (3.6-4.4) 14 


3.9 (3.7-4.2) 6 


BCW 


11.1(10.8-11.7) 13 


11.0(10.4-11.4)7 


NL 


7.9 (7.2-9.0) 14 


7.6 (6.8-8.0) 6 


IFL 


3.5(3.1-3.8) 15 


3.6 (3.4-3.8) 8 


AL 


2.9(2.7-3.1) 14 


2.8 (2.8-3.0) 7 


RW 


3.0 (2.8-3.3) 12 


3.0 (2.9-3.0) 3 


BL 


7.4 (6.8-9.0) 12 


7.2 (7.0-7.3) 5 


SH 


9.1 (8.5-9.7) 9 


9.2 (8.9-9.4) 6 



patch, swelling of the anterior lip of the external auditory meatus, 
small subarcuate fossa, large medial superior posterior mastoid 
cavity, and presence of an accessory tympanum. Dipodillus henleyi 
can be distinguished from all other Egyptian species by its smaller 
dimensions, particularly alveolar length of upper molars; occlusal 
pattern of molariform teeth; swelling of the auditory meatal lip; 
interparietal shape; and additional characters in Figure 47 and 
Table 14. 

Collection.— Dug from burrows in sand, trapped alive, and caught 
by hand at night using a spotlight. 

Habitats.—Sinai Peninsula: Not well defined. Collected by Haim 
and Tchnemov (1974) in stoney, gravelly wadis vegetated with 
Anabasis sp. and Lygos raetam and Anabasis articulata, together 
with Zilla spinosa. 

Eastern Desert: Plains, vegetated wadis (fig. 5), coastal marshes, 
cultivated areas (Hoogstraal, 1963), sand among rocks, in hard 
gravel beside the large shrub, Salvadora persica, and sand under 
stands of bunch grass, Panicum turgidum. 

Western Desert: Newly reclaimed land; sparsely vegetated, hard 
sand, and densely vegetated patches of soft sand (Hoogstraal, 
1963); coastal vegetation on clay and stoney land (fig. 8); Psam- 



OSBORN & HELMY: MAMMALS OF EGYPT 179 

momys obesus burrows in salt marshes; and beside Atriplex 
halimus and Limoniastrum monopetalum (fig. 7). Ranck (1968) 
found it in Libya in salt marshes and confined to coastal plain and 
littoral deserts. 

Burrows.— Burroy/s with plugged entrances were found in loose 
sand in stands of Panicum turgidum on the Red Sea coastal plain 
near Gebel Elba (Hoogstraal et al., 1957b). Burrows are simple and 
shallow. 

Associates.— Yonndi in the same habitat with D. simoni, D. 
amoenus, Psammomys obesus, and sometimes Gerbillus gerbillus 
and G. pyramidum. 

Reproduction.— Females with four newborn young were collected 
in Wadi Digla, southeast of Cairo, and near Bahig in the Western 
Mediterranean Coastal Desert in mid-June and mid-August, 
respectively. 

Sex ratio.— In a sample of 45 museum specimens of D. henleyi, 
there were 26 (58 per cent) males and 19 females. 

Key to Egyptian Subspecies of Dipodillus henleyi 

1. Brownish hairs on posterior dorsum not prominent. Dark-tipped hairs of hind 

limb reaching heel. Tail average 10 per cent shorter. (Western Desert) 

henleyi, p. 179. 

2. Brownish hairs on posterior dorsum prominent. Dark-tipped hairs of hind limb 
not reaching heel. Tail average 10 per cent longer. (Sinai Peninsula, Eastern 
Desert) mariae, p. 180. 

Dipodillus henleyi henleyi De Winton, 1903 

Type /oco/ity.— Egypt. BEHEIRA: Wadi el Natroun, Zaghig. 

Distribution in Egypt— Figure 54. Western Nile Delta and 
Mediterranean Coastal Desert to Libyan frontier. 

External characters.— Slightly darker than D. mariae, but 
brownish tipped hairs on posterior dorsum less prominent. Ven- 
trolateral strip of hairs with white bases narrower. Whitish rump 
patch smaller. Dark-tipped hairs of hind leg reaching heel. In 
general, there is less extension of white in D. henleyi than in D. 
mariae. 

Measurements.— Table 17. Dimensions average about the same as 
in D. h. mariae, except for slightly shorter tail. 

Specimens examined.— Total 35. 

BEHEIRA: Bir Victoria (2), Wadi el Natroun (2), Zaghig (Type), Gebel Muluk (1). 



180 FIELDIANA: ZOOLOGY 

ALEXANDRIA: Alexandria 4.8 km. W (1). Amiriya (2). 

MATRUH: Cairo-Alexandria desert road km. 55 (2): Mariut (1): Bahig (1); Burg el 
Arab (1): Abu Mena 1.6 km. E (1|: El Imayid (1): Ras efHekma (5): Mersa Matruh (1); 
Sidi Barrani 19 km. S (2). 51 km. W (2|: Salum 12.8 km. SE (2). 18 km. SE (2): Bir 
Bosslanga (Bir Wair) (3). 

GIZA: Abu GhaUb (1), Abu Rawash (1). 

Published records.— Records are from De Winton (1903), Wassif 
(1956a). Setzer (1958d), and Hoogstraal (1963). 

BEHEIRA: Bir Victoria. Wadi el Natroun. Gebel Muluk. Zaghig. 

MATRUH: Cairo-Alexandria desert road km. 55; Mariut, Burg el Arab; Alexan- 
dria 48 km. W; Mersa Matruh; Sidi Barrani, 52.8 km. W, 10 km. S; Bir Bosslanga 
(Bir Wair). 

GIZA: Abu GhaUb. 

Dipodillus henleyi mariae (Bonhote, 1909) 

Dipodillus mariae Bonhote, 1909, Proc. Zool. Soc., London, pt. 2, p. 792. 
Dipodillus henleyi makrami Setzer, 1958, J. Egyt. Publ. Health Assn., 33, No. 6, 
p. 212. 

Type locality.— Egypt. CAIRO: Gebel Mokattam. 

Distribution in Egypt— Figure 54. Sinai Peninsula and Eastern 
Desert. 

External characters.— Figure 55. Slightly paler thanZ). h. henleyi, 
but dark brownish hairs on posterior dorsum more prominent. Ven- 
trolateral strip of hairs with white bases, wider. Whitish rump area 
larger. Band of dark-tipped hair on side of head extending from 
mystacinal area beneath eye to base of ear, narrower and paler. 
Dark-tipped hairs of hind leg not reaching heel. In general, there is 
more extension of white in D. h. mariae than in D. h. henleyi. 

Cranial characters.— Although reported by Setzer (1958d) to have 
larger bullae than D. h. henleyi, measurements of same and skull 
height are not confirmative (table 17). 

Measurements.— Table 17. About the same average dimensions as 
D. h. henleyi, except for slightly longer tail. 

i?emarAfs.— Specimens from the Eastern Desert previously allot- 
ted toD. h. henleyi (Wassif, 1956; Hoogstraal et al., 1957b) andD. h. 
makrami (Setzer, 1958d) are here identified as subspecies D. h. 
mariae because of similarity in size and color and proximity of 
distribution. 

Specimens examined— Total 19. 

ISMAILIA: El BaUah (1), El Qantara (3). 

SUEZ: Cairo-Suez road km. 22 <1), Wadi el Rokham <1). 



OSBORN&HELMY: MAMMALS OF EGYPT 181 

CAIRO: Cairo (1). Gebel Mokattam (1). Gebel el Ahmar (1), Heliopolis 8 km. E (2), 
Cairo-suez road 2 km. E of Cairo (1). Maadi (1), Wadi Digla 3.2 km. E of Maadi (1). 

RED SEA: Wadi Araba, St. Anthony Monastery area (1); Wad Bali (1). 

SUDAN ADMINISTRATIVE: Wadi Ibib (1). Bir Kansisrob 3.2 km. N (1). Halaib 
20.8 km. NW (1). 

Published records.— Records are from Allen (1915), Wassif 
(1956a), Setzer (1952, 1958d), and Haim and Tchernov (1974). 

SINAI: Wadi el Feiran, Gebel Maghara, Bik'at Hayareach southeast of Ras el 
Naqb. 

ISMAILIA: El BaUah. El Qantara. 

SHARQIYA: Faqus. 

CAIRO: Gebel Mokattam, Heliopolis, Heliopolis 8 km. E, Ain Shams, Maadi, 
Wadi Digla. 

SUEZ: Cairo 35 km. E, 22 km. E. 

SUDAN ADMINISTRATIVE: Bir Kansisrob 3.2 km. N; Halaib 20.8 km. NW, 3.2 
km. W. 

Genus Sekeetamys EUerman, 1947 

Monotypic genus of dipodil-like rodent with naked palm and sole; 
long, fluffy fur; and black, bushy tail with white tip. Tubercles and 
pads of palm, and tubercles of sole as in genus Dipodillus. 

Bulla greatly inflated; lateral accessory mastoid chamber present; 
medial superior posterior mastoid cavity relatively large, visible 
from behind and not concealed by exoccipital as in genus Meriones. 
Meatal lip swollen slightly ventrally. Accessory tympanum absent. 
Suprameatal triangle small. Superior wall of parapterygoid fossa 
perforated. Tubercles and configuration of upper first molar as in 
Dipodillus. Upper third molar with large, transient posterolabial 
fold. 

Sekeetamys calurus (Thomas, 1892) 

Gerbillus calurus Thomas, 1892, Ann. Mag. Nat. Hist., (ser. 6), 9. p. 76. 

Type locality. -Egypt. SINAI: "Unknown" (Thomas, 1892b, p. 
77); Tor (Chaworth-Musters and EUerman, 1947, p. 482). 

General distribution.— V^estern portions of Arabian Peninsula, 
Jordan, southeastern Israel, Sinai Peninsula, Eastern Desert of 
Egypt. 

Common names.— Bushy Tailed Dipodil, Bushy Tailed Jird, Abu 
Ya. 

Distribution of subspecies in Egypt. — Figure 56. Sekeetamys 
calurus calurus: Sinai Peninsula; Sekeetamys calurus makrami: 
Eastern Desert. 



182 



FIELDIANA: ZOOLOGY 



2 5* 2 6* 2 7* 28* 2 9* 30* 



32* 33* 34* 35* 36* 37* 




Fui 56. Collection localities of Sekeetamys calurus calurus (circles) and 
makrami (dots) and sight records (S). 



.S. c. 



Diagnosis.— harge, dipodil-like rodent with naked palm and sole, 
fur long and fluffy, dorsal hairs brownish yellow with blackish tips, 
sides yellowish to orangish, underparts and feet white. Tail bushy, 
blackish with white tip. 

Skull with greatly inflated bulla, small suprameatal triangle, 
mastoid with lateral accessory chamber, meatal lip not swollen, 
accessory tympanum absent, medial superior mastoid cavity visible 
posteriorly. 

Adult head and body length average 114 mm.; tail 145 mm., 128 
per cent of head and body length; foot 32 mm,; ear 20 mm.; 
occipitonasal length 35.6 mm,; weight 41 gm. 

External characters. — Figure 57. Upper parts dark brownish 
yellow. Dorsal hair tips black, subterminal bands yellowish. Side 



OSBORN & HELMY: MAMMALS OF EGYPT 



183 




Fig 57. Live specimen of Sekeetamys calurus makrami. 

with prominent line of yellowish to orangish extending to wrist and 
ankle. Hairs of back and side with gray bases, hairs of belly and feet 
white. Mystacial area orangish, suborbital and subauricular areas 
pale, postorbital and postauricular spots white. White rump patch 
absent. Ear prominent, sparsely haired, pigmented. Tail bushy and 
"squirrel-like" (Allen, 1915, p. 6), basal one-fifth color of back, distal 
four-fifths fuscous to blackish, not bicolored, usually with a con- 
spicuous white tip. 

Palatal ridges.— Figyire 33, Diastemal ridges rather straight; first 
to fourth intermolar ridges recurved, and all about the same length; 
fifth directed medially, slightly shorter, and thicker. 

Glans penis and baculum.— As in Dipodillus sp. (Wassif et al., 
1969). 

Feet.— Figure 34. Palm and sole hairless. Sole pigmented. Hand 
with three postdigital tubercles and two distinct palmar pads. Foot 
with three postidigital, one posthallucal, and two tarsal tubercles. 




Flu. 58. Skull of Sekeetamys calurus makramL 



184 



OSBORN & HELMY: MAMMALS OF EGYPT 185 

Cranial characters.— Figure 58. Skull elongate with prominent 
supraorbital ridges and conspicuous cranial ridges, nasals pointed 
posteriorly, interparietal outline ovoid, bulla markedly inflated. 
Anterior surface of tympanic bulla extending almost to level of 
anterior margin of foramen ovale. Posterior margin of mastoid 
chambers extending beyond level of paroccipital and supraoccipital. 
Medial superior posterior mastoid cavity visible from behind. Ac- 
cessory mastoid chamber posterior to auditory meatus. 
Suprameatal triangle small, or "vestigial" (Ellerman, 1941, p. 527), 
closed posteriorly by vertical extension of L-shaped suprameatal 
process of temporal bone. Accessory tympanum absent. Auditory 
meatus inflated slightly ventrally. Width across meatuses slightly 
greater than zygomatic width. Parapterygoid fossa with large per- 
foration in superior wall. Zygomatic arch slender. Zygomatic plate 
broad and high, anterior margin gradusdly rounded, not reaching 
level of premaxillary-maxillary suture. 

Teeth.— Figure 59. Upper incisor grooved on anterior surface. 
Upper first molar with alternate first lingual and labial cusps. 
Cusps of mi become confluent shortly before m^ as in other Ger- 
billinae. M^ with transient posterolabial fold. 

Measurements.— Table 18. Male and female dimensions subequal. 

Age determination.— Adults have m' with anterior cusps con- 
fluent and cranial sutures closed, as in gerbils and dipodils. 

Variation.— The tail lacks a white tip in a few specimens from 
Sinai and the Eastern Desert, including the type of S. c. makrami. 
Specimens of S. c. makrami from the Eastern Desert are generally 
darker than S. c. calurus from Sinai Peninsula and, except for five 
pale individuals from sandstone and limestone habitats in the 
northern part of the Eastern Desert, have a narrower line of clear 
color on the side. 

The posterolabial fold on m^ is visible in 10 (50 per cent) of 20 
specimens from the Eastern Desert and 21 (70 per cent) of 30 
specimens from Sinai Peninsula. Observations indicate that the 
posterolabial fold is transient and disappears at an earlier age in 
Eastern Desert animals than in those from the Sinai Peninsula. The 
type specimen of S. c. makrami has m^ peglike and lacking a 
posterolabial fold, Sekeetamys c. makrami average dimensions are 
mostly slightly smaller than S. c. calurus (table 18). Intergrades be- 
tween the two subspecies occur in the northern part of the Eastern 
Desert. 



186 FIELDIANA: ZOOLOGY 

Comparisons.— Sekeetamys calurus differs strikingly from all 
other Egyptian Gerbillinae by its bushy tail with white tip. One 
other rodent, Eliomys quercinus, has a black bushy tail, but is 
distinguished by a black facial mask. Cranially, Sekeetamys differs 
from the genus Meriones in that the medial superior jwsterior 
mastoid cavity is not covered by the exoccipital. The molar pattern 
also differs in the two genera (fig. 59). Feet (fig. 34), teeth (figs. 38, 
59), and bulla are most similar to genus Dipodillus. Palatal ridges of 
Sekeetamys are distinctive (fig. 33). The diploid chromosome 
number is 38, as in Egyptian Gerbillus pyramidum (Wassif et al., 
1969). 

Remarks.— TYas rodent, like Pachyuromys duprasi, is an aberrant 
form requiring special taxonomic consideration. Previously, S. 
calurus has been considered to be a separate genus as well as a 
subgenus, or a species of Gerbillus, Dipodillus, or Meriones (Ander- 
son, 1902; Allen, 1915, 1939; Innes, 1932; Flower, 1932; Ellerman, 
1941, 1948, 1949; Chaworth-Musters and Ellerman, 1947; Ellerman 
and Morrison-Scott, 1951; Wassif, 1954; Wassif and Hoogstraal, 
1953; Petter, 1956; Wassif et al., 1969). 

Collection.— Trapped alive in rocky habitats, shelving limestone 
(fig. 14), pockets in sandstone cliffs, crevices in granite, or among 
boulders; sometimes far from vegetation. 

//a6i tats.— Strictly a rock-adapted rodent, S. calurus is rarely 
trapped away from rocks or cliffs (fig. 14) and is occasionally cap- 
tured in abandoned stone huts. Reported from mountain tops in 
Sinai Peninsula (Haim and Tchernov, 1974). 

Behavior.— Easily excited when first captured, extremely agile, 
and difficult to handle; becomes docile after several weeks of 
repeated handling. Specimens taken from live traps invariably have 
the snout injured from striking the wire mesh in attempting to 
escape. 

Climbing ability, according to Zahavi and Wahrman (1957), sur- 
passes that of Dipodillus dasyurus, which live in the same habitat. 
The bushy tail, they think, may be of some advantage. 

Nocturnal, becoming active at dusk, Sekeetamys is beautifully 
graceful in motion. The tail held squirrel-like in an upright, curved 
position, is never allowed to touch the ground. 

Food— Near Fawakhir mine, S. makrami was trapped in crevices 
containing parts of seed capsules of Zilla spinosa, seed coats of 









^S) ^^^) _y:-^ ^^T-^J •^») (i^k 

^ ^ ^ ^ 1^ ^ 





A 
^ 

'^%!) 
^'W 



S.CAlUffUS 



M.CRASSUS 



Fk; 59. Crown views of right upper (U) and left lower (L) molars of mature (M) 
and immature (I) Sekeetamys calurus, Meriones crassus, Psammomys obesus, and 
Pachyuromys duprasi. 



Table 18. — Means (and ranges) of measurements, ratios, and weight of adult 
Sekeetamys calurus. 





S. c. calurus 


S. c. makrami 


HBL 


118.9(98-128)25 


110.1 (98-119)20 


TL 


144.4(131-164)20 


146.2(131-164) 19 


TL/HBL% 


123.6(110-148)20 


133.3(116-149) 19 


FL 


33.1 (31-35) 25 


31.3 (29-33) 21 


EL 


21.4 (20-23) 25 


19.2(17.0-20.5)21 


Wt 


~ 


41.4 (26.6-49.8) 17 


ONL 


35.8 (34.5-37.4) 20 


35.4 (33.0-37.5) 18 


ZW 


18.2(17.2-19.1) 10 


18.0(16.9-19.0) 18 


lOW 


5.5 (5.2-5.9) 26 


5.4 (5.0-5.7) 20 


NL 


14.4 (13.5-15.5) 20 


13.6(12.4-15.5) 17 


IFL 


6.4 (6.1-6.8) 18 


5.8 (5.2-6.4) 20 


AL 


5.2 (4.8-5.8) 17 


4.8 (4.5-5.1) 20 


RW 


4.6(4.3-5.1)26 


4.6(4.2-5.1) 19 


BL 


11.7(11.0-12.7)25 


11.6(11.0-12.3)20 


SH 


13.8(13.0-14.7)22 


13.6(13.2-14.3)20 



187 



188 FIELDIANA: ZOOLOGY 

Citrullus colocynthis, and parts of succulent Zygophyllum 
coccineum. In Wadi Fatira, Aerva javanica was eaten and carried to 
dens. Branches of Z. coccineum and Cleome droserifolia given to 
captive Sekeetamys were eaten with apparent relish. Captive 
animals also ate cockroaches and crickets. 

Associates.— Eliomys quercinus, Acomys cahirinus, A. russatus, 
and Dipodillus dasyurus are rock-adapted and live in the same 
habitat as Sekeetamys. 

Reproduction.— No information from nature is available. Flower 
(1932) reported litters from captive animals every month of the year 
except September. Number of young averaged 2.8 in 47 litters; two 
litters of six were recorded. 

Sex ratio.— A sample of 24 from the Eastern Desert consisted of 
13 (54 per cent) males and 11 females. 

Key to Egyptian Subspecies of 
Sekeetamys calurus 

1. Color pale, side stripe wider; dimensions slightly larger, particu- 

larly incisive foramina length. (Sinai Peninsula) calurus, p. 188. 

2. Color dark, side stripe narrower; dimensions slightly smaller, 

particularly incisive foramina length. (Eastern Desert) 

makrami, p. 189. 

Sekeetamys calurus calurus (Thomas, 1892). 

Type /oca/i(y. -Egypt. SINAI: Tor. 

Distribution in Egypt— Figure 56. Northern, central, and 
southern parts of Sinai Peninsula. 

External characters. —See species description. Sekeetamys c. 
calurus has a narrower dorsal stripe, broader strip of clear color on 
side, and is paler than S. c. makrami. 

Cranial characters.— See species description. 

7>e(/r.— Figure 59, and see discussion under species. Posterolabial 
folds on m^ were found in 70 per cent of S. c. calurus and 50 per cent 
of S. c. makrami specimens. 

Measurements.— Table 18. Sekeetamys c. calurus average dimen- 
sions are slightly larger than S. c. makrami, especially incisive 
foramina length. 

Specimens examined.— Total 30. 



OSBORN&HELMY: MAMMALS OF EGYPT 189 

SINAI: Sinai (locality not stated) (2); Wadi el Sheikh (6). 4.8 km. W (5); St. 
Catherine Monastery area (9); Wadi Raha (2); Tor (5); Umm Bugma (1). 

Published records.— Records are from Anderson (1902), Allen 
(1915), Flower (1932), Wassif and Hoogstraal (1953), Setzer (1961a), 
and Haim and Tchernov (1974). 

SINAI: Abu Zenima, Umm Bugma (Ambogma), Wadi Saal, Tor, St. Catherine 
Monastery area, Gebel Yiallaq (Yelleq). 

Sekeetamys calurus makrami (Setzer, 1961) 

Sekeetamys makrami Setzer, 1961, J. Egypt. Pub. Health Assn., 36, No. 3, p. 90. 

Type locality.-Egypt. RED SEA: Wadi Gumbiet. 

Distribution in Egypt.— Figure 56. Eastern Desert. 

External characters.— See species description. Sekeetamys c. 
makrami has a broader dorsal stripe, narrower strip of clear color on 
side, and is slightly darker than the nominate subspecies in the 
southern part, but not in the northern part, of the Eastern Desert. 

Cranial characters.— See species description. 

Teeth.— Figure 59, and see discussion under species and S. c. 
calurus. 

Measurements.— Table 18. Sekeetamys c. makrami average 
dimensions are mostly smaller than S. c. calurus, especially incisive 
foramina length. 

Remarks.— Some differentiation, probably due to isolation, has 
occurred between populations of S. calurus in Sinai and the Eastern 
Desert. The degree of difference is not considered sufficient to 
warrant full species rank for the Eastern Desert population, as pro- 
posed by Setzer (1961a). Setzer's decision was based on a single 
atypical specimen. 

Specimens examined.— Total 25. 

SUEZ: Ain Sukhna cliffs (2). Wadi Qiseib (1), Wadi Dom (1). 

CAIRO: Wadi Hof (2). 

RED SEA: Wadi Fatira, Abu Kharif mine area (2); Wadi Abu Sheeh (1); mouth of 
Wadi Atalla (3); Bir Seyala (1); Wadi el Hammamat, Fawakhir mine area (8); Wadi 
Abu Qraiya (1); Wadi Sikait (1); Bir Gumbiet (Type). 

SUDAN ADMINISTRATIVE: Gebel Nesla (1). 

Published records. — Records are from Anderson (1902), 
Hoogstraal et al. (1957b), and Setzer (1961a). 

RED SEA: Wadi Sikait. Bir Gumbeit. 



190 FIELDIANA: ZOOLOGY 

Sight record of D. J. Osbom.— 

SUDAN ADMINISTRATIVE: Gebel Elba. Wadi Akwamtra tribuUry. 

Genus Meriones Illiger, 1811 

Slender to stocky rodents with dorsum yellowish brown to 
brownish, venter white. Ear large, sparsely haired. Tail fully haired, 
tip bicolored with a black dorsal brush. Length of tail greater than 
85 per cent of head and body length. Palm bare, sole partly haired. 
Hand with three postdigital tubercles and two large palmar pads. 
Foot with one large postdigital pad, a hallucal tubercle, and no 
plantar tubercles (fig. 34). 

Skull angular in some species, usually with supraorbital ridge well 
developed. Tympanic bulla prominently inflated, mastoid and 
meatal swelling variable. Accessory tympanum absent in most 
species. Interparietal and exoccipital slightly modified by expan- 
sion of bullae. Medial superior posterior mastoid cavity relatively 
small and concealed by exoccipital. 

Upper incisor with single groove on anterior surface. Molars 
hypsodont when immature, always with prismatic crowns; never 
tuberculate. First upper molar three-rooted in adults. Third upper 
and lower molars usually simple, peglike, occasionally with single 
transient fold. 

Key to Egyptian Species of 
Meriones 

1. Posterior surface of mastoid bulla inflated beyond level of paroccipital process. 
External auditory meatus swollen to or almost to level of zygomatic process of 
temporal. Tail with long, conspicuous black brush. 

a. Suprameatal triangle open posteriorly. Accessory tympanum absent. Tail 
base buffy. Claws pale. 

i. Bulla excessively inflated. Exoccipital and basioccipital constricted (fig. 
60). Ear not pigmented. Feet white. (Sinai Peninsula. Eastern and 
Western Deserts) crassus, p. 191. 

ii. Bulla not excessively inflated. Exoccipital and basioccipital not con- 
stricted (fig. 61). Ear pigmented. Feet partly colored. (Northeastern Sinai 
Peninsula) sacramenti, p. 204. 

b. Suprameatal triangle closed posteriorly. Accessory tympanum present. Tail 

base orangish. Ears not pigmented. Claws black. (Western Desert) 

libycus, p. 207. 

2. Posterior surface of mastoid bulla not inflated beyond level of paroccipital pro- 
cess. Auditory meatus not greatly swollen. Tail without long, conspicuous 
black brush. 

a. Suprameatal triangle partially closed posteriorly. Belly hairs with gray 
base. Ear pigmented. Tail brush large. (Western Desert) shawi, p. 214. 



OSBORN&HELMY: MAMMALS OF EGYPT 191 

b. Suprameatal triangle completely closed posteriorly. Belly hairs usually lack- 
ing gray base. Ear pigmented. Tail brush very small. (Northeastern Sinai 
Peninsula) tristrami, p. 218. 

Meriones crassus Sundevall, 1842 

Meriones crassus Sundevall, 1842, Kongel. Svenska Vet.-Akad. Handl. 
Stockholm, pi. II, fig. 4a,b,c,d, p. 233. 

Type locality.— Egypt. SINAI: Ayun Musa. 

General distribution.— West Pakistan, Afghanistan, southern 
Russian Turkestan, Iran, Iraq, Syria, Lebanon, Jordan, Israel, 
Saudi Arabia, Sinai Peninsula, Egypt, northern Sudan, Libya, 
Algeria, northern Nigeria. 

Common names.— Silky Jird, Sundevall's Jird. 

Distribution of subspecies in Egypt— Figure 62. Meriones 
crassus crassus: Sinai Peninsula and greater part of Eastern Desert; 
Meriones crassus pallidus: Southern part of Eastern Dessert; 
Meriones crassus perpallidus: Western Desert. 

Diagnosis.— harge jird with long, soft dorsal pelage. Dorsum pale 
yellowish brown, side with narrow, buff-colored areas, hairs of 
venter and feet white. Mystacial and circumorbital areas pale; 
postauricular patch conspicuous, white. Tail either faintly or not 
bicolored, white or buffy below, upper surface as dorsum, with con- 
spicuous black apical brush. Ear, sole, and claws not pigmented. 
Sole partly haired. 

Skull angular, strongly ridged in adults. Bulla inflated posteriorly 
beyond level of paroccipital process; anterior lip of external auditory 
meatus conspicuously swollen to level of zygomatic process of tem- 
poral. Accessory tympanum absent. 

Adult head and body length average 136 mm.; tail 133 mm., 98 
per cent of head and body length; hind foot 34 mm.; ear 18 mm.; 
occipitonasal length 38.6 mm.; weight 80 gm. 

External characters.— Figure 63. Dorsal pelage long, soft. Dor- 
sum pale brownish yellow, finely marked with black. Dorsal color 
separated from white of venter by narrow, clear buff area on side 
and thigh. All dorsal hairs, except white postauricular patch, with 
gray bases. Belly hairs with or without gray bases. Mystacial and 
circumorbital areas pale. Dark color of upper portion of head not 
extending below level of nose, eye, and ear. Postauricular patch con- 
spicuous, white. Ear with long, pale buff to whitish hairs on anterior 




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192 






Fk; 61. Skull of Meriones sacramenti. 



193 



194 



FIELDIANA: ZOOLOGY 



,. 25* 26* 27* 26* 2 9* 30* 31* 32* 33* 34 



35 36 37* 




Fici 62. Collection localities of Meriones crassus crassus (squares), M. c. per- 
palUdus (dots), and M. c. pallidus (circles). 

margin; whitish hairs sparsely covering inner and outer surfaces of 
pinna. Foot and hand white. Tail with upper surface color of 
dorsum; underside usually white, distinctly or indistinctly 
bicolored. Apical brush prominent, black, about one-third length of 
tail. Underside of tip clear white or with scattered black hairs. 

Palatal ridges. — Figure 33. Diastemal ridges thick, straight. 
First, second, and third intermolar ridges recurved, almost reaching 
midline. Fourth and fifth intermolar ridges directed medially. 

Glans penis and baculum.— The minute spines of the penis are in 
pentagonal sockets. The three cartilaginous processes of the 
baculum are not separate, but joined together at their bases and 
separated from the shaft by connective tissue. (See under Gerbillus 
pyramidum and Wassif et al., 1969.) 



OSBORN & HELMY: MAMMALS OF EGYPT 



195 




Fig. 63. Live specimen of Meriones cmssus. 



Feet— Figure 34. Palm bare, sole partly haired. Fore foot with 
three distinct postdigital pads and two distinct proximal plantar 
pads. Hind foot with large, lobed postdigital pad and a single 
hallucal pad. 

Cranial characters.— Figure 60. Skull angular, strongly ridged in 
adults. Supraorbital ridge prominent. Posterior margin of nasals 
variable in position, but usually anterior to posterior level of fronto- 
premaxillary suture (table 20). Interparietal narrow with posterior 
margin angular. Zygomatic plate rarely reaching level of 
premaxillary-maxillary suture, completely or partially covering 
infraorbital foramen in lateral view. Small notch in base of 
zygomatic plate allows exposure of the foramen. Posterior mastoid 
chambers of bulla swollen markedly beyond level of paroccipital 
process and supraoccipital. Suprameatal triangle open posteriorly; 



196 FIELDIANA: ZOOLOGY 

not enclosed by occipital or temporal processes. Tympanic bulla 
conspicuously swollen with anterior margin encroaching onto the 
parapterygoid fossa and reaching level of anterior margin of 
foramen ovale. Auditory meatal area swollen dorsally, ventrally, 
and anteriorly, usually contacting zygomatic process of temporal 
(fig. 60). Width across external auditory meatal swellings greater 
than zygomatic width. Accessory tympanum absent. 

TWetA.— Anterior surface of upper incisor grooved. Molariform 
teeth hypsodont in immatures, cusps prismatic, never tubercular 
(fig. 59). Upper and lower third molars peglike. 

Measurements.— Table 19. Males consistently larger than 
females. Measurements are of adult males. Means (and ranges) of 
occipitonasal length (in millimeters) of 10 adult males and 11 adult 
females are 39.7 (37.7 to 42.2) and 38.1 (36.2 to 39.3), respectively. 

Age determination.— Adult specimens have lateral folds of m' not 
extending below level of alveolus, and tooth roots exposed (fig. 73 of 
Psammomys obesus). 

Variation.— Dark and pale individuals exist in all samples 
designated as M. c. pallidas from the southern Eastern Desert (fig. 
62). Samples from Sinai Peninsula, northern Eastern Desert, and 
Western Desert are consistently darker than M. c. pallidas. Samples 
from the Western Desert show a clinal increase from north to south 
in darkness of dorsal color and in percentage of uniformly colored 
vs. bicolored tails. 

Samples from the northern Western Desert were named M. c. per- 
pallidus by Setzer (1961a) and said to be paler than M. c. crassus. 
These subspecies cannot be distinguished by this character alone. 
Gray basal bands on belly hairs predominate in perpallidus and 
increase in width and darkness from north to south. Exposure of the 
infraorbital foramen occurs in a greater percentage of perpallidus 
than of Eastern Desert subspecies (table 20). The frequency of this 
character also increases slightly from north to south in the Western 
Desert. 

The average dimensions of samples from the Western Desert are 
smaller than those from the Eastern Desert, except in the southern 
part of the range of M. c. crassus. The mean of most dimensions 
increases from north to south in the Western Desert and decreases 
from north to south in the Eastern Desert (table 21). 

Comparisons.— Meriones crassus differs from other Egyptian 



Table 19. — Means (and ranges) of measurements, ratios, and weight of adult 
Meriones crassus. 



M. 



c. crassus 



HBL 

TL 

TL/HBL% 

FL 

EL 

Wt 

ONL 

ZW 

low 

NL 

IFL 

AL 

RW 

BL 

SH 

PAW 



136.5 

136.9 

101.3 

34.5 

18.6 

83.3 

39.1 

21.2 

6.1 

15.1 

6.8 

5.6 

5.4 

15.3 

15.9 

10.8 



(114-153)47 
(105-158) 40 
(86.0-116.2)40 
(31-37) 48 
(14-22) 46 
(54.0-112.8)26 
(36.6-42.2) 47 
(18.0-22.9) 38 
(5.3-7.0) 45 
(12.7-16.6)45 
(5.2-8.2) 46 
(4.8-6.4) 45 
(4.8-6.1) 45 
(12.9-16.6) 39 
(14.1-17.0)44 
(9.5-12.5) 32 



M. c. pallidus M. 

134.8(131-138)4 134.1 

135.2(133-140)4 126.1 

100.4 (96.4-104.4) 4 93.7 

35.5 (35-36) 4 32.2 

17.8(16-19)4 18.4 

85.2 (70.7-93.4) 4 76.0 

38.4 (37.3-38.9) 4 37.9 

20.7, 21.0 20.6 

5.9 (5.8-6.0) 4 6.0 

14.6(13.8-15.2)4 14.6 

6.7 (6.3-7.2) 4 6.6 

5.6 (5.3-5.8) 4 5.8 

5.2 (5.1-5.3) 4 5.2 

15.4(14.9-15.7)4 14.8 

15.9 (15.8-16.0) 3 15.5 

10.5(9.9-11.2)4 11.6 



c. perpallidus 

(116-152) 28 
(115-135) 22 
(85.7-103.1)22 
(30-35) 29 
(16-21) 29 
(51.0-99.0) 22 
(34.5-40.4) 25 
(19.1-22.2) 23 
(5.5-6.6) 27 
(12.8-15.5) 23 
(6.0-7.3) 27 
(5.1-6.3) 27 
(4.4-5.8) 27 
(13.8-15.8) 26 
(14.6-16.4) 28 
(10.7-12.2)7 



Table 20. — Color and cranial variations in Meriones crassus. 



Sinai Peninsula 

and Eastern Desert 

M. c. crassus and M. c. pallidus 



Western Desert 
M. c. perpallidus 

1. Basal bands of belly hairs: 

Gray White Gray 

Number 45 13 8 

Percent 81 19 6 

2. Coloration of base of tail: 

Bicolored Not bicolored Bicolored 

Number 34 23 94 

Percent 60 40 78 

3. Position of posterior margin of nasals with respect to posterior level of fronto- 
premaxillary suture: 

Anterior Intermediate Equal Anterior 

Number 39 10 8 110 

Percent 68 18 14 91 

4. Exposure of infraorbital foramen in lateral view: 

Exposed Not exposed Exposed 

Number 38 22 19 

Percent 63 37 23 



White 

120 

94 

Not bicolored 
27 
22 



Intermediate 
6 
5 

Not exposed 
64 

77 



Equal 
5 
4 



197 



198 FIELDIANA: ZOOLOGY 

Tablk 21. — Means (and ranges) of head and body and occipitonasal lengths of 
adult Meriones crassus from Sinai Peninsula and northern to southern localities in 
Eastern and Western Deserts. 

HBL ONL 

Sinai Peninsula 139.0(130-153)8 39.3(36.6-41.3)8 

Eastern Desert 

Wadis Gafra and Iseili 141.3 (132-151) 12 39.7 (37.7-42.2) 10 

Red Sea localities 140.0 (122-148) 8 39.7 (38.4-42.1) 9 

Wadi Asyuti 130.6(120-143)7 37.8(36.7-40.8)7 

Wadi Qena system 131.0 (114-140) 12 38.8 (36.8-40.9) 13 

Wadi AUaqi system 134.8 (131-138) 4 38.4 (37.3-38.9) 4 

Western Desert 

Northern localities 128.2(116-136)6 36.8(34.5-39.2)7 

Hatiyet el Sunt 131.0 (119-144) 12 37.8 (36.3-39.0) 9 

Farafara Oasis 141.3 (123-152) 10 38.9 (35.0-40.4) 9 

species in having paler color, band of clear color on side narrower, 
feet with white hairs only, mastoid chamber and meatus much more 
inflated, exoccipital and basioccipital constricted, and mean of 
paroccipital width (PAW) actually and relatively smaller (tables 19, 
22). The infraorbital foramen is much less exposed in M. crassus 
than in M. shawi and M. tristrami, but about the same as in M. 
sacramenti (figs. 60, 61). Additional comparative characters are in 
Table 23. 

The diploid chromosome number and FN of M. crassus are 60 and 
72; of M. libycus, 44 and 74; M. shawi, 44 and 78; and M tristrami, 
72 and 74 (Nadler and Lay, 1967). 

Collection.— Meriones crassus readily enters live traps placed 
near burrows or in vegetation. Digging is sometimes difficult due to 
hardness of terrain. Occasionally, specimens have been captured by 
hand at night on barren desert. 

Habitats.— TYds species is found in wadis and coastal areas of 
Sinai Peninsula and the Eastern Desert (figs. 5, 15) where there is 
vegetation or human habitation or past activity. Burrows may be in 
barren, stoney, gravelly, or mud terraces around or beneath 
buildings or tents; under trash heaps (Briscoe, 1956) and straw 
piles, but not always in the immediate vicinity of a food source. 
Hoogstraal (1963) observed burrows 0.2 km. from vegetation in 
Wadi Digla east of Maadi. Similar situations exist in the Western 
Desert (fig. 64). 

Shrubs with which M. crassus is usually associated in wadis of the 



OSBORN & HELMY: MAMMALS OF EGYPT 



199 




Fui 64. Western Desert. Pebble desert interspersed with soft sand and gypsum 
(nafash). About 7 km. SE of km. 208 on Cairo-Bahariya road. Tree in background 
{Acacia raddiana) is over 100 m. from the burrow. Habitat of Meriones crassus per- 
pallidus. At 3 km. beyond, there is an acacia grove in a depression at about 28°55' N 
iat.. 29°31' E long. 

Eastern Desert are Cassia senna (Hoogstraal et al., 1957b), C 
italica, Lygos raetum, Acacia sp.; Leptadenia pyrotechnica; 
Calligonum comosum; Zilla spinosa; Citrullus colocynthis, also men- 
tioned by Lewis et al. (1965) in northern Saudi Arabia; and Panicum 
turgidum, also noted by Briscoe (1956). Patches of date palms 
{Phoenix dactylifera) and rushes ijuncus sp.) may also harbor col- 
onies of this jird. Near Gulf of Suez shores, M. Crassus has been 
trapped beside Nitraria retusa. 

In the Western Desert, this species is found under acacias and in 



200 FIELDIANA: ZOOLOGY 

surrounding desert (fig. 64), beside Calligonum comosum (fig. 9), 
near human camps and trash heaps in otherwise barren desert; in 
shallow depressions beneath stands of Hyoscyamus muticus; in 
outlying areas around oases beneath f4itraria retusa (fig. 17), and in 
sand beneath date palms. The western-most locality was barren 
desert between Siwa and Qara. Ranck (1968) referred to Libyan 
habitats of M. crassus as marginal, since he did not collect it in date 
palm groves or oasis vegetation. 

Activity.— Lewis et al. (1965) found that, in northern Saudi 
Arabia, M. crassus became active at twilight, and the period of 
greatest activity was for about an hour after dark. In Wadi Umm 
Karayiet, a tributary of Wadi AUaqi, M. c. pallidas was taken in our 
live traps between 1200 and 1400 hours (universal time) in March. 

Captive behavior.— Meriones crassus, except for Pachyuromys 
duprasi, is the most docile wild rodent in Egypt. Specimens grasped 
and removed from live traps do not bite and make little effort to 
escape. 

BurroM;s.— Burrows of M. crassus are shallow with numerous 
openings (Briscoe, 1956), often in very hard ground. This species is 
not known to plug openings with sand or earth. 

Burrow microclimate.— According to Briscoe (1956), burrow 
temperatures during late September 1953 ranged from 77 ° to 97 °F., 
and relative humidity, from 24 to 78 per cent. Air temperatures at 
this time were 78° to 103°F., and relative humidity, 21.5 to 77 per 
cent. 

Food— Fruits and/or seeds of one or two plant species listed 
above (figs. 9, 15) are often the only food source for this rodent. 
Green vegetation is eaten when available. Bitter, green fruits as well 
as dry pulp and seeds of Citrullus colocynthis, for examples, are 
staple food in much of the Eastern Desert. Vegetable material from 
camel dung around campsites provides food in otherwise sterile 
environments. Lewis et al. (1965, p. 73), stated from observations in 
northern Saudi Arabia: "Perhaps one of the major reasons for its 
success as a species lies in its ability to exist on any food available." 

Tortonese (1948) fed captive Men'ones (probably M. crassus) from 
the vicinity of Tel el Kebir in Wadi Tumilat, leaves, fruit peels, 
bread, cheese, and small pieces of meat. We have fed it raw potatoes, 
raw carrots, dried bread, and various seeds. 

Seeds of Zilla spinosa and seeds and stalks of Anabasis articulata 



OSBORN&HELMY: MAMMALS OF EGYPT 201 

were regularly found near burrow openings in Sinai (Haim and 
Tchernov, 1974). 

Reproduction.— The average number of young in a sample of 10 
reproducing females, including embryos, fetal scars, and nestlings 
was 3.3 (range, 1 to 5). An eight-month breeding period is indicated 
by data from November to June. 

Sex ratio.— In a museum sample of 73 specimens, there were 36 
(49 per cent) males and 37 females. 

Associates.— Rodents living in the same habitat as M. crassus are 
Jaculus jaculus, Gerbillus gerbillus, G. perpallidus, and occasional- 
ly, M. shawi and M. lybicus. 

Key to Egyptian Subspecies of Meriones crassus 

1. Belly hairs usually with gray bases, infraorbital foramen exposed (table 20). 
(Western Desert) perpallidus, p. 202. 

2. Belly hairs usually white to base, infraorbital foramen not exposed (table 20). 

a. Dorsum slightly darker than in pallidus. (Northern Eastern Desert and 
Sinai Peninsula) crassus, p. 201. 

b. Dorsum slightly paler than in crassus. (Southern Eastern Desert) 

pallidus, p. 203. 

Meriones crassus crassus Sundevall, 1842 

Meriones crassus asyutensis Setzer, 1961, J. Egypt. Publ. Health Assn., 36, No. 3, 
p. 82. 

Type locality.— Egypt. SINAI: Ayun Musa. 

Distribution in Egypt— Figare 62. Sinai Peninsula and greater 
part of Eastern Desert. 

External characters. —See under species description. Meriones c. 
crassus specimens usually lack gray bases on belly hairs (table 20). 

Cranial characters. — Infraorbital foramen not exposed in lateral 
view in a larger proportion of individuals (table 20) than in other 
subspecies. 

Measurements.— Table 19. Average dimensions of M. c. crassus 
are larger than pallidus and perpallidus. 

Remarks.— Data from M. c. crassus and pallidus are combined in 
Table 20. Meriones c. asyutensis described by Setzer (1961a, p. 82) 
from three specimens is considered synonymous with M. c. crassus. 
Seven additional specimens from the type locality do not exhibit the 
paleness described by Setzer. See also remarks under M. shawi. 



202 FIELDIANA: ZOOLOGY 

Specimens examined.— Total 193. 

SINAI: Bir Thai (1): El Quseima (2): Gebel el Bruk t2); Feiran Oasis (2). 1.6 km. E 
(4): Wadi el Sheikh (5); St. Catherine Monastery area (8); IsmaiUa 80 km. E (1|. 

ISMAILIA: Fayid 4.8 km. NW (2|. 

SUEZ: HeUopoUs 12.8 km. E (1): Cairo-Suez road km. 18 (3), km. 29 (1|, km. 34 (1). 
km. 35 (1). km. 40 <4): El Dar el Bayda (2); Wadi el Gafra (14|; Wadi IseiU (23); Bir Gin- 
dali (1): Gebel el Katamiya observatory area (2); Wadi Dom (3|; Wadi Abu Sanduq (6|. 

RED SEA: Wadi el Nil (4); St. Paul Monastery area (1); Bir Zafarana (2); Ras 
Zafarana (6): Wadi Bali (2); Wadi Umm Delfa (2); Bir Abu Zawal (4|; Wadi Fatira (4); 
Abu Kharif mine area (3); Wadi Atalla mouth (7); Wadi Abu Sheeh 96 km. E of Qena 
(1): Wadi Graygar (4); Wadi Semna (2); Wadi Abu Shaar (2); El Ahiah 10 km. N (3): 
Safaga (1); Quseir (3); Qusur el Banat (3); Fawakhir mine 4.8 km. E (2); Wadi Umm el 
Seniyet (2); Wadi Abu Ziran (4); Wadi Abu Qraiya (1); Wadi Sukari (2); Bir Shalatein 
(3). 

QENA: Wadi el Sheikh Isa mouth (4). 

SHARQIYA: Bilbeis (2). 

CAIRO: Gebel el Ahmar (1); Heliopolis 4.8 km. E (1). 8 km. E (8); Cairo-Suez road 
km. 11 (1); Wadi Digla 3.2 km. E of Maadi (4); Wadi Hof (2): Wadi Garawi 10 km. SE 
of Helwan (3). 

ASYUT: Wadi Asyuti (10). 

Published records.— Records are from Bonhote (1912), Allen 
(1915), Flower (1932), Cha worth-Musters and Ellerman (1947), 
Setzer (1952, 1961a), and Haim and Tchernov (1974). 

SINAI: Ayun Musa; Gebel el Bruk, Nakhl, Kossaima (El Quseima), Tor, Wadi 
Feiran, Feiran Oasis, Feiran Oasis 1.6 km. E, St. Catherine Monastery area, Bir 
Thai. Mt. Sinai, Umm Shomer, Gebel El Igema (Gebel Egma). 

ISMAILIA: Fayid 4.8 km. NW. 

SUEZ: Cairo-Suez road km. 12.8, km. 18, km. 35; Bir Gindali. 

CAIRO: Gebel el Ahmar; Cairo-Suez road km. 8, km. 11; Maadi 3.2 km. E. 

RED SEA: Ras Zafarana, Quseir. 



Meriones crassus perpallidus Setzer, 1961 

Meriones crassus perpallidus Setzer, 1961, J. Egyp 
p. 86. 

Type locality.— Egypt. GIZA: Cairo- Alexandria desert road km. 
4. 



Meriones crassus perpallidus Setzer, 1961, J. Egypt. Publ. Health Assn. 36, No. 3, 
p. 86. 



Distribution in Egypt— Figure 62. Western Desert south of 
Western Mediterranean Coastal Desert vegetation. 

External characters. — Under species description. Belly hairs of M. 
c. perpallidus usually have a gray base (table 20). 

Cranial characters.—See species description. A larger proportion 



OSBORN&HELMY: MAMMALS OF EGYPT 203 

of perpallidus have the infraorbital foramen exposed in lateral view 
(table 20) than in other subspecies. 

Measurements.— Table 19. 

Sanation.— Samples from the northern part of the Western 
Desert are paler, include a smaller percentage of individuals having 
belly hairs with gray bases, and have smaller average dimensions 
than samples from further south. 

Comparisons. —Meriones c. perpallidus differs from M. c. pallidas 
in the same way that it differs from the nominate subspecies: darker 
color, belly hairs with gray bases, and slightly smaller dimensions. 

i?emar/js.— Specimens of M. c. perpallidus examined by Setzer 
(1961a) differed little from the nominate subspecies, except in size, 
but clinal variation from north to south in size, color, exposure of 
infraorbital foramen, and other characters (table 20) are distinctive 
enough to warrant retention of the trinomen. 

Meriones crassus selysi (Pomel, 1856) is an Algerian form. The 
name was applied to specimens from Wadi el Natroun by Schwann 
(1905) and Bonhote (1912b). Other subspecies have since been 
described from intermediate areas. 

Specimens examined.— Total 75. 

GIZA: Giza (1); Cairo-Alexandria desert road km. 10 (1), km. 4 (Type); Gebel el 
Ghigiga (3); Abu Rawash (1); Kirdasa (3); Cairo-Bahariya road km. 208, Acacia 
grove, 7 km. E (6). 

FAIYUM: Lake Qarun 3 km. NW (1). 

EL MINYA: el Bahnasa (3). Hatyet el Sunt (16). 

BEHEIRA: Bir Victoria (7), Wadi el Natroun (2). 

MATRUH: Camel Pass Dune area (4); El Maghra 27 km. W (1); Raqabet el Rala (1); 
Bir Nahid (1); Siwa-Qara road 46 km. NE of Siwa (1); Qara (1). French Camp No. 2 (1). 

EL WADI EL GEDEED: Farafara Oasis, El Khanafis (1); Bir Qokshira (2). 4.8 
km. NE (1); Abu Minqar (14), 80 km. S (2); Kharga Oasis, Ain Amur (1). 

Published records.— Records are from Schwann (1905), Innes 
(1932), and Setzer (1961a). 

BEHEIRA: Bir Victoria, Wadi el Natroun, Abu Makkar Monastery. 
GIZA: Cairo-Alexandria desert road km. 4.6, km. 10; Kirdasa. 
FAIYUM: Lake Qarun 3 km. NW. 
EL MINYA: El Bahnasa. 

Meriones crassus pallidas Bonhote, 1912 

Meriones crassus pallidas Bonhote, 1912, Abstr. Proc. Zool. Soc., London, No. 
103, p. 3. 



204 FIELDIANA: ZOOLOGY 

Type locaiity. -Sudan. NORTHERN: Atbara. 

Distribution.— Figvire 62. Southern part of Eastern Desert and 
northeastern Sudan. 

External characters.— Meriones c. pallidus is slightly paler than 
other subspecies in Egypt. 

Cranial characters. — See species description. In cranial 
characters, M. c. pallidus does not differ from the nominate form. 

Measurements.— Table 19. 

Remarks.— Meriones c. pallidus differs very little from the 
nominate subspecies except in paler color, as noted by Bonhote 
(1912), and slightly smaller dimensions. 

Specimens examined.— Total 24. 

RED SEA: Wadi Gumbiet (1). Bir Abraq (8). 
SUDAN ADMINISTRATIVE: Abu Ramad (1). 
ASWAN: Wadi Umm Karayiet (11). Bir Haimur (3). 

Published records.— Records are from Hoogstraal et al. (1957b) 
and Setzer (1961a). 

RED SEA: Bir Abraq. Wadi Naam. 

Meriones sacramenti Thomas, 1922 
Meriones sacramenti Thomas, 1922, Ann. Mag. Nat. Hist., (ser. 9), 10, p. 552. 
Type locality. — Israel: Beersheba 16 km. S. 

General distribution.— Southern Israel and northeastern Sinai 
Peninsula. 

Common name.— Negev Jird. 

Distribution in Egypt.— Figure 65. Northeastern Sinai Peninsula. 

Diagnosis.— Large jird with relatively coarse pelage. Dorsum 
dark cinnamon brown, side with line of clear cinnamon, venter 
white. Ear prominent, tip pigmented. Tail with line of black or scat- 
tered black hairs along upper surface, not bicolored. Tail brush 
black, conspicuous. Sole with small bare area. Claws pale. 

Skull angular with strongly developed supraorbital ridge. 
Posterior margin of nasal anterior to posterior level of fronto- 
premaxillary suture. Infraorbital foramen exposed. Mastoid bulla 
inflated beyond level of paroccipital. Suprameatal triangle open 
posteriorly. Auditory meatus swollen to level of zygomatic process 
of temporal. Accessory tympanum absent. 




c 



205 



206 FIELDIANA: ZOOLOGY 

Head and body length average 155 mm.; tail 151 mm., 97 per cent 
of head and body length; hind foot 36 mm.; ear 18 mm.; 
occipitonasal length 41.6 mm. 

External characters.— Dorsum dark cinnamon brown, side with 
conspicuous line of clear cinnamon extending from wrist to heel and 
sometimes onto side of foot. Venter white. Hairs of dorsum and side 
with gray bases, belly hairs with gray bases in about 50 per cent of 
individuals examined. Feet white, except as noted above, claws pale. 
Band extending from mystacial area beneath eye to base of ear 
slightly paler than upper surface of head. Supraorbital patch small, 
indistinct. Postauricular patch small, whitish. Ear with distal one- 
third pigmented. Anterior margin of pinna with long buffy to cin- 
namon hairs, inner hairs white, outer cinnamon. Upper surface of 
tail paler than back, with conspicuous black brush about one-third 
of tail length and a line of black or scattered black hairs nearly 
reaching the base. Tail otherwise buffy and not clearly bicolored 
except for tip. 

Palatal ridges.— Not observed. 

Glans penis and baculum.— Not observed. 

Feet.— Palm bare; sole with small naked area. Claws pale. 

Cranial characters.— Figure 61. Cranium somewhat angular, 
supraorbital ridge strongly developed. Posterior margin of nasals 
rounded, not reaching posterior level of frontopremaxillary suture. 
Anterior margin of zygomatic plate reaching or almost reaching 
level of premaxillary-maxillary suture and not completely covering 
infraorbital foramen. Interparietal with round posterior margin. 
Tympanic bulla markedly swollen, anterior margin level with or 
beyond middle of foramen ovale. Mastoid chambers conspicuously 
swollen, posterior surface extending slightly posterior to paroc- 
cipital process. Suprameatal triangle open posteriorly. External 
auditory meatus swollen anteriorly to level of zygomatic process of 
temporal. Distance across meatuses usually slightly greater than 
zygomatic width. Accessory tympanum absent. 

Teeth. — Upper incisor grooved. Molar pattern prismatic as in 
other species of Meriones. 

Measurements.— Table 22. Male and female dimensions subequal. 

Age determination.— Adults have the same features as M. 
crassus. 



OSBORN&HELMY: MAMMALS OF EGYPT 207 

Variation.— Some individual variation was observed in meatal 
swelling. Specimens from Israel have a continuous black line on the 
upper tail surface from tip to base. In Sinai specimens, the line is 
indefinite. 

Comparisons.— Meriones sacramenti is distinguishable from M. 
crassus by darker color, pigmented ears, color on feet, less inflated 
mastoid chambers and meatus of bulla, and less modification of 
interparietal and occipital. It differs from M. lybicus in having ear 
pigmented, paler color, claws pale, posterior margin of nasals not 
reaching posterior level of frontopremaxillary suture, infraorbital 
foramen exposed, and suprameatal triangle open posteriorly. From 
M. shawi, M. sacramenti differs in having tail brush larger, infra- 
orbital foramen less exposed, and mastoid chamber and meatus of 
bulla much more inflated. Comparison with M. tristrami is under 
that species. Further comparisons are in Tables 22 and 23 and 
Figure 60. Most dimensions of M. sacramenti average slightly 
larger, except for bulla length in M. crassus, than those of other 
Egyptian species of Meriones. 

Remarks.— Meriones sacramenti appears to be more closely 
related to M. shawi than to other species in Egypt. No local informa- 
tion on natural history is available. In Israel, it inhabits sandy 
localities and is unevenly distributed (Zahavi and Wahrman, 1957). 

A single specimen of M. sacramenti from Bir Lehfan, north- 
eastern Sinai Peninsula, called M. shawi shawi by Wassif (1953b), 
was assumed to be M. tristrami by Fetter (1957) because of 
geographic location, and accepted as such by Zahavi and Wahrman 
(1957), Setzer (1961a), and Hoogstraal (1963). 

Specimens of M. sacramenti from Rafa in Giza Zoological 
Museum were marked ''Psammomys. " 

Specimens examined.— Total 21. 

Egypt: SINAI: Bir Lehfan (1). Rafa (9). 

Israel: Beersheba (2), 16 km. S (2); Zahr el Rubin (2); Rishon el Zion (1); Ramleh (2); 
"Palestine" (2). 

Published records. -SINAI: Bir Lehfan (Wassif, 1953b). 

Meriones libycus (Lichtenstein, 1823) 

Gerbillus libycus Lichtenstein. 1823, Verzeich. Doubl. Zool. Mus. Berlin, No. 9, 
p. 5. 

Type locality.— Egypt: Libyan Desert of describer taken to mean 



208 FIELDIANA: ZOOLOGY 

"near Alexandria" by Chaworth-Musters and Ellerman (1947, p. 
485). 

General distribution. — Iran, Azerbaijan S.S.R., Iraq, Syria, Jor- 
dan, Israel, Western Desert of Egypt, Libya. 

Common name.— Libyan Jird. 

Subspecies in Egypt — 

Meriones Ubycus libycus Lichtenstein, 1823 

Type locality.— Egypt. Western Desert, probably south of Alex- 
andria. 

Distribution in Egypt.— Figure 65. Northern part of Western 
Desert. 

Diagnosis.— harge jird with soft pelage, dorsum dark yellowish 
brown, side with line of clear orangish, venter white. Ear not 
pigmented. Tail color of back, not distinctly bicolored, and with an 
orangish base. Tail brush black, conspicuous. Sole partly haired, not 
pigmented. Claws black. 

Skull not prominently angular, but with well-developed supra- 
orbital ridge. Posterior margin of nasals level with or behind 
posterior margin of frontopremaxillary suture. Infraorbital foramen 
not visible in lateral view. Mastoid bulla inflated beyond level of 
paroccipital process. Suprameatal triangle usually closed posterior- 
ly. Auditory meatus swollen to level of zygomatic process of tem- 
poral bone. Accessory tympanum present. 

Adult head and body length average 142 mm.; tail 145 mm., 102 
per cent of head and body length; hind foot 35 mm.; ear 20 mm.; 
occipitonasal length 38.6 mm.; weight 84 gm. 

External characters.— Dorsum dark yellowish brown; side with 
narrow but conspicuous line of clear orangish extending from wrist 
to heel and sometimes onto side of foot; venter white, occasionally 
with cream-colored areas on chest and belly. 

Hairs of dorsum, side, and greater part of belly with gray bases. 
Feet white, except for color on side as noted and with blackish 
claws. Mystacial, preorbital, suborbital, and subauricular areas 
grayish. 

Postauricular patch small, whitish. Ear not pigmented, long buffy 
hairs on anterior margin, pinna sparsely covered with buffy white 
hairs, producing a whitish border. Upper surface of tail color of back 
with scattered black hairs. Tail brush on upper surface of tip black. 



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212 FIELDIANA: ZOOLOGY 

very conspicuous, about one-third tail length. Tail not distinctly 
bicolored; proximal portion of underside dark orangish. 

Palatal ridges. — Identical with that of M. crassus (fig. 33). 

Glans penis and baculum.—See under M. crassus. 

Feet — Palm bare, sole partly naked proximally. Claws black. 

Cranial characters. — Figure 60. Skull somewhat angular with pro- 
minent supraorbital ridge. Posterior margin of nasals rounded or 
pointed extending to or slightly beyond level of frontopremaxillary 
suture. Anterior margin of zygomatic plate reaching or almost 
reaching level of premaxillary-maxillary suture and covering in- 
fraorbital foramen. Interparietal with round posterior margin. Tym- 
panic bulla conspicuously swollen, with anterior margin level with 
or beyond middle of foramen ovale. Posterior surface of mastoid 
bulla slightly posterior to paroccipital process. Suprameatal 
triangle closed or almost closed posteriorly by ascending portion of 
suprameatal process of temporal. External meatal swelling reaching 
level of zygomatic process of temporal. Distance across meatuses 
greater than zygomatic width. Accessory tympanum present. 

Teeth. — Figure 59. Upper incisor grooved. Molar pattern 
prismatic as in other species of Meriones. M with transient 
posterior fold. 

Measurements.— Table 22. Male and female dimensions subequal. 
Means (and ranges) of occipitonasal length (in millimeters) of 25 
adult males and 28 adult females, respectively are: 38.8 (36.4 to 
41.4) and 38.3 (35.8 to 40.9). 

Age determination.— Adults have the same features as M. 
crassus. 

Variation.— No distinct variation was observed in Egyptian 
material. 

Compansons.— Table 23. Meriones libycus is distinguishable 
from other species of Egyptian jirds by its darker color, orangish 
base of tail, dark claws, more anterior position of margin of 
zygomatic plate, and presence of an accessory tympanum. Dif- 
ferences are discussed under other species and listed in Table 23. 
Chromosomal comparisons are under M. crassus. 

Remarks.—Specimens referred to here as M. libycus are identical 
with material called M. caudatus by Ranck (1968). The only 
specimens of M. libycus identified by Setzer (1961a, p. 87) and 



OSBORN&HELMY: MAMMALS OF EGYPT 213 

discussed by Hoogstraal (1963, p. 18) are from Salum on the Libyan 
frontier. All others from coastal locaHties between Alexandria and 
Salum were M shawi. Hayman (1948, p. 40) misidentified M. 
libycus from Bahrein as Psammomys obesus. 

Confusion of taxonomists was also discussed by Lay and Nadler 
(1969), who succeeded in hybridizing M. libycus and M. shawi in 
captivity. Further discussion is given under M shawi. 

Specimens examined.— Total 110. 

BEHEIRA: Wadi el Natroun, Gebel Muluk (5), Zaghig (3). 

MATRUH: Qur el Hilab (2); El Maghra (28). Hatiyet Labaq (5); Abu Dweiss (4); Sir 
Abd el Nabi (2); Qara (18); Bahrein (16); El Malfa (2); Ain Qureishit (4); Abu Shuruf 
(1); Ain el Baqar (7); Salum 16 km. E (4), 17 km. E (1), 19 km. E (3); Bir Sidi Omar (1); 
Bir Shafarzin (1). 

EL FAIYUM: Wadi MuweUih, Bir Dakaar (3). 

Published records.— Records are from Chaworth-Musters and 
Ellerman (1947), Hayman (1948, as Psammomys obesus), and 
Setzer (1961a). 

BEHEIRA: Wadi el Natroun. 
MATRUH: Salum, Bahrein. 

Collection.— Meriones libycus, like other species of the genus, 
readily enters live traps placed beside burrows or in vegetation. Dig- 
ging for it is sometimes difficult, because burrows penetrate be- 
tween roots of plants. 

Habitats.— This species is found commonly in sand mounds 
formed around Nitraria retusa (fig. 17) in Wadi MuweUih, El 
Maghra, Qara, and Bahrein. One habitat in Wadi MuweUih included 
the grass Desmostachya bipinnata. Dead fronds under clusters of 
date palms harbor the species, and similar cover was provided by a 
pile of dead olive branches at Qara which hid burrows of this rodent. 
Specimens obtained from a rush-reed {Juncus sp.-Phragmites 
australis) association at El Maghra were probably only foraging 
there. One specimen was dug from a hole in hard barren ground 
under an acacia tree in the Maghra area. 

A colony at Bahrein burrowed in damp, salty sand beneath 
Tamarix sp. and Nitraria retusa. 

East of Salum, M. libycus was trapped beneath Lycium sp. and 
Nitraria retusa on sandy ridges above a salt marsh. 

Two individuals were trapped beside holes in hard ground near 
scattered vegetation (Hammada scoparia, Pityranthus tortuosus, 



214 FIELDIANA: ZOOLOGY 

Zilla spinosa, Artemisia inculta, Peganum harmala, Carthamus sp.) 
at Bir Sidi Omar and Bir Shafarzin near the Libyan border. 

The occupation by M. libycus of both mesic and dry habitats in 
the Western Desert of Egypt is in contrast to the restricted 
ecological distribution reported by investigators in other countries. 

Lewis et al. (1965, p. 73) found M. libycus syrius in northern Saudi 
Arabia only in "association with more or less permanent vegetation 
or with water." Ranck (1968, p. 165) observed that, in Libya, M. 
caudatus (M. libycus) "is never found associated with mesic 
habitats." 

Activity.— This species has been observed among shrubs after 
sunrise at El Maghra. Lewis et al. (1965) remarked that M. I. syrius 
was active during the day in northern Saudi Arabia. Ranck (1968, p. 
169) recorded an individual foraging in full daylight in Libya. In the 
vicinity of Benne-Abbes, Algeria, M libycus was active during the 
day for periods long enough to allow visual studies (Daly and Daly, 
1975a). 

Captive behavior.— Meriones libycus, unlike shawi and crassus, is 
very aggressive, difficult to handle, and bites readily. 

Burrows.— Burrov/s with numerous openings are dug in mounds 
around vegetation or hidden beneath the detritus under wild date 
palms. One burrow under an isolated acacia tree contained a store of 
pods and seeds. 

/Reproduction.— Data from four females taken in April and May 
averaged three young, with a range of two to four. 

Sex ratio.— In a sample of 82 museum specimens, males and 
females were equal in number. 

Meriones shawi Rozet, 1833 
Meriones shawi Rozet, 1833, Voyage dans la Regence dAlger. 1, p. 243. 
Type locality.— Algeria. ORAN: Oran. 

General distribution.— Egypt, Libya, Tunisia, Algeria, and 
Morocco. 

Common name.— Shaw's Jird. 

Subspecies in Egypt — 



OSBORN&HELMY: MAMMALS OF EGYPT 215 

Meriones shawi isis (Thomas, 1919) 
Meriones isis Thomas, 1919, Ann. Mag. Nat. Hist., (ser. 9), 3, p. 271. 

Type locality.— Egypt. ALEXANDRIA: Alexandria, Ramleh. 

Distribution in Egypt.— Figyire 65. Coastal desert, from type 
locality west to Salum on Libyan frontier, and El Maghra. 

Diagnosis.— Large jird with soft pelage. Dorsum brownish yellow, 
side with conspicuous line of clear yellowish or orangish extending 
to wrist and heel. Venter white with pale yellowish areas. Ear prom- 
inent and pigmented. Tail with upper surface paler than dorsum, 
not distinctly bicolored, underside cinnamon colored. Tail brush 
blackish, about one-fourth of tail length. Sole partly haired, not 
pigmented. Claws pale. 

Skull somewhat angular but not strongly ridged. Mastoid bulla 
moderately inflated but not extending beyond level of paroccipital 
process. Infraorbital foramen large and conspicuous in lateral view. 
Posterior margin of nasals anterior to posterior level of frontopre- 
maxillary suture. Accessory tympanum absent. 

Adult head and body length average 143 mm.; tail 140 mm., 97 
per cent of head and body length; hind foot 34 mm.; ear 19 mm.; 
occipitonasal length 38.8 mm.; weight 90 gm. 

External characters.— Dorsum dark brownish yellow; side with 
narrow but conspicuous line of clear yellowish or orangish extend- 
ing to wrist and heel and sometimes onto side of foot; venter white 
with pale yellowish area on upper chest and sometimes middle of 
belly. Hairs of dorsum, side, and greater part of belly with gray 
bases. Feet white except for coloration on side as noted. Band from 
mystacial area beneath eye to base of ear very slightly paler than 
upper surface of head. Preorbital and postorbital spots conspicuous, 
grayish. Postauricular patch small and whitish. Ear sparsely 
covered with short, buffy hairs; long hairs fringing anterior 
margin. Distal one-third of pinna pigmented. Upper surface of tail 
with blackish hairs but paler than dorsum; blackish brush on upper 
surface of tip about one-fourth of tail length. Tail not distinctly 
bicolored, underside of base near cinnamon color. 

Feet.— Palm bare; sole partly naked proximally. Claws pale. 

Palatal ndg^es.— Similar to that of M. crassus (fig. 33). 

Glans penis and baculum.— Not observed. 



216 FIELDIANA: ZOOLOGY 

Cranial characters.— Figure 60. Cranium not prominently angular 
or ridged. Posterior margin of nasals not extending to posterior 
level of frontopremaxillary suture. Anterior margin of zygomatic 
plate not reaching level of premaxillary-maxillary suture and not 
concealing large infraorbital foramen. Interparietal with round 
posterior margin. Tympanic bulla conspicuously swollen; posterior 
surface of mastoid bulla extending beyond level of exoccipital, but 
not beyond level of paroccipital process. Suprameatal triangle 
almost closed posteriorly by descending process of supraoccipital. 
External auditory meatal swelling not reaching level of zygoma and 
distance across meatuses slightly less than zygomatic width. 
Accessory tympanum absent. 

Teeth.— Figure 59. Upper incisors grooved. Molar pattern 
prismatic as in other species of genus, except lingual and labial folds 
of m^ are often equal in depth. M^ with transient posterior fold. 

Measurements.— Table 22. Male and female dimensions subequal. 
Means (and ranges) of occipitonasal length (in millimeters) of eight 
adult males and seven adult females are 38.8 (37.1 to 41.5) and 38.6 
(35.3 to 40.0), respectively. 

Age determination.— Adults have same features as M. crassus. 

Vanafion.— Specimens from near Alexandria have the meatal area 
slightly more swollen than those from other parts of Egypt. Occa- 
sional specimens have posterior margin of nasals about level with 
the posterior border of the frontopremaxillary suture. 

Comparisons. —Meriones shawi differs from M. crassus in darker 
color, more prominent strip of clear color on side, ear pigmented, 
and mastoid chamber and meatus considerably less inflated. It dif- 
fers from M. libycus in having base of tail paler, tail brush much 
smaller, ear pigmented, claws pale, posterior margin of nasals 
anterior to level of frontopremaxillary suture, infraorbital foramen 
exposed, and mastoid bulla and meatus less swollen. From M 
sacramenti, M. shawi differs in having belly hairs with gray bases, 
more conspicuous tail brush, more inflated mastoid bulla, sUghtly 
more exposed infraorbital foramen, and slightly smaller dimensions 
(table 22). Characters can be compared in Figure 60 and Table 23. 
Refer also to M. crassus and M. libycus. 

Remarks.— Controversies and errors (Flower, 1932; Setzer, 1961a; 
Ranck, 1968) on the status of M. shawi in Egypt were resolved by 
Lay and Nadler (1969). Morphological characters presented in Table 



OSBORN&HELMY: MAMMALS OF EGYPT 217 

23, in addition to those from Petter (1953, 1961) and Lay and Nadler 
(1969), confirm M. shawi as a species distinct from M. libycus. 

The name ''melanurus'' was applied by Riippell (1842, p. 95) to 
representatives of M. crassus from sandy areas "near Alexandria," 
Egypt and "Petraischen Arabien" (Stony Arabia or Arabia Petra is 
actually the core of the Sinai Plateau according to Abu Al-Izz, 
1971), near Tor, Sinai. Fitzinger and Heuglin's (1866, p. bl4)Rhom- 
bomys melanurus from Tor could only be M. crassus on geographic 
grounds and, likewise, Nehring's (1901, p. 170) M. {Rhombomys) 
melanurus from Koseir (Quseir) on the Gulf of Suez. Nehring's 
descriptions of meatal and buUar inflations also indicate M. crassus. 
Anderson (1902, p. 267) and Bonhote (1912, p. 227) applied 
''melanurus'" as a trinomen to M. shawi, whereas Thomas (1919a, p. 
264) thought the name was best applied to the form with a promi- 
nent black tail tuft, M. libycus, in agreement with Riippell's figure 
and description. Unfortunately, Riippell's "types" did not represent 
either species. Such confusion places ''melanurus'' in obsolescence. 

Meriones shawi isis Thomas (1919a), however, is a valid trinomen, 
and "isis'' has apparently been applied to specimens of M. shawi 
only. 

Specimens examined.— Total 59. 

ALEXANDRIA: Ramleh (1), near Alexandria (2), El Amiriya 5 km. S (2). 

MATRUH: Lake Mariut (7); Bahig 8 km. S (1), 12-13 km. NE (11); Burg el Arab (6); 
Abu Mena (2), 4.8 km. E (1); El Daba (1); El Alamein (6); El Maghra (2); Abu Haggag 
1.6 km. E (1); Ras el Hekma (1); Mersa Matruh (8); Sidi Barrani 22.8 km. E (1). 3.2 
km. S (1), 19.2 km. S (1), 48 km. W (2), 52.8 km. W (1); Salum (1). 

Published records.— Records are from Anderson (1902), Thomas 
(1919a), Chaworth-Musters and Ellerman (1947), and Setzer (1961a 
under M. /. libycus). 

ALEXANDRIA: Ramleh, Alexandria, Mex. 

MATRUH: Mersa Matruh; Burg el Arab; El Daba; El Alamein; Sidi Barrani 22.8 
km. E. 3.2 km. S, 8 km. S, 19.2 km. S, 48 km. W, 52.8 km. W; Salum. 

Collection.— Meriones shawi, like other jirds, readily enters live 
traps. Digging it from burrows in hard clay or within the roots of 
shrubs is often difficult. 

Habitats.— Meriones shawi burrows in hard clay of Western 
Mediterranean Coastal Desert, particularly beneath the shrubs 
Anabasis articulata and Lycium sp. (fig. 8), where its burrows may 
be mistaken for those of Psammomys obesus and vice versa. It has 
also been collected from Bedouin barley fields, fields overgrown 



218 FIELDIANA: ZOOLOGY 

with thistles, beneath Lycium sp. (fig. 48) on rocky slopes and 
coastal dunes, and in mounds of sand around Nitraria retusa in the 
eastern part of El Maghra (fig. 17). Specimens referred to M L 
libycus from fig groves (fig. 7) on the Mediterranean coast west of 
Alexandria (Hoogstraal, 1963) were M. s. isis. 

Activity.— According to Petter (1961), M. shawi is active outside 
of burrows during part of the day in Algeria. 

Captive 6e/ia uior.— Although not initially as docile as M. crassus, 
M. shawi responds favorably to frequent handling. 

Burro u;s.— Burrows are usually dug beneath shrubs and have 
numerous openings which are not closed. 

Food — Information on food plants of this species is lacking. It 
does, however, frequent Bedouin barley fields in the coastal desert 
and no doubt takes a toll from the annual crop. 

Reproduction.— No information was obtained on the Egyptian 
form. 

Sex ratio. — In a sample of 46 museum specimens, there were 22 
(48 per cent) males and 24 females. 

Meriones tristrami Thomas, 1892 
Meriones tristrami Thomas, 1892, Ann. Mag. Nat. Hist. (ser. 6), 9, p. 148. 
Type locality.— Israel: Dead Sea Region. 

General distribution. — Iran, Iraq, Azerbaijan S.S.R., Turkey, 
Syria, Lebanon, Palestine, and northeastern Sinai Peninsula. 

Common name.— Tristram's Jird. 

Subspecies in Egypt— Probably the nominate subspecies. 

Meriones tristrami tristrami Thomas, 1892 

Distribution in Egypt— Figure 65. Known in Egypt from 
specimens reported from El Arish, northeastern Sinai Peninsula 
(Zahavi and Wahrman, 1957). 

Diagnosis.— Rather small jird with soft pelage, dorsum yellowish 
brown, side with line of clear yellowish extending onto wrist and 
heel. Venter white. Ear prominent, tip pigmented. Tail upper sur- 
face as back, bicolored; underside orangish to cinnamon. Tail brush 
blackish and poorly developed. Sole partly haired, not pigmented. 
Claws pale. 



OSBORN & HELMY: MAMMALS OF EGYPT 219 

Skull rounded, weakly ridged. Mastoid bulla moderately inflated 
but not extending beyond level of exoccipital or paroccipital pro- 
cess. Infraorbital foramen partly exposed in lateral view. Posterior 
margin of nasals anterior to posterior level of frontopremaxillary 
suture. Lip of auditory meatus very slightly swollen. Accessory 
tympanum absent. 

Adult head and body length average 129 mm.; tail 133 mm., 103 
per cent of head and body length; hind foot 34 mm,; ear 20 mm.; oc- 
cipitonasal length 36.0 mm. 

External characters.— Dorsum varying from pale to dark 
yellowish brown; side with narrow but conspicuous line of clear 
yellowish to orangish extending to wrist and heel and sometimes 
onto foot; venter white, and feet white, except as noted. Hairs of 
dorsum and side with gray bases. Belly hairs usually all white, 
except for occasional individuals with very faint gray bases. 
Mystacial, preorbital, suborbital, and subauricular areas slightly 
paler than upper surface of head. Postorbital spot conspicuous, 
grayish. Postauricular spot conspicuous and white. Ear with long 
whitish hairs on anterior margin, pinna sparsely covered with 
whitish hairs; distal one-third pigmented. Tail with color of upper 
surface as dorsum; bicolored, underside at base orangish. Blackish 
brush on upper surface of tip of short hairs inconspicuous, about 
one-fourth length of tail. 

Palatal ridges.— Not observed. 

Feet.— Palm bare, sole partly naked proximally. Claws pale. 

Cranial characters.— Figure 60. Cranium not prominently angular 
or ridged. Posterior margin of nasals not extending to posterior 
level of frontopremaxillary suture. Anterior margin of zygomatic 
plate not reaching level of premaxillary-maxillary suture and only 
partially concealing infraorbital foramen. Interparietal broad, 
somewhat ovoid. Tympanic bulla conspicuously swollen; anterior 
surface level with middle of foramen ovale. Posterior surface of 
mastoid bulla not extending beyond level of exoccipital and paroc- 
cipital process. Suprameatal triangle closed posteriorly by union of 
descending process of supraoccipital and ascending portion of 
suprameatal process of temporal. Lip of external auditory meatus 
slightly swollen. Accessory tympanum absent. 

Teeth.— Upper incisors grooved, molars as in other species. 

Measurements.— Table 22. 



220 FIELDIANA: ZOOLOGY 

Age determination.— Adults have same features as M. crassus. 

Vanation.— Considerable variation in shade of color exists in this 
species. Zahavi and Wahrman (1957) noted that coloration often 
resembled shade of the soil in which it lived. 

Comparisons.— Table 23. Meriones tristrami is distinguishable 
from all other Egyptian species of Meriones by the very poorly 
developed tail brush; small mastoid bulla, combined with closure 
posteriorly of the suprameatal triangle; and the broad, ovoid inter- 
parietal (fig. 60). 

Specimens examined.— Six from various localities in Lebanon and 
Syria. 

Published records.— Record from Zahavi and Wahrman (1957). 

SINAI: El Arish. 

//a6i fats.— According to Zahavi and Wahrman (1957), this is a 
widely ranging species in Israel; living in sand along the Mediterra- 
nean coast, alluvial soils, clay soils, and suitable situations in 
mountains. 

Economic importance.— This species, according to Zahavi and 
Wahrman (1957), shows fluctuations in numbers and in some years 
is a pest to agriculture. 

Genus Pachyuromys La taste, 1880 

Monotypic genus of jird-like rodent with long, fluffy fur. Tail 
relatively short, club-shaped, lacking apical brush. Palm and sole 
partly haired; pads and hallucal tubercle like Gerbillus and 
Meriones. 

Bulla extremely inflated. Meatal lip swollen to level of zygomatic 
process of temporal. Accessory tympanum present. Suprameatal 
triangle very large, but closed posteriorly. Parapterygoid fossa with 
superior wall perforated. Enamel pattern of molars laminated. 

Pachyuromys duprasi Lataste, 1880 

Pachyuromys duprasi Lataste, 1880, Naturaliste, Paris, 1, p. 314. 

Type locality.— Algeria. GHARDAIA: Loghouat. 

General distribution.— Northv/estern Egypt, Libya, Tunisia, 
Algeria. 

Common names. — Fat Tailed Jird, Abu Lya. 

Subspecies in Egypt. — 



OSBORN&HELMY: MAMMALS OF EGYPT 221 

Pachyuromys duprasi natronensis De Winton, 1903 

Pachyuromys duprasi natronensis De Winton, 1903, Nov. Zool., 10, p. 285. 

Type locality. -Egypt. BEHEIRA: Bir Victoria. 

Distribution in Egypt— Figiire 66. Northern part of desert west 
of the Nile Delta. 

Diagnosis.— Jird-\ike rodent with fur long and fluffy, palm and 
sole partly haired, dorsal hairs pale cinnamon with blackish tips, 
sides pale cinnamon, underparts and feet white. Tail shorter than 
head and body, club-shaped, lacking a brush. 

Skull with extremely inflated auditory bulla, large suprameatal 
triangle, meatal lip swollen, accessory tympanum present, paroc- 
cipital process elongate and adnate to bulla. 

Adult head and body length average 108 mm.; tail 58 mm., 54 per 
cent of head and body length; foot 23 mm.; ear 14 mm.; 
occipitonasal length 34 mm.; weight 36.5 gm. 

External characters.— Figure 67. Upper parts pale cinnamon, dor- 
sal hairs tipped with black, side with narrow strip of pale cinnamon 
extending almost to heel, but not onto forelimb. Hairs of back and 
side with dark gray bases; hairs of belly, underparts, and feet white. 
Mystacial area partly pigmented, circumorbital area color of sides, 
white supraorbital spot faint or lacking, white postauricular patch 
small. White rump patch absent. Ear pigmented, sparsely haired, 
anteroventral margin with tuft of long cinnamon hairs. Tail thick 
and club-shaped, bicolored, dorsal surface color of side, ventral sur- 
face white, apical brush lacking. 

Palatal ridges.— Not observed. 

Glans penis and baculum.— Not observed. 

Feet.— Palm and sole partly haired, pads and tubercle similar to 
Gerbillus and Meriones in Figure 34. 

Cranial characters.— Figure 68. Skull elongate despite triangular 
outline due to enormously swollen auditory bullae. Supraorbital 
ridge poorly developed, not reaching level of posterior margin of 
lacrimal. Parietal ridge inconspicuous. Interparietal narrow, 
angular in outline posteriorly. Anterior surface of tympanic bulla 
anterior to level of foramen ovale. Posterior margin of mastoid 
chambers extending well beyond posterior margin of supraoccipital 
and paroccipital. Medial superior mastoid cavity absent. Subar- 
cuate fossa small. Suprameatal triangle very large, closed posterior- 




o 



222 



OSBORN & HELMY: MAMMALS OF EGYPT 



223 




FiCi 67. Live specimen of Pachyuromys duprasi natronensis. 

ly by union of descending process of supraoccipital and ascending 
part of suprameatal process of temporal bone. External auditory 
meatus swollen, contacting zygomatic process of temporal. 
Accessory tympanum present. Supraoccipital and basioccipital nar- 
row, constricted by swelling of bullae. Paroccipital process elongate, 
thin, adnate to wall of bulla. Parapterygoid fossa deep, crowded by 
bulla, and with large perforation in superior wall. Zygomatic plate 
with rounded anterior margin not reaching level of premaxillary- 
maxillary suture. 

TeetA.— Figure 59. Upper incisors grooved on anterior surface. 
Molars rooted. M and mj appear to be tuberculate in very young 
animals, becoming laminate in immatures. M^ and m2 show no 
indication of tubercles. Enamel pattern more similar to that of 
Meriones crassus rather than adult Gerbillus, as indicated by Fetter 
(1956). Third molars simple, lacking folds. 

Measurements.— Table 24. Male and female dimensions subequal. 

Age determination.— Adults have laminae of moleirs worn and 
skull sutures closed. 

Variation. — Individual variation in presence or absence of black- 
tipped hairs on the side and in shade of color is noticeable and was 
mentioned by Setzer (1963). 



224 FIELDIANA: ZOOLOGY 

Table 24. — Means (and ranges) of measurements, ratios, and weight of adult 
Pachyuromys duprasi natronensis. 



HBL 


108.3 (93-121) 4 


ZW 


19.3(17.5-20.2)4 


TL 


58.2 (55-62) 4 


low 


6.2 (5.8-6.4) 4 


TL/HBL% 


54.5 (47.9-66.6) 4 


NL 


13.0(11.7-13.8)4 


FL 


23.3 (22-24) 4 


IFL 


6.4 (6.2-6.8) 4 


EL 


14.0(12-16)4 


AL 


5.2 (4.8-5.7) 4 


Wt 


36.5 (22.0-44.6) 3 


RW 


4.9 (4.4-5.0) 4 


ONL 


24.9 (32.4-36.5) 4 


SH 


14.7(14.2-15.0)4 



Comparisons.— Egyptian specimens of P. cL natronensis differ 
from Libyan samples, the nominate form, and P. duprasi faroulti of 
Algeria in paler color. Pachyuromys duprasi differs externally from 
all other rodents by its short, thick tail; cranially by its enormously 
inflated auditory bulla and shape of interparietal and paroccipital 
process (fig. 68). Color in Egyptian specimens is similar to Meriones 
crassus rather than M. libycus, as was suggested by Setzer (1963). 

Specimens examined.— Total 14. 

BEHEIRA: El Khatatba (3). Bir Victoria (Type and 2). 1.6 km. E (1). Wadi el 
Natroun (3). 

TAHREER: Cairo-Alexandria desert road km. 163 (1). 

MATRUH: Cairo-Alexandria desert road km. 26 (1). Abd el Mawla (1). Bir Shafar- 
zin 25.6 km. E (1). 

Published records.— Records are from Setzer (1952, 1963) and 
Hoogstraal (1963). 

BEHEIRA: Kom Hamada, between Kom Hamada and Bir Victoria, El Khatatba. 
Bir Victoria. 

TAHREER: Cairo- Alexandria desert road km. 179. 

MATRUH: Cairo- Alexandria desert road km. 17, Abar el Dafa (36 km. W of Mersa 
Matruh). 

GIZA: El Qatta, Abu Ghalib W of. Abu Rawash W of. near Cairo, Cairo- 
Alexandria desert road km. 10.5. 

Co//ec(ion.— Trapped alive and dug from burrows. 

//a6i(a(s.— Vegetated sand sheets (fig. 10) south of the Western 
Mediterranean Coastal Desert and southern limits of the coastal 
desert vegetation (fig. 20); sometimes in rocky desert. One was 
trapped in a stand of Anabasis articulata in an isolated sandy 
depression east of Bir Shafarzin, and another was dug from barren 
gravel 26 km. NW of Cairo. The type locality, Bir Victoria, is a 
small, shallow sandy depression sparsely vegetated vnt\i Artemisia 






Fk; 68. Skull of Pachyuromys duprasi natronensis. 



225 



226 FIELDIANA: ZOOLOGY 

monosperma and patches of Hyoscyamus muticus. These habitats 
are comparable with what Ranck (1968, p. 157) called "transitional 
deserts which run roughly parallel to the more lush coastal plain" 
and where P. duprasi was "most abundant." 

Burrows. — We have dug fat tailed rats from simple burrows about 
1 m. in depth in hard sand. Petter (1961) illustrated a very complex 
burrow of the Algerian subspecies. We have observed that this 
rodent moves about considerably and may occupy burrows of other 
species. 

Activity.— VJe have observed that, in the wild, fat tailed rats 
become active at dusk. 

Captive behavior.— The most docile of Egyptian rodents. Never 
bites and makes little effort to escape when handled. In captivity, 
strangely enough, this lethargic animal is cannabalistic (Flower, 
1932), and females have eaten their young. 

Food.—Pachyuromys duprasi no doubt utilizes a variety of 
plants, but we have only observed it feeding on Anabasis articulata 
and Artemisia monosperma. 

In the laboratory, Petter (1961) fed Algerian fat tailed rats grain, 
chopped meat, cheese, milk, lettuce, and lucerne (Medicago sativa). 
He also mentioned its affinity for live crickets. Suggestion has been 
made that, in Libya, terrestrial snails were eaten by P. duprasi 
(Setzer, 1957, p. 60). Thus far, we have been unable to check this 
possibility, but have observed what appeared to be "gnawed" snail 
shells in Pachyuromys habitat in the Western Desert. 

Associates.— Meriones crassus, Gerbillus gerbillus, G. andersoni, 
0. perpallidus, 0. pyramidum (possibly), and Jaculusjaculus live in 
the same habitats as P. duprasi. 

Reproduction.— Flower (1932) reported litters of three to five, 
seven, and nine born in captivity and during the months of April, 
May, July, October, and November. 

Young at birth were said to be naked, blind, and helpless, like 
those of Rattus sp. 

Genus Psammomys Cretzschmar, 1828 

Stocky rodents, dorsum blackish to reddish orange, side and 
venter yellowish. Ear small, rounded. Tail less than 85 per cent of 
head and body length, fully haired; tip black with dorsal brush. 
Palm bare; sole partly haired. 



OSBORN&HELMY: MAMMALS OF EGYPT 227 

Skull angular and strongly ridged. Bulla including meatus greatly 
inflated. Interparietal outline squarish. Exoccipital broad and flar- 
ing. Paroccipital process very large, extending laterally. Upper 
incisor smooth on anterior surface. Molars hypsodont when 
immature, crowns prismatic. 

Psammomys obesus Cretzschmar, 1828 

Psammomys obesus Cretzschmar, 1828, in Riippell, Atlas zu der Reise im 
nordliche Afrika, Saugeth., pi. 22, 23, p. 58. 

Type /oca/ity.— Egypt. ALEXANDRIA: Alexandria. 

General distribution. — Israel, Arabia, Sinai Peninsula, Egypt, 
Sudan, Libya, Tunisia, Algeria, Morocco. 

Common names.— Fat Sand Rat, Jarada. 

Distribution of subspecies in Egypt.— Figure 69. Psammomys 
obesus terraesanctae: northern and southern parts of Sinai Penin- 
sula and northern part of Eastern Desert. Psammomys obesus 
nicolli: northeastern part of Nile Delta; Psammomys obesus obesus: 
northwestern part of Nile Delta and northern part of Western 
Desert. 

Diagnosis.— Large blackish to reddish orange rodent with 
yellowish side and belly. Tail thick, shorter than head and body; tip 
black, prominent. Ears short, rounded, densely haired. Sole partly 
haired. Skull angular and strongly ridged. Bulla greatly inflated, 
anterior lip of external auditory meatus swollen to level of 
zygomatic process of temporal bone. Exoccipital and paroccipital 
broad, prominent. Upper incisor smooth on anterior surface. 
Crowns of molariform teeth prismatic, never tuberculate. 

Adult head and body length average 170 mm.; tail 130 mm., 76 
per cent of head and body length; hind foot 38 mm.; ear 15 mm.; 
occipitonasal length 42.6 mm.; weight 146.8 gm. 

External characters.— Figure 70. Dorsum blackish yellow to 
brownish or reddish orange. Width of color bands on dorsal hairs 
v£iry with subspecies and habitat (fig. 71). Side brownish to 
yellowish. Venter pale to dark yellow. All hairs with gray base 
except white hairs in axilla, groin, and behind ear. Foot with upper 
surface yellowish. Hair tuft on sole yellowish to whitish. Ear, foot, 
and claws pigmented. Ear densely haired, whitish or yellowish. 
Whitish postorbital spot absent. Postauricular spot white, small. 
Entire tail tip black, not bicolored; black brush prominent. 




228 




229 



230 FIELDIANA: ZOOLOGY 

Palatal ridges. — Fig[ire 33. Diastemal ridges thick and slightly 
curved. First to third intermolar ridges recurved; fourth intermolar 
ridge short; fifth, long, curving anteriorly. 

Glans penis and baculum.— Not observed. 

Fec^— Figure 34. Palm bare, sole with tuft of plantar hairs. 
Tubercles and pads of palm as in other Gerbillinae. Sole with three 
subdigital tubercles, a hallucal tubercle, and single plantar tubercle. 

Cranial characters. — Figure 72. Skull massive, strongly ridged, 
angular. Zygomatic arch heavy and wide. Supraorbital and parietal 
ridges prominent. Anterior margin of zygomatic plate usually 
reaching level of premaxillary-maxillary suture. Notch at level of 
antorbital foramen prominent. Interparietal outline squarish. Exoc- 
cipital and paroccipital broad and flaring. Anterior margin of tym- 
panic bulla slightly beyond posterior margin of foramen ovale. 
Posterior margin of mastoid bulla usually beyond level of occipital 
condyle, but not beyond level of paroccipital process. Anterior lip of 
auditory meatus swollen to level of zygomatic process of temporal 
bone. 

Teeth.— Figure 59. Upper incisor without anterior groove except 
distal 2.5 mm. in nestlings. Molariform teeth hypsodont in 
immatures, rooted in adults (fig. 73), cusps prismatic, never tuber- 
culate even before eruption. First upper molar with two large and 
one small root in adults. M^ with transient fold, m, simple. 

Measurements.— Table 25. 

Age determination.— Adult specimens have lateral folds of upper 
first molar not extending below alveolar level and roots of teeth 
usually exposed (fig. 73). Strongly developed parietal ridges and 
closure of the suprameatal triangle posteriorly by fusion of the 
hamular process of the squamosal and the supraoccipital process 
are additional criteria used in adult selection. 

Variation.— Samples from salt marsh habitats in the Nile Delta 
designated as subspecies nicolli show much more extension of 
melanin than obesus and terraesanctae from coastal salt marsh and 
desert habitats (Thomas, 1908). Color variations within subspecies 
are listed in Table 26, and variation in width of color bands of hairs 
is shown in Figure 71. Tips of hairs are blackish in darker 
individuals, brownish in paler ones. Subterminal bands are 
yellowish orange and basal bands dark gray. Figure 71 illustrates 
hairs with color bands of average widths. Means (and ranges) of tip. 



20 



IB 



10 



8 . 





mm 



NICOLLI 



TERRAE' 
SANCTAE 



OBESUS 



Fig. 71. Diagrams of middorsal hairs from subspecies of Psammomys obesus. 
Each figure represents average of five hairs. Black tip represents blackish or 
brownish color; clear subterminal part, yellowish orange; and punctate basal part, 
dark gray. 



231 






Fl(i 72. Skull of Psammomys obesus. 



232 



OSBORN & HELMY: MAMMALS OF EGYPT 



233 





Fig. 73. Lateral view of right upper molars of adult (above) and immature (below) 
Psammomys obesus. Note that lateral folds do not extend below alveolar level in 
adult or mature molars, but do so in immatures. The same age criterion is used for 
Meriones sp. 

subterminal, and basal bands of five middorsal hairs (in millimeters) 
from a specimen of nicolli are: 1.8 (1.5 to 2.0), 2.9 (2.6 to 4.0), and 8.9 
(8.2 to 10.0), respectively; terraesanctae: 0.6 (0.4 to 1.1), 8.0 (7.0 to 
8.9), and 6.8 (6.3 to 7.3); obesus: 0.8 (0.6 to 1.2), 5.6 (5.3 to 5.8), and 
4.8 (3.6 to 5.6). 

Specimens of obesus and terraesanctae from desert habitats are 
essentially alike in color and width of hair bands. Specimens of 
obesus from coastal salt marshes differ, as shown in Figure 71. 

In comparison with these subspecies, samples of nicolli are darker 
and average larger in all measurements. Subspecies nicolli and ter- 
raesanctae are similar in having the posterior nasal margin reaching 
the level of the posterior edges of the frontopremaxillary suture, but 
in obesus, nasal margin is anterior to this point (fig. 74). There are 
few exceptions within samples examined, although some have 
posterior nasal margin in an intermediate position (table 27). Nasal 
length (table 25) does not reflect this morphological difference. 

Comparisons.— The only Egyptian rodents with which Psam- 
momys obesus might be confused are Meriones libycus, M. crassus, 



234 FIELDIANA: ZOOLOGY 

Table 25. — Means (and ranges) of measurements, ratios, and weight of adult 
Psammomys obesus. 

P. o. obesus P. o. nicoUi P. o. terrae sane toe 

HBL 168.4(151-187)73 178.6(160-199)41 157.5(144-168)20 

TL 125.4(100-144)69 143.5(122-157)38 122.8(115-131)19 

TL/HBL% 73.5(60.9-82.5)71 80.2(69.1-90.1)38 78.0(73.6-89.6)19 

FL 36.9 (32-40) 78 40.2 (38-43) 46 36.8 (35-40) 21 

EL 14.8(14-16)77 16.0(14-18)41 14.6(13-17)21 

Wt 141.8(116.3-205.1)37 130.0(106.6-223.0)27 114.5(92.1-135.3)12 

ONL 41.7(38.8-45.4)68 45.4(42.8-48.2)40 41.0(37.7-44.9)22 

ZW 24.9 (23.7-27.4) 59 26.7 (25.4-28.8) 31 24.4 (22.2-26.6) 18 

lOW 6.7(6.0-7.3)68 7.1(6.3-8.2)38 6.6(5.8-7.3)20 

NL 16.2(14.0-17.6)67 18.6(17.0-20.9)38 16.6(15.0-18.5)21 

IFL 6.4(4.6-7.2)71 7.2(6.3-8.0)39 6.4(5.7-7.1)21 

AL 7.2(6.8-7.9)77 7.7(7.2-8.4)43 7.1(6.8-8.0)23 

RW 6.2(5.8-6.8)74 6.6(6.3-7.4)32 6.2(5.7-6.8)22 

BL 13.3(12.3-14.4)73 14.2(13.1-15.3)40 13.6(12.2-14.6)23 

SH 16.2(14.7-17.7)72 17.5(15.2-19.2)37 15.9(14.7-17.5)23 

POW 14.7 (12.8-16.5) 36 16.4 (15.2-17.9) 26 14.4 (12.9-15.9) 15 

and M. shawi; however, slightly longer tails and ears, bicolored tail 
tips, whitish bellies, and grooved upper incisors in Meriones 
distinguish them from Psammomys. Cranial characters are dis- 
cussed under species of Meriones. 

Remarks.— No specimens of Psammomys are known from the 
Delta between east and west branches of the Nile, except near Ras el 
Bar in the northeast and Quweisna in the south (fig. 69). No 
evidence of fat sand rats has been found in the extensive salt 
marshes and stands of halophytic plants south of Lake Burullus or 
between Kafr el Sheikh and Baltim. Differences in position of 
posterior margin of the nasals (fig. 74, table 27) in eastern and 
western populations illustrate the effectiveness of this hiatus as a 
barrier to gene flow. 

Color and size differences (tables 26, 25), useful as they are in 
separating subspecies, are environmentally influenced. Desert P. o. 
obesus are almost identical in size and coloration with desert P. o. 
terrae sanctae. 

Specimens identified as P. obesus from Bahrein (Hayman, 1949) 
are Meriones libycus, and those from Wadi el Natroun, El Beida 
(Setzer, 1963) and "hard surface sand desert near Bir Hooker" 
(Hoogstraal, 1963, p. 20) are M. crassus. 

Convexity of parietals suggested by Setzer (1963) as a diagnostic 



OSBORN & HELMY: MAMMALS OF EGYPT 



235 



Table 26. — Color patterns of Psammomys obesus. 



Region P. o. obesus 

Crown without or with a 

scattering of 
black-tipped hairs 

Dorsum orange yellow, 

brownish yellow, 
brownish orange 

Side narrow strip of pale 

yellowish to yellowish 
brown 

Belly pale to dark yellow 

Tail black dorsal hairs 

75% from tip 



P. o. nicoUi 

with all hairs black 
tipped 

blackish to 
brownish yellow 

narrow strip of 
yellowish to brown 

dark yellow 

black hairs on entire 
dorsal surface 



P. o. terraesanctae 

without black-tipped 
hairs 



orange yellow to 
brownish orange 

broad strip of clear, 
pale yellow 

pale yellow with 
some white 

with black dorsal 
hairs 40-50% from tip 



character of P. o. nicolli is a normal condition of all three subspecies. 

Collection.— Digging is the only effective method of obtaining 
quantities of fat sand rats alive. Daly and Daly (1974) captured 
them in wire cage traps baited with Suaeda vermiculata (=S. 
mollis). 

Habitats.— HaibitaXs of Psammomys are saline soils and salt 
marshes with stands of succulent halophytic vegetation chiefly of 
family Chenopodiaceae. Coastal salt marshes (fig. 7) are subject to 
flooding in winter rainy season, and sand rats must sometimes 
abandon burrows and move to higher land (Wassif, 1953b). Inland, 
fat sand rats depend chiefly upon two plant species, Anabasis ar- 
ticulata and Hammada elegans, and occur to the southern limits of 
these species, except along the Red Sea coast and on the limestone 
plateau of the Western Desert. 

On the coastal plain, plants suitable for fat sand rats occur on 



Table 27. Variation in position of posterior margins of nasals in Psammomys 
obesus. 



Subspecies 


No. examined 


P. o. obesus 


104 


P. o. nicolli 


40 


P. o. terraesanctae 


26 



No. with nasal margin 
Anterior Posterior Intermediate 

89 



4 


11 


36 


2 


21 


2 






UJ 

O 

6 

o.* 



236 



OSBORN & HELMY: MAMMALS OF EGYPT 237 

loamy and sandy soils, usually where rainwater accumulates. Short 
wadis of northern Eastern and Western Deserts are suitable sites, 
but are subject to severe flooding. 

Psammomys burrows beside and beneath the Cairo-Alexandria 
desert road in the Nubareia area, behind stone cribbing under 
highways in the Suez Canal area, in piles of stone around poles of 
power lines in the rocky desert west of Mersa Matruh, and in 
embankments along the Alexandria-Salum railway. 

The most southern colony in the Western Desert was discovered 
in the salt marsh of El Maghra in the northeastern part of Qattara 
Depression. East of there, a small colony subsisted in an isolated 
stand of Hammada elegans on hard gravelly desert near Qaret el 
Mashruka. There is another colony to the west in Wadi Labaq. 

A typical Psammomys colony will have a burrow beneath nearly 
every shrub and a maze of trails running between burrows and food 
supply. 

Behavior.—Sand rats are mainly diurnal and can be seen climbing 
into low shrubs, cutting branches, and dragging them into burrows. 
Food is not actively stored in burrows, but large accumulations of 
waste give this impression. The desire for fresh food and trampling 
of uneaten portions can be observed in captive animals. 

Natural curiosity of the fat sand rat gives the impression of docili- 
ty. Quite the contrary, it is a vicious fighter when confined in groups 
and will bite readily if restrained. 

Antisocial behavior has been studied in attempts to find com- 
patable pairs for laboratory colonies (Prange et al., 1968). Daly and 
Daly (1957b) found P. obesus to be the most aggressive and soliteiry 
of gerbils studied thus far. 

Persons who have seen wild fat sand rats standing upright and 
alert beside their burrows, likened them to prairie dogs of western 
U.S.A. (Flower, 1932). 

Burrows.— ¥aX sand rats dig tunnel networks in earth or sand 
mounds around shrub bases. Rock piles and stone cribbing are also 
attractive burrow sites. An occupied burrow has signs of recent dig- 
ging, fresh tracks, waste food in or beside openings, and usually 
toilets near openings. The toilets, slight excavations from covering 
feces and urine (fig. 75), are characteristic of Psammomys and 
Meriones. 

Tunnels of a burrow system are seldom deeper than 0.5 m., but 



238 



FIELDIANA: ZOOLOGY 










'■>«^ -r»s v_ 



Fig. 75. Burrow and toilet of Psammomys obesus. If the hillock is damp or con- 
tains fresh feces, a sand rat is "at home." 

may be several meters in length. Burrow systems in salt marshes 
are more complex than in desert, probably because of soil condi- 
tions. Burrow systems in mounds beneath desert plants often have 
a central chamber, and in wadis subject to flood, tunnels do not 
extend below the base of the mound. 

The number of openings in 19 burrow systems near Damanhour, 
Hafs, ranged from 4 to 11, with an average of seven. Shallow bur- 
rows with one, two, or three openings were considered temporary 
feeding or hiding places. In another area near Ras el Ish, the number 
of openings in 14 burrow systems ranged from 6 to 21, with an 
average of 11. 

Plants in salt marshes beneath which burrows are dug, in addition 
to the species listed above, are: Salicornia fruticosa, Halocnemon 
strobilaceum, Arthrocnemon glaucum, Salsola kali, exotic Atriplex 
nummularia, and other food plants of family Chenop)odiaceae. 
Desert populations burrow beneath Anabasis articulator Hammada 
elegans, and Lycium sp. (figs. 8, 48) in the Mediterranean Coastal 
Desert. Petter (1961) listed additional plants in Algeria. 



OSBORN&HELMY: MAMMALS OF EGYPT 239 

Associates.— Rodents living in the same habitats as Psammomys 
are: Pachyuromys duprasi, Jaculus orientalis, J. jaculus, Allactaga 
tetradactyla, Mus musculus, Dipodillus amoenus, D. simoni, D. 
henleyi, D. campestris, and Meriones shawl The following have 
been removed from Psammomys burrow systems: J. orientalis, D. 
amoenus, D. henleyi, M. musculus, Hemiechinus auritus, cobras 
{Naja sp.), and scorpions. 

Food and other uses of plants.— Fat sand rats require large quan- 
tities of food because of the high water content of vegetation on 
which they feed (Schmidt-Nielsen, 1964); therefore, succulent 
species of family Chenopodiaceae are their major natural food. Of 
these, the following are, from personal observation, known to be 
eaten in large quantities: Salicornia fruticosa, Halocnemon 
strobilaceum, Salsola tetrandra, Atriplex inamoena, A. halimus, 
Anabasis articulata, and Hammada elegans. Eaten in small quanti- 
ty are: Zygophyllum coccineum, Z. album, Frankenia hirsuta, and 
when green, Zilla spinosa. Desert populations feed chiefly on 
Anabasis articulata and Hammada elegans. 

The following list includes species found inside burrows: leaves 
and stems of Mesembryanthemum crystallinum and Limoniastrum 
monopetalum and calyces of Hyoscyamus muticus. Thirteen species 
listed by Wassif (1953b) added Bassia muricata, Salsola inermis, Ar- 
throcnemon glaucum, Nitraria retusa, Spergularia diandra, Cakile 
maritima, Parapholis marginata, and Sphenopus divaricatus. 

Three main food species recorded by Daly and Daly (1973) in Wadi 
Saoura, Algeria, were Suaeda vermiculata (=S. mollis), Traganum 
nudatum, and Salsola foetida. Zygophyllum album, if eaten to ex- 
cess, caused mild poisoning. They also noted that in some areas fat 
sand rats must compete with camels for food. 

Anderson (1902), upon removing 500 heads of barley from a bur- 
row, concluded that P. obesus was very destructive to grain. Obser- 
vations indicate this species is not a seed eater, although according 
to Petter (1961), it can be brought to accept sunflower seeds, along 
with carrots, lettuce, figs, cherries, etc. He, too, has found stalks of 
wheat within burrows of Psammomys. Grain might well have been 
brought into burrows by commensals such as Jaculus orientalis. 

Nests found in burrows in coastal deserts were made from shred- 
ded woody stems of Zilla spinosa and Anabasis articulata, Stipa 
capensis and various other Graminae, Ifloga spicata and Filago 





2| 




^ ^ S2 


« 




w^ 




cd 




s 





^ I I 



> 

9 



I I 



I I I 



2 I 



§3 

-a 3 



& I 



s 



-, 6 



CO 00 o cc 



ll 



OS U3 t^ M 
^ 'H t~ O 



en s 
t 

Q 



2 I 



I 2 I 



I I 



I I 2 



Hi 



eo « S 



3 



*- c 5 

o c CS 

cu < Q 



is * 
2 "^ 
2 c 

>. c O 
n 2 — 
•3 6 ■« 

e « CB 
C OB >~ 

< Q ^ 



240 



OSBORN&HELMY: MAMMALS OF EGYPT 241 

desertorum, Medicago sp., and a combination of paper and grass. In 
salt marshes near Damietta, nests were of fragmented Frankenia 
revoluta, Salicornia fruticosa, and rice straw; near the sea at Ras el 
Hekma, of long grass-like leaves of marine Posidonia oceanica and 
Cymodocea major. 

Included in nests were flower stalks of Plantago ovata, pods of 
Astragulus sp., stems of Phragmites australis, barley straw, 
feathers, dry camel dung, and an assortment of fragmented 
cigarette boxes, bits of plastic, gum wrappers, and cigarette filters 
gleaned from roadside areas. 

Reproduction.— Data on reproduction are limited, but indicate 
that the breeding period is from September to May, about eight 
months. Ten litters, fetuses, and nestlings, averaged 3.2, range 1 to 
8. Gestation in the Algerian subspecies is 23 to 25 days (Daly and 
Daly, 1975b). 

Popa/af ions.— Difference in numbers of animals inhabiting dense- 
ly vegetated salt marshes (fig. 7) compared with desert situations 
(fig. 5) are considerable. Data collected over a period of years from 
several salt marshes and one desert locality, Wadi Gindali, are in 
Table 28. Accurate estimates of areas are, however, lacking. 

On November 6, 1964, the population of a 200- by 8-m. strip of 
vegetation (Salicornia fruticosa and Halocnemon strobilaceum) on 
the eastern shore of Lake Manzala near Ras el Ish occupied 14 bur- 
row systems and consisted of six adult males, seven adult females, 
and one juvenile of each sex. There was approximately one mature 
animal per burrow system. Females with nestlings, juveniles, and 
sometimes subadults of mixed sexes are found occupying the same 
burrow system, but adult males and females, never. Daly and Daly 
(1974) studied spatial distribution in an area near Benne-Abbes, 
Algeria. 

Sex ratios.— Number of males and females in samples where 
removal was complete indicate about equal numbers or slightly 
more males in younger age groups and fewer males in older groups. 
Data in Table 29 indicate a predominance of females in all age 
groups. A greater turnover among males was noted by Daly and 
Daly (1974). 

Predators.— Diurnal hawks doubtlessly prey upon fat sand rats. 
The skin of a juvenile impaled on a shrub in Wadi Labaq, west of 
Maghra Oasis, indicated predation by a shrike. Cobras [Naja sp.) 



242 FIELDIANA: ZOOLOGY 

Tahi.k 29. — Number and per cent of males and females in three age classes of 
museum samples of Psammomys obesns. 





Adult 


Subadult 


Juvenile 




Males Females 


Males Females 


Males Females 


Number 


76 101 


30 54 


10 17 


Per cent 


43 57 


36 64 


37 63 



have, on several occasions, been removed from burrows in salt 
marshes. 

Economic importance.— The discovery by Schmidt-Nielsen et al. 
(1964) that Psammomys obesus develops diabetes mellitus syn- 
dromes from dietetic changes, has resulted in demands for large 
numbers of this species from diabetes research laboratories (Brunk 
and Strasser, 1967; DeFronzo et al., 1967), consequently researchers 
were challenged to develop techniques of maintaining laboratory 
colonies (Prange et al., 1968; Strasser, 1968). Further studies are by 
Hackel, Labovitz et al. (1967); Hackel, Mikat el al. (1967); Brodoff 
and Zeballos (1970); and Brodoff et al. (1971). 

Bedouins near Amiriya dig fat sand rats for food. 

Notation. — A maxillary with two molars of Psammomys obesus 
from the Upper Paleolithic site at Khor el Sil, Kom Ombo, was iden- 
tified by P. Turnbull (personal communication, 1975). This was from 
Late Pleistocene collections discussed by Reed et al. (1967) and 
Reed and Turnbull (1969). 

Key to Egyptian Siihspkciks of 
Psammomys obesus 

1. Posterior margin of nasals not reaching posterior limits of frontopremaxillary 
sutures (fig. 74). Dorsum brownish (table 26). (Northwestern Nile Delta and 
Western Desert) obesus, p. 242. 

2. Posterior margin of nasals reaching posterior limits of frontopremaxillary 
sutures (fig. 74). 

a. Size large, dorsum blackish (table 26). (Northeastern Nile Delta). nico//i, p. 243. 

b. Size smaller, dorsum reddish orange (table 26). (Eastern Desert and Sinai 
Peninsula) terraesanctae, p. 244. 

Psammomys obesus obesus Cretzschmar, 1828 

Type locality. -Egypt. ALEXANDRIA: Alexandria. 

Distribution in Egypt. — Figure 69. Northwestern part of Nile 
Delta and northern part of Western Desert. 

External characters. — Dorsum reddish to brownish orange, sides 
clear yellow, belly pale yellow with white hairs in axilla and groin in 



OSBORN&HELMY: MAMMALS OF EGYPT 243 

paler individuals. Tail with black hairs along about 75 per cent of 
dorsal side (table 26). 

Cranial characters.— Figure 72. Skull large and strongly ridged in 
majority of adults. Posterior margin of nasals not reaching level of 
posterior edges of frontopremaxillary suture (fig. 74). 

Measurements.— Table 25. Intermediate in dimensions between 
other two subspecies. 

Variation.— This subspecies varies considerably in color depend- 
ing upon habitat. Those from the southern limits of distribution are 
as pale as specimens of terraesanctae, but differ in having shorter 
basal bands on the hairs. Specimens from salt marshes within the 
Delta are sometimes almost as dark as nicolli, but lack the extensive 
amount of blackish hairs, particularly on the dorsal surface of the 
tail (table 26). 

Comparisons.— Psammomys o. obesus can be distinguished from 
nicolli and terraesanctae by the anterior position of the posterior 
margin of nasals (fig. 74) and, in most cases, by color (tables 26, 27). 

Specimens examined.— Total 221. 

MINUFIYA: Quweisna (1). 

BEHEIRA: Idku 2 km. W (1). Hafs (38). Abu el Matamir (2). 

TAHREER: Cairo- Alexandria desert road km. 153 (1). Nubareia (2). 

ALEXANDRIA: Mandara (1). Mex (1), El Amiriya (56). 

MATRUH: Lake Mariut (21); Bahig (2), 19 km. S (1), 25 km. S (2), 48 km. SE (1); 
Burg el Arab (9); Abu Mena (2): El Hammam (4), 6 km. S (5); El Afritat (5); Qaret el 
Mashruka 1 km. N (1); Qur el Hilab (1); El Maghra (1): Wadi Labaq. 30 km. W of El 
Maghra (2); El Alamein (9); El Daba (7); Ras el Hekma (9); Mersa Matruh (6), 22 km. 
E (1); Sidi Barrani (3), 42 km. W (2); Salum 3.2 km. S (7). 4.8 km. S (2). 10 km. SE (7). 
18 km. E (1). 22 km. E (6), 24 km. E (3). 48 km. E (1). 

Published records.— Records are from Anderson (1902), Bonhote 
(1909), Wassif (1953b), and Hoogstraal (1963). 

BEHEIRA: Rosetta. Idku 2 km. W, Hafs, Abu el Matamir, Busili, Abu Hommos. 
MUNIFIYA: Quweisna. 
ALEXANDRIA: Mandara. El Amiriya. 

MATRUH: Lake Mariut. Burg el Arab. Sidi Abd el Rahman. El Alamein, El 
Daba, Mersa Matruh. 

Psammomys obesus nicolli Thomas, 1908 

Psammomys obesus nicolli Thomas, 1908, Ann. Mag. Nat. Hist., (ser. 8), 2, p. 92. 
Type locality.— Egypt. DAMIETTA: Damietta. 



244 FIELDIANA: ZOOLOGY 

Distribution in Egypt — FigMre 69. Northeastern part of Nile 
Delta. Salt marshes, shores of Lake Manzajla. 

External characters. — Dorsum blackish to brownish yellow (fig. 
71), sides brownish, belly dark yellow. Tail with black hairs along 
entire dorsal surface. 

Cranial characters. —Skull large, strongly ridged. Posterior 
margin of nasals reaching level of posterior edges of frontopre- 
maxillary suture (figs. 72, 74; table 27). 

Measurements.— Table 25. Largest of subspecies. 

Co mpanson.— External features of subspecies listed in Table 26 
and hair color bands in Figure 71 indicate P. o. nicolli is much darker 
and with greater extension of melanin than either obesus or ter- 
raesanctae. 

Cranially, nicolli can be distinguished from obesus by slightly 
larger dimensions and position of posterior margin of nasals (fig. 
74). From terraesanctae, distinguishing characters are color and size 
(fig. 71; tables 25, 26). 

/Jemarfes. — Intergrades between nicolli and terraesanctae from El 
Ballah are listed under both names. A series somewhat paler than 
typical nicolli and having dimensions comparable with terraesanc- 
tae was placed in the latter category. 

Specimens examined— Total 90. 

PORT SAID: Port Said (12). Ras el Ish (26). Bayadeia (2). 
ISMAILIA: El BaUah (4). IsmaiUa (1). 
SHARQIYA: Tel Abu Ekaim (1). 

DAMIETTA: Ras el Bar (1), Shatt Gheit el Nasara (23). DamietU (19). Fariskur 
(1). 

Published records.— Records are from Thomas (1908), Hoogstraal 
(1963), and Setzer (1963). 

PORT SAID: Port Said. Ras el Ish. 

ISMAILIA: El Ballah. El Qantara. 

SHARQIYA: Between Gezira Seud and San el Haggar. Tel Abu Ekaim. 

DAMIETTA: Damietta. Shata, Shatt Gheit el Nasara. Fariskur. 

Psammomys obesus terraesanctae Thomas, 1902 

Psammomys obesus terraesanctae Thomas. 1902, Ann. Mag. Nat. Hist., (ser. 6). 9, 
p. 363. 

Type locality.— Palestine, Dead Sea. 



OSBORN&HELMY: MAMMALS OF EGYPT 245 

Distribution in Egypt — Figure 69. Northern and southern parts 
of Sinai Peninsula and northern part of Eastern Desert. 

External characters. — Palest subspecies of fat sand rat in Egypt. 
Dorsum reddish orange, sides clear yellow, and belly pale yellow 
with white hairs in axilla and groin (table 26). Dorsal hairs (fig. 71) 
with very narrow blackish or brownish terminal bands. 

Cranial characters. —Skull fairly strongly ridged. Posterior 
margin of nasals reaches level of posterior edges of frontopre- 
maxillary suture (fig. 74). 

Measurements.— Table 25. Smallest of subspecies in Egypt. 

Comparison.— Psammomys o. terraesanctae can be distinguished 
from nicolli and obesus by paler coloration (table 26) and smaller 
dimensions (table 25), and from the latter by difference in position of 
posterior margin of nasals (fig. 74). Though similarity in coloration 
exists between desert specimens of obesus and terraesanctae, the 
latter differs in having shorter tips, wider subterminal and basal 
color bands on dorsal hairs (fig. 71). 

i?emar/js.— Specimens of terraesanctae from dark, saline soils 
approach nicolli in coloration, but those from adjacent pale soils are 
light colored as are those from typical desert situations. The 
posterior margin of the nasals, unlike the majority from Egypt, is in 
the anterior position in the type of terraesanctae and a few addi- 
tional specimens from Palestine and Saudi Arabia. This may well be 
an example of character displacement. 

Specimens examined.— Total 37. 

SINAI: El Arish 137 km. W (2), Wadi Abu Aweigila (1), Wadi Gedeiret (1). 
ISMAILIA: El Ballah (12). 

SUEZ: Wadi el Rokham (1). Wadi el Gafra (2). Wadi Gindali (16). Wadi el Nasouri 
(2). 

Published records. — Records are from Flower (1932), Wassif 
(1953b), and Hoogstraal (1963). 

.SINAI: Rafa. Wadi Gedeirat. Khabra Abu Guzoar, El Hamda. Awlad Ali, El Qan- 
tara, Rumani. Quseima, El Arish. 

SUEZ: Wadi el Rokham. Wadi el Nasouri. 

Family 2. Spalacidae 
Genus Spalax Giildenstaedt, 1770 

Blind fossorial rodents lacking external ear. Tail not visible. 
Pelage soft, nondirectional. Supraoccipital broad, flat, and sloping 



246 FIELDIANA: ZOOLOGY 

forward. Interparietal absent. Median sagittal and lambdoidal 
ridges present, skull otherwise smooth. Upper incisor orthodont, 
smooth on anterior surface. Molar outline round, enamel pattern 
S-shaped. Dental formula: |. 5. sJ x 2=16. 

Spalax ehrenbergi Nehring, 1898 

Spalax ehrenbergi Nehring, 1898. S. B. Ges. Nat. Fr. Berlin (for 1897). pi. 2. p. 178. 

Type locality. — Israel: Jaffa. 

General distribution. — Syria, Lebanon. Israel. Egypt, Libya. 

Common names.— Mole Rat, Abu Amma. 

Subspecies in Egypt.— 

Spalax ehrenbergi aegyptiacus (Nehring, 1898) 

Spalax aegyptiacus Nehring, 1898, S. B. Ges. Nat. Fr. Berlin (for 1897). p. 180. 

Type locality. -Egypt. ALEXANDRIA: Ramleh. 

Distribution in Egypt — Figure 76. Northern part of Western 
Mediterranean Coastal Desert. 

Diagnosis.— Blind, mole-like, fossorial rodent. Pelage brownish, 
soft, nondirectional. Eye vestigial, covered with hairy skin. Pinna 
absent, meatal opening prominent. Tail not visible. Snout broad, 
flat, with band of soft, stiff bristles extending from nostril to level 
of eye. 

Supraoccipital large, flat, sloping forward. Median sagittal and 
lambdoidal ridges prominent, supraorbital and tempoparietal ridges 
absent. Interparietal absent. Infraorbital foramen large, incisive 
foramen small, palatal foramen minute. Upper incisor orthodont. 
anterior surface smooth. Molars round in outline, enamel pattern 
S-shaped. 

Adult total length average 184 mm., foot 25 mm., condylonasal 
length 43 mm. 

External characters. — Figure 77. Upper parts reddish to pale 
brown, venter grayish. All hairs with blackish bases; basal bands 
narrower on belly than on dorsal and side hairs. Fur velvety, soft, 
and nondirectional. Feet pale silver gray above. Pinna absent, 
meatal opening with a prominent cartilaginous tube. Snout broad, 
flat, with band of short, stiff, pale bristles extending from nostril to 
level of eye. Normal vibrissae absent. Tail not visible externally. 

Cranial characters. — Figure 78. Braincase triangular in shape. 




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247 



248 



FIELDIANA: ZOOLOGY 




Fig. 77. Live specimen of Spuiux fhr^nov/f-i ut-^ v/vnui u.- 



Zygomatic arch wide, especially at the temporal process. Supra- 
orbital and tempoparietal ridges lacking. Median sagittal and lamb- 
doidal ridges prominent. Interparietal absent. Posterior nasal 
margin narrow, irregular. Zygomatic plate small. Infraorbital 
foramen very large, incisive foramen small, palatal foramen minute. 
Parapterygoid fossa small, roof nearly obliterated by large foramen. 
Posterior palatal margin about level with m^, bearing a median 
spine. Palate constricted, deeply grooved and ridged. Tympanic 
bulla moderately inflated. Occipital condyle almost completely 
behind level of supraoccipital. Supraoccipital large and sloping for- 
ward to level of zygomatic process of temporal. Lambdoidal crest 
high and prominent. Mandible with well-developed coronoid pro- 
cess. Alveolar process prominent and protruding to height of the 
condylar process. Angular process turned outward. 

Teeth. — Figure 79. Incisors long. Upper incisor broad, orthodont, 




Fk; 78. Skull of Spalax ehrenbergi aegyptiacus. 



249 



250 



FIELDIANA: ZOOLOGY 



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K NOKVIGICUS 



A.NHOTICUi 



Fk; 79. Crown views of right upper (U) and left lower (L) molars of Egyptian 
Muridae and Spalax ehrenbergi. 

anterior surface smooth. Molars round in outline, inner and outer 
reentrant folds forming an S-shaped pattern. M'. m* subequal. 
slightly larger than m'. Folds in adult m' become isolated into two 
inlets. 

Measurements.— Tdhle 30. Males average slightly larger than 
females. 

Age determination. — Adults have folds of m' isolated into islets, 
cranial sutures closed, and median sagittal ridge prominent. 

Vanaf ton. — Measurements of specimens from Libya average 
slightly smaller than those from Egypt (Ranck, 1968). 



OSBORN & HELM Y: MAMMALS OF EGYPT 251 

Table 30. — Means (and ranges) of measurements, and weight of adult Spalax 
ehrenbergi aegyptiacus. 





Male 


Female 


Total length 


189.0 (163-203) 7 


178.0(155-204) 15 


FL 


25.0 (23-26) 7 


24.9 (23-26) 15 


CNL 


44.2 (42.5-46.5) 7 


41.8(37.9-44.5) 11 


ZW 


31.7 (29.5-33.8) 6 


28.8 (26.3-31.9) 13 


lOW 


7.1 ( 6.8- 7.5) 7 


7.1 ( 6.6- 7.5) 14 


IFL 


4.0 ( 3.8- 4.1)4 


3.8 ( 3.4- 4.0)4 


AL 


7.6 ( 7.4- 7.7) 4 


7.5 ( 7.4- 7.5) 4 


RW 


8.7 ( 8.4- 9.2) 7 


8.1 ( 7.5- 8.8) 14 


SH 


17.6 (16.4-18.9) 7 


17.3 (16.5-18.4) 12 


Wt 





107.2. 120.4 



Comparisons.— Spalax ehrenbergi differs from S. leucodon in 
smaller size and having two instead of one islet in adult m^. Spalax 
e. aegyptiacus differs from the nominate subspecies in smaller 
average measurements. 

Specimens examined.— Tots\ 35. 

MATRUH: Mariut (9); Bahig (9); Burg el Arab (10). 12.8 km. W (2); Zawiyet el 
Mithniyan (1); Sidi Barrani 28.8 km. E (1). 32 km. E. (1); Buq Buq 8 km. SW (1). 9.6 
km. SW (1). 

Sight record of D. J. Osborn.— 
MATRUH: El Hammam 6 km. S (mounds). 

Published records.— Records are from Anderson (1902), Flower 
(1932), and Hoogstraal et al. (1955). 

ALEXANDRIA: Ramleh. El Amiriya. 

MATRUH: Mariut, Burg el Arab, Mersa Matruh, Sidi Barrani 28.8 km. E. 

Collection.— Spalax is caught easily in Macabee pocket gopher 
traps. Hoogstraal (1963, p. 21) took specimens "with onion bait and 
a noose hung in an opened burrow." We have also captured it alive 
by closing off the burrow behind an animal when it came to plug the 
opening. A pit is dug 25 to 30 cm. behind the opening to within a few 
centimeters of the tunnel so that a shovel or hoe can be quickly 
driven across the tunnel to prevent retreat. 

Habitats.— VJestern Mediterranean Coastal Desert plain in deep 
sandy and loamy soils where A sphodelus microcarpus predominates 
and Bedouins have established barley fields. The most western site 
of collection in Egypt is 9.6 km. SW of Buq Buq in sandy soil under 



252 FIELDIANA: ZOOLOGY 

a plant community dominated by Thymelaea hirsuta. Not found in 
low areas subject to flooding nor in the higher rocky areas. 

Ranck (1968, p. 197) reported the species from the coastal plain 
and uplands of the Cyrenaican Plateau in Libya, and "larger valleys 
and more fertile soils of the higher tablelands." 

Burnoo's.— Subterranean tunnels 30 to 40 m. in length and 46 cm. 
below the surface were exposed near Mariut (Anderson, 1892). Tun- 
nels are usually branched and have food storage, sanitary, and 
nesting chambers. Excavated soil from several passages is pushed 
to the surface forming small mounds 10 to 20 cm. high. Breeding 
mounds, according to Nevo (1961) who studied S. e. ehrenbergi in 
Israel, average about 40 cm. in height, 160 cm. long, and 135 cm. 
wide. These mounds, constructed at the beginning of the rainy 
season (October-November) contain nesting and food storage 
chambers and numerous passages. 

BeAauior.— Extremely alert and aggressive in captivity, biting 
readily. 

Hoarding instinct has been observed in captive S. ehrenbergi 
(Anderson, 1892), but not in S. leucodon (Watson, 1961). Storage 
chambers containing tubers or bulbs are usually found in burrows of 
S. ehrenbergi (Anderson, 1902 and pjersonal observations of 
authors). 

Spalax initiates burrowing by scratching with the foreclaws and 
thrusting with the head. The incisors do the major work of excava- 
tion in hard ground. The head is also used to push and pack soft or 
loose soil (Montagu. 1924; Reed, 1958; Watson; 1961). During 
winter and spring rainy seasons, Spalax is active and mounds of ex- 
cavated earth are common, but in the dry season few are visible. 

Various aspyects of the behavior of S. ehrenbergi have been studied 
by Nevo (1961, 1969, p. 485). In the latter publication, mating 
behavior was said to consist of "three distinct stages— agnostic, 
courtship, and copulation" and to vary between chromosome forms. 
Copulation in nature, according to Nevo, takes place in the females' 
breeding mounds. 

Although adapted to live subterraneanly, S. ehrenbergi is known 
to be active on the ground surface during night or day for the occa- 
sional purposes of collecting green food, finding a mate, and in the 
case of dispersing young, searching for future territory. Spalax 
skulls found in barn owl pellets (Bate, 1945; Dor, 1947) were the 
original indirect evidence of nocturnal surface activity. 



OSBORN & HELMY: MAMMALS OF EGYPT 253 

Food— Bulbs, tubers, and roots are gathered by Spalax and 
stored in the burrow system. Sixty -eight bulbs were once removed 
from two storage chambers (Anderson, 1902). An important food of 
S. e. aegyptiacus is the tubers of a lily, Asphodeles microcarpus. 
Most of the plants listed by Nevo (1961) which are eaten by S. e. 
ehrenbergi in Israel, occur in the habitat of S. e. aegyptiacus. 
Among these are bulbs and corms of Narcissus tazetta, Belevalia 
flexuosa, Gladiolus italicas, Oxalis pes-caprae, Arisarum vulgare; 
roots of Alhagi mannifera; and foliage of Asphodelus microcarpus, 
Urginea maritima, Medicago sp., Hordeum sp., and Eryngium sp. 

Captive animals have been fed potatoes, onions, carrots, beets, 
and shelled broad beans (Watson, 1961). We found that S. e. aegyp- 
tiacus could be maintained satisfactorily for several days on an 
onion diet. 

Associates.— ^MTTOvrs of Spalax occur in habitats occupied by 
Gerbillus andersoni, Meriones shawi, Jaculus jaculus, J. orientalis, 
Allactaga tetradactyla, Psammomys obesus, and other species in- 
habiting sandy and clay soils of the Western Mediterranean Coastal 
Desert. 

Reproduction.— No data from Egypt. Nevo (1961) recorded one 
litter per year, pregnant females from January-March, most fre- 
quent number of young three to four, and range of litter size one to 
nine in Israel. 

Sex ratio.— A sample of 27 museum specimens of S. e. aegyptiacus 
included seven (26 per cent) males and 20 females. Fewer males than 
females were also found in S. e. ehrenbergi. In 18 litters comprising 
a total of 67 specimens, 27, or 40.3 per cent, were males, and in 106 
adults, 37, or 34.9 per cent, were males (Nevo, 1961). 

Remarks.— Lay and Nadler (1972) postulated that the ancestral 
stock of North African Spalax originated from northern Sinai and 
southern Israel populations, which, even though separated by a 
hiatus of 400 km., and some 10,000 to 25,000 years, have the same 
2n=60 karyotypes. 

Wahrman et al. (1969) discovered that in Israel, distinct, but 
homogeneous, populations occurred with decreasing diploid 
numbers northward in correlation with decreasing aridity. 



Family 3. Muridae 
Fur soft to relatively harsh and spinous. Tail without apical 



264 FIELDIANA: ZOOLOGY 

brush, annulations not concealed by hair or bristles. Supraorbital 
spots not prominent except in genus Acamys. Supraorbital and 
tempoparietal ridges prominent except in genus Mus. Infraorbital 
foramen relatively large, incisive foramen long, except in genus 
Nesokia; palatine foramen minute. Tympanic and mastoid bullae 
slightly inflated. Upper incisor variable in curvature, anterior sur- 
face smooth. Cheek teeth tuberculate (tubercles in three 
longitudinal rows) or laminate, never prismatic. Dental formula: {, 5, 
gjx2=16. 

KkY T() ECYITIAN GkNKRA OF MllRIDAK 

1. Dorsal pelage spinous. Tempoparietal suture following cranial ridge. Inter- 
parietal very large, semicircular in outline, occupying one-half or more of area be- 
tween cranial ridges Acomys, p. 285. 

2. Dorsal pelage not spinous. Tempoparietal suture turning downward and caudad 
above base of zygomatic process of temporal bone. Interparietal with variable 
outline, occupying less than one-half of area between cranial ridges. 

a. Size large, head and body length greater than 150 mm. 

i. Dorsum brownish. Palatal margin level with or behind posterior edge of m . 
(a) Incisive foramen relatively long. Zygomatic arch not bowed laterally. Up- 
per incisor compressed, opisthodont. Mandible with small alveolar 

process Rattus, p. 263. 

(b) Incisive foramen actually and relatively very short. Zygomatic arch 
bowed laterally. Upper incisor not compressed, proodont. Mandible with 

large alveolar process Nesokia, p. 309. 

ii. Dorsum speckled black and yellowish. Palatal margin anterior to posterior 
edge of m' Arvicanthus, p. 254. 

b. Size small, head and body length less than 100 mm Mus, p. 273. 



Genus Arvicanthis Lesson, 1842 

Large murid, pelage relatively harsh, speckled black and 
yellowish dorsally with black middorsal stripe. Tail shorter than 
head and body, distinctly bicolored, annulations almost concealed 
by hair. 

Skull massive, strongly ridged. Zygomatic arch thickened in mid- 
dle. Interparietal ovoid to rectangular in outline. Postpalatal 
margin anterior to posterior edge of m"*. M' seven-rooted, slightly 
longer but narrower than m''. 

Arvicanthis niloticus (Desmarest, 1822) 

Arvicola niloticus Desmarest. 1822, Kncylcopedie Methodique Mammalogie. pt. 
2. p. 281. 

Type locality.— Egypt. 



OSBORN&HELMY: MAMMALS OF EGYPT 255 

General distribution.— ^ontYivrestern Arabia, Egypt, Sudan, 
Ethiopia, Uganda, Kenya, Tanzania, Chad, and from Nigeria west 
to Senegal. 

Common names.— Nile Kusu, Grass Rat, Field Rat, Far el Gheiti. 

Subspecies in Egypt — 

Arvicanthis niloticus niloticus (Desmarest, 1822) 

Type locality.— Egypt. 

Distribution in Egypt— Figure 80. Nile Valley and Delta, El 
Faiyum, Dakhla and Kharga Oases, El Maghra, and canals extend- 
ing into Western Mediterranean Coastal Desert. 

Diagnosis.— harge, slender rat with dorsum speckled black and 
yellow; belly white. Ear longer than one-half hind foot length, 
covered with orangish hairs. Tail thin, shorter than head and body 
length, bicolored, blackish above, yellowish below. 

Supraorbital ridges prominent and forming a postorbital process 
in adults; tempoparietal ridges curving and high on the braincase. 
Incisive foramen relatively long and extending to level of anterior 
root of m'. M' lacking cingulum on anterior border of crown. 

Adult head and body length average 180 mm.; tail 142 mm., 78 
per cent of head and body length; foot 36 mm.; ear 21 mm.; con- 
dylonasal length 38.5 mm.; weight 140 gm. 

External characters.— Figure 81, Table 32. Dorsal pelage coarse, 
speckled black and yellow. Hairs of dorsum with narrow black tip, 
broad yellow subterminal band, and black base. Middorsal stripe 
black, indistinct or distinct, extending from crown to base of tail. 
Belly hairs whitish with black base. Mystacial and circumorbital 
area, ear, and small postauricular patch orangish. Long yellowish to 
orangish hairs on rump. Foot orangish to blackish above. Palm and 
sole bare, pigmented. Tail bicolored, blackish above, whitish to 
yellowish below; hairs almost concealing annulations. 

Cranial characters.— Figure 82 and Table 32. Dorsal skull outline 
convex in lateral view. Braincase relatively broad. Zygomatic arch 
thickened in middle, not bowed laterally. Rostrum broad, short in 
appearance. Nasals tapering gradually posteriorly, posterior margin 
abrupt and divided. Supraorbital ridge prominent and forming a 
postorbital process in adults. Frontoparietal suture U-shaped. Tem- 
poparietal suture turning abruptly downward and caudad above 



266 



FIELDIANA: ZOOLOGY 



. 25* 26* 27* 26* 2 9* 30* 3l' 32* 33* 34* 35* 36* 37* 




Fig. 80. Collection localities of Arvicanthis niloticus niloticus. 

base of zygomatic process of temporal. Temjjoparietal ridge high on 
cranium, curving slightly laterad. Parietal part of ridge less 
developed than temporal. Interpeirietal irregularly ovoid to almost 
rectangular in outline. Lambdoidal and median supraoccipital 
ridges prominent. Occipital condyle protruding slightly beyond 
posterior level of supraoccipital. 

Zygomatic plate sharply rounded above, anterior margin nearly 
vertical, not reaching level of premaxillary-maxillary suture. In- 
fraorbital foramen relatively large. Incisive foramen long and 
narrow, posterior margin almost level with anteromedial root of m'. 
Postpalatal foramen minute and situated on or just anterior to the 
maxillopalatine suture. Posterior palatine margin anterior to 
posterior level of m^. As in Anderson's (1902, p. 280) description, 
there is "no post dental shelf." Palate constricted slightly. 
Parapterygoid fossa narrow and deep. Tympanic bulla moderately 



OSBORN & HELMY: MAMMALS OF EGYPT 



257 




Fig. 81. Dorsal views of Muridae. Left to right: Arvicanthis niloticus niloticus 
(two specimens), Rattus rattus (two specimensi, R. noruegicus, and Nesokia indica. 



inflated, compressed laterally. Mandible relatively deep, coronoid 
process prominent, alveolus of incisor small. 

Teeth.— Yig^ive 79 and Table 32. Incisor compressed, opisthodont, 
anterior surface smooth. Molars relatively heavy, cuspidate, becom- 
ing laminate. Laminae crescent shaped, particularly on m'. M' with 
five long and two short roots, cingulum lacking, and anterolateral 
cusp present. M^ broader than m', anterolateral cusp variable; pre- 
sent on mj. M^ with two laminae and a distinct anteromedial cusp. 



Measurements.— Tsihle 31. 
subequal. 



Male and female measurements 







Fii; 82. Skull of Arvicanthis niloticus niloticus. 



258 



OSBORN&HELMY: MAMMALS OF EGYPT 259 

Table 31. — Means (and ranges) of measurements, ratios, and weight of adult 
Rattus rattus, R. norvegicus, and Arvicanthis niloticus. 

R. rattus R. norvegicus A. niloticus 

HBL 180.1 (156-208) 19 219.9 (196-254) 13 180.6 (159-202) 20 

TL 219.3 (188-244) 17 196.0 (145-234) 13 142.3 (125-173) 15 
TL/HBL% 121.4 (113.3-134.1) 17 88.4 (64.2-99.0) 12 78.7 (70.9-90.2) 15 

FL 36.2 (32-39) 21 43.4 (40-51) 13 36.7 (33-42) 20 

EL 23.7(21-26)21 20.8(20-23)13 21.0(19-23)20 

Wt 137.3(87.0-174.0)17 259.3(208.3-360.0)4 139.8(102.0-201.2)8 
CNL 41.4 (38.5-44.6) 18 46.8 (43.2-52.2) 12 38.5 (35.1-41.8) 19 

ZW 19.8(17.7-22.4)18 23.2(20.9-26.9)7 19.6(18.2-20.8)18 

low 5.8 ( 5.3- 6.7) 18 6.5 ( 5.9- 7.3) 12 5.6 ( 5.0- 5.9) 19 

RW 7.5 ( 6.7- 8.6) 17 8.9 ( 8.2-10.5) 11 7.2 ( 6.5- 7.9) 13 

NL 15.1 (13.2-16.9) 17 17.5 (16.1-19.3) 12 15.0 (13.0-16.5) 19 

IFL 7.4 ( 6.8- 8.4) 18 7.9 ( 7.0- 8.8) 12 8.6 ( 8.2- 9.5) 19 

AL 6.9 ( 6.3- 7.6) 18 7.4 ( 6.8- 7.8) 12 7.6 ( 6.8- 8.2) 19 

SH 14.6(13.3-16.3)18 16.3(15.1-18.2)10 14.8(14.0-16.0)19 

Age determination.— Adults have well-developed postorbital pro- 
cess and cranial sutures closed. 

Variation. — Dorsal stripe distinction and amount of yellow or 
orange hair on rump vary individually. 

Comparisons.— Arvicanthis niloticus differs from other large 
Egyptian murids in speckled color and orangish marking, post- 
palatal margin anterior to posterior edge of m\ and other characters 
in Table 32. Arvicanthis n. niloticus differs from Sudanese 
subspecies testicularis in darker color and proportionately shorter 
tail. 

Specimens examined.— Total 130, 

DAQAHLIYA: Minshat el Ikhwa (1). Mit Ghamr (3). 

BEHEIRA: Wadi el Natroun (1). Rosetta (6), Kafr el Dawar (2). 

MINUFIYA: Quweisna (1), El Ghunamiya (1). Nadir (1). Birket el Sabh (1). 

GIZA: Giza (3), Wardan (1), El Baragil (1), Abu Rawash (3), Kirdasa (3). Zawyet 
Abu Mussalam (1). Sakkara (4). Bulaq el Dakrur (1). El Marazig (2). 

CAIRO: Cairo (12). Maadi (1). 

EL FAIYUM: Shooting Club (1). Shakshuk (2). Lake Qarun (2), Gharah (1). 

SOHAG: Awlad Hamza (3). Sohag (2). 

QENA: Qena (4). Dandara (4), Isna (5). Isna E. (1). Wadi Nassim (1). Dishna (4), 
Abu Shusha (6). Luxor (1). 

ASWAN: East Aswan (1). 

MATRUH: Bahig (26). El Maghra (3). 

EL WADI EL GEDEED: Dakhla Oasis. Mut (13): Kharga Oasis, El Kharga (1). 



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262 FIELDIANA: ZOOLOGY 

Published records. — Records are from Anderson (1902) and Setzer 
(1952, 1963). 

DAQAHLIYA: Minshat el Ikhwa. 
ALEXANDRIA: Alexandria. 
BEHEIRA: Wadi el Natroun. 
MINUFIYA: Quweisna. El Ghunamiya. 
QALYUBIYA: Qalyub. Qalama. 

GIZA: Wardan. El Baragil. El Magedla. Zawyet Abu Massalam, Nahya. Abu 
Rawash. Sakkara. Bulaq el Dakrur. 

EL FAIYUM: Shooting Club. Lake Qarun. 
MINYA: Minya 
QENA: Wadi Nassim. 
ASWAN: Aswan. 

Habitats. — Flov/er (1932) said A. niloticus was never known to 
enter houses. Hoogstraal, (1963, p. 22) remarked that, although 
these animals live in close proximity to human structures, they are 
"seldom if ever encountered inside buildings." 

Setzer (1952) likened A. niloticus in appearance with the North 
and South American cricetine genus Sigmodon, which also occupies 
a similar ecological niche. 

In nature, habitats are canal banks with grass cover of 
Desmostachya bipinnata and Imperata cylindrical gardens, fields, 
railway embankments, and almost any damp area under vegetative 
cover or piles of crop wastes. Recent collections (1967) are from oHve 
groves and gardens in Bahig following completion of a freshwater 
canal into that area of the Western Mediterranean Coastal Desert. 

One specimen of .4. niloticus was found dead under a date palm in 
the palm-reed-rush community of El Maghra in 1965 and two were 
trapped there in 1974. How the animals arrived in such an area sur- 
rounded by many kilometers of waterless desert is a mystery, unless 
they were carried in accidently in some kind of camel load, as has 
also been assumed in the case of Mus musculus. 

Burrows— Burrows are shallow, long, and winding with numerous 
openings and never closed. Short burrows are used as temporary 
hiding or feeding places. 

Activity and 6e/iaf tor. — Diurnal and nocturnal. Usually seen and 
trapped during hottest period of the day. Can be seen crossing 
highways that run between canal banks or other areas of suitable 
habitat. 



OSBORN & HELMY: MAMMALS OF EGYPT 263 

Hassan and Hegazy (1968) reported one mature male and one 
mature female per burrow, except during the winter when these rats 
congregate for warmth. 

Arvicanthis niloticus, like Rattus sp., is aggressive and difficult 
to handle. 

Reproduction. — hccoTdSxig to Hassan and Hegazy (1968), the 
main breeding period is June-November. They found 60 per cent of 
females pregnant in October and five to six per cent in January. 
Gestation period was 20 days, litter size five to six, number of lit- 
ters per female three to four per year, and females matured in three 
months. The life span in nature was estimated at two and one-half to 
three years. 

Happold (1966b) reported young were seen at all times in North 
Sudan, and pregnant females with two to ten young were collected 
in February, July, and September. The rainy season had no effect on 
reproduction. 

Sex ratio.— A museum sample of 40 consisted of 17 (42.5 per cent) 
males and 23 females. 

Economic importance.— Hassan and Hegazy (1968) place con- 
siderable emphasis on the importance of A. niloticus as a pest to 
agriculture. They considered it to be the most numerous of murids, 
because its natural enemies, snakes and mongoose, have been killed 
off by man, and because the climate is favorable. 

It reportedly eats grains, vegetables, and fruits; gnaws the base of 
sugar cane, damaging approximately 30 per cent in upper Egypt; 
damages trees by gnawing the bark; and takes cotton bolls to make 
nests. 

i?emar/es. — Possibly A. niloticus is Klunzinger's (1878, p. 148) 
"large thick-headed field mice which frequent the fields and earthen 
dikes, and in many quarters are esteemed as dainties by the 
peasants." It was also eaten by the Romans. 

Genus Rattus Fischer, 1803 

Large murids with relatively harsh pelage, brownish dorsally. Tail 
length variable, color variable, annulations conspicuous. 

Skull massive, strongly ridged. Zygomatic arch slender. Inter- 
parietal semicircular to shield-shaped in outline. Postpalatal margin 
posterior to m\ M' five-rooted, crown longer than m^. 



264 FIELDIANA: ZOOLOGY 

Key to Egyptian Spkciks of Rattua 

1. Back brownish; belly gray, buff or cream. Tail length more than 100 per cent of 
head and body, slender, not bicolored. Kar length more than one-half hind foot. 
Tempoparietal ridges curving laterally, low on braincase. M without a cingulum, 
m with anterolateral cusp (fig. 79) rattus, p. 264. 

2. Back brownish: belly gray. Tail length less than 100 per cent of head and body, 
thick, bicolored. Ear length less than one-half hind foot. Tempoparietal ridges 
parallel, high on braincase. M with a cingulum, m without anterolateral cusp 
(fig. 79) norvegicus, p. 269. 

Rattus rattus (Linnaeus, 1758) 
Mus rattus. Linnaeus, 1758, Syst. Nat., 10th ed., p. 61. 
Type locality. —Sv/eden: Upsala. 

General distribution.— Almost cosmopolitan due to accidental 
transportation by man. May occur in the wild state in parts of 
southeastern Asia. 

Common names.— House Rat, Black Rat. 

Distribution in Egypt — Figure 83. Nile Valley and Delta, coastal 
towns, and certain oases in Western Desert. 

Diagnosis.— Large, slender murid. Dorsum brownish; venter 
gray, buff, or white. Muzzle sharp. Ear longer than one-half hind 
foot length, sparsely haired. Tail thin, longer than head and body 
length, not bicolored. 

Skull elongate. Tempoparietal ridges low on cranium, curving 
laterally; area between ridges convex. Zygomatic arches slender, 
not flaring laterally. Occipital condyle not protruding beyond level 
of supraoccipital. Incisive foramen reaching level of anterior root of 
m'. M' lacking a cingulum on anterior border of crown. 

Adult head and body length average 180 mm.; tail 219 mm., 120 
per cent of head and body length; foot 36 mm.; ear 24 mm.; con- 
dylonasal length 41.4 mm.; weight 137 gm. 

External characters. — Figure 81. Three color phases occurring in 
Egypt, with previously recognized subspecific names in paren- 
theses, are: 

1. Dorsum blackish brown; venter, hand, and foot gray (rattus). 

2. Dorsum grizzled brown; venter hairs gray with buff tip; hand 
and foot brownish, toes whitish ialexandrinus). 

3. Dorsum pale grizzled brown; venter hairs white with pale 



OSBORN&HELMY: MAMMALS OF EGYPT 265 

, 25* 26* 27* 28* 2 9* 30* 31* 32* 3 3' 34* 35* 36* 37* 



I 30 




Fig 83. Collection localities of Rattus rattus (circles) andR. norvegicus (dots). 

yellowish or cream tip; hand and foot pale grayish to whitish 
ifrugivorous). 

Ear relatively long and sparsely haired. Tail thin, relatively long, 
brownish, not bicolored. Palm and sole naked. 

Cranial characters.— Figure 84. Posterior margin of nasals round 
or truncate, not reaching level of posterior margin of frontopre- 
maxillary suture. Supraorbital and parietal ridges strongly 
developed, the latter curving laterally, low on side of braincase; area 
between ridges convex. Median supraoccipital ridge prominent. 
Occipital condyle not protruding beyond level of supraoccipital. 
Zygomatic plate projecting forward slightly, gradually rounded 
above, narrow in proportion to height. Incisive foramen usually 
reaching level of anterior root of m'. 






FlU. 84. Skull of Rattus rattus. 



266 



OSBORN&HELMY: MAMMALS OF EGYPT 267 



Teeth. — Figure 79. Upper incisor smooth on anterior surface. M' 
without cingulum on anterior border of crown and lacking accessory 
outer tubercle. First laminae of m' crescentric, cusps normal. M^ 
with posterior outer tubercle, m.^ with or without accessory 
anterolateral tubercle. 

Measurements.— Table 31. Male and female dimensions subequal. 
Means (and ranges) of condylonasal length (in millimeters) of nine 
adult males and nine adult females are 40.7 (38.5 to 46.8) and 42.1 
(39.4 to 44.6), respectively. 

Note that tail length in R. rattus is more than 100 per cent of head 
and body length, ear length more than one-half hind foot length, and 
hindfoot length less than 40 mm. 

Age determination.— Adults have cusps of molars worn to 
laminate pattern, cranial sutures closed. 

Variation.— Color phases listed above were previously given 
subspecific names, but all three and intermediates may occur in any 
one locaHty in Egypt (Setzer, 1952). Caslick (1956) found different 
color phases in the same litter in R. rattus in U.S.A. 

Comparisons.— Rattus rattus differs externally from R. 
norvegicus in having head and body length averaging shorter, tail 
longer rather than shorter than head and body, ear more than one- 
half hind foot length, and hind foot actually shorter; cranially, in 
having the tempoparietal ridges curving outward instead of being 
parallel, occipital condyle not protruding rather than protruding 
beyond the level of the supraoccipital, and m' lacking a cingulum 
(table 32). 

Both R. rattus and R. norvegicus can be distinguished from 
Arvicanthis niloticus on the basis of color, since the latter has the 
dorsum speckled black and yellow, whereas in Rattus, the dorsum is 
brownish. Cranially Rattus sp. have the postpalatal margin behind 
m\ whereas in Arvicanthis, it is anterior to m'. Other differences are 
in Table 32. Differences between R. rattus and Nesokia indica are 
listed under the latter. 

Specimens examined.— Total 96. 

SINAI: El Arish (1). 
PORT SAID: Ras el Ish (5). 
ISMAILIA: IsmaiUa (4). 
SUEZ: PortTawfik(l). 
GHARBIYA: Mehalla el Kubra (1). 



268 FIELDIANA: ZOOLOGY 

DAMIETTA: Shatt Gheit el Nasara (3). Kafr el Battikh (1). 

KAPR EL SHEIKH: Kafr el Sheikh (1). Baltim |5k El Burg 111. 

BEHEIRA: Rosetta (1). Wadi el Natroun (1). Beni Salami (1). 

ALEXANDRIA: Alexandria (5). 

MINUFIYA: Birket el Sabh (2). 

QALYUBIYA: El Khanka (1). 

GIZA: Giza (4); Giza Zoological Gardens (1); El Badrshein, El Maraziq (4): Abu 
Rawash (6); Abu Ghalib (1); Tanash (5); Sakkara (1). 

CAIRO: Cairo (3). Abassia Fever HospiUl (1). Gebel Mokattam (1). Bulaq (1). 
Heliopolis (1). Maadi (1). 

EL FA I YUM: Seila (2). Abu Gandir (1). Qalamsha (4|, Gharah (4). Kom O Shim (1). 

ASYUT: Asyut (1). 

QENA: Luxor. Valley of the Kings (1); Isna, Wadi Nassim (1|, Dandara Temple HI, 
El Taramsa (1|. 

ASWAN: Kom Ombo (5|. 

MATRUH: BahigdI. 

EL WADI EL GEDEED: Kharga Oasis, Bulaq (2|, Farafara Oasis, Abu Minqar 
(7). 

Published records. — Records are from Anderson (1902), Bonhote 
(1910), Flower (1932), Setzer (1952, 1963). and Hoogstraal (1963). 

SINAI: El Arish. 

ISMAILIA: Fayid. 

DAQAHLIYA: Simbillawein 8 km. W. 

DAMIETTA: Shatt Gheit el Nasara. Kafr el Battikh. 

KAFR EL SHEIKH: El Burg. 

ALEXANDRIA: Alexandria (Alexandria harbor area. Karmouz. El Manshiya. El 
Atarein. El Labban. Mina el Basall; Ramleh. 

BEHEIRA: Rosetta, Damanhour. Fuwa, Wadi el Natroun. 

GIZA: Giza; Abu Rawash; Abu Rawash 3.2 km. W. 4.8 km. W; Tanash; Mena 
suburb; Sakkara; Kafr Teharmes; Talbia; Atf. 

CAIRO: Cairo. Heliopolis. Bulaq. Abbasia, Maadi. 

EL FAIYUM: Faiyum. Kom O Shim, Qasrel Gebali. 

MINYA: El Minya. 

ASYUT: Asyut, Beni Adi, El Wilidiya, El Badari. 

QENA: Isna, Wadi Nassim. 

ASWAN: Aswan. 

MATRUH: Mersa Matruh. 

Sudan. NORTHERN: Wadi Haifa. 

Collection. — Dug from burrows and live-trapped. Conibear traps 
are effective in capturing rats that avoid live traps. 

//a6i7ats.— Commensal with man, R. rattus is common in houses 



OSBORN&HELMY: MAMMALS OF EGYPT 269 

and buildings throughout the Nile Delta and Valley and in coastal 
towns. Hoogstraal (1963) mentioned finding it in desert towns along 
the Alexandria-Mersa Matruh railway. In the Abu Minqar area 
south of Farafara Oasis, R. rattus was trapped in houses and grain 
storage areas. In the Nile Valley and Delta, R. rattus inhabits canal 
banks and cultivated fields. According to Hassan and Hegazy 
(1968), this species prefers a drier habitat than R. norvegicus such 
as higher parts of buildings and seed and grain stores. The 
frugivorous phase or "palm rat" they wrote, is harmful to date palm 
trees. 

Burrow's.— Burrows are like those of/?, norvegicus, shallow and 
with many openings, but not as close to water. 

Activity.— Y)i\xrTia\. and nocturnal. 

Captive behavior.— \A]&^e R. norvegicus, R. rattus is aggressive, 
difficult to handle, and bites readily. 

Food.— Rattus rattus is known to feed on tomatoes, egg plant, 
and other vegetables in rural Egypt. Bonhote (1910) noted that this 
rat ate the fruit and seeds of the plane tree {Ficus sycomorus). 
Hassan and Hegazy (1968) reported it feeding in grain and seed 
stores. 

Populations.— Rattus rattus appears to be more numerous than 
R. norvegicus (Hoogstraal, 1963). 

Associates.— Wthongh Hassan and Hegazy (1968) stated thati?. 
rattus avoids the habitats of R. norvegicus, the two species are 
often collected in the same local area. Further comments are under 
R. norvegicus and Arvicanthis niloticus. 

Sex ratio.— A museum sample of 24 contained 11 (46 per cent) 
males and 13 females. 

Rattus norvegicus (Berkenhout, 1769) 

Mus norvegicus Berkenhout 1769, Outlines Nat. Hist., Gt. Britain and Ireland, 
Vol 1, p. 5. 

Type locality.— Great Britain. 

General distribution. — Nearly cosmopolitan due to accidental 
transportation by man. May occur in original wild state in north- 
eastern Asia. 

Common names. — Norway Rat, Brown Rat, Sewer Rat. 



270 FIELDIANA: ZOOLOGY 

Distribution in Egypt — Figure 83. Coastal towns, Nile Delta and 
Valley. ^ 

Diagnosis.— "Large, stocky rat. Dorsum dark brown, venter gray. 
Muzzle blunt. Ear shorter than one-half hind foot length, densely 
haired. Tail thick, shorter than head and body length, bicolored. 

Skull elongate. Tempoparietal ridges parallel, high on side of 
cranium, area between ridges almost flat. Zygomatic arches slender, 
not flaring laterally. Occipital condyle protruding beyond level of 
supraoccipital. Incisive foramen not reaching level of anterior root 
of m'. M' with a cingulum on anterior border of crown. 

Adult head and body length average 220 mm.; tail 196 mm., 88 
per cent of head and body length; foot 43 mm.; ear 20 mm.; con- 
dylonasal length 46.8 mm.; weight 259 gm. 

External characters. — Figure 81. Dorsum dark brown, side brown 
mixed with gray; belly hairs grayish, with white or cream tips. All 
hairs with gray base. Ear relatively short, densely covered with 
short hairs. Tail thick, relatively short, sparsely haired; bicolored, 
brownish above, paler or whitish below. Feet sparsely covered with 
whitish hairs above, palm and sole naked. 

Cranial characters. — Figure 85. Nasals with posterior margin 
rounded or pointed, usually level with posterior margin of frontopre- 
maxillary suture. Supraorbital ridges strongly developed, tem- 
poparietal ridges parallel, high on side of braincase, area between 
ridges relatively flat. Median supraoccipital ridge prominent. 

Occipital condyle protruding beyond level of supraoccipital. 
Zygomatic plate projecting forward slightly, sharply rounded 
above, wide in proportion to height. Incisive foramen seldom 
reaching level of anterior root of m'. 

Tee (A. — Figure 79. Upper incisor smooth on anterior surface. M' 
with cingulum on anterior border of crown, sometimes with small 
accessory outer tubercle on first lamina; first lamina distorted by 
supression of outer cusp. M^ without posterior outer tubercle, mj 
with accessory transient anterolateral tubercle. 

A/easarements.— Table 31. Male and female dimensions subequal. 
Means (and ranges) of condylonasal length (in millimeters) of four 
adult males and eight adult females are 47.9 (45.0 to 52.2) and 46.2 
(43.2 to 50.7), respectively. 

Note that tail length in R. norvegicus is less than 100 per cent of 







Fig. 85. Skull of Rattus norvegicus. 



271 



272 FIELDI ANA: ZOOLOGY 

head and body length, ear length less than one-half hind foot length, 
and hind foot length is 40 mm. or more. 

Age determination. — Adults have cusps of molars worn to 
laminate pattern, cranial sutures closed. 

Comparisons.— Rattus norvegicus and R. rattus are compared 
under the latter and with Arvicanthis in Table 32. 

Specimens examined— Total 41. 

PORT SAID: Port Said (2). El Ghamil Beach (3), Ras el Ish (1). 
KAFR EL SHEIKH: Baltim (1). El Hamul (1). 
ALEXANDRIA: Alexandria (6). 

GIZA: Giza (1), Giza Zoological Gardens (4). Abu Rawash (12). Abu Ghalib (1). 
Imbaba 1.6 km. W (2). 

BEHEIRA: Wadi el Natroun (1). 

ASYUT: Asyut Fever HospiUl (1). Beni Adi (1). 

QENA: Isna (2). 

SUDAN ADMINISTRATIVE: Abu Ramad (2). 

Published records.— Records are from Anderson (1902), Bonhote 
(1910), Flower (1932), Setzer (1952, 1963), and Hoogstraal (1963). 

PORT SAID: Port Said. 
ISMAILIA: Ismailia. 
SUEZ: Suez. 

ALEXANDRIA: Alexandria (Alexandria harbor area. Mina el Basal, El Labban. 
Karmouz, Sharia France, Customs area. El Atarien). 

BEHEIRA: Abu Hommos. Damanhour. Fuwa. Wadi el Natroun. 

GIZA: Abu Rawash. Abu Ghalib. Kafr Hakim. Imbaba 1 km. W. Atf. 

CAIRO: Cairo. 

EL FAIYUM: Faiyum. 

ASYUT: Beni Adi. 

QENA: Isna. 

ASWAN: Aswan. 

Collection.— Dug from burrows and live-trapped. Conibear traps 
are effective in capturing this species. 

Egyptian rat-catchers, if not cautioned, will break off the incisors 
to lessen the chances of being bitten or having the rats chew 
through a sack and escaping. 

Habitats.— Commensal with man, R. norvegicus has been trapped 
in village houses and buildings in the Nile Valley and Delta and in 
buildings in coastal towns. In the wild state, habitat requirements 



OSBORN & HELMY: MAMMALS OF EGYPT 273 

are definitely mesic. Burrows in canal banks are near water and the 
species has also been dug from burrows beside the sea. Hassan and 
Hegazy (1968) remarked that R. norvegicus burrowed beneath 
buildings because of dampness. They also found it in stables and 
chicken houses. 

^urroifs.— Burrows usually have several openings that are never 
closed. Burrows are commonly dug around and under buildings and 
near water. 

Behavior.— Mainly nocturnal. Rattus norvegicus readily enters 
water and often escapes native rat-catchers by diving into canals 
and swimming under the surface. Captive rats are aggressive and 
difficult to handle without being bitten. 

Food— Diet of/?, norvegicus appears to be more omnivorous than 
that ofR. rattus. Burrows at Gamil Beach, Port Said, contained fish 
and a crab. Hassan and Hegazy (1968) reported that it attacked 
chicks, ate eggs, and killed and ate the black rat. 

Populations. — Recent observers (Hoogstraal, 1963) suggest that 
R. norvegicus is not as abundant as R. rattus. Earlier observations 
(Bonhote, 1910) indicated that the former was gradually replacing 
the latter. Bonhote also maintained that R. norvegicus had driven 
Arvicanthis niloticus out of Giza Zoological Gardens. 

Associates.— Rattus norvegicus, R. rattus, Mus musculus, and 
Acomys cahirinus are all commensal with man, but their relation- 
ships to one another in buildings are not clearly known. In nature, 
there is a certain amount of habitat sharing between all these 
species along with A. niloticus. 

Reproduction. —The only record we have from Egypt is six fetuses 
from a female caught in February. 

Sex ratio.— \ museum sample of 20 contained seven (35 per cent) 
males and 13 females. 

Genus Mus Linnaeus, 1758 

Small murids with soft pelage, grayish to brownish dorsally. Tail 
usually slightly longer than head and body, indistinctly bicolored, 
annulations almost concealed by hair. 

Skull fragile, rounded; rostrum short, ridges weakly developed. 
Interparietal ligulate in outline. Zygomatic plate with prominent 
masseteric tuberosity. Postpalatal margin posterior to level of m\ 



274 FIELDIANA: ZOOLOGY 

Upper incisor with prominent subapical notch. M' three-rooted, 
crown longer than m^ and m"* combined. 

Mu8 musculus Linnaeus. 1758 

Mus musculus Linnaeus. 1758, Syst. Nat., 10th ed., p. 62. 

Type locality.— Sweden: Upsala. 
General distribution.— Cosmopolitan. 
Subspecies in Egypt — 

Mus musculus praetextus (Brants, 1827) 

Mus praetextus Brants. 1827, Het geslacht der Muizen, p. 125. 

Type locality. -Syria. 

Common names.— House Mouse, Far, Sisi. 

Distribution in Egypt — Figure 86. Northeastern and Western 
Sinai Peninsula, Suez Canal area. Gulf of Suez and Red Sea Coast 
south to Mersa el Alem, Nile Delta and Vgilley, Western Mediterra- 
nean Coastal Desert, oases of the Western Desert. 

Diagnosis.— Small murid with dorsum gray or brownish, venter 
white to buffy. Tail usually slightly longer than head and body, 
indistinctly bicolored. 

Skull fragile, rostrum relatively short, cranium rounded, ridges 
weakly developed. Interparietal broad, ligulate in outline. Upper 
incisor with prominent subapical notch. M' as long as m^ and m^ 
combined. 

Adult head and body length average 84 mm.; tail 84 mm., 100 per 
cent of head and body length; foot 19 mm.; ear 14 mm.; 
occipitonasal length 22.1 mm.; weight 15.0 gm. 

External characters. — Pelage soft. Dorsal color varying from gray 
and tawny to light and dark brown. Side with or without narrow, 
clear tawny border. Belly white to buffy. Dorsum darker than side 
due to greater number of black hairs and black-tipped hairs. Dorsal 
and side hairs with dark gray base. Belly hairs with or without gray 
base. General color determined by subterminal bands of pale gray, 
tawny, or cinnamon on back and side. Hair of ear, suborbital spot, 
and postauricular patch slightly paler than color hairs. Mystacial, 
suborbital, and subauricular areas color of side. Tail indistinctly 
bicolored, brownish above, whitish below. 

Setzer (1952, p. 362) recognized three main color phases in Egyp- 



OSBORN & HELMY: MAMMALS OF EGYPT 



275 



,. 25* 26- 27' 28* 2 9* 30* 3l' 32* 3 3* 34* 35* 36* 37 




Fig. 86. Collection localities of Mus musculus praetextus. 

tian Mus musculus. A fourth from Alexandria he referred to as a 
light phase that did not correspond to "any of the named kinds sup- 
posed to be in the area." The four color phases are listed in Table 33 
with Setzer's color description (1952, p. 362) in parentheses. 

Cranial characters.— Figure 87. Skull fragile. Rostrum relatively 
short. Braincase rounded. Supraorbital and tempoparietal suture 
turning abruptly downward and caudad above base of zygomatic 
process of squamosal; interparietal broad, ligulate in outline. Lamb- 
doidal and median supraoccipital ridges not prominent. Posterior 
nasal margin truncate or bluntly rounded. Zygomatic plate small, 
sharply rounded above; bearing a conspicuous masseteric tuberosi- 
ty on lower border. Incisive foramen long and extending to level of 
medial root of m'. Palatine foramen minute. Parapterygoid fossa 
long, broad, and shallow. Postpalatal margin posterior to level of 



276 



FIELDIANA: ZOOLOGY 



Table 33. Color phases of Mus musculus praetextus. 





Intermixture 


» 




Dorsal color/distribution 


of black 


Side and flank 


Belly hairs 


Gray (pallid neutral gray) 


strong 


pale gray 


white to base; 


W. Coastal Desert 






white, gray base 


Tawny (light phase) 


very light 


pale tawny 


white to base 


Individual variant 








Light brown (cinnamon brown) 


light 


buffy to brown 


white to base; 


Common 






white or buffy, 
gray base 


Dark brown (mummy brown) 


strong 


buffy to brown 


buffy, gray base 


Common 









m^. Tympanic bulla moderately inflated, mastoid bulla slightly 
inflated. Occipital condyle not protruding beyond level of supraoc- 
cipital. Mandible with well-developed coronoid and alveolar 
processes. 

Teeth.— Figure 79. Upper incisor compressed, opisthodont, 
anterior surface smooth, cutting edge with prominent subapical 
notch cut into it by action of the lower incisor. M' with three roots. 
Crown of m' longer than m^, m^ together. First lamina of m' 
distorted by backward displacement of outer cusp. M*^ lacking addi- 
tional anteriolateral cusp. M^ with two laminae. 

Measurements.— Tables 34, 35. Male and female measurements 
subequal. Means (and ranges) of occipitonasal length (in 
millimeters) of 10 adult males and 10 adult females are 21.9 (20.6 to 
23.5) and 22.0 (21.1 to 22.9), respectively. 

Age determination.— Adults have cusps of molars worn to 
laminate pattern, cranial sutures closed. 

Variation.— Color phases, variation in size and in proportion of 
tail to head and body length are summarized in Tables 33 and 35. 

Typical Mus m. praetextus has a light or dark brown back; white 
or buffy belly; belly hairs usually with gray bases; and commonly a 
narrow tawny border on the side. It is found in northeastern Sinai 
Peninsula; Suez Canal zone; Gulf of Suez and Red Sea coasts south 
to Mersa el Alem; Nile Delta and Valley; oases of Siwa, Qara, 
Bahariya, Farafara, Dakhla, and Kharga; and the area around 
Salum. Both dark and light phases occur in these areas. Darkest 
individuals are usually found in the more mesic habitats. Tawny 




Fig. 87. Skull of Mus musculus praetextus. 



277 



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OSBORN&HELMY: MAMMALS OF EGYPT 281 

individuals occur occasionally in the Nile Delta. One was found on 
the Gulf of Suez coast. 

Two populations distinct from all other praetextus are: (1) from 
the Western Mediterranean Coastal Desert (Bahig, Abu Mena, Ras 
el Hekma, and Mersa Matruh) and have dorsum pale gray or brown 
and belly white (table 33) and (2) from El Maghra with dorsum light 
brown and belly dark buffy brown (table 33). Two specimens from 
Wadi Muwellih have equally dark bellies. 

In Libya, coastal populations are dark, and inland populations 
pale (Ranck, 1968). The reverse occurs in the Western Desert of 
Egypt, excepting specimens from Salum, which may represent the 
eastern limit of the dark coastal types mentioned by Ranck. 

Adult measurements (table 34) indicate that the Nile Valley and 
Delta populations average slightly smaller than those from other 
parts of Egypt. Data on all ages (table 35) show considerable varia- 
tion in tail length, but no sharp differences between samples of feral 
and commensal mice. In fact, samples from Egypt have no set of 
characters that can be considered typically commensal or feral. 

Setzer (1957c) and Ranck (1968) maintained that, in Libya, M. 
musculus segregated into commensal and feral types, although not 
with all the distinctive features set forth by Schwarz and Schwarz 
(1943). 

Comparison.— Mus musculus can be distinguished from most 
other Egyptian mice by small size; lack of contrasting head, side, 
and rump markings; tail lacking a brush (in comparison with mice 
other than murids); interparietal ligulate in outline; subapical notch 
on upper incisor; and prominent masseteric tuberosity on lower 
border of zygomatic plate. 

Remarks.— Hhere has no doubt been continuous transportation of 
house mice up and down the Nile Valley and Delta and between the 
Nile and the Suez Canal zone and the more accessible oases for a 
long period of time; the result is a rather uniform population in 
these areas. Mus was probably originally transported by camel 
caravans to the oases. Oases with continued contact with the Nile 
by modern transportation have house mouse populations most like 
the Nile Valley populations. In less accessible areas such as El 
Maghra, populations of house mice have followed the trend of 
natural subspeciation and, under mesic conditions, become dark. 
House mice of the relatively xeric coastal desert of pale soils dif- 
ferentiated into a lighter form. 



282 FIELDIANA: ZOOLOGY 

Samples from Gulf of Suez and Red Sea coasts are too small for 
satisfactory analysis. >. 

Specimens examined.— Total 392. 

SINAI: EI Arish (18). Ayun Musa (1). EI Tor (2). 

PORT SAID: El Gamil beach (5). 

ISMAILIA: Ismailia (5). EI Ballah (3). 

SUEZ: Suez (1). Ain Sukhna (5). Wadi Abu Seyala (2). Wadi Dom (1). Wadi Abu 
Sanduq 4 km. S (1), Wadi Yesein (1). 

RED SEA: Safaga 6.4 km. S (3). Hurghada (3). Mersa el Alem (3). 

SHARQIYA: Tel Abu Ekaim near El Salhiya (1). 

DAQAHLIYA: Simbillawein (1). 

GHARBIYA: El Mehalla el Kubra (1). 

DAMIETTA: Shatt Gheit el Nasara (2). Shatt el Mel (3). Kafr el Battikh (1). 

KAFR EL SHEIKH: Baltim (22). El Burg (6). El Hamul (11). BuruUus (6). Kafr el 
Sheikh (2). El Hamra (5). Tel Khamis (9). 

BEHEIRA: Kom Hamada (1). Wadi el Natroun (9). 

ALEXANDRIA: Alexandria harbor area (1). 

MINUFIYA: Birket el Sabh (1). 

GIZA: Abu Rawash (26); Giza (4); Giza Zoological Gardens (2): Awlad Hawra (1): 
Bahariya Oasis. Bawiti (4). Bir Qasr (9). 

CAIRO: Abassia Fever Hospital (3). Maadi (1). Helwan (6). 

EL FAIYUM: Abu Gandir (1). Ezbet Ayub Ali (1). Seila (1), Minshat TanUwi (1). 
Shooting Club (1). Lake Qarun SW end (1). Wadi Muwellih (2). 

ASYUT: Asyut (2). 

QENA: Dishna (1). Isna (1). El Deir (1). 

ASWAN: Kom Ombo. El Biyara (8); Aswan (1); West Aswan (3); Aswan Dam 
Hospital (2); Amada Temple (1); Armina West (4); Armina Temple 10 km. NW (1): EI 
Siboua Temple (1); Gebel Adda (2); Qustul West (1). 

MATRUH: Bahig (8). 10 km. S (1); Abu Mena 3.2 km. E (4). 6.4 km. E (1): El Ham- 
mam (3): Ras el Hekma (24); Mersa Matruh (4). 10 km. E (1); El Maghra (13): Abu 
Dweiss (2); Siwa (4); Siwa Oasis, Abu Shuruf (1); Bahrein (16); Nuweimisa (3); Qara 
(10); Salum (21). 1.6 km. E (1); Bir el Qattara (3). El Zeitun (1). 

EL WADI EL GEDEED: Farafara Oasis (1); Dakhla Oasis. Mut (17). 4.8 km. N 
(3). 5 km. N (1). 10 km. N (6); Bir el Nokta (4); Gharb el Mawhoub (3); Abu Minqar (1); 
Bir No. 2(1); Kharga Oasis. El Gezira (5). 

Published records. — Records are from Anderson (1902), Schwann 
(1905). Bonhote (1909. 1912). De Winton (1903). Flower (1932). 
Ellerman (1948), Hayman (1948), Setzer (1952), and Wassif (1953b, 
1960b). 

SINAI: EI Arish, Ayun Musa. 
PORT SAID: Port Said. 
SUEZ: Suez. 



OSBORN&HELMY: MAMMALS OF EGYPT 283 

RED SEA: Shad wan Island. 

DAQAHLIYA: Simbillawein 8 km. W. 

KAFR EL SHEIKH: Baltim. 

ALEXANDRIA: Alexandria, Dikheila airfield 0.8 km. W. 

BEHEIRA: Idku; Wadi el Natroun; Wadi el Natroun, Zaghig. 

MINUFIYA: Nadir. 

GIZA: Kafr Teharmes. El Kunaiyisa, Talbia, Giza Zoological Gardens. 

CAIRO: Cairo. Heliopolis, Maadi. 

EL FAIYUM: Faiyum. 

ASWAN: Aswan. 

MATRUH: Siwa Oasis. 

EL WADI EL GEDEED: Kharga Oasis. Kharga. 

Sudan. NORTHERN: Wadi Haifa. 

Collection.— Trapped alive and dug from burrows. Disturbance at 
one or more openings of a communal burrow can cause house mice to 
leave by another entrance one by one where they can be picked up 
by hand. 

//a 6i tats.— Throughout Egypt, M. musculus has been collected 
from houses and tents. It also inhabits grain stores and is common 
in gardens and fields. In nature, the habitats of M us are as follows: 

Gulf of Suez: Beach sand at Ain Sukhna under dense growth of 
Salicornia fruticosa and in rocky wadi mouths near the sea. 

Port Said, Gamil Beach: Coarse sand with scattered Suaeda salsa. 

Nile Delta and Nile Valley: Sand flats at El Borg near Baltim 
covered with Alhagi mannifera and Carthamus glaucus. 

In salty waste land at El Hamra and Tel Khamis, house mouse 
trails ran between scattered Suaeda pruinosa and burrows were 
located under this shrub. Other halophytic Chenopodiaceae in the 
latter area were Halocnemon strobilaceum and Arthrocnemon 
glaucum, together with Mesembryanthemum nodiflorum. These 
areas of black salty silt were identical with Psammomys obesus 
habitat in the western part of the Delta near Hafs, but no mammals 
other than Mas were collected. 

Near Abu Rawash, house mice were dug from burrows in grassy 
hummocks in a meadow. 

Bauer (1963) collected Mus from canal banks and beside the Nile 
in Upper Egypt. 

Western Mediterranean Coastal Desert: Palm and olive groves, 
edges of barley fields in Anabasis articulata (fig. 8) in vicinities of 



284 FIELDIANA. ZOOLOGY 

Bahig and Abu Mena, respectively. On Ras el Hekma. Mus was col- 
lected at cliff bases and in sand dunes near the sea, and in rocky ter- 
rain and salt marshes. 

Near Salum at Bir Qattara, house mice were taken under dense 
vegetation and among rocks at a cliff base spring near the sea. 

Western Desert Oases: Mus musculus and no other rodent, with 
the exception of Arvicanthus niloticus from Maghra, inhabits the 
palm-reed-rush {Phoenix-Phragmites-^J uncus) community of salt- 
encrusted lake shores and salty sand (fig. 22). In Farafara Oasis, 
Mus was also trapped beneath wild date palms in dry, windblown 
sand. One specimen was dug from a shallow burrow under a carton 
in dry sand south of the coastal vegetation. The area had been a Pan 
American Oil Company campsite two weeks previously. 

In Libya, Ranck (1968) reported Mus from elevated patches of 
Phragmites in areas of open water; sedges and other mesophytic 
plants encircling saline lakes; and mesic pockets in the coastal 
escarpment. 

Happold (1967bd) found Mus in houses and stores in Khartoum in 
winter months. It was not found in villages away from the Nile 
Valley. 

Be/iauior.— Communal. Nocturnal in nature, but commensal in- 
dividuals appear to be less so. Although easily handled, Mus bites 
readily. 

Burro m;s.— Shallow, usually under shrubs or in grass or rush- 
covered hummocks. Several short tunnels lead to a large nest 
chamber. 

Associates.— Mus shares habitats with nearly every other Egyp- 
tian rodent except those confined to dry, barren desert. Relation- 
ships between species are not known, although evidence from collec- 
tions from houses and buildings in the Nile Valley and Delta 
indicates that Acomys cahirinus becomes the dominant species, 
forcing Mus to be feral (see commensalism notes under y4. cahirinus). 

Food— Various crop plants and stored agricultural products are 
eaten by Mus. Presence of house mice in sebakhas in the Western 
Desert was discovered from cuttings of Juncus sp. Habitat data 
indicate that Mus, like Psammomys obesus, can survive on 
halophytic Chenopodiaceae. 



OSBORN & HELMY: MAMMALS OF EGYPT 285 

Reproduction.— No evidence of a breeding season. Gravid females 
and young in nests were found the year around. Average and range 
of embryos and fetuses from six females were six (three to seven). 
Communal nests contained 10, 18, and 19 young. 

Sex ratio. — In a sample of 59 museum specimens of Mus 
musculus, males numbered 26 (44 per cent) and females 33. 

Economic importance.— Hassan and Hegazy (1968) mentioned 
that Mus ate and lived in grain stores. 

Genus Acomys I. Geoffroy St. Hilaire, 1838 

Small- to medium-size murids with prominent pigmented ears. 
Dorsal pelage spinous. Spines V-shaped in cross section. Tail with 
broad conspicuous annulations alternating with bristles. Skull 
strongly built; braincase broad, conspicuously convex dorsally. 
Supraorbital and tempoparietal ridges well developed, the latter low 
on the braincase and curving outward. Median supraoccipital ridge 
prominent. Interparietal very large, semicirculeir. Tempoparietal 
suture following cranial ridge. Posterior nasal margin divided. 
Zygomatic plate relatively large, gradually rounded above, 
masseteric tuberosity on lower border inconspicuous. Incisive 
foramen long and extending to level of medial root of m\ Palatine 
foramen minute. Parapterygoid fossa very long, broad, and shallow. 
Palatines forming a shelf closing the mesopterygoid fossa to the 
level of or anterior to the level of the basisphenoid-presphenoid 
suture. Tympanic bulla moderately inflated, mastoid ossified. 
Occipital condyle not protruding beyond level of supraoccipital. 
Mandible with very small coronoid and alveolar processes. 

Upper incisor compressed, opisthodont, anterior surface smooth, 
cutting edge normal. M^ three-rooted; crown not longer than m^, m^ 
together. First lamina of m' somewhat distorted backward as in 
Mus. M^ with an additional but transient anterolateral cusp as in 
Rattus rattus. M^ with two laminae as in Mus. 

Key TO Egyptian Species of y4comys 

1. Dorsal color reddish; palm, sole, and tail black. Tail shorter than head and body. 
Palate without a median keel. Apex of mesopterygoid shelf anterior to basi- 
sphenoid-presphenoid suture russatus, p. 286. 

2. Dorsal color brownish, blackish, or slate, not reddish. Palm, sole, and tail not 
black. Tail usually longer than head and body. Palate with a median keel. Apex of 

mesopterygoid shelf at level of basisphenoid-presphenoid suture 

cahirinus, p. 293. 



286 FIELDIANA: ZOOLOGY 

Acomys russatus (Wagner, 1840) 

Mus russatus Wagner. 1840. Abh. Bayer Akad. Wies. Math.-Naturwiss. KL. 3. 
pi. 3. Hg. 2. p. 195. 

Type locality. — Egypt. SINAI: probably Nohel. 

General distribution. —Saudi Arabia, Israel, Sinai Peninsula, 
Egypt. 

Common name.— Golden Spiny Mouse. 

Distribution of subspecies in Egypt. — Figure 88. Acomys 
russatus russatus: southern part of Sinai Peninsula; Acomys 
russatus aegyptiacus: northern part of Eastern Desert of Egypt. 

Diagnosis. — Dorsum reddish orange; venter pale yellowish white; 
palm, sole, ear, and tail black. Spinous pelage on head, back, side, 
and rump. Tail shorter than head and body, not bicolored; annula- 
tions fairly conspicuous, alternating with white bristles. 

Skull with apex of mesopterygoid shelf anterior to level of 
basisphenoid-presphenoid suture. Palate without median keel. 
Zygomatic arch noticeably thickened anteriorly. 

Adult head and body length average 111 mm.; tail 70 mm., 62 per 
cent of head and body length; hind foot 20 mm.; ear 19 mm.; oc- 
cipitonasal length 28.9 mm.; weight 36.0 gm. 

External characters. — Figure 89. Pelage spiny on head, back, side, 
and rump. Dorsal color reddish to reddish orange. Side and rump 
yellowish to yellowish brown. Belly and underparts whitish to 
yellowish, darkened by the grayish bases of the hairs. All dorsal 
hairs and spines with a minute black tip, broad orangish to 
yellowish subterminal band, and pale gray base. Legs grayish. 
Palms, soles, ear, and tail black. Tail not bicolored. 

Ear covered with white and buffy hairs. Mystacial area dark due 
to white hairs not quite concealing the black skin. Pre- and subor- 
bital region color of side. Suborbital spot small, white, and con- 
spicuous. Ear with white basal and posterior patches. 

Bacw/um. — Baculum terminates in a completely ossified trifid, 
lateral ossicles equal in size with medial (Atallah. 1967). 

Cranial characters.— See Figure 91 of Acomys cahirinus skull and 

Opposite: 

Fi(i 88. Collection localities oi Acomys russatus russatus (circles) and A. r. aegyp- 
tiacus (dots). 




287 



FIELDIANA: ZOOLOGY 




-% 




Fig. 89. Live specimen of Acomys russatus. 



description under genus. Braincase broad; interparietal very large, 
semicircular; supraorbital and parietal ridges well developed, the 
latter curving outward. Apex of mesopterygoid shelf anterior to 
level of basisphenoid-presphenoid suture. Palate lacking a median 
keel. Zygomatic arch noticeably thickened anteriorly, compared 
with A. cahirinus. 

Teeth.— Molars are identical to A. cahirinus. Upper incisor 
opisthodont, with anterior surface smooth. 

Measurements.— Table 36. Tail shorter than head and body 
length. Male and female dimensions equal. 

Age determination. — Adult specimens have well-developed 
cranial ridges, basioccipital-basisphenoid suture closed, and molar 
cusps showing some amount of wear. 

Comparisons.— Acomys russatus and A. cahirinus differ from all 
other Egyptian rodents in having spiny dorsal pelage. Acomys 
russatus differs from A. cahirinus in having dorsal color reddish, 
rather than blackish, slate, or brownish; tail much shorter than head 



OSBORN & HELMY: MAMMALS OF EGYPT 289 

Table 36. — Means (and ranges) of measurements, ratios, and weight of adult 
Acomys russatus. 





Sinai Peninsula 


Eastern Desert 




A. r. russatus 


A. r. aegyptiacui 


HBL 


113.0(106-122) 15 


109.2(90-117)6 


TL 


74.8 (68-81) 8 


64.4 (56-75) 5 


TL/HBL% 


67.0 (60.5-76.4) 8 


58.9 (50.0-70.5) 5 


FL 


20.4 (20-22) 16 


19.2(19-20)6 


EL 


20.3 (19-22) 16 


19.3(16-20)6 


Wt 





37.0 (24.0-53.2) 3 


ONL 


28.7(27.6-31.1) 11 


29.2 (27.8-31.1) 5 


ZW 


14.3 (13.9-14.8) 14 


14.4 (13.9-14.8) 4 


low 


4.5 ( 4.3- 4.9) 18 


4.6 ( 4.4- 4.9)5 


BCW 


13.1 (12.4-13.7) 17 


13.1 (12.4-13.6) 4 


NL 


11.2(10.0-12.0) 11 


11.5(11.0-12.0)5 


IPL 


6.2 ( 6.4- 7.4) 18 


6.9 ( 6.6- 7.4) 4 


AL 


5.0 ( 4.8- 5.2) 18 


5.2 ( 5.0- 5.2) 5 


SH 


10.4 ( 9.9-10.9) 16 


10.4 ( 9.9-10.9) 4 



and body length (tables 36, 37); palate without a median keel; apex 
of mesopterygoid shelf anterior to basisphenoid-presphenoid suture; 
and zygomatic arch thickened anteriorly. 

Collection.— Golden spiny mice will enter live traps. None have 
been dug from burrows. 

Habitats.— Acomys russatus is known only from rocky hillsides, 
cliffs, and boulder-strewn canyons. 

Behavior.— This species has been seen active during morning and 
afternoon in southern Sinai (Wassif and Hoogstraal, 1953). In some 
areas where it shares the habitat with A. cahirinus, it is diurnal. 

Acomys russatus, like other spiny mice, is agile and difficult to 
handle, but not as aggressive as A. cahirinus. 

Commensalism.—We do not know if ^. russatus enters buildings. 

Associates. — In rocky habitat in the Sinai Peninsula, A. russatus 
is found in the same habitat as A. cahirinus, Dipodillus dasyurus, 
Sekeetamys calurus, and Eliomys quercinus. In the Eastern Desert, 
it occurs with the same species except for E. quercinus. 

Populations.— Acomys russatus is rare in nature and in museum 
collections compared with A. cahirinus. It also appears to be discon- 
tinuously distributed, whereas A. cahirinus shows continuous 
distribution, at least in mountainous areas, in the Eastern Desert. 

Reproduction.— The only records from Egypt are two young bom 
in May and three fetuses collected in June. 



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292 FIELDIANA: ZOOLOGY 

Sex ratio. — In a museum sample of 31, there were 12 (38 per cent) 
males and 19 females. 

/JemarAs.— Shkolnik and Borut (1969) showed that A. russatus 
was better adapted for a diurnal existence than A. cahirinus because 
of its ability to withstand much higher ambient temperatures, to 
utilize salty water, and to extract water from halophytic plants. 
Acomys cahirinus, however, can withstand much colder situations 
which accounts for its existing at much higher elevations than A. 
russatus. 

Spiny mice were found to possess a "kidney with an outstanding- 
ly efficient mechanism for urine concentration" (Shkolnik and 
Borut, 1969, p. 254). Both species were thought to obtain additional 
water by eating desert snails, and A. russatus was often observed 
eating Atriplex halimus, Hammada scoparia, and Anabasis 
articulator 

See further notes on food under A. cahirinus. 

Kky to Egyptian Subspkciks ok Acomys russatus 

1. Dorsum dark reddish orange. Tail average 20 per cent longer. Hind foot slightly 
longer. (Southern part of Sinai Peninsula) russatus, p. 292. 

2. Dorsum pale reddish orange. Tail average 20 per cent shorter. Hind foot slightly 
shorter. (Northern part of Eastern Desert) aegyptiacus, p. 293. 

Acomys russatus russatus (Wagner, 1840) 

Type locality.— Egypt. SINAI. 

Distribution in Egypt— Figure 88. Southern part of Sinai 
Peninsula. 

External characters. —See species description. Acomys r. russatus 
is slightly darker or more reddish than A. r. aegyptiacus. 

Cranial characters. —See species description. There are no cranial 
differences between the Egyptian subspecies of A. russatus. 

Measurements.— Table 36. Tail length is about 20 per cent, and 
hind foot length slightly longer in A. r. russatus than in A. r. 
aegyptiacus. 

Specimens examined.— Total 21. 

SINAI: Wadi el Sheikh (9). St. Catherine Monastery area (6), Wadi el Arbaein (1), 
Feiran Oasis (2). Tor (3). 

Published records. — Records are from Flower (1932), Wassif and 
Hoogstraal (1953). and Setzer (1959c). 



OSBORN&HELMY: MAMMALS OF EGYPT 293 

SINAI: Feiran Oasis, St. Catherine Monastery area, and inland from Tor. 
Acomys russatus aegyptiacus Bonhote, 1912 

Acomys russatus aegyptiacus Bonhote, 1912, Abstr. Proc. Zool. Soc., London, 
No. 103, p. 3; Proc. Zool. Soc.. London, p. 230. 

Type /oca/ity.— Egypt. CAIRO: Helwan, Wadi Hof. 

Distribution in Egypt— Figure 88. Northern part of Eastern 
Desert. 

External characters. —See species description. Acomys r. aegyp- 
tiacus is slightly paler or more orangish than A. r. russatus. 

Cranial characters.— See species description. 

Measurements.— Table 36. Differences in dimensions of the two 
subspecies, aegyptiacus and russatus, are discussed under the 
latter. 

Remarks.— Differences between the subspecies are very slight, 
and further sampling may indicate that no real differences exist. 

Specimens examined.— Total 15. 

SUEZ: Wadi Abu Seyala (2), Bir Abu Seyala (1). Wadi Qiseib (6), Bir Qiseib (3). 
CAIRO: Gebel Mokattam (1), Wadi Hof (Type). 
RED SEA: Wadi Atalla mouth (1). 

Published records.— Records are from Bonhote (1909, 1912) and 
Setzer (1959e). 

CAIRO: Gebel Mokattam. Wadi Hof. 
SUEZ: Wadi Sayal (Wadi Abu Seyala). 

Acomys cahirinus (Desmarest, 1819) 

Mus cahirinus Desmarest, 1819, Nouveau Dictionnaire Hist. Nat., Vol. 29, p. 70. 

Type locality.— Egypt. CAIRO: Probably Cairo. 

Common names. — Egyptian Spiny Mouse, Abu Shoaka. 

General distribution.— V/estern Sind, southern Iran, southern 
Asia Minor, Arabian Peninsula, Jordan, Israel, islands of Cyprus 
and Crete, Sinai Peninsula, Egypt, Libya, Mauritania, Morocco; 
Sudan south through Ethiopia, Somalia, Kenya, and Tanzania to 
Rhodesia; and west to Nigeria and southern Algeria. 

Distribution of subspecies in Egypt— Figure 90. Acomys 
cahirinus cahirinus: Nile Delta and Nile Valley south to Aswan, 
Suez Canal area, Bahariya Oasis; Acomys cahirinus dimidiatus: 



294 FIELDIANA: ZOOLOGY 

„ 25' 26' 27' 26* 2 9' 30* 3l' 32' 3 3* 34* 35* 36* 37* 




Fici. 90. Collection localities of Acomys cahirinus cahirinus (circles), A. c. 
megalodus (dots). A. c. dimidiatus (open squares). A. c. hunteri (solid squares), A. c. 
helmyi (triangles), and A. c. viator (hexagons). 

southern part of Sinai Peninsula; Acomys cahirinus megalodus: 
northern part of Eastern Desert south to Wadi Araba; ^4 corny s 
cahirinus hunteri: Eastern Desert south of Wadi Araba, Nile Valley 
south of Aswan; Acomys cahirinus helmyi: Farafara, Dakhla, and 
Kharga Oases; Acomys cahirinus viator: Gebel Uweinat and 
possibly Gilf el Kebir. 

Diagnosis.— Color ranging from brownish cinnamon dorsally, 
with white belly and feet, to overall slate with white toes, Spinous 
pelage extends from behind shoulders onto rump. Tail usually 
longer than head and body length, bicolored or not, annulations 
broad, alternating with whitish or brownish bristles. 

Skull with apex of mesopterygoid shelf at level of basisphenoid- 
presphenoid suture. Palate with median keel. Zygomatic arch un- 
modified. 



OSBORN&HELMY: MAMMALS OF EGYPT 295 

Adult head and body length average 108 mm.; tail 109 mm., 104 
per cent of head and body length; foot 20 mm.; ear 21 mm.; oc- 
cipitonasal length 30.1 mm.; weight 41.6 gm. 

External characters. — Pelage spiny from behind shoulder onto 
rump, but not on side. Color of Nile Valley and Delta populations: 
overall slate with white feet; no suborbital, sub- or postauricular 
markings; tail not bicolored. Color of desert populations: dorsum 
pale to dark brownish; side, rump, and limbs pale to dark cinnamon; 
belly, underparts, and feet white. Dorsal hairs and spines with black 
tip, cinnamon subterminal band, and pale gray base in shoulder 
region; white base in lumbar and sacral region. Width of tip, subter- 
minal, and basal color bands variable. Ear pigmented, covered with 
whitish hairs. Tail bicolored, brownish above, whitish below. 
Mystacial area partly pigmented. Pre- and suborbital region color of 
side. Suborbital spot small, white, and conspicuous. Ear with white 
basal and posterior patches. 

Bacu/um.— Baculum terminates in a trifid, with middle ossicle 
ossified, lateral ossicles small and cartilaginous (Atallah, 1967). 

Cranial characters. — Figure 91. See description under genus. 
Braincase broad; interparietal very large, semicircular; supraorbital 
and parietal ridges well developed, the latter curving outward. Apex 
of the mesopterygoid shelf at level of the basisphenoid-presphenoid 
suture. Palate with a median keel. Zygomatic arch not thickened 
anteriorly, compared with A. russatus. 

Teeth. — Upper incisor opisthodont, with anterior surface smooth. 
M' not longer than m^ m^ together; m^ with transient anterolateral 
cusp; m^ with two laminae. 

Measurements.— Table 37. Tail usually as long as head and body 
length. Male and female dimensions subequal, as noted by Setzer 
(1959e). 

Age determination. — Adult specimens have well-developed 
cranial ridges, basioccipital-basisphenoid suture closed, teeth with 
cusps showing some amount of wear. 

Variation.— The main color differences between melanistic com- 
mensal subspecies cahirinus and other Egyptian subspecies have 
been dealt with. Acomys c. dimidiatus from Sinai Peninsula is the 
palest of desert subspecies; megalodus of the North Galala Plateau 
is slightly darker, and pigmentation increases through the southern 




Flu. 91. Skull of Acomys cahirinus. 



296 



OSBORN&HELMY: MAMMALS OF EGYPT 297 

part of the range of this subspecies into hunteri. Samples from 
Gebel Mokattam near Cairo and cliffs near Helwan show in- 
tergradation in color with cahirinus and are comparable in dimen- 
sions. Southward along the Nile Valley, intergradation between 
cahirinus and hunteri has been observed. 

In a sample of hunteri from Fawakhir mine area, those from 
buildings approach cahirinus in having side and belly grayish. 
Specimens from surrounding hills became more similar to typical 
hunteri the further they were trapped from dwellings. 

Color of cahirinus specimens from houses in Bahariya Oasis 
ranges from overall slate with white feet of typical commensal, to 
blackish brown dorsally, side brownish or grayish, belly grayish to 
white; similar to a sample of cahirinus from Gebel el Ghigiga in the 
Cairo area. Specimens from houses in Kharga and Dakhla Oases are 
slate color dorsally, pale on the underside, and are probably com- 
mensals of A. c. helmyi. Samples of helmyi from isolated areas in 
Farafara Oasis are about as pale dorsally as megalodus. 

Acomys c. viator from Gebel Uweinat is about the same color as 
hunteri. 

Dimensions vary considerably in A. cahirinus (table 37). 
Specimens with the largest dimensions are of subspecies megalodus 
from Gebel Katamiya in the northern Eastern Desert. In general, 
size diminishes east from there into dimidiatus, west into cahirinus, 
and south into hunteri. Another subspecies of large dimensions is 
helmyi from Farafara Oasis. Acomys c. viator specimens from 
Gebel Uweinat are representative of a race with smaller dimensions 
than Egyptian subspecies. 

Comparisons.— Acomys cahirinus and A. russatus differ from all 
other Egyptian rodents in having spiny dorsal pelage. Comparison 
between these two species is under the latter. 

Remarks. —Setzer (1959) retained A. cahirinus as a monotypic 
species and referred the other named Egyptian forms to A. 
dimidiatus. He argued that cahirinus differed from dimidiatus in 
that the majority were either partly or totally melanistic, had 
smaller dimensions, especially the nasal length, braincase more flat- 
tened, foramen magnum more rounded and extending further dor- 
sally, upper incisor more nearly vertical, and pterygoid fossa longer 
and broader. 

Melanism is prominent in cahirinus, but intergradation occurs 



298 FIELDI AN A: ZOOLOGY 

between Nile Valley and desert forms, as has also been noted by 
Bauer (1963). With regard to measurements, data in Table 37 in- 
dicate considerable geographical variation in dimensions in A. 
cahirinus in Egypt. Acomys c. cahirinus appears to be significantly 
smaller than megalodus, but there are gradations in dimensions be- 
tween all populations of this species. 

Atallah (1967). following Setzer (1959). recognized A. cahirinus 
and A. dimidiatus as distinct species and listed nasal length and 
cranial conformation as the discriminating characters. Incidentally, 
nasal length according to Setzer. is the median length. In Table 37. 
it is the greatest length of the nasals (NL). 

Skull height (SH) in Table 37 gives no indication of any dispropor- 
tionately flattened or rounded braincase. Shape of the foramen 
magnum does not show any consistant variations in shape. When 
compared by camera lucida drawings, upper incisors show no dif- 
ferences in curvature. Neither could any differences be discerned in 
the parapterygoid fossae. Thus, we are convinced that a single 
species, A. cahirinus, occurs in Egypt. 

Happold (1969) concluded that A. cahirinus cineraceus was the 
only subspecies in gebels north of Khartoum and probably A. 
cahirinus was the only species occurring in that region. Intergrada- 
tion between subspecies hunteri and cineraceus has not been 
demonstrated. 

Collection. —Spiny mice readily enter live traps and occasionally 
are dug from burrows. 

Habitats. — In the Nile Delta and Valley, the melanistic A. c. 
cahirinus is considered to be almost completely commensal (Setzer, 
1959e), because it is the commonest mouse in buildings and houses. 
Some have been taken in gardens, date groves, and rocky hills and 
cliffs bordering the Nile Delta and Valley. The subspecies is 
numerous in tombs and temples where, incidentally, melanism 
tends to become diluted. 

Desert subspecies usually inhabit rocky hillsides, cliffs, and 
boulder-strewn canyons (fig. 16), but they may also be found living 
commensally in settlements and native huts (Hoogstraal et al.. 
1957). Concentrations of spiny mice occur in the vicinity of date 
palms in the Eastern and Western Deserts. 

Altitude appears to have no effect on distribution as long as food 
is available (Hoogstraal et al., 1957b). 



OSBORN&HELMY: MAMMALS OF EGYPT 299 

Burrows.— Spiny mice were reported from burrows in sand in the 
southeastern part of Egypt (Hoogstraal et al., 1957b). One was dug 
from a simple burrow about 0.5 m. in length in a hard gravel terrace 
in Wadi Araba. 

Behavior.— This species has been seen active at all hours of the 
day, but mostly in early morning and late afternoon (Hoogstraal et 
al., 1957b). 

Spiny mice are extremely agile, difficult to handle, and bite 
readily. 

Economic importance.— The fact that A. cahirinus lives in close 
association with man leads one to assume that it feeds on crops and 
food stores. The possibility of Acomys as a reservoir for com- 
municable arthropod-borne diseases has been considered, and 
samples from Wadis in the Gulf of Suez area were tested for typhus 
group antibodies. Results did not show clearly the presence of either 
epidemic or murine typhus (Hoogstraal et al., 1967b). 

Diabetes mellitus has been described from A. c. dimidiatus, and 
A. c. cahirinus is under investigation (Strasser, 1968; Strasser and 
Brunk, 1968; Brunk and Strasser, 1969). A review of the literature 
is given by Strasser (1968). 

Food.— Acomys cahirinus utilizes a variety of plants and seeds for 
food. Dates are a staple diet in some areas. Those living in barren 
cliffs lacking plants probably forage for windblown plant remains 
trapped in crevices. We are quite certain that undigested organic 
matter in human feces supplements the diet of A. cahirinus in the 
vicinity of Egyptian Frontier Corps camps and houses along the 
Gulf of Suez coast of Egypt. 

The dried flesh and bone marrow of mummified humans is a 
source of food for A. cahirinus in the tombs of Gebel Drunka south- 
west of Asyut. In tombs of west Aswan, Maser (1966) observed A. 
cahirinus scavenging the feces of fruit bats {Roussettus a. aegyp- 
tiacus). Apparently, the spiny mice living in these tombs do not 
forage outside for food, since we and Maser noted the lack of 
Acomys traffic in and out of the tombs. We noted also that the mice 
were active during the day in the tombs. Maser found only spiders 
in the stomachs of A. cahirinus collected from barren sandstone 
hills beside the Nile River in Egyptian Nubia (unpublished field 
notes of Christopher O. Maser). 

Another interesting find was mice with stomachs filled with green 



300 FIELDIANA: ZOOLOGY 

algae, the only plant food available in the spring of 1965, which ap- 
peared along the shores of Lake Nasser following a rising and lower- 
ing of the water level (Dr. C. A. Reed, personal communication). 

We have tested the proclivity of A. cahirinus for insects such as 
crickets, locusts, butterflies, and moths. Locust legs and wings are 
often found in rock crevices inhabited by A. cahirinus in the Eastern 
Desert. Captive A. cahirinus in Oman (Harrison, 1972) ate berries, 
biscuits, lizards, any large insects, camel spiders, and scorpions. 

The eating of desert snails by A. cahirinus was reported by Flower 
(1932), Bodenheimer (1935), Cloudsley -Thompson and Chadwick 
(1964), and Shkolnik and Borut (1969). Middens of shells oi Eremica 
desertorum are a common feature in canyons of the limestone 
plateaus east of Cairo. Unlike jerboas and jirds, spiny mice are 
unable to subsist on a diet of dry food alone. High evaporative water 
loss is probably compensated for by the water obtained from eating 
land snails (Shkolnik and Borut, 1969). 

In the Eastern Desert of Egypt, snails are found only in the north- 
ern limestone plateau (excepting the Gebel Elba area). In areas lack- 
ing snails, insects, other invertebrates, and possibly small 
vertebrates probably provide the necessary water, or as was men- 
tioned under A. russatus, salty, succulent plants may suffice. 

Acomys cahirinus from sandstone and granite habitats pay no 
attention to proffered snails. Spiny mice from limestone areas, 
however, quickly dispose of snails in the following manner: A snail 
that is sealed onto a branch or another snail is "deftly" pried off 
with the incisors, unattached snails are readily accepted. The seal is 
pierced, and the tongue and lower incisors are worked into the open- 
ing. The liquid behind the seal is lapped, and then the mouse bites 
off bits of the shell until it reaches the snail, which is quickly pulled 
out and devoured. Slime presents no problem to a spiny mouse, for 
second and third snails are accepted as readily as the first. Shells 
gnawed by captive spiny mice are in Figure 92. 

Commensalism. — In Egypt, A. cahirinus lives in houses in desert 
outposts and crowded urban areas. Bodenheimer (1935) observed 
that this spiny mouse invaded houses in Jerusalem at the beginning 
of cold weather. It has been referred to as the "house-mouse" of 
Cairo (HeugUn, 1877) and said to outnumber Mus musculus in 
human dwellings (Bonhote, 1910; Flower, 1932). In marked contrast 
are the statements of Ranck (1968) and Happold (1969) that com- 
mensalism was unknown in A. cahirinus in Libya and Sudan. Such a 



OSBORN & HELMY: MAMMALS OF EGYPT 



301 




Fio. 92. Shells of snails (Eremica desertorum) gnawed by a captive Acomys 
cahirinus. 



reversal of habitat, Happold (1969, p. 141) suggested, "may be a 
result of competition between commensal species, or because of the 
denser human habitation in Egypt, which results in 'jebel-like' 
habitats." This does not explain commensalism in the desert, 
however. 

Egyptian houses are constructed either of brick, stone, and mor- 
tar or of mud and brick and therefore are similar to the natural cliff 
and rock habitat of A. cahirinus. Availability of food and absence of 
predators in desert houses furnishes spiny mice with an ideal 
situation. 

In the Nile Valley and Delta, Acomys may simply be better 
adapted to living in houses than is Mus, which is usually found in 
the more mesic, vegetated habitats. The short, mild Egyptian 
winter probably does not force Mus indoors as is the case in more 
rigorous climates. 

Populations.— The first NAMRU-3 expedition to Gebel Elba 
(Hoogstraal et al., 1957a) in the spring of 1954 reported the follow- 



302 FIELDIANA: ZOOLOGY 

ing catches of i4. c. hunteri (as A. dimidiatus hunteri): In Wadi Kan- 
sisrob. 3 March, 42 spiny mice were taken an 72 live traps, and 30, 
42, 38, 35, 19, and 15 per night were trapped in the general area over 
a period of six days. 

The NAMRU-3 expedition to Gebel Elba in the spring of 1967 
recorded the following catches: In Wadi Kansisrob, 18 February, 
three spiny mice were taken in 50 live traps, and three, five, three, 
one, and seven per night were trapped in other areas around Gebel 
Elba with an average daily setting of 24 traps for five days. The 
total catch of this expedition, including that from Wadi Kansisrob, 
was 22 spiny mice per 170 trap nights. 

We have no idea of what factors might have caused such fluctua- 
tion in the spiny mouse populations of Gebel Elba. In 1967. as in 
1954, there was no indication of drought, and mountain slopes and 
valleys abounded in green vegetation. In fact, this area supports the 
richest flora in Egypt and has a more stable climate than other 
parts of the Eastern Desert. 

One other locality from which we have data on numbers of spiny 
mice is a cliff area around Bir Qiseib, 5.8 km. inside Wadi Qiseib 
from the Gulf of Suez, where we obtained 5 to 12 ^. cahirinus in 
single night catches with 50 live traps. 

Catches of commensal A. cahirinus always contain fewer adult 
animals than collections from nature, and as a result, A. c. cahirinus 
samples in Table 37 are small. 

Associates. — In rocky habitat in the Sinai Peninsula, A. cahirinus 
can be found with Eliomys melanurus, A. russatus, Dipodillus 
dasyurus, and Sekeetamys calurus. In the Eastern Desert, it occurs 
with the same species, except for Eliomys quercinus. In south- 
eastern Egypt, during a period of extremely high population, A. 
cahirinus was collected from sandy habitat with Gerbillus gerbillus 
(Hoogstraal et al., 1957a). In the Western Desert, A. cahirinus 
occurs with Dipodillus campestris patrizii in rocky areas and with 
G. gerbillus in sandy areas. As a commensal, A. cahirinus appears to 
share the same areas with Rattus, and to some extent with Mus 
musculus. 

Reproduction. — Number of young based on fetal scars, embryos, 
and fetal counts from 13 noncommensal females ranged from one to 
six (average, three). The females were collected in the months of 
February. March, April, June, September, and October. 



OSBORN&HELMY: MAMMALS OF EGYPT 303 

Sex ratio.— A sample of 122 museum specimens of ^. cahirinus, 
contained 57 (46 per cent) males and 65 females. 

Notation.— Msiny wild spiny mice lack either part of or the entire 
tail. Not only do the vertebrae separate as readily as those in a 
lizard's tail, but the skin slips off like a loose sheath. Preparing 
study skins of these mice is one of the most frustrating tasks of the 
collector. The tail invariably drops off, and the skin tears at the 
slightest tug. Extreme care must be taken when handling live spiny 
mice to prevent injuries. 

There is a possibility that tail autotomy is an escape mechanism 
(Michener, 1976), but how does one differentiate between tails 
damaged by predators or in fighting? The focus of repeated attacks 
by aggressive captive adult A. cahirinus is the tail (personal obser- 
vations of D. Osborn). 

Kky to EciYPTiAN SuBSPKClKS OF Acomys cahirinus 

1. Dorsum blackish, belly slate. (Nile Valley and Delta) cahirinus, p. 303. 

2. Dorsum brownish, belly white. 

a. Dorsum dark brown. 

i. Head and body length average less than 100 mm. (Southwestern part of 
Western Desert) viator, p. 305. 

ii. Head and body length average more than 100 mm. (Southern part of 
Eastern Desert) hunteri, p. 305. 

b. Dorsum pale brown. 

i. Tail distinctly bicolored. 

(a) Color paler, dimensions average slightly smaller. (Sinai Peninsula) 

dimidiatus, p. 306. 

(b) Color darker, dimensions average slightly larger. (Northern part of 
Eastern Desert) megalodus, p. 307. 

ii. Tail indistinctly bicolored. (Western Desert oases except Bahariya) 

helmyi, p. 308. 

Acomys cahirinus cahirinus (Desmarest, 1819) 

Type /oca/ity.— Egypt. CAIRO: Cairo. 

Distribution in Egypt— Figure 90. Nile Delta, Nile Valley south 
to Aswan, Suez Canal area, Bahariya Oasis. 

External characters.— See species description. Acomys c. cahi- 
rinus specimens from houses and agricultural areas are usually 
melanistic, but those from tombs and cliffs may have the dorsum 
blackish brown, side grayish or brownish, belly pale, and tail indis- 
tinctly or distinctly bicolored. 



304 FIELDIANA: ZCX)LOGY 

Cranial characters. — Figure 91. See species description. 

Measurements.— Table 37. Dimensions of i4. c. cahirinus are com- 
parable with A. c. hunteri from the southern part of the Eastern 
Desert, but significantly smaller on the average than in subspecies 
dimidiatus , megalodus, and helmyi. 

Variation.— Variation in color and intergradation with other sub- 
species have been discussed. An albinistic specimen was collected 
from Abu Rawash, Giza Governorate. 

Comparisons.— Acomys c. cahirinus can usually be distinguished 
from other subspecies on bases of its melanistic or slate color and 
tail not bicolored or indistinctly bicolored. 

Specimens examined.— Total 275. 

SUEZ: Suez (9). 

SHARQIYA: Faqus (1). 

DAQAHLIYA: Aga Minshat el Ikhwa (2). 

GHARBIYA: Tanta (3). Sherbin (3). 

ALEXANDRIA: Port of Alexandria (2). Alexandria (1). 

MINUFIYA: Birket el Sabh (1). 

GIZA: Abu Rawash (35); Tanash (2); Mena Village (3): Giza (18); Gebel al Ghigiga 
(4); Saqiyet Meki (1); Abassia (15); Kafr Taharmes (1); Bahariya Oasis, Mandisha (5); 
Bawiti (4); Ain Marun (1); El Aguz (1). 

CAIRO: Abassia (15), Abassia Fever Hospital area (II). Citadel (3), Gebel Mokat- 
tam (5), Sharabiya (2). Maadi (9). Helwan (13), Bab el Sharia (1). 

ASYUT: Asyut (2). Tombs in Gebel Drunka (10), Asyut Fever Hospital (1), Ben 
Adi (4). 

QENA: Qena (1). Dandara Temple (21). Kaiman el Matana (1). Luxor (5), Valley of 
the Kings (14), Habou city temple (1). 

ASWAN: Muneiha (9); Kom Ombo (1); Aswan Dam Hospital, west bank (5); West 
Aswan tombs (5); West Aswan, Nile bank (16): Aswan, 19.2 km. N (2). 16. km. N (4); 
El Koror area (2). 

Published records.— Records are from Flower (1932), Setzer (1952, 
1959e), and Wassif (1960a). 

SUEZ: Suez. 

SHARQIYA: Faqus. Mina el Qamh. 

DAQAHLIYA: Aga Minshat el Ikhwa. Simbillawein 8 km. W. 

GHARBIYA: Tanta, Sherbin. Mehalla el Kubra. 

ALEXANDRIA: Alexandria. 

BEHEIRA: Wadi el Natroun. 

MINUFIYA: Quesna (Quweisna). 

QALYUBIYA: Basus, Nawa. 



OSBORN&HELMY: MAMMALS OF EGYPT 305 

GIZA: Giza; Abu Rawash; Kafr Taharmes; Saqyet Meki; Tanash; Bahariya Oasis, 
Bawiti and Mandisha. 

CAIRO: Abassia. Abassia Fever Hospital, Cairo Citadel, Sharabiya, Maadi, Bab 
el Sharia. 

ASYUT: Asyut. 

QENA: Kaiman el Matana. 

Acomys cahirinus viator (Thomas, 1902) 

Acomys viator Thomas, 1902, Proc. Zool. Soc, London, 2, p. 10. 

Type locality.-Uhysi. TRIPOLITANIA: Socna, Wadi Sultan. 

Distribution in Egypt— Figure 90. Gebel Uweinat and possibly 
Gilf el Kebir. 

External characters.— Dorsum dark brownish; side, rump, and 
outer leg surface dark cinnamon. Belly, underparts, and feet white. 
Tail distinctly bicolored, brownish above, white below. Acomys c. 
viator is similar in color to hunteri. 

Cranial characters.— See species description. 

Measurements.— Table 37. Acomys c. viator is significantly 
smaller in average dimensions than other Egyptian subspecies, 
except for foot length. 

Specimens examined.— Total 18. 

Sudan. NORTHERN: Gebel Uweinat, Karkur Murr (8). 
Libya. CYRENAICA: Cufra Oasis, El Giof (10). 

Acomys cahirinus hunteri (De Winton, 1901) 

Acomys hunteri De Winton, 1901, Nov. Zool., 8, p. 401. 

Type locality. -Sudan. KASSALA: Tokar. 

Distribution in Egypt — Figure 90. Eastern Desert from Wadi 
Araba southward. West side of Nile River south of Aswan. 

External characters.— See species description. Dorsum dark 
brown; side, rump, and outer leg dark cinnamon. Belly, underparts, 
and feet white. Tail bicolored, brownish above whitish below. There 
is a tendency toward melanism in commensals. 

Cranial characters.— See species description. 

Measurements.— Table 37. Acomys c. hunteri has smaller dimen- 
sions than the northern megalodus subspecies. 

Variation.— Color variation in hunteri was discussed above. There 



306 FIELDIANA: ZOOLOGY 

are pale individuals in the northern part of the range and there is 
considerable difference in size between northern and southern popu- 
lations of hunteri (table 37). Intergradation Has been demonstrated 
between subspecies hunteri and megalodus, and hunteri and 
cahirinus. 

Comparisons.— Acomys c. hunteri is generally darker and smaller 
than megalodus, about equal in dimensions to cahirinus, but does 
not show the extreme melanism nor dark blackish back and brown- 
ish or grayish sides of nonmelanistic cahirinus. 

Specimens examined.— Total 171. 

RED SEA: Wadi Araba (4). Ras Zafarana (1). Wadi el Deir (9). Ras Gharib (1). 
Wadi Bali (5). Wadi Fatira near Abu Kharif mined). Wadi Sikait (1), Wadi Ghadir (6). 
Mersa Alem 20 km. SW (1), Fawakhir Mine area (28), Wadi Sukari (4). Bir Abraq (7). 
Wadi Hodein (2). Bir Gumbiet (1), Wadi Gumbiet (1). Qusur el Banat (1). 

SUDAN ADMINISTRATIVE: Gebel Hamra Dom (2). Wadi Kansisrob (7). Bir 
Kansisrob (11), Wadi Akwamtra (16). 

ASYUT: Wadi Asyuti (2). 

ASWAN: Kom Ombo (1); Kom Ombo, El Kagug cave (8); Gebel Magal Gabril area 
(4): Bir Umm Hibal in Wadi Dihmit (1); Wadi AUaqi (2); AUaqi Village 11.2 km. S (1); 
Bir Murra (2); Wadi Nagib (4); Wadi Haimur mine area (2); Bir Haimur (1); Wadi 
Abusku (1); Wadi Quleib (4); Armena East pumping station (4); Gebel Adda (1); 
Ballana East (9): Nag Misaw (4); Kalabasha 4 km. S (1); Seyala West (3); Madiq (3); 
Dakka 2 km. N (1). 

Sudan. NORTHERN: Wadi Haifa (3). 

Published records. — Records are from Flower (1932), Hoogstraal 
et al., (1957b), Setzer (1959), Hoogstraal (1963), and Bauer (1963). 

RED SEA: Wadi Sikait, Wadi Hodein, Wadi Gumbiet. Bir Abraq. 
SUDAN ADMINISTRATIVE: Bir Kansisrob. Wadi Kansisrob. 
ASWAN: Abu Simbel. Wadi Allaqi. 
Sudan. NORTHERN: Wadi Haifa. Khor Musa Pasha. 

Acomys cahirinus dimidiatus (Cretzschmar, 1826) 

Mus dimidiatus, Cretzschmar, 1826, in Riippel, Atlas zu der Raise im nordliche 
Afrika, Saugeth. pi. 13, fig. a, p. 37. 

Type locality. -Egypt. SINAI. 

Distribution in Egypt — Figure 90. Southern part of Sinai Penin- 
sula. 

External characters.— See species description. Dorsum pale 
brownish. Side, rump, and outer leg surface pale cinnamon. Belly, 
underparts, and feet white. Tail bicolored, pale brownish above. 



OSBORN & HELMY: MAMMALS OF EGYPT 307 

white below. Acomys c. dimidiatus is the palest Egyptian sub- 
species. 

Cranial characters. —See species description. 

Measurements.— Table 37. Acomys c. dimidiatus averages slight- 
ly smaller in most dimensions, except ear length, than the largest 
Egyptian subspecies, megalodus. 

Comparison.— Acomys c. dimidiatus is paler and slightly smaller 
than A. c. megalodus from which it is barely distinguishable. From 
other species, except the Western Desert helmyi, dimidiatus can be 
distinguished by much larger average dimensions, especially ear 
length (table 37), and conspicuously paler coloration. See under ^. c. 
helmyi for comparison with that subspecies. 

Intergradation between dimidiatus and megalodus has not been 
demonstrated. 

Specimens examined.— Total 33. 

SINAI: Wadi el Sheikh (20), Feiran Oasis (3), Wadi el Arbaein (3), St. Catherine 
Monastery area (6), Tor (1). 

Published records.— Records are from Wassif and Hoogstraal 
(1954) and Setzer (1959). 

SINAI: St. Catherine Monastery area, Feiran Oasis, Bir Thai. 

Acomys cahirinus megalodus (Setzer, 1959) 

Acomys dimidiatus megalodus Setzer, 1959, J. Egypt. Publ. Health Assn.. 34, No. 
3, p. 98. 

Type locality. -Egypt. SUEZ: Wadi Sayal (Wadi Abu Seyala). 

Distribution in Egypt— Figure 90. Northern part of Eastern 
Desert north of Wadi Araba. 

External characters.— See species description. Dorsum pale to 
dark brownish. Side, rump, and outer leg surfaces pale to dark cin- 
namon. Belly, underparts, and feet white. Tail bicolored, pale 
brownish above, white below. Acomys c. megalodus is slightly 
darker than dimidiatus and considerably paler than darker hunteri. 

Cranial characters.— See species description. 

Measurements.— Table 37. Acomys c. megalodus averages con- 
siderably larger in most measurements than other Egyptian 
subspecies. 

Comparisons.— Acomys c. megalodus differs only slightly in size 



308 FIELDIANA: ZOOLOGY 

and color from dimidiatus, but on the basis of geographical separa- 
tion, these two subspecies are being retained. From typical hunteri 
and cahirinus, A. c. megalodus can be distinguished on the bases of 
larger size (table 37) and paler color. Intergradation between these 
subspecies has been demonstrated. Comparison with helmyi is 
given under that subsi)ecies. 

Specimens examined.— Total 118. 

SUEZ: Gebel Katamiya (15); Wadi Iseili (3|: Ain Sukhna (11). 3.5 km. S (1): Gebel 
Sukhna (2); Wadi Abu Seyala (17. Type); Wadi Nasr (1); Wadi Nak! (5): Wadi Yesein 
(5): Wadi Qiseib (27): Wadi Dom (22); Wadi Abu Sanduq (2). 

CAIRO: Wadi Hof (6). 

Published records. — Records are from Setzer (1959) and Hoog- 
straal(1963). 

SUEZ: Ain Sukhna. Wadi Sayal (Wadi Abu Seyala). 

Acomys cahirinus helmyi ssp. nov. Osborn 

Type. — Field Museum Natural History No. 108279, original No. 
13901 in H. Hoogstraal catalog. Adult female skin and skull. Col- 
lected April 21, 1969, by Ibrahim Helmy. 

Type locality.-Egypt. EL WADI EL GEDEED: Farafara Oasis, 
Ain el Wadi (56 km. NNE of Qasr Farafara). 

Distribution in Egypt— Figure 90. Farafara, Dakhla, and Kharga 
Oases. 

External characters.— Dorsum pale brownish, side, rump, and 
outer leg surface pale cinnamon. Belly, underparts, and feet white. 
Tail not distinctly bicolored, pale brownish above, whitish below. 
Acomys c. helmyi is almost identical in color with megalodus except 
for the obscure bicoloring of the tail. 

Cranial characters. —See species description. 

Measurements.— Table 37. Acomys c. helmyi averages slightly 
less in some measurements than the large pale subspecies megalo- 
dus of the Eastern Desert. Measurements of the type specimen are: 
Head and body length 127 mm.; tail 113 mm., 88 per cent of head 
and body length; hind foot 20 mm.; ear 22 mm.; occipitonasal length 
31.4 mm.; weight 31.4 gm. 

Habitats.— Acomys c. helmyi was trapped at Ain el Wadi in damp 
sand under wild date palms (Phoenix dactylifera). The only other 
vegetation in the area was Tamarix sp. Five kilometers N in Wadi 



OSBORN&HELMY: MAMMALS OF EGYPT 309 

Hennis, A. c. helmyi was trapped in dry sand under wild date palms. 
Two and three specimens of Gerbillus g. gerbillus were trapped in 
the same habitats, respectively. Commensal Acomys from Dakhla 
and Kharga oases are tentatively considered as belonging to this 
subspecies. 

Specimens examined.— Total 31. 

EL WADI EL GEDEED: Farafara Oasis. Ain el Wadi (Type. 14), Wadi Hennis (2); 
Dakhla Oasis. Mut (6). 10 km. S (5): El Kharga. Hibis Temple (3). 

Published records.— Wassif (1960b) referred spiny mice from 
Dakhla Oasis, Mut to ^4. c. cahirinus. 

Genus Nesokia Gray, 1842 

Monotypic genus of large, rat-like rodent with tail considerably 
shorter than head and body length. 

Skull slightly modified for fossorial life. Incisive foramen short, 
palatal foramen minute, palate constricted and deeply grooved, 
occipital region broad and sloping forward slightly. Upper incisor 
proodont. Cheek teeth with thin, transverse ridges. 

Nesokia indica (Gray and Hardwicke, 1832) 

Arvicola indica Gray and Hardwicke, 1832, 111. Ind. Zool.. Vol. 1, pi. XL 

Type locality. — India. 

General distribution. —Southwestern Asia from Chinese Turke- 
stan into Turkey, Syria, Israel, and northern Saudi Arabia; northern 
Egypt. 

Common names.— Bandicoot Rat, Girdi, Abu Emaya, Abu A fan. 

Subspecies in Egypt — 

Nesokia indica suilla (Thomas, 1907) 

Nesokia suilla Thomas, 1907, Ann. Mag. Nat. Hist., (ser. 7). 20, p. 203. 

Type locality. -Egypt. SUEZ: El Shallufa (Shaluf). 

Distribution in Egypt — Figure 93. El Shallufa near Suez, north- 
western margins of the Nile Delta, Wadi el Natroun, Bahariya 
Oasis. 

Diagnosis.— Large, stocky rat with dorsum pale brown, venter 
buffy, feet white, muzzle blunt. Ear large, sparsely haired. Tail con- 
siderably shorter than head and body length, thinly haired. 



310 FIELDIANA: ZOOLOGY 

"•■ "■' ^^' ^7' ^^' ^^' ^°' ^'' ^^' ^3- 34- 35- 36* 37- 




Fu; 93. Collection localities of Nesokia indica suilla and sites of Paleolithic 
remains (X). 

Skull angular, strongly ridged. Tempoparietal ridges curving 
laterally; area between ridges flat. Zygomatic arch thickened anteri- 
orly, bowing laterally. Occipital condyle protruding beyond level of 
supraoccipital. Supraoccipital large, sloping forward. Incisive and 
palatal foramina short. Molars laminate, cusps lacking. 

Adult head and body length average 184 mm.; tail 121 mm., 66 
per cent of head and body length; foot 39 mm.; ear 20 mm.; condylo- 
incisive length 44.2 mm.; weight 244 gm. 

External characters. — F'\g[\re 81. Pelage comparatively soft. Dor- 
sal hairs with pale brown tips, side and venter hairs buffy. All hairs 
with gray base. White patch on throat variable. Feet white. Palm 
and sole naked. Ear relatively long, sparsely haired. Tail thick, 
brownish, sparsely haired, shorter than head and body length. Ex- 
ternal fossorial modifications lacking, except for long claws. 



OSBORN&HELMY: MAMMALS OF EGYPT 311 

Cranial characters. — Figure 94. Skull large, strong, and angular. 
Braincase large, convex, strongly ridged. Rostrum relatively short, 
broad. Nasals spatulate in outline, posterior margin narrow, irregu- 
larly rounded. Zygomatic arch thickened anteriorly, curving lateral- 
ly. Supraorbital and cranial ridges strongly developed, the latter 
high on the braincase and curving laterally. Interparietal relatively 
small, shape varying from broadly triangular to ovoid with suture 
extremely irregular. Lambdoidal crest high and prominent. Supra- 
occipital large and sloping forward slightly. Median supraoccipital 
ridge not prominent. Occipital condyle protruding well beyond level 
of supraoccipital. Zygomatic plate large, extending forward to level 
of premaxillary-maxillary suture. Infraorbital canal large, conspicu- 
ous. Tempoparietal suture turning abruptly downward and caudad 
above base of zygomatic process of temporal. Auditory bulla rela- 
tively small. Incisive foramen actually and relatively short, palatine 
foramen minute. Root of upper incisor forming a hillock behind the 
palatine foramen. Mandible with well-developed coronoid and alve- 
olar processes. Palate constricted and deeply grooved. Postpalatal 
margin about level with m^. Parapterygoid fossa narrow and deep. 
Pterygoid process long and extending below level of tympanic bulla. 

Teeth. — Figure 79. Incisors long. Upper incisor proodont, broad, 
not compressed; anterior surface smooth and flat. Molars lacking 
evidence of cusps. Occlusal surfaces with thin transverse laminae. 
M' with five roots, crown longer than that of m^. 

Measurements.— Table 38. Male and female dimensions subequal. 

Age determination.— Adults have cranial sutures closed and 
laminae of upper molars broadly crescent-shaped or transverse. 

Variation. — Individual variation in presences or absence of white 
throat patch. 

Comparisons.— Nesokia i. suilla appears to differ from N. i. 
bacheri in having slightly smaller dimensions. Nesokia indica dif- 

Table 38. — Means (and ranges) of measurements, ratios, and weight of adult 
Nesokia indica suilla. 



HBL 


183.6 (165-197) 5 


CIL 


44.2 (42.0-46.1) 8 


TL 


121.0(110-134)5 


ZW 


27.7 (26.1-29.3) 7 


TL/HBL% 


66.2 (62.0-69.0) 5 


low 


6.6 ( 6.3- 7.2)8 


FL 


39.0 (36-42) 5 


NL 


15.6 (14.8-16.3) 5 


EL 


20.4 (20-21) 5 


IFL 


6.2 ( 5.7- 6.8) 8 


Wt 


244.0 (205.5-280.0) 3 


AL 


9.7 ( 9.0-10.4)8 







Fk; 94. Skull of Nesokia indica suiUa. 



312 



OSBORN&HELMY: MAMMALS OF EGYPT 313 

fers from other larger murids in having less contrast between color 
of dorsum and venter, entire upper surface of feet white, tail rela- 
tively shorter than head and body length, zygomatic arches flaring, 
incisive foramina relatively short, supraoccipital proportionately 
larger and sloping forward, lambdoidal ridge prominent, incisors 
proodont, and molars lacking cusps. 

Specimens examined.— Total 53. 

SUEZ: El Shallufa (2). 

BEHEIRA: Kom Hamada. Dissht el Ashraf (6|: Kafr el Dawar (1): Kom el Hanash 
(3): Wadi el Natroun (10). 

CAIRO: Cairo (3). 

EL FAIYUM: Faiyum (6); Lake Qarum (6). 1.6 km. N (1); Tamiya (1); Ezbet Ayub 
AU (1). 

GIZA: Bahariya Oasis, Bir Wigaba (1); Bir Qasr (1); Mandisha (8); El Ghaba (3). 

Published records.— Records are from Anderson (1902), Flower 
(1932), Hoogstraal et al. (1955), and Wassif (1960b). 

SUEZ: Shaluf (El Shallufa). 

BEHEIRA: Kom el Hanash. Wadi el Natroun. 

FAIYUM: Lake Qarun; Lake Qarun SE end. 1.6 km. N; Tamiya; Ezbet Ayub Ali. 

GIZA: Bahariya Oasis, Mandisha, El Aguz. 

Paleolithic sites. — References are under Remarks. 

ASWAN: Khor el SU. 

Sudan. NORTHERN: Wadi Haifa. 

Collection.— Dug from burrows. 

Habitats.— Damp soil in cultivated and wasteland (fig. 21), 
borders of saline lakes, and canal banks. 

Burrou;s. — Burrows are usually less than 0.5 m. below the surface 
and, according to Briscoe (1956), consists of a network of corridors 
varying from about 2.5 to 9 m. in length. Hoogstraal (1963) found 
nest chambers to be at a lower level than subsurface tunnels. 

In El Faiyum in late August, 1953. when air temperatures ranged 
from 86° to 106° F. and relative humidity, from 29.7 to 47 per 
cent, temperature and humidity in bandicoot rat burrows varied 
from 84° to 104° F. and 54.5 to 75.0 per cent, respectively 
(Briscoe, 1956). 

Captive behavior.— Extremely aggressive and bites without 
hesitation. One animal which we reared by hand and kept for seven 



314 FIELDIANA: ZOOLOGY 

months never tamed. Lay (1967, p. 190) remarked that, in Iran, "all 
bandicoots . . . captured alive fought fiercely." 

We have not observed this rat outside a burrow, but it may go out 
to obtain food. Farmers have reported damage to young water- 
melons where it was present. 

Food — Fleshy roots of A Ihagi mannifera and Typha elephantina 
are eaten by Nesokia and stored in underground chambers. 
Hoogstraal (1963) remarked that farmers complained of damage to 
corn, barley, and vegetables by this species. In the laboratory, we 
reared Nesokia on a diet of raw carrots. 

Associates.— Nesokia has been collected in the same areas as 
other Murine rats and burrows in dense vegetation occupied by Ger- 
billus pyramidum (fig. 21). Whether any other rodents utilize bur- 
rows of Nesokia is not known. 

/Reproduction.— Flower (1932) reported six litters containing one 
to four young born February to June in Giza Zoological Gardens. 
We collected three naked young from a nest in Bahariya Oasis in 
mid-October. A female from Wadi el Natroun was lactating in 
October. Data from other sources indicate that there is no estab- 
lished breeding season in the species (Lay, 1967). 

iiemar^s. — Populations of A^. indica in Egypt are scattered (fig. 
93) and isolated. Evidence that the former range was greater than it 
is today is indicated by British Museum specimens (4.8.4.1, 4.8.4.2, 
4.8.4.3) collected in 1904 from the vicinity of Cairo, where the 
species no longer exists, and fossil jaws recovered from Upper 
Paleolithic sites of Late Pleistocene in the Nile Valley at Khor el Sil 
near Kom Ombo (P. F. Turnbull, personal communication; Reed et 
al., 1967; Reed and Turnbull, 1969) and Wadi Haifa in northern 
Sudan (Robinson, 1966). 

The problem of determining the origin of relict populations of ban- 
dicoot rats in the oases of Bahariya and Wadi el Natroun is pro- 
vocative, since these depressions are surrounded by barren desert 
and developed partly as a result of deflation by wind (p. 21). Ob- 
viously, N. indica was there before the period of aridity and wind 
action began. 

The migration of Nesokia into Egypt was probably via riverine or 
deltaic habitats across northern Sinai, which are now defunct or in- 
undated. A number of geological facts presented by Abu al-Izz 
(1971) can be used to explain the once widespread distribution (fig. 



OSBORN&HELMY: MAMMALS OF EGYPT 315 

93). During the Miocene, the western end of the old Nile Delta was 
at what is now known as Wadi el Natroun. The eastern end was in 
the area of the Suez Canal. At the end of the Pliocene, the Nile 
Valley to Kom Ombo was an elongated estuary. In the Pleistocene, 
Wadi Tumilat functioned both as a drain and a tributary of the Nile. 
There were, of course, suitable habitats and ample opportunities for 
population expansion during the wetter periods of Egypt's 
geological history. During Oligocene and Miocene, lakes occurred 
between Bahariya depression and Maghagha in the Nile Valley. 
After the Oligocene, the land rose and the lakes shrank. Dessication 
and excavation of the Bahariya depression resumed. In the 
Pleistocene, the Nile River cut through the eastern edge of El 
Faiyum depression filling it with water. Subsequent dry periods 
resulted in isolation. 

Disappearance of Nesokia from the Nile Vfilley in Upper Nubia 
could i>ossibly have been due to down-cutting of the river and 
dessication and disappearance of habitats. In lower Nubia, the rest 
of the Nile Valley and most of the Delta, intensification of 
agriculture and the flood method of irrigation could have accounted 
for its disappearance. 

The Suez population of Nesokia was believed by Aharoni (1932) to 
have come there by ship; a rather fanciful and presumptuous 
conclusion. 

Family 4. Muscardinidae 
Characters are given under the genus. 

Genus Eliomys Wagner, 1840 

Monotypic, polymorphic genus of a squirrel-like rodent. Fur soft, 
dense. Black facial mask prominent and distinctive. Tail bushy, 
almost round, black with tip white and base of short gray hairs. 
Palm and sole bare. 

Skull smooth, round, lacking ridges. Zygomatic plate small, infra- 
orbital canal very large. Incisive foramen relatively long, broaden- 
ing posteriorly. Palatine foramen small, ovoid. Tympanic and 
mastoid bullae conspicuously inflated. Tympanic bulla with three 
prominent partitions. Superior wall of parapterygoid fossa per- 
forated. Angle of lower jaw with single large foramen. Upper incisor 
orthodont, anterior surface smooth. Crowns of cheek teeth squarish, 
concave, and transversely ridged. Dental formula: \, 5, i, i x 2=20. 



316 FIELDIANA: ZOOLOGY 

EUomys quercinus (Linnaeus, 1766) 

Mus quercinus Linnaeus, 1766, Syst. Nat., 12th ed.,^. 84. 

Type locality.— Germany. 

General distribution.— Europe, Western U.S.S.R., Turkey, Syria, 
Israel, northern Saudi Arabia, Sinai Peninsula, northwestern 
Egypt, Libya, Tunisia, Algeria, Morocco, Spanish Sahara, and 
islands in the western Mediterranean Sea. 

Common names.— Garden Dormouse, Abu Khol 

Distribution of subspecies in Egypt— Figure 95. EUomys quer- 
cinus melanurus: Sinai Peninsula; EUomys quercinus cyrenaicus: 
western part of Mediterranean Coastal Desert. 

Diagnosis.— Squirrel-like rodent. Tail black, partly bushy with 
gray base and white tip. Face with black mask from whiskers to 
base of ears. Dorsum brownish, side gray, underparts and feet 
white. 

Skull smooth, auditory bulla greatly inflated, with three con- 
spicuous partitions. Suprameatal triangle large, meatal lip expand- 
ed anteriorly, accessory tympanum absent. Upper incisor smooth. 
Cheek teeth four, crowns squarish, surface concave, transversely 
ridged. 

Adult head and body length average 124 mm.; tail 114 nmi., 92 
per cent of head and body length; foot 26 mm.; ear 28 mm.; 
occipitonasal length 35 nmi.; weight 52 gm. 

External characters.— Figure 96. Dorsum brownish, side gray and 
belly whitish to cream. All hairs with dark gray bases except 
whitish area of throat and cheek. Rostrum and crown orangish. 
Black facial marking begins posterior to mystacial area, encircles 
eye, and continues to base of ear. White area of cheek extends to 
shoulder region. Ear with long, white hairs around opening; inner 
surface thinly covered with short, white hairs; outer surface almost 
naked. Black patch medial to pinna. Postauricular patch white. 
Base of tail, about one-fourth of tail length, covered with short 
whitish to grayish hairs; rest of tail black and bushy, except for 
small white tip. Feet thinly covered with white hairs above, palm 
and sole naked. 

Cranial characters.— Figure 97. Rostrum elongate; cranium 
smooth, rounded; nasals truncate or emarginate posteriorly; fronto- 




.h 



CO 
?^ 

•2 



317 



318 FIELDIANA: ZOOLOGY 




Fig. 96. Live specimen of Eliomys quercinus cyrenaicus. 

parietal suture U-shaped due to encroachment of frontals posterior- 
ly between parietals; interparietal broad, ligulate in outline. 
Zygomatic plate small, but with prominent masseteric tuberosity; 
infraorbital canal very large. Incisive foramen relatively long, nar- 
row anteriorly, and broad posteriorly. Posterior palatine foramen 
very small, ovoid in outline, and diverging posteriorly. 
Parapterygoid fossa broad, shallow; superior wall perforated. 
Postpalatal margin level with posterior edge of molars. 

Tympanic bulla greatly inflated and with three distinct partitions 
visible through the external auditory meatus. Suprameatal triangle 
large and open posteriorly. Mastoid bulla with large anterior 
chamber, subarcuate fossa not conspicuous, medial inferior 
posterior mastoid chamber very large, accessory mastoid chamber 
present. Accessory tympanum absent. Anterior lip of external 
auditory meatus broadened. Angle of lower jaw large, inflected, 
with single large foramen. 

Bacu/um.— Baculum flattened and curved dorsoventrally, taper- 
ing from base to tip with small lateral projections at middle (Didier, 
1953). 

Teeth.— Upper incisor orthodont, smooth on anterior surface. 
Cheek teeth brachydont; crowns squarish, concave, and transverse- 
ly ridged. 

Measurements.— Table 39. Male and female dimensions subequal. 






FlU. 97. Skull of Eliomys quercinus. 



319 



320 FIELDIANA: ZOOLOGY 

Age determination.— Adult animals have cusps of molars worn 
and cranial sutures closed. 

Vana (ion.— Tails of some specimens lack white tips. Eliomys q. 
cyrenaicus from the Western Desert is somewhat darker than E. q. 
malanurus from Sinai and has a slightly shorter ear and longer 
alveolus (table 39). Smaller auditory bulla and shorter tail in 
cyrenaicus were reported by Hoogstraal et al. (1955), Hoogstraal 
(1963), and Ranck (1968), but data in Table 39 do not support these 
conclusions. Bulla length, incidentally, is a highly variable measure- 
ment. Skull height, an indicator of the degree of tympanic bulla 
inflation, is a reliable check against erroneous assumptions about 
bulla size in small samples. Size of the foramen in the angle of the 
lower jaw is individually variable and therefore not consistently 
smaller in cyrenaicus as the aforementioned authors assumed. The 
most obvious difference between E. q. cyrenaicus and E. q. 
melanurus, which occupy the ends of a series of circum- 
Mediterranean subspecies, is the proportion of the length of the 
grayish, short-haired portion of the tail to the length of the rest of 
the tail: about one-sixth in cyrenaicus and one-third in melanurus. 
Eliomys melanurus was formerly considered to be a distinct species, 
but Niethammer (1959) recognized it as a subspecies of E. quer- 
cinus. He found differences between the subspecies mainly in the 
color pattern of the tail base. Corbet (1966, p. 208) supported the 
latter's decision of combining the various forms into a single species 
because of the lack of demonstration of "an abrupt and absolute 
discontinuity in variation." Ranck (1968) also accepted Nietham- 
mer's conclusions. 

Comparisons.— Eliomys quercinus differs from all other Egyptian 
rodents in having black facial markings; black, bushy tail with 
short-haired base; premolars in upper and lower jaws; and cheek 
teeth with concave surfaces. Sekeetamys calurus is the only other 
Egyptian rodent with a black, bushy tail. 

Collection.— Eliomys readily enters live traps. 

//a6ifafs.— Trapped alive in limestone cliffs of Western Mediter- 
ranean Coastal Desert. In Sinai, it has been taken in gardens, in the 
mountains, and in a Bedouin tent (Wassif and Hoogstraal, 1953). 
Flower (1932) reported one caught in a new stone building at El 
Kossaima. Ranck (1968) reported E. q. denticularis from loose sand 



OSBORN & HELM Y: MAMMALS OF EGYPT 321 

Table 39. — Means (and ranges) of measurements, ratios, and weight of adult 
Eliomys quercinus. 





E. q. melanurus 


E. q. cyrenaicus 


HBL 


121.6(104-135)5 


124.2(110-140)25 


TL 


112.2(109-117)4 


114.9(104-127)22 


TL/HBL% 


89.4 (82.6-96.5) 4 


92.9 (77.2-100.9) 22 


FL 


27.5 (26-29) 5 


26.3 (24.0-29.0) 26 


EL 


31.2 (29-33) 5 


27.1 (25-31) 26 


Wt 





51.8 (38.4-63.0) 16 


ONL 


34.6 (33.6-36.0) 5 


35.3 (34.3-36.6) 17 


ZW 


19.6(19.1-20.0)3 


20.0 (19.0-21.5) 17 


lOW 


4.6 ( 4.6- 4.7) 5 


4.5 ( 4.3- 4.8) 17 


NL 


12.6(12.3-12.8)5 


13.5 (12.3-14.5) 17 


IFL 


4.3 ( 3.8- 4.8)4 


4.8 ( 4.4- 5.2) 14 


AL 


4.3 ( 3.8- 5.1)3 


5.5 ( 5.2- 5.6) 16 


BL 


11.6(10.9-12.1)5 


10.7(10.1-11.4)17 


SH 


14.4 (14.0-14.8) 4 


14.5(13.6-15.4) 17 



around the base of unpruned date palms and tamarix clumps in the 
Fezzan district of Libya. 

Captive fteAafior.— Extremely wild and agressive. We have not 
kept this species in captivity long enough to study its responses 
further. 

Associates. — Irv Sinai Peninsula, E. quercinus lives in rocky 
habitats with Sekeetamys calurus, Dipodillus dasyurus, Acomys 
cahirinus, and A. russatus. In the Western Desert, it is found in 
coastal cliffs with D. campestris. 

Reproduction.— No information is available on reproduction in 
Egyptian subspecies. 

Sex ratio.— A sample of 30 specimens from the Western Mediter- 
ranean Coastal Desert contained 12 (40 per cent) males and 18 
females. 

Key TO Egyptian Subspecies of E/tomys quercinus 

1. Basal one-third of tail grayish. Color pale. (Sinai Peninsula) melanurus, p. 321. 

2. Basal one-sixth of tail grayish. Color darker. (Northwestern part of Western 
Desert) cyrenaicus. p. 322. 

Eliomys quercinus melanurus (Wagner, 1840) 

Eliomys (Myoxus) melanurus Wagner, 1840, Abh. Bayer Akad. Wiss. Math.- 
Naturwiss. Kl., pi. 3. fig. 1. p. 176. 

Type locality. -Egypt. SINAI. 



322 FIELDIANA: ZOOLOGY 

Distribution in Egypt. — Figure 95. Sinai Peninsula. 

External characters. — Slightly paler and with short-haired, gray 
basal part of tail about twice as long as in E. q. cyrenaicus. 

Measurements.— Table 39. Ear length averages slightly longer 
and alveolar length shorter in melanurus than in cyrenaicus. See 
under variation. 

Specimens examined.— Total six. 

SINAI: Wadi Arbaein (3), Wadi el Raba (1), Gebel Musa (1). Tor (1). 

Published records.— Records are from Anderson (1902), Flower 
(1932), Wassif and Hoogstraal (1953), Hoogstraal (1963), and Haim 
and Tchernov (1974). 

SINAI: Nakhl; St. Catherine Monastery area; Gebel Musa; Wadi el Arbaein; El 
Raba; Wadi Rahaba, Quo Monastery area; EI Kossaima (El Quseima): Ain Sudr; 
Nohel; Wadi Dalma. 

Eliomys quercinus cyrenaicus (Festa, 1921) 

Eliomys cyrenaicus Festa, 1921, Boll. Mus. Zool. Anat. Comp. Univ. Torino, 36, 
No. 740, p. 4. 

Type locality.-Uhya. CYRENAICA: Gheminez. 

Distribution in Egypt— Figure 95. Northwestern part of Western 
Desert. 

External characters.—Slightly darker and with short-haired, gray 
basal part of tail about one-half as long as in melanurus. 

Measurements.— Table 39. Ear length averages slightly shorter 
and alveolar length longer in cyrenaicus than in melanurus. See 
under variation. 

Specimens examined.— Total 30. 

MATRUH: Ageeba 24 km. W of Mersa Matruh (18), Salum area (11), Salum 16 
km. E (1). 

Published records.— Records are from Hoogstraal et al. (1955) and 
Hoogstraal (1963). 

MATRUH: Salum. 

Family 5. Dipodidae 

Relatively small to large rodents, head and body length average 
105 to 148 mm. Modified for bipedal locomotion. Tibia and fibula 
fused, hind foot elongate, functional toes three, forelimb reduced, 
neck short. Tail about 150 per cent of head and body length, with 



OSBORN & HELMY: MAMMALS OF EGYPT 



323 



broad, feathered black subterminal band and white tip. All species 
have conspicuous whitish to grayish hip bands. Infraorbital 
foramen greatly enlarged. Upper root of zygomatic process 
posterior to lower root. Maxillary plate minute. Postpalatal margin 
posterior to level of m^. Bulla inflation variable. Angular process of 
lower jaw perforated. Upper incisors smooth on anterior surface in 
genus Allactaga, grooved in Jaculus. Dental formula: |, 



1x2=16-18. 



o» 



Key to Egyptian Genera of Dipodidae 

1. Ear longer than one-half hind foot length. Hind foot with three functional and one 
vestigial toe. Whitish hip bands not converging above base of tail. Mastoid bulla 
not inflated, tympanic bulla slightly inflated and not fused anteroventrally. 
Rudimentary premolar in upper jaw. Enamel pattern of m and m E-shaped (fig. 
98. table 40) Allactaga, p. 323. 

2. Ear shorter than one-half hind foot length, hind foot with three functional toes. 
Whitish hip bands converging above base of tail. Mastoid and tympanic bullae 
greatly inflated, the latter fused anteroventrally. Premolar lacking in upper jaw. 
Enamel pattern of m and m Z-shaped (fig. 98, table 40) Jaculus, p. 333. 

Genus Allactaga Cuvier, 1836 

Small jerboa. Dorsum speckled black and orange. White hip 
bands not converging above base of tail. Hind foot with three toes 
functional and one vestigial. Large digital and plantar pads not 
concealed by hair. 







ATeiRADACTYLA 



J.ORIENMIIS 



J.JACULUS 



Fui 98. Crown views of right upper (U) and left lower (L) molars of mature Allac- 
taga tetradactyla, Jaculus orientalis. and./, jaculus. 



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OSBORN&HELMY: MAMMALS OF EGYPT 327 

Zygomatic process of temporal without prominent lateral process, 
but extending beyond level of lip of external auditory meatus. Bony 
plate of lower root of zygomatic process not fused with maxilla 
(foramina maxillaris open dorsaUy). Jug^al with vertical and 
horizontal parts narrow. Interparietal broad, triangular. 
Postpalatal process with single spine. Mastoid bulla not inflated. 
Tympanic bulla slightly inflated, apices not fused. Upper incisor 
proodont, anterior surface smooth. Upper jaw with one vestigial 
premolar, three molars. M' and m^ with E-shaped enamel pattern. 
Lower jaw with angle uninflected, perforated by single, large 
foramen. Fossa between m^ and outer side of jaw, deep. Baculum 
absent. 

Allactaga tetradactyla (Lichtenstein, 1823) 

Dipus tetradactyla Lichtenstein, 1823, Verzeichniss der Doubletten des 
Zoologischen Museums der Berlin., p. 2. 

Type locality.— Egypt: "Libyan Desert between Siwa and Alexan- 
dria" (describer) taken to mean "Egypt, near Alexandria" by Eller- 
man (1940, p. 584). 

General distribution.— Mediterranean Coastal Desert of Egypt 
and eastern Libya. 

Common names.— Four-toed Jerboa, Gerbouh. 

Distribution in Egypt— Figure 99. Northern part of Western 
Mediterranean Coastal Desert. 

Diagnosis-SmaW jerboa with dorsum speckled black and orange, 
side grayish, and venter white. Ear pigmented, longer than one-half 
of hind foot length. Toe and tarsal pads large, not concealed by hair. 
Three functional toes, one vestigial. 

Skull inflated, mastoid bulla not expanded, tympanic bulla slight- 
ly expanded. Posterior margin of nasals truncate with a small 
median "V." 

Adult head and body length average 110 mm.; tail 169 mm., 155 
per cent of head and body length; hind foot 56 mm.; ear 40 mm.; 
occipitonasal length 29 mm.; weight 52 gm. 

External characters. — Figure 100. Dorsum dark, speckled black 
and orange; rump orangish, sides grayish. Dorsum hairs with black 
tips, orangish subterminal bands, and gray bases. Side hairs black 
tipped, white to base. Belly, underparts, and forelimb white. Hind 
foot with blackish hairs on underside of metatarsal and base of toes. 




I 



e 
o 

at 

■s 

"So 



OSBORN & HELM Y: MAMMALS OF EGYPT 



329 




i 

^ 



Fig. 100. Live specimen of AUactaga tetradactyla. 



Distal part of toes white. Mystacial area buffy. Suborbital area 
white. Postorbital and postauricular patches buffy. Ear pigmented, 
covered with whitish hairs. Whitish posterolateral hip bands not 
converging above base of tail. Tail paler than dorsum on upper sur- 
face, gradually becoming whitish on underside, with blackish, 
feathered subterminal band and white tip. 

Feet.— Hind foot with three functional toes, one vestigial. Sole 
naked. Plantar and digital pads large, naked. 

Cranial characters.— Figure 101. Most characters were described 
under genus and listed in Table 40. Cranium broadly triangular, 
parietal region inflated. Interparietal broad, triangular. Nasals 
truncate posteriorly with small median "V" (fig. 102). Zygomatic 
process of temporal extending beyond level of lip of external 
auditory meatus. Meatal lip flaring laterally. Mastoid bulla not in- 
flated, tympanic bulla slightly inflated. Lower jaw with angle 
uninflected, perforated by a single, large oval foramen. Fossa be- 
tween posterior molar and outer side of jaw deep. 



330 



FIELDIANA: ZOOLOGY 




Fiu 101. SkuW of Allactaga tetradactyla. 



TeetA.— Upper incisor slender, proodont, smooth on outer surface 
(fig. 101). Upper jaw with one vestigial premolar and three molars. 
M , m^ with E-shaped enamel pattern (fig. 98). 

Measurements.— TsihXe 41. Male dimensions average very slightly 
larger than female. Means (and ranges) of occipitonasal length (in 
millimeters) of 11 adult males and nine adult females are 29.0 (28.1 
to 30.1) and 28.9 (27.3 to 30.4), respectively. 

Age determination,— \d\x\ts have enamel cusps of upper molars 








«o 



O 




g 
III 



331 



332 FIELDIANA: ZOOLOGY 

Table 41. — Means (and ranges! of measurements, ratios, and weight of adult 
AUactaga tetradactyla and Jaculus orientalis. 





A. tetradactyla 


J. orientalis 


HBL 


109.7(102-119) 19 


148.4 (137-160) 31 


TL 


169.0(154-180)17 


224.2(195-243)31 


TL/HBL% 


154.7 (138.7-169.5) 17 


146.1 (127.5-177.3)31 


FL 


56.1 (51-59) 19 


74.7 (71-78) 31 


EL 


40.6 (37-43) 19 


32.9 (28-35) 30 


Wt 


48.0, 52.8, 56.0 


134.5(108.6-147.0) 17 


ONL 


28.9 (27.3-30.4) 20 


36.9 (36.2-38.0) 26 


ZW 


20.9(19.1-22.6) 16 


28.3(27.1-30.0)24 


low 


9.8 ( 9.1-10.5)20 


14.5 (13.9-15.3) 25 


NL 


10.7(10.0-11.6) 19 


15.1 (13.9-17.0) 25 


IFL 


5.6 ( 5.1- 6.0)20 


5.6 ( 5.1- 6.2)26 


AL 


5.9 ( 5.2- 6.2) 20 


6.7 ( 6.2- 7.4) 25 


SH 


13.4 (12.8-13.8) 19 


17.7 (17.0-18.6) 26 



united into lamellae, cranial sutures closed. In addition, degree of 
development of the dorsolateral inflections of the anterior part of 
the zygomatic arches correlates with tooth wear and suture closure 
(Lewis et al., 1965). 

Comparisons.— Allactaga tetradactyla can be distinguished from 
other Egyptian jerboas, Jaculus jaculus and J. orientalis, by its 
longer ears, darker color, vestigial fourth toe, rudimentary upper 
premolar, proodont incisor with smooth anterior surface, lack of in- 
flation of auditory bulla, and other characters in Table 40 and 
Figure 102. Wassif (1960d) should be consulted for additional 
osteological and other differences between Egyptian jerboas. 

Specimens examined.— Total 25. 

MATRUH: Burg el Arab (8): Bahig S of (5); Bahig 18 km. S (2); Abu Mena (3), 11.2 
km. Ed); Mersa Matruh (5); Sidi Barrani (1). 

Published records. — Records are from Setzer (1958a) and 
Hoogstraal (1963). 

MATRUH: Burg el Arab, Mersa Matruh, Sidi Barani 8 km. E, Abar el Dafa. 

Collection.— Dug from burrows and collected with butterfly nets 
at night using spotlight. 

Habitats.— Allactaga tetradactyla inhabits salt marshes and ad- 
jacent areas in coastal valleys of the Western Desert. It is also 
found inland on flat, clay desert in the vicinity of Bedouin barley 
fields and/or in areas where the predominant vegetation is Anabasis 
articulata (Hg. 8). 



OSBORN&HELMY: MAMMALS OF EGYPT 333 

Burrows.—Simple, 60 to 150 cm, deep. "Burrows appear often to 
be occupied only briefly and empty burrows oiJaculus orientalis are 
sometimes utilized." Winter rains often flood their burrows 
(Hoogstraal, 1963, p. 29). 

Associates.— Allactaga tetradactyla lives in salt marshes with 
Psammomys obesus, Jaculus orientalis, and Dipodillus sp. (fig. 7). 
Inland, in hard clay desert, it is again found with these species and 
with Meriones shawi (fig. 8). 

Populations.— Hoogstraal (1963) remarked on the small numbers 
of this species and its ecological limitation, that it had disappeared 
from certain coastal valleys near Alexandria, and that desert 
reclamation threatened it with extinction. 

Genus Jaculus Erxleben, 1777 

Large and small jerboas. Dorsal color varying from dark brownish 
orange to orangish or cinnamon. White hip bands converging above 
base of tail. Hind foot with three functional toes. Digital and plan- 
tar pads small, concealed by long stiff hairs. 

Zygomatic process of temporal with lateral process extending 
sUghtly or not beyond level of lip of external auditory meatus. Bony 
plate of lower root of zygomatic process fused with outer surface of 
maxilla closing foramina maxillaris dorsally. Jugal with broad ver- 
tical part and narrow horizontal part. Interparietal narrow, shield- 
shaped. Postpalatal process with two spines. Posterior margin of 
mastoid bulla inflated beyond level of supraoccipital. Tympanic 
bulla greatly inflated, apices fused medially below basioccipital and 
basisphenoid. Upper incisor orthodont or opithsodont, grooved on 
anterior surface. Upper jaw with three molars, first and second with 
Z-shaped enamel pattern. Lower jaw with angular process inflected 
and a shallow fossa between posterior molar and outer side of jaw. 
Baculum present. 

Skin folds can be brought over nostrils for protection when the 
animals are burrowing and pushing soil with the short, broad snout. 

Key to Egyptian Species of Jaculus 

1. Size large, head and body length 135-160 mm., ear length 28-35 mm., hind foot 
length more than 70 mm. Anterior mastoid chamber not completely filling 
suprameatal triangle. Posterior margin of nasals with V-shaped division (fig. 102) 
orientalis, p. 334. 

2. Size small, head and body length 99-120 mm., ear length 18-28 mm., hind foot 
length less than 70 mm. Anterior mastoid chamber completely filling 



334 FIELDIANA: ZOOLOGY 

suprameatal triangle. Posterior margin of nasals with U-shaped division (fig. 102) 
jaculus, p. 339. 

Jaculus orientalis Erxleben. 1777 

Jaculus orientalis Erxleben, 1777. Systema Regni Animalis. p. 404. 

Type locality.— Egypt. SINAI: "In the mountains separating 
Egypt from Arabia" (according to the describer on p. 404). 

General distribution. —Sinai Peninsula, Egypt, Libya, Tunisia, 
Algeria, and Morocco. 

Common names.— Greater Egyptian Jerboa, Gerbouh. 

Subspecies in Egypt — 

Jaculus orientalis orientalis Erxleben, 1777 

Type locality.— Given under species. 

Distribution in Egypt.— Figure 99. Northern Sinai, Southwestern 
Sinai Peninsula, Western Mediterranean Coastal Desert. 

Diagnosis.— Large jerboa with dorsum brownish orange, side and 
hip band grayish. Ear pigmented. Hind foot with sole haired. Heel, 
metatarsus, and base of toes black. Tail long, with black subter- 
minal band and white tip. 

Skull inflated. Anterior mastoid chamber partly filling 
suprameatal triangle. Posterior margin of nasals with V-shaped 
division. Angle of lower jaw inflated, perforated with single large, 
round foramen. 

Largest species of genus in Egypt. Adult head and body length 
average 148 mm.; tail 224 mm., 146 per cent of head and body 
length; hind foot 74 mm.; ear 32 mm.; weight 134 gm. 

External characters.— Dorsum brownish orange, sides grayish 
due to black-tipped white hairs, becoming buffy to orangish on flank 
and thigh. Hip band of white hairs with grayish tips. Dorsum hairs 
with gray bases, orangish subterminal bands, and brownish tips. 
Underparts and forelimb white. Side of snout and head buffy. 
Supraorbital and postauricular spots small, whitish. Ear 
pigmented, whitish hairs on inner surface, buffy hairs on outer sur- 
face and lower anterior margin. Hind limb brownish orange on outer 
side. Foot white above, hair of sole and base of toes blackish, toe 
tips white. Tail with upper surface paler than dorsum, bicolored, 
underside white. Distal part of tail with white, feathered tip and 



OSBORN & HELMY: MAMMALS OF EGYPT 



335 



black, subterminal band. Dorsal tail surface whitish proximal to 
black band. 

Feet— Palm naked. Sole haired. Plantar and digital pads small 
and concealed by brush of long, stiff hairs. 

Cranial characters. — Figure 103. Posterior nasal margin bifur- 
cated into a broad "V" (fig. 102). Anterior mastoid chamber not fill- 
ing suprameatal triangle. Lateral extension of zygomatic processs 
of temporal bone extending slightly beyond level of lip of external 
auditory meatus. Lip of meatus flaring laterally. Lower jaw with 
angular process inflected and perforated by one large round 






Fk; 103. Skull of Jaculus orientalis orientalis. 



336 FIELDIANA: ZOOLOGY 

foramen. Characters in common with J. jaculus were discussed 
under the genus. 

Tee t/i. — Figure 98. Incisors orthodont, upper with groove on 
anterior surface. Molars three in upper and lower jaws, enamel pat- 
tern of m', m Z-shaped. 

Measurements.— Table 41. Males average slightly larger in most 
dimensions than females, except occipitonasal length, the averages 
(and ranges) of which, in 14 adult males and 12 adult females, are (in 
millimeters): 36.7 (36.2 to 37.8) and 37.1 (36.3 to 38.0), respectively. 

Age determination.— Adults have enamel cusps of upper molars 
united into lamellae and skull sutures closed. Degree of develop- 
ment of the dorsolateral inflection of the anterior end of zygomatic 
arch correlates with tooth wear and suture closure as in Alloc taga. 

Variation. — Individual variation in color was noted by Setzer 
(1958a). 

Comparisons.— Jaculus orientalis differs from A. tetradactyla in 
larger dimensions, except for ear length, slightly paler color, hair on 
sole, hair concealing tarsal and toe pads, greater inflation of 
auditory bulla, and other characters listed under the descriptions of 
genera and in Table 40. From J. jaculus, J. orientalis differs in larger 
dimensions, esp>ecially ear and hind foot length, darker color, com- 
pletely pigmented ear, less swollen anterior mastoid chamber, 
V-shaped, rather than U-shaped, posterior nasal margin (fig. 102), 
and other characters Hsted in Table 40. 

Remarks. — Immature specimens from Ras Abu Rudeis on the 
Gulf of Suez coast of Sinai Peninsula are treated as J. o. orientalis 
since no adult specimens are available, as also noted by Setzer 
(1958a) and Hoogstraal (1963). 

Specimens examined.— Total 83. 

SINAI: Ras Abu Rudeis (2). 

BEHEIRA: Beheira Nakhla (1). 

ALEXANDRIA: Abu Qir (3). Ramleh (1). Amiriya (12). 

MATRUH: Uke Mariut (8); Bahig (6). 18 km. S (2). 25.6 km. S (10); Abu Mena (4). 
6.4 km. Ed): Burg el Arab (4); Raqabet el HaUf (1); El Alamein (3); Ras el Hekma (5); 
Maatin el Garawla (1): Mersa Matruh (3), 1.6 km. NE (3). 2.4 km. NE (1 1), 8 km. NE 
(2). 

Published records. — Records are from Anderson (1902), Setzer 
(1958a), Hoogstraal (1963), and Haim and Tchernov (1974). 



OSBORN&HELMY: MAMMALS OF EGYPT 337 

SINAI: Ras Abu Rudeis, Nakhl, Kuntila, Gebel Maghara. 

BEHEIRA: South side of Lake Idku (Abu Hommos area). 

ALEXANDRIA: Ramleh, Alexandria. 

MATRUH: Burg el Arab; El Hammam; El Daba; El Alamein; Mersa Matruh. L6 

km. NE, 2.4 km. NE. 8 km. NE; Sidi Barrani. 

Collection.— Dug from burrows and captured at night under a 
spotlight with butterfly nets. Occasionally taken in live traps. 

Habitats.—Sinai Peninsula: Seashore area near Ras Abu Rudeis 
on the Gulf of Suez coast and northern and eastern desert areas. 

Western Mediterranean Coastal Desert: Salt marshes with domi- 
nant plant usually Salicornia fruticosa (fig. 7); limestone slopes 
above salt marshes supporting Suaeda fruticosa (fig. 7); gardens; 
olive groves; Bedouin barley fields; clay desert in the Thymelaea- 
Anabasis association (fig. 8); and sandy or rocky slopes supporting 
Thymelaea hirsuta, Noaea mucronata, Lycium sp., and Echinops 
spinosissimus. One specimen was collected near Artemisia 
monosperma in a sandy area 6.4 km. E of Abu Mena. 

Behavior.— KiTmiz s (1965) remark that some burrows led into 
large underground recreation parlors or "jerboa clubs," where these 
rodents gather to frolic, is no doubt a Bedouin fabrication. 

Jaculus orientalis is strictly nocturnal, becoming active at dusk. 
It is a sociable species, and solitary individuals are rarely found as is 
common with J. jaculus. 

Greater jerboas are relatively docile and not prone to bite. They 
do not struggle to escape if held by the tail. 

.Burro It; s.— Burrows are usually in hard ground, slanting to a 
depth of about 2 m. Openings are closed when occupied, at least in 
summer, with one or two sand plugs. One or two escape tunnels may 
be present and difficult to locate on the surface. There is usually a 
sleeping or nest chamber containing camel hair or shredded plant 
material and a food storage chamber (Hoogstraal, 1963). Burrow 
locations vary seasonally from hillsides in the winter rainy season to 
near margins of fields or close to vegetation in summer (Kirmiz, 
1965). In the latter habitat, J. orientalis may be found in burrows of 
Meriones shawi. In salt marshes, J. orientalis shares burrows with 
Psammomys obesus. Allactaga tetradactyla is occasionally re- 
moved from burrows of J. orientalis. 

Food— Sprouting vegetation, plant roots, or barley grains 
planted by Bedouins are Usted as food of wild J. orientalis by Kir- 



338 FIELDIANA: ZOOLOGY 

miz (1962, p. 32) and are the preferred foods of captive jerboas, but 
"they will eat bread, rice, and vegetables, such as fresh com, green 
peas, green beans, carrots, potatoes, lentils, etc. They also eat 
peanuts and melon seeds. They do not eat dates, dry fruits, bananas 
or tomatoes." In Kirmiz's laboratory, J. orientalis survived on 
wheat and barley grains alone for one to three years. Hoogstraal 
(1963) found quantities of dates (contrary to Kirmiz, 1962), barley, 
and other seeds stored in burrows. With the variety of perennial and 
ephemeral plants in the Western Mediterranean Coastal Desert 
flora, there is no lack of food for this and other rodent species. A col- 
lection of seed capsules and dry fruits from the burrow of a greater 
jerboa near Bahig in April, 1964, included mostly Malva aegyptia, 
and Calendula micrantha, Trigonella stellata. Astragalus hamosus, 
Medicago sp., Lophochloa pumila, and Parapholis marginata. 
Various succulent shrubs, such as Salicornia fruticosa and Suaeda 
fruticosa, are probably browsed by J. orientalis inhabiting salt mar- 
shes or the periphery. 

Although J. orientalis appears to be physiologically capable of 
surviving in true desert along with J. jaculus, it does not. Perhaps 
availability of food limits this larger species to the littoral 
semidesert. 

Water.— Kirmiz (1962, 1965), in studies of the physical, 
behavioral, and physiological adaptations of J. orientalis to the 
desert environment of the Western Mediterranean Coastal Desert, 
emphasized its ability to live in the laboratory on dry food without 
water. Our experience with this species and J. jaculus indicates that 
succulent vegetation and new growth (e.g., barley sprouts) provide 
available water in nature. 

Popu/af ions.— Hoogstraal (1963, p. 32) recorded 1 to 50 or more 
greater jerboas per 0.8 km., "depending both on availability of food 
and nature of soil." 

Associates.— As mentioned, J. orientalis is found with A. 
tetradactyla and P. obesus. The species has also been collected in 
the same habitat with Meriones shawi, Gerbillus andersoni, J. 
jaculus, and various Dipodillus sp., all inhabitants of the Western 
Mediterranean Coastal Desert and, some of them, the coastal salt 
marshes. 

Reproduction,— Jaculus orientalis does not breed in captivity, 
according to Kirmiz (1962). Reproductive data from wild sp>ecimens 



OSBORN&HELMY: MAMMALS OF EGYPT 339 

are scanty. One postpartum female had two fetal scars in March, 
another was found lactating in June, and a third carried two fetuses 
in August. Flower (1932) said three appeared to be the average litter 
size, noted one case of four or five, and gave months of birth as 
February, April, and early July. These data indicate that the 
breeding season begins after the winter rains (November-February) 
and lasts about five or six months. 

Sex ratio. — In a sample of 64 museum specimens of greater jer- 
boas, males numbered 30 and females 34. 

Economic importance.— Thighs and lumbar regions are roasted 
and eaten by Bedouins. 

These rodents probably cause some loss to Bedouins by feeding 
on sprouting barley and ripe grain. 

Jaculus jaculus (Linnaeus, 1758) 

Mus jaculus Linnaeus, 1758, Syst. Nat., 10th ed., p. 62. 
Type locality.— Northern Egypt. 

General distribution. — Iraq, Syria, Lebanon, Israel, Jordan, Saudi 
Arabia, Sinai Peninsula, Egypt, Libya, Algeria, Mauritania, 
Spanish Sahara, Chad, Niger, and Somalia. 

Distribution of subspecies in Egypt— Figure 104. Jaculus jaculus 
schlueteri: Sinai Peninsula, northern part of Eastern Desert; 
Jaculus jaculus butleri: southern part of Eastern Desert; Jaculus 
jaculus flavillus: Western Mediterranean Coastal Desert; Jaculus 
jaculus jaculus: Western Desert from the Mediterranean Coastal 
Desert southward. 

Common names.— hesser Jerboa, Gerbouh. 

Diagnosis.SmaW jerboa with dorsum orangish to brownish cin- 
namon. Side orangish to grayish, hip band whitish to grayish. Ear 
tip pigmented. Hind foot with or without dark hair on heel or 
metatarsus. Toes three. Tail long, with black subterminal band and 
white tip. 

Parietal portions of skull inflated. Anterior chamber of mastoid 
bulla completely filling suprameatal triangle. Posterior margin of 
nasals with a U-shaped division. Lower jaw with angle inflected and 
perforated with one or two foramina of varying size. 

Smallest species of the genus in Egypt. Adult head and body 
length average 107 mm.; tail 181 mm., 166 per cent of head and 



340 FIELDIANA: ZOOLOGY 

,,. 25* 26* 27* 26' 2 9* 30* 31* 32* 3 3* 34* 35* 36* 37* 




Fig. 104. Collection localities of ,/aculus jaculus jaculus (circles), ./. / flaviUus 
(dots). J. j. schlueteri (closed squares), ./. / butleri (open squares), and Hystrix 
cristata (triangles). 



body length; hind foot 62 mm.; ear 22 mm.; occipitonasal length 32 
mm.; weight 55.0 gm. 

External characters.— Figyire 105. Dorsum orangish to brownish 
cinnamon. Sides orangish to grayish, hip band whitish to grayish 
due to varying widths of blackish hair tips. Thighs orangish to cin- 
namon. Dorsal hairs with gray bases. Side, belly, and hip band hairs 
with white bases. Underparts and forelimb white. Hind foot 
sometimes with blackish or buffy hairs on heel, metatarsus, and 
base of toes. Distal part of toes and upper side of foot white. Ear tip 
pigmented. Whitish hairs on inner surface of ear denser than on 
outer surface. Anterior margin of ear with whitish fringe. Mystacial 
and preorbital areas white, suborbital areas white or partly 



OSBORN & HELMY: MAMMALS OF EGYPT 



341 



pigmented. Supraorbital spot large, white. Postauricular spot 
white, inconspicuous. Upper surface of tail about color of dorsum, 
gradually becoming whitish on underside. Tail cylindrical except for 
broad blackish or brownish feathered subterminal band and white 
tip. Subterminal band often preceeded by white ring which may or 
may not connect on underside with the white of tip. The black band 
is thereby "complete" or "incomplete" (table 42). 

Feet— Palm naked, sole haired. Plantar and digital pads small 
and concealed by brush of long, stiff hairs. 

Cranial characters.— See Figure 103 of Jaculus orientalis. Most 
characters were described under genus and are listed in Table 42. 
Skull triangular in dorsal outline, parietal region inflated. Inter- 
p£irietal broad, shield-shaped. Anterior mastoid chamber completely 
filling suprameatal triangle. Posterior margin of nasals with a 
U-shaped division (fig. 102). Angle of lower jaw inflected, perforated 
by one or two round foramina of varying size (fig. 106). 

TeefA.— Figure 98. Upper incisor opisthodont, anterior surface 
grooved. Three molars in upper and lower jaws. Enamel pattern of 
m\ m^ Z-shaped. 





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Fk; 105. Live specimen oiJaculus jaculus jaculus. Note that longest vibrissae are 
contacting ground. 



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F'u; 106. Lower jaw (upper) and posteriors of lower jaws (lower two figures) in 
ventral view of Jaculus jaculus showing variations in foramina in angles of jaw. 
Single foramina in each jaw is not shown. 

342 




343 



344 FIELDIANA: ZOOLOGY 

Measurements.— Table 43. Dimensions of males and females sub- 
equal. Means (and ranges) of occipitonasal length (in millimeters) of 
10 adult males and eight adult females, are: 31.4 (30.7 to 32.2) and 
31.4 (31.0 to 31.8), respectively. 

Age determination.— Adults have enamel cusps of upper molars 
united into lamellae and cranial sutures closed. In addition, degree 
of development of the dorsolateral inflections of zygomatic arches 
correlates with tooth wear and suture closure as in Allactaga and J. 
orientalis. 

Variation,— Y)oTss\ color in J. jaculus varies from brownish orange 
to brownish cinnamon in Sinai Peninsula, Eastern Desert, and 
Western Mediterranean Coastal Desert subspecies schlueteri, 
butleri, and flavillus, respectively, to orangish in Western Desert 
jaculus. Individuals of either color phase can be found randomly 
distributed within the geographical range of the other. Immatures 
and molting individuals cannot always be segregated into one or the 
other color phase. The black tail band is usually complete, hip bands 
more grayish, and hairs of sole more commonly pigmented in the 
brownish races; whereas in orangish J. jaculus, the black tail band is 
usually incomplete, hip bands less grayish, and hairs of sole less 
commonly pigmented (table 42). The data in Table 42 were initially 
segregated on the basis of dorsal color. 

Thomas (1922) noted that color was more uniformly darker and 
tail band usually complete in J. / butleri in Sudan compared with J. 
/ jaculus of Egypt. 

Number and size of foramina in the angle of the lower jaw vary 
through one large, one large and one small, two large, and two small 
(fig. 106). Asymmetry in number and size is common, although in 
Table 42, the largest number in either side of the jaw only was 
recorded. 

In a sample of 12 orangish J. j. jaculus from the semidesert near 
Giza, four were asymmetrical, five had one foramen in each side, and 
three had two foramina in each side. In another sample of 12 of 
mixed colors from Bir Victoria, one was asymmetrical, six had one 
foramen in each side, and five had two foramina in each side. Three 
of the latter were dark in color. Data in Table 42 show that in the 
northern subspecies, schlueteri and flavillus, 60 per cent have two 
foramina in one side of the lower jaw, and 40 per cent have a single 
foramen in the lower jaw. In subsi)ecies butleri and jaculus, approx- 



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346 FIELDIANA: ZOOLOGY 

imately the reverse occurs, 40 per cent have two foramina and 60 
per cent have one foramen (table 42). Thomas (1922) recorded a 
single foramen only in the lower jaw of J. j. butleri in Sudan. 
Number of foramina in the lower jaw is of limited use in 
distinguishing subspecies and of some value in separating species of 
jerboas. 

Jaculus j. butleri is smaller in most dimensions than other Egyp- 
tian subspecies (table 43). In J. j. schlueteri, the hind foot averages 
slightly longer than in other subspecies (fig. 107). In the northern 
subspecies, schlueteri and flavillus, ears average longer than in 
subspecies butleri and jaculus to the south. Other measurements 
show no significant clinal variation. Skull height, which is an in- 
dicator of variation in parietal swelling and/or bulla inflation, gives 
no indication of proportional differences among subspecies. 

Comparison.— Jaculus jaculus can be distinguished from J. orien- 
talis by smaller external and cranial dimensions, especially the 
shorter hind foot and ear; paler color; less pigmentation on ear; more 
swollen anterior mastoid chamber, completely filling the 
suprameatal triangle; and U-shaped, rather than V-shaped, 
posterior bifurcation of the nasals (fig. 102). These and additional 
characters are listed in Table 42. Jaculus jaculus differs from Allac- 
taga tetradactyla in paler color; much shorter and less pigmented 
ears; hair on sole; pads of foot concealed by hair; lack of the nonfunc- 
tional fourth toe; much greater inflation of the auditory bulla; upper 
incisor opisthodont and grooved, rather than proodont and smooth; 
and lack of a premolar. Additional characters are listed in Table 42. 

Remarks.— The two color phases, orangish and brownish cin- 
namon, according to Ranck (1968), represent two sympatric species, 
J. jaculus and J. deserti. The latter, he maintains, can be 
distinguished from J. jaculus by the following characters: (1) darker 
hair on dorsum, side, and sole of feet; (2) smaller, more compact 
skulls; (3) more inflated auditory bulla; and (4) two distinct foramina 
of equal or unequal size in the angular process of the mandible as 
opposed to a single foramen. 

Four "distinct" Libyan populations of J. deserti recognized by 
Ranck (1968) consisted of two specimens allotted to J. deserti 

Opposite: 

Fk; 107. F'requency diagrams of ear length and hind foot length of subspecies of 
.laculus jaculus. Overlapping is indicated by double lines. 



35 



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WES TERN DESERT : 

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JACULUS Q 



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SINAI AND EASTERN DESERT 



SCHLUETERI Q 

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18 20 22 24 26 28 
EAR LENGTH 



56 58 60 62 64 66 68 mm 
HIND FOOT LENGTH 



347 



348 FIELDIANA: ZOOLOGY 

flavillus, and five, one, and one for each of three "new" subspecies. 
He also considered schlueteri a subspecies of J. deserti. The data 
presented herein on variation (table 42, fig. 107) are not in accord 
with Ranck's conclusions and do not support the application of J. 
deserti to Egyptian and Libyan subspecies. 

Collection.— Dug from burrows and caught at night under a 
spotlight with an insect net, Jaculus jaculus enters live traps, par- 
ticularly when placed beside feeding stations (fig. 108) or in tracks 
made in sand by a vehicle. Incidently, trap lines placed in car tracks 
are also effective in capturing other species that frequent sandy 
habitats. 

Habitats.— SinsA. Peninsula: "Loose sand hills and hillocks" in the 
north (Wassif, 1953b, p. 115); "plains and wide, sandy wadis, often 
near tamarisk trees, from the Red Sea coasts to high altitudes" 
(Hoogstraal, 1963, p. 30). "It exists up to an altitude of 1,500 meters 
on the plateau of St. Catherine Monastery" (Haim and Tchernov, 
1974, p. 216). 

Eastern Desert: Sand and gravel accumulations around trees and 
shrubs; hard, sandy, and gravelly terraces and deltas of mountain 
wadis; coastal plains; and intersecting wadis. Reported from acacia 
parkland at the foot of Gebel Elba (Hoogstraal, 1963) where bur- 
rows were found in sandy areas near vegetation. 

Western Desert: Mediterranean Coastal Desert on clay soils in 
and around Bedouin barley fields and in Thymelaea-Anabasis com- 
munities (fig. 19); and sandy areas supporting /I rtemisia monosper- 
ma (fig. 20), margins of oases, and depressions supporting one or 
several species of grasses and/or flowering plants (fig. 10). South of 
the littoral vegetation, burrows are usually in hard, barren sand 
slopes as far as several hundred meters from the nearest vegetation. 

Behavior.— Jaculus jaculus, as other Egyptian jerboas, is strictly 
nocturnal. Unlike the more sociable J. orientalis, solitary in- 
dividuals are commonly found wandering at night in barren desert. 
This proneness to wander was mentioned by Harding-King (1925) 
who once followed a jerboa's track in the Western Desert for about 
14 km. 

Jaculus jaculus is nervous, struggles to escape, does not bite when 
handled, and emits plaintive cries. If held by the tail, jerboas cease 
struggling. 



OSBORN & HELMY: MAMMALS OF EGYPT 



349 




Fig. 108. Western Desert 3 km. SSE of Camel Pass Dune. Solitary specimen of 
Comulaca monocantha, an important browse plant of Gazella sp. and a food source 
for Jaculus jaculus. Note foot and tail markings of the latter indicating activity 
while feeding. Live trap at left caught one specimen of J. jaculus jaculus. 



Driving a car around or over a burrow, even in daytime, 
stimulates a jerboa to leave the burrow. After an erratic chase, if the 
jerboa is not captured, it will return and disappear into the burrow 
via the single, small opening. If attempt is made to dig out the jer- 
boa, it may suddenly emerge from the ground via an escape passage. 
If chased, the jerboa may again lead an erratic flight and return to 
the burrow site. If, in its movements, another hole was discovered, 
the jerboa would enter it. A jerboa, well in advance of its pursuers, 
may stop suddenly and remain motionless, sometimes avoiding fur- 



350 FIELDIANA: ZOOLOGY 

ther discovery. For additional information on J. jaculus behavior, 
Happold (1967a, 1968a) should be consulted. 

fiurrou;s. — Burrows dug in hard, barren sand or beneath serir are 
usually difficult to find, because all excavated material is carried 
away by wind. Occupied burrows are plugged with sand in summer, 
open in winter. Burrows are simple, slanting, a meter or more in 
depth, and according to Hoogstraal (1963), usually deeper in winter 
than in summer. One or two escape exits are usually dug vertically 
just beneath the ground surface. The nest chamber near the end of 
the burrow is furnished with shredded plant material. 

West of Cairo, in hard, barren clayey-sand slopes that descended 
gradually for as much as 64 m., Briscoe (1956) found shallow to ex- 
ceptionally deep burrows. The burrows extended from 30 to 168 cm. 
below the surface and were 60 to 196 cm. long. In early September, 
1953, ground temperatures varied from 78° to 104° F. and relative 
humidity, 33.5 to 48 per cent. Within burrows, temperature varied 
from 80° to 100° F. and relative humidity 35.5 to 49.5 per cent. 

Food. — Happold (1968a, p. 433) found that "seeds, dried desert 
grasses, and roots" were the normal food of the lesser jerboa in 
Sudan. He (Happold, 1967a) fed captive animals grass, cubes of 
cucumber, and sunflower seeds as a basic diet and said it cannot eat 
large, hard seeds and will reject them. We have maintained J. 
jaculus in captivity on a diet of dry bread, sorghum grain, and raw 
carrots. Tortonese (1948) fed captive J. jaculus leaves, fruit peels, 
crumbs of bread, and cheese. 

In the Eastern Desert, lesser jerboas were trapped beside the 
pungent-smelling Cleome droserifolia and, in captivity, ate the 
harsh, glandular foliage. Aerva javanica was also eaten. Jerboas 
taken in Salsola baryosma on the Red Sea coastal plain were 
assumed to feed on the succulent leaves. At El Maghra in the 
Western Desert, a stand of Zygophyllum album was found to be a 
feeding area of J. jaculus. Captive animals readily accepted and ate 
the fleshy leaves. A specimen or two of lesser jerboa can usually be 
caught beside spiny Comulaca monocantha, where foot and tail 
markings indicate jerboas have been feeding (fig. 108). Seeds and 
new growth of various grasses, such as the hummock-forming 
Stipagrostis scoparia and S. vulnerans, are eaten as is the large, 
common bunch grass, Panicum turgidum (fig. 20). Seeds of 
Calligonum comosum (fig. 9) are also eaten. Stipagrostis plumosa, a 



OSBORN&HELMY: MAMMALS OF EGYPT 351 

small grass of runnels and sandy depressions, is often kept closely 
clipped by grazing lesser jerboas. 

The inability of J. jaculus to cope with large, hard seeds probably 
accounts for the absence of the species from certain acacia groves 
where no other plants exist. Meriones crassus, M. libycus, Gerbillus 
gerbillus, and G. perpallidus can utilize acacia seeds and are found 
near trees on otherwise barren gravel or sand. Jerboas that we 
found at night in barren desert were assumed to be "foraging" for 
small, windblown seeds trapped in crevices or hollows. 

In the Western Mediterranean Coastal Desert, food of the lesser 
jerboa no doubt includes barley and sprouting grain and other 
plants also eaten by the greater jerboa. Jaculus jaculus collected in 
Thy melaea- Anabasis associations (fig. 19) and in the vicinity of 
Artemisia monosperma (fig. 20) probably eat some parts of these 
plants, particularly the buds of the latter. Beyond this, we have no 
exact information of foods of the lesser jerboa in this area. 

Observations of Happold (1967a) in Sudan and of Hoogstraal et 
al. (1957b) in Egypt indicate that J. jaculus does not store food in its 
burrows. Our investigations in Egypt have resulted in the same 
conclusion. 

Water.— Water is metabolized from the natural food of J. jaculus, 
although experimenters have shown that this species will lose 
weight rapidly on a diet of dry grain alone (Schmidt-Nielsen, 1964, 
p. 182). Tortonese (1948) and Happold (1968a) observed that J. 
jaculus drinks water in captivity. In nature, jerboas may utilize dew 
(Bagnold, 1954). 

Details of the physiologic features of water economy in J. jaculus 
have been discussed by Shalaby (1962) and Gabr and Shalaby (1962, 
1964). 

Populations.— hesser jerboas are never abundant, and usually no 
more than three or four are seen in any one locality. The distribution 
in Sudan is "patchy" (Happold, 1967a), and like J. orientalis, 
numbers vary with availability of food and type of terrain. 

Associates.— Jaculus jaculus shares the same environments with 
Allactaga tetradactyla, J. orientalis, Meriones libycus, M. shawi, M. 
crassus, Gerbillus gerbillus, G. andersoni, G. pyramidum, G. per- 
pallidus, Pachyuromys duprasi, and some Dipodillus sp. Burrow 
sites, except for areas within the Western Mediterranean Coastal 



352 FIELDIANA: ZOOLOGY 

Desert vegetation, are isolated from most other species, except 
perhaps M. crassus. 

Reproduction. — Flower (1932) reported four young born in May 
and examined nestlings in August. Additional records of pregnant 
and lactating females occur from February to September. Number 
of young range from 4 to 10. 

Happold (1967a) gave the reproductive period at Khartoum from 
October to November in 1964 and October to February in 1965. 
Average number of young in 18 litters which he examined was 
three, range two to five. 

Later, he (Happold, 1970) succeeded in breeding J. j. butleri in cap- 
tivity by providing adequate space and suitable nesting conditions. 
He described in detail courtship and mating behavior and develop- 
ment of the young. 

Sex ratio. — In a sample of 144 museum specimens of lesser jer- 
boas, males numbered 75 (52 per cent) and females 69. 

Economic importance.— These rodents probably cause some loss 
to Bedouins by feeding on sprouting barley and grain. Hoogstraal 
(1963) observed that this jerboa never invaded established 
cultivated areas. We have a single specimen from a peanut field in 
the desert edge near Abu Rawash. 

Key to Egyptian Subspecies of Jaculus jaculus 

1. Dorsum brownish, black tail band usually complete on the underside. 

a. Ear length averaging more than 23 mm. 

i. Hind foot length averaging more than 63 mm. (Sinai Peninsula and northern 
part of Eastern Desert) schlueteri, p. 352. 

ii. Hind foot length averaging less than 63 mm. (Western Mediterranean 
Coastal Desert) flavillus, p. 353. 

b. Ear length averaging less than 23 mm. (Southern part of Eastern Desert) 
butleri, p. 355. 

2. Dorsum orangish, black tail band usually incomplete on the underside. (Western 
Desert) jaculus, p. 356. 

Jaculus jaculus schlueteri (Nehring, 1901) 

Dipus schlueteri Nehring. 1901, S.B. Ges. Nat. Fr. Berlin, p. 163. 
Type locality. — Israel: Jaffa. 

Distribution in Egypt — Figure 104. Sinai Peninsula and northern 
part of Eastern Desert. 

External characters. — Dorsum brownish cinnamon, side and hip 
band white with cinnamon and grayish or blackish tipped hairs. 



OSBORN&HELMY: MAMMALS OF EGYPT 353 

Mystacial and preorbital area white; supraorbital spot huffy, in- 
conspicuous; postauricular spot small, whitish. Black tail band com- 
plete on underside in 89 per cent of specimens (table 42). 

Cranial characters.— See species description. Angle of lower jaw 
with two foramina in 62 per cent and one foramen in 38 per cent of 
specimens (table 42). 

Measurements.— Table 43, Figure 107. 

Kanation.— Orangish individuals appear sporadically in the 
population. Those in the Eastern Desert near Cairo were considered 
by Setzer (1959a) to be intergrades with J. j. jaculus of the Western 
Desert. 

Intergradation cannot be demonstrated between schlueteri and 
butleri because of lack of material. 

Comparisons.— Jaculus j. schlueteri differs from other Egyptian 
subspecies chiefly in greater average hind foot length (table 43, fig. 
107). Other differences are mentioned under the following 
subspecies. 

Specimens examined.— Total 64. 

SINAI: Ras Abu Rudeis (10), Wadi Raha (2). Feiran Oasis (2). 

ISMAILIA: Fayid (10). 4.8 km. NE (5). 

SUEZ: Wadi Ghuweibba (2); Ain Sukhna area (3); Wadi el Katamiya mouth (2); 
Wadi el Gafra (1); Wadi GindaU (1); Cairo-Suez road km. 28 (2). km. 24 (2), km. 20 (1). 
km. 18 (1). 

CAIRO: Wadi Digla (1), Gebel el Ahmar (4). HeUopolis 8 km. E (4). 

RED SEA: Wadi el Nil (1). Wadi Abu Shaar (1), Abu Kharif mine area in Wadi 
Fatira (1), Fawakhir mine (1), 16 km. E in Wadi Abu Ziran (1); Wadi Umm Selemat 
(1). 

ASYUT: Wadi el Asyuti 20 km. SE of Asyut (3). 

QENA: Luxor (2). 

Published records.— Records are from Allen (1915), Wassif 
(1953b), and Setzer (1958a). Some were listed by these authors as J. 
j. jaculus. 

SINAI: Ras Abu Rudeis, Wadi Feiran, Feiran Oasis, Wadi Raha. 
SHARQIYA: El Salhiya. 
ISMAILIA: Nefisha. El Ferden. 
SUEZ: Wadi Ghuweibba. Ain Sukhna. 

Jaculus jaculus flavillus Setzer, 1955 

Jaculus jaculus flavillus Setzer. 1955, Proc. Biol. Soc. Wash. 68, p. 184. 



354 FIELDIANA: ZOOLOGY 

Type locality.— Egypt. MATRUH: Salum. Bir Bosslanga (Bir 
Wair). 

Distribution in Egypt.— Figure 104. Western Mediterranean 
Coastal Desert. 

External characters.— Dorsum brownish cinnamon, side and hip 
bar white with cinnamon and blackish tipped hairs. Mystacial and 
preorbital areas white; supraorbital spot whitish, not prominent; 
postauricular spot small, white. Black tail band complete on under- 
side in 80 per cent of specimens (table 42). 

Cranial characters. —See species description. Angle of lower jaw 
with two foramina in 58 per cent and one foramen in 42 per cent of 
specimens (table 42). 

Measurements.— Table 43, Figure 107. 

Variation.— Orangish individuals appear sporadically in the 
population. Intergradation between J. / flavillus and J. j. jaculus oc- 
curs in the areas of Wadi el Natroun, Bir Victoria (Setzer, 1958a) 
and El Birigat (fig. 104) and could no doubt be demonstrated all 
along the southern limits of the Mediterranean Coastal Desert 
vegetation. 

Co mpansons.— Longer average ear length (fig. 107) and brownish 
rather than orangish color in flavillus are the main characters that 
can be used to distinguish the subspecies from jaculus. From 
schlueteri, flavillus differs in slightly smaller average hind foot 
length. In coloration, these two subspecies are nearly identical. 

Specimens examined.— Totai 49. 

BEHEIRA: Bir Victoria (12); Wadi el Natroun. Abu Makkar Monastery (Deir 
Makaryus) (1): Wadi el Natroun (7). 15 km. W (4); El Hamra (1|: EI KhaUtba (4); Kom 
Hamada (1): El Birigat (2). 

MATRUH: El Amiriya (2); Bahig (1). 18 km. SE (2); Burg el Arab (1); Raqabet el 
Halif (1); Abu Mena (2): El Hawa 20 km. S of El Hamman (1): El Daba (1); Mersa 
Matruh (1); Sidi Barrani 32 km. W (4); Salum. Bir Bosslanga (Bir Wair) (Type). 

Published records. — Records are from Setzer (1955, 1958a), and 
some were listed by him as J. j. jaculus. 

BEHEIRA: El Khatatba. Kom Hamada. 

MATRUH: Burg el Arab, EI Daba, Mersa Matruh, Sidi Barrani 32 km. W, Bir 
Bosslanga (Bir Wair). 



OSBORN&HELMY: MAMMALS OF EGYPT 355 

Jaculus jaculus butleri Thomas, 1922 

Jaculus jaculus butleri Thomas, 1922, Ann. Mag. Nat. Hist., (ser. 9), 9, p. 296. 
Jaculus jaculus elbaensis Setzer, 1955, Proc. Biol. Soc., Wash., 68, p. 183. 

Type locality. -Sudan. KHARTOUM: Khartoum. 

Distribution in Egypt— Figure 104. Southern part of Eastern 
Desert. 

External characters.— Dorsum brownish orange; side and hip 
band white with pale orangish and grayish tipped hairs. Mystacial 
and preorbital areas white; supraorbital spot whitish, not promi- 
nent; postauricular spot small, white. Black tail band complete on 
underside in 90 per cent of specimens (table 42). 

Cranial characters.—See species description. Angle of lower jaw 
with two foramina in 40 per cent or one foramen in 60 per cent of 
specimens (table 42). 

Measurements.— Table 43, Figure 101 . Jaculus j. butleri averages 
slightly smaller in most dimensions than other Egyptian 
subspecies. 

Variation.— A few specimens approach subspecies jocu/us in hav- 
ing more orangish coloration. Intergradation cannot be 
demonstrated between butleri and schlueteri because of lack of 
material. 

Comparisons.— Jaculus j. butleri differs from J. j. schlueteri main- 
ly in shorter hind foot and ear (fig, 107) and slightly more orangish 
color. From subspecies jaculus, butleri differs in less orangish and 
more brownish color, and much larger frequency of specimens with 
black tail band complete on underside (90 per cent vs. 36 per cent) 
(table 42). 

Remarks.— We have synonymized J. / elbaensis Setzer under J. j. 
butleri Thomas, owing to similarity in color; equal numbers with 
black tail band complete on underside; comparable cranial 
characters, including frequency of two foramina in the lower jaw; 
and equal dimensions, particularly hind foot and ear length, of 
respective samples. 

Specimens examined.— Total 31. 

RED SEA: Wadi Naam (1), Bir Abraq (4). 

ASWAN: Aswan 1.6 km. SE (1), Wadi el AUaqi (1). 

SUDAN ADMINISTRATIVE: Wadi Ibib (2); Wadi Adeib, 4.8 km. N Bir Kan- 



356 FIELDIANA: ZOOLOGY 

sisrob (4). 4 km. N of Bir Kansisrob <3); Bir Sarrara (1); Abu Ramad 4 km. N (1). Wadi 
Darawena (1). »^ 

Sudan. KASSALA: Wadi Onib (1). 

KHARTOUM: Khartoum 16 km. N (11). 

Published records.— Records are from Setzer (1955, 1958a) and 
Hoogstraal et al. (1957b) and were listed as J. j. elbaensis. 

RED SEA: Bir Abraq. Wadi Naam. 

SUDAN ADMINISTRATIVE: Wadi Darawena; Bir Sarrara; Bir Kansisrob 4 km. 
N. 4.8 km. N. 

Jaculus jaculus jaculus (Linnaeus, 1758) 

Type locality.— Northern Egypt, probably near Giza Pyramids. 

Distribution in Egypt— Figure 104. Western Desert south of 
Mediterranean Coastal Desert. 

External characters.— Dorsum orangish; side and hip band white 
with pale orangish and grayish tipped hairs. Mystacial and pre- 
orbital area white; supraorbital spot white, conspicuous; 
postauricular spot small, white. Black tail band complete on under- 
side in 36 per cent of specimens (table 42). 

Cranial characters.—See species description. Angle of lower jaw 
with two foramina in 34 per cent and one foramen in 66 per cent of 
specimens (table 42). 

Measurements.— Table 43, Figure 107. 

Variation,— Brownish individuals appear sporadically in the 
population. An albino specimen and three with white areas on the 
back were collected from the desert near Giza Pyramids. In- 
tergradation mentioned under J. j. flavillus occurs between J. j. 
jaculus and J. j. flavillus in the areas of Wadi el Natroun, Bir Vic- 
toria (Setzer, 1958a), and El Birigat. Setzer also considered the few 
orangish specimens of schlueteri east of Cairo as intergrades. 

Comparisons.— Jaculus j. jaculus differs from other Egyptian 
subspecies by greater percentage of individuals with dorsum 
orangish (table 42). From Mediterranean Coastal Desert subspecies 
flavillus, it can be distinguished by shorter ear length, and from 
schlueteri, by shorter ear and hind foot (table 43, fig. 107). Dimen- 
sions of J. / jaculus average slightly larger than in J. / butleri (table 
43. fig. 107). 

Specimens examined.— Total 155. 



OSBORN&HELMY: MAMMALS OF EGYPT 357 

EL TAHREER: El Tahreer 3.2 km. W (1). Cairo-Alexandria desert road km. 165 
(1). 

BEHEIRA: Bir Victoria (3). Wadi el Natroun (3). El Birigat 2 km. W (2), Abu el 
Matamir (1). Wadi el Farigh (1). 

GIZA: El Mansuriya (2); Abu Rawash (10); Abu Ghalib (6); Giza Pyramids 8 km. 
W (1). 8 km. NW (31; Sakkara (1); Faiyum road km. 5 (1); El Qatta (2); Bahariya 
Oasis. Bir Qasr (4). Bawiti (3), Ain Guffara (1). El Hara (1). 

EL FAIYUM: Faiyum (2); Kom O Shim (5), 3.2 km. N (2); west end of Lake 
Qarum, 3 km. N (3); Wadi MuweUih (12). 

MINYA: Hatiyet el Sunt (4), El Bahnasa (1). 

ASYUT: Beni Adi (5). 

QENA: Wadi Nassim (2), Dandara 6 to 8 km. S (1). 

MATRUH: Cairo-Alexandria desert road km. 17 (1), km. 30 (1), km. 35 (1). km. 102 
(1); Bahig 35 km. S (1), 42 km. S (1); Ilwat Hawa (5); Bir Nahid (1); Qaret el Ided (1); 
Qaret el Mashruka (2); Qasr el Qatagi (1); El Maghra (23); Siwa Oasis (1), Ain el 
Dakrur (1); Camel Pass Dune area (5). 

EL WADI EL GEDEED:tFarafara Oasis 4.8 km. NE (3); Dakhla Oasis, Balat (1); 
Kharga Oasis. Baris (10). El Gezira (2). El Kharga 8 km. S (1); Nasser Village (2); Bir 
Quiseiba (3). 

Sudan. NORTHERN: Gebel Uweinat. Karkur Murr (3). 

Published records.— Records are from Wassif (1953b) and Setzer 
(1958a). 

BEHEIRA: El Birigat 2 km. W. Abu el Matamir. El Khatatba 1.6 km. W. Kom 
Hamada, Bir Victoria, Wadi el Natroun, Zaghig. 

GIZA: Imbaba, Abu Ghalib, Abu Rawash. Sakkara. 

FAIYUM: Kom O Shim. Kom O Shim 1.6 km. N, near Lake Qarun. 

QENA: Wadi Nassim. 

MATRUH: Cairo-Alexandria desert road km. 30. Siwa Oasis. 

EL WADI EL GEDEED: Kharga Oasis. 

Sudan. NORTHERN: Gebel Uweinat. (one specimen of two was erroneously 
labelled J. deserti rams Ranck. 1968 by Osborn and Krombein. 1969). 

Family 6. Hystricidae 
Characters under species. 

Genus Hystrix Linnaeus, 1758 

Hystrix cristata Linnaeus, 1758 

Hystrix cristata Linnaeus. 1758, Syst. Nat., 10th ed., p. 56. 

Type locality.— Italy: Rome. 

General distribution. — Italy, Sicily, Mauritania, Morocco, 
Algeria, Tunisia, Libya, Egypt, Northern Sudan, Asben, Senegal. 



358 FIELDIANA: ZOOLOGY 

Common name— Crested Porcupine. 

Distribution in Egypt — Figare 104. Probably limited to cliffs 
north of Salum, if not extinct. 

Diagnosis.— Dorsum and tail pelage of long, round, hollow, black 
and white quills. Head and body length average about 600 mm. 
Toes four, five. Nasofrontal region of skull inflated, nasals extreme- 
ly long and broad. Angular process of mandible arising from outer 
side of alveoli. Crowns of cheek teeth flat, complexly folded. Dental 
formula: \,i [,1x2 = 20. 

Historical notes.— A Bedouin in Salum recalled having killed a 
porcupine in the cliffs north of there in the 1950's. He referred to it 
as "Pore epic." 

Carvings of porcupine occur in the Fifth and Sixth Dynasty 
mastaba of Pehenouka at Sakkara. 

Bisharin tribesmen claimed that porcupine occur in the Red Sea 
or Kassala District of Sudan and called it "hanhan" (Keimer, 1949). 

Quills reported by Wassif (1953b) and Hoogstraal (1963) from Ain 
Gudairat (Ain el Gedeirat), 90 km. SE of El Arish in northeastern 
Sinai may have been of H. indica which ranges from Saudi Arabia, 
Israel, Syria, and Turkey eastward into India and Ceylon. 



ORDER CARNIVORA 

Key to Egyptian Families of Carnivora 
Cranial and Dental Characters 

1. Upper tooth row longer than one-half skull length. 

a. Alisphenoid canal present. Rostrum narrow. Paroccipital process not extend- 
ing below bulla Family 1 . Canidae, p. 359. 

b. Alisphenoid canal lacking. Rostrum broad. Paroccipital process extending 
below bulla Family 4. Hyaenidae, p. 422. 

2. Upper tooth row shorter than one-half skull length. 

a. Lower cheek teeth three. Rostrum short, broad. Cranium short, rounded. 
Postpalatal foramen on maxillopalatine suture Family 5. Felidae, p. 434. 

b. Lower cheek teeth four or more. Cranium elongate. Postpalatal foramen on 
maxilla. 

i. Rostrum relatively short. Bulla not constricted nor divided by septum 

Family 2. Mustelidae, p. 395. 

ii. Rostrum relatively long. Bulla constricted and divided by septum 

Family 3. Viverridae, p. 410. 

External Characters 

1. Limbs long. Features dog- or cat-like. Hind foot with four toes. 

a. Dog-like. Muzzle long. Tail bushy. Claws blunt, nonretractile. 

i. Pelage never striped or spotted. Muzzle slender. Fore and hind limb length 
equal Family 1 . Canidae, p. 359. 

ii. Pelage striped. Muzzle broad. Hind limb shorter than fore 

Family 4. Hyaenidae, p. 422. 

b. Cat-like. Muzzle short, broad. Tail cylindrical, never bushy. Pelage usually 
with stripes and/or spots. Claws sharp, curved, and retractile (except in 
Acinonyx) Family 5. Felidae, p. 434. 

2. Limbs short. Features weasel-like. Hind foot with five toes. 

a. Pelage plain or striped. Tail slender or bushy, two-thirds or less than length of 
head and body Family 2. Mustelidae, p. 395. 

b. Pelage coarsely grizzled or spotted and striped. Tail cylindrical to somewhat 
bushy, longer than two-thirds of head and body. . Family 3. Viverridae, p. 410. 

Family 1. Canidae 
Carnivores with dog-like features. Muzzle elongate; ears large, 
erect, pointed; legs long in proportion to body length. Feet 

359 



360 FIELDIANA: ZOOLOGY 

semidigitigrade, toes 5-4, inner toe of forefoot vestigial; claws blunt, 
nonretractile. Tail long, two-thirds of or more than length of head 
and body, bushy, and with scent gland near dorsal base. 

Rostrum and nasals elongate, upper tooth row length equal to or 
greater than one-half skull length. Paroccipital process prominent, 
protruding. Tympanic bulla conspicuously inflated, septum lacking. 
Alisphenoid canal present. Baculum well developed, grooved on 
underside. 

Incisors unspecialized; canines long, powerful; premolars sharp; 
carnassials well developed, modified for cutting and crushing; re- 
maining molars are the crushing type. Dental formula: |, \, \, 
fx2=42. 

Key to Egyptian Genera of Canidae 

1. Dorsum blackish, side yellowish. Tail brush-like, t<p black. Frontal region of skull 
strongly elevated. Postorbital process convex above. Postorbital swelling pre- 
sent. Cranium broadest at bases of zygomatic processes of temporals 

Canis, p. 360. 

2. Dorsum reddish, side grayish to buffy. Tail brush-like or bushy and club-shaped, 
tip black or white. Frontal region of skull not elevated or slightly elevated. 
Postorbital process concave above. Postorbital swelling absent. Cranium 
broadest on sides, narrower at bases of zygomatic processes of temporals. 

a. Tail relatively long (62 per cent of head and body length), cylindrical, tip white. 
Dorsal stripe broad, prominent. Side grizzled gray and yellowish. Skull ridged 
in adults. Frontal region not elevated Vulpes, p. 371. 

b. Tail relatively short (55 per cent of head and body length), brush-like, tip black. 
Dorsal stripe narrow, inconspicuous. Side pale yellowish buff. Skull smooth or 

with inconspicuous lyre-shaped ridges. Frontal region elevated slightly 

Fennecus, p. 387. 

Genus Canis Liimaeus, 1758 

Dog-like carnivores with broad dorsal mane. Tail relatively short, 
brush-like, tip black. Pupil of eye round. Frontal region of skull in- 
flated. Postorbital process convex dorsally, lacking posterior ridge. 
Postorbital region swollen. Posterior end of nasals at level of or 
posterior to frontomaxillary suture. Cranial ridges high and promi- 
nent. External occipital protuberance extends caudad of occipital 
condyle. Tip of zygomatic process thin. Cranium broadest at base of 
zygomatic process. Canines relatively short, thick; point of upper 
canine does not reach level of mental foramen when jaws are closed. 
Cheek teeth heavy, with or without cingula. 

Canis aureus Linnaeus, 1758 

Canis aureus Linnaeus, 1758, Syst. Nat., 10th ed., p. 40. 



OSBORN&HELMY: MAMMALS OF EGYPT 361 

Type locality. -Iraji, LARISTAN (now FARS). 

General distribution.— IhaiXsind and Burma west throughout 
India and Ceylon, Pakistan, Afghanistan, southern Turkestan, 
Iran, Iraq, Transcaucasia, Turkey, southern Russia and 
southeastern Europe, Syria, Lebanon, Jordan, parts of Saudi 
Arabia, Israel, Sinai Peninsula, Egypt, Sudan, Ethiopia, Somalia, 
and Kenya. Libya west to Morocco and Rio de Oro and south to 
Senegal. 

Common names.— Jackal, Deeb, Abu Soliman. 

Subspecies in Egypt — 

Canis aureus lupaster (Hemprich and Ehrenberg, 1833) 

Canis lupaster Hemprich and Ehrenberg, 1833, Symbolae Physical Mamm., Dec. 
2. foUo ff. 

Type locality.— Egypt. EL FAIYUM. 

Distribution in Egypt— Figure 109. Sinai Peninsula, Nile Delta 
and Valley and bordering deserts. Western Mediterranean Coastal 
Desert, and oases of the Western Desert. 

Diagnosis. —Size and appearance hke a large blackish yellow dog 
with a dorsal mane. Tail relatively short, brush-like, black dorsally 
and on tip. Black marking on anterior of forelimb. 

Skull dog-like, frontals inflated. Cranial ridges prominent. Exter- 
nal occipital protuberance extends caudad of occipital condyle. 
Postorbital process large, convex dorsally. Nasal bones taper 
gradually posteriorly, terminating at level of or posterior to fronto- 
maxillary suture. Bulla rounded, smooth. Jaw teeth in line, not 
crowded. Last lower premolar with two posterior cusps and a 
cingulum. 

Adult head and body length average 872 mm.; tail 312 mm., 36 
per cent of head and body length; foot 200 mm.; ear 112 mm.; con- 
dyloincisive length 185 mm.; weight 13 kg. 

External characters.— Dorsal mane of long, coarse, black-tipped 
hairs with yellowish subterminal bands and buff to whitish bases, 
extends from crown to base of tail and onto shoulder and hip. 
Agouti nature of hairs on hip gives an impression of broken stripes. 
Side yellowish, with scattering of black- and white-tipped hairs. 
Chin grayish; throat, belly, and inside of legs whitish to yellowish. 
Chest with medial strip of black-tipped hairs. Axillary, inguinal, and 



362 



FIELDIANA: ZOOLOGY 



,. 25* 26* 27* 28* 2 9* 30* 31* 32* 33* 34* 35* 36* 37 




Fici 109. Collection localities of Canis aureus lupaster and sight records (S). 

perineal areas whitish. Hairs on side of neck whitish, black-tipped. 
Muzzle rufous grizzled with white. Frontal area and cheek grizzled 
white, yellowish, and black. Crown rufous or grizzled rufous, 
yellowish, and black. Etir rufous behind, buffy inside with black- 
tipped hairs on anterior margin. Legs deep buff to rufous on outer 
side. Feet orangish buff; hair of palm and sole rufous, not covering 
pads. Prominent black stripe along anterior of foreleg. 

Pelage of young pale brownish above from snout to rump with 
narrow black dorsal stripe. Throat and belly buff. Short, faint black 
stripe on anterior of lower foreleg. 

Cranial characters. — Figyire 110. Skull large, elongate; frontal, 
sagittal, and lambdoidal ridges strongly developed. External oc- 
cipital protuberance prominent, extending considerably posterior to 
level of occipital condyle. Frontal region inflated. Postorbital pro- 



OSBORN & HELMY: MAMMALS OF EGYPT 



363 





•i 




Fig. 1 10. Skull of Canis aureus lupaster. 



364 FIELDIANA: ZOOLOGY 

cess convex above, posterior ridge lacking. Nasals long, tapering 
gradually posteriorly, terminating at level of or posterior to level of 
frontomaxilltiry suture. Postorbital region swollen. Paroccipital pro- 
cess large, projecting. Bulla large, rounded, and smooth. 

Bacu/um.— Baculum similar to that of Vulpes sp., but smoother, 
longer, and relatively slenderer. Deep, thin-edged ventral channel 
extends about four-fifths of length from base. Cross-section 
triangular for that distance, then changes to elongate, rounded tip 
(Didier, 1946). Total lengths of two adult bacula, 70 and 73 mm. 

Teeth.— Teeth, similar to Vulpes sp., but much larger and heavier, 
crowns relatively higher, canines heavier and shorter. Upper canine 
tip does not reach level of mental foramen when jaw is closed; P4 has 
two posterior tubercles and a cingulum. Cingula on outer borders of 
m' wide and conspicuous, but cusps narrow. Lower carnassial has 
large metaconid, heel area nearly one-half that of rest of tooth, and 
cusps strongly developed (fig. 110). 

Measurements.— Table 44. Male and female dimensions appear to 
be subequal, but Anderson (1902) said females were much smaller 
than males. 

Age determination.— Adults have frontoparietal ridges fused 
posterior to frontoparietal suture, forming a high sagittal ridge, 
cranial sutures closed, teeth worn. Teeth show varying degrees of 
wear, greatest in desert specimens. 

Variation. — Width of dorsum varies individually and in- 
dependently of width of mane. Tone of rufous color on snout, back of 
ears, and feet also varies individually. Specimens from desert 
localities slightly paler than those from the Nile Delta and Valley. 
Flower (1932) said jackals from northwestern Egypt were smaller 

Table 44. — Means (and ranges) of measurements, ratios, and weight of adult 
Cams aureus lupaster. 



HBL 


871.6(822-923)9 


RW 


33.8 (31.2-37.8) 14 


TL 


312.8 (290-347) 9 


POW 


34.8(31.8-38.9) 13 


TL/HBL% 


35.8 (33.7-39.5) 9 


BCW* 


54.4 (51.8-59.1) 13 


FL 


199.8(190-212)9 


NL 


72.0 (65.9-84.8) 14 


EL 


112.4(104-121)9 


M'-M' 


58.2 (54.2-63.0) 14 


Wt (kg.) 


13.0(10.0-15.0)4 


C-M' 


80.3 (75.6-86.8) 14 


CIL 


185.2 (173.5-196.0) 13 


SH 


66.8 (62.0-74.0) 9 


zw 


101.4(93.5-111.4) 14 







*At level of tempoparietal suture. 



OSBORN&HELMY: MAMMALS OF EGYPT 365 

than those from the Nile Delta and considered them as subspecies 
tripolitanus. Setzer (1957c) did not concur, nor do the data herein. 
Determination of intergradation in this area awaits further 
collecting. 

Comparisons.— Canis aureus is distinguishable from other Egyp- 
tian Canidae in having the dorsum blackish and maned, frontal 
region of skull elevated, a prominent postorbital swelling, cranium 
broadest at bases of zygomatic processes, and larger dimensions 
(tables 44, 45, 46). Canis a. lupaster is considered to be the largest 
and darkest of North African and Southwest Asian subspecies 
(Pocock, 1941). 

Large canid skulls found in the desert are usually dog. Some have 
been deposited in museums and labelled "jackal." Following are 
characters that distinguish dog from jackal skulls. No single 
character, however, is always reliable. 

(1) Greater inflation of frontal region between postorbital pro- 
cesses. 

(2) Forehead steeper due to lesser angle between rostrum and 
cranium. 

(3) Dorsal surface of braincase lower relative to postorbital pro- 
cesses. 

(4) Postorbital region more inflated and elongated. 

(5) Auditory bulla flattened, less inflated, and surface rough 
rather than smooth. 

(6) Zygomatic arch with greater upward curvature. 

(7) Snout and palate usually shorter and broader. 

(8) Jugular or paroccipital process heavier and more protruding. 

(9) Hypoglossal foramen on transverse ridge of above usually 
more anterior and ventral in position. 

(10) Tuberosities on basioccipital larger. 

(11) Lower jaw shorter, thicker and more curved. 

(12) Posterior margin of ramus concave rather than straight. 

(13) Level of last lower molar above others. 

(14) Canines and other teeth generally larger. 

(15) Cingula lacking on first and second upper premolars. 

(16) Teeth usually crowded and set at angles rather than in line. 

(17) Fourth lower premolar with one rather than two posterior 
tubercles and lacking cingulum (fig. 111). 

(18) Posterior heel of lower carnassial relatively narrower and 
cusps less strongly developed. 



366 FIELDIANA: ZOOLOGY 

Table 45. — Means (and ranges) of measurements, ratios, and weight of adult 
male (M) and female (F) Vulpes vulpes aegyptiaca. 







Giza 


Bahig 


HBL 


M 


602.1 (541-652)9 


554.4 (533-592) 8 




F 


589.6 (552-634) 24 


513.4 (465-570) 7 


TL 


M 


374.6 (343-401) 9 


349.5 (326-398) 8 




F 


361.6(307-391)23 


320.8 (302-368) 7 


TL/HBL% 


M 


62.9 (58.7-67.2) 8 


62.8 (57.3-67.2) 8 




F 


61.4 (58.2-74.4) 23 


62.8 (52.9-72.8) 7 


FL 


M 


149.8(136-165)9 


137.5(127-149)8 




F 


148.7(134-160)24 


123.3(105-134)6 


EL 


M 


99.0 (93-109) 9 


93.1 (91-106)8 




F 


97.7 (91-106) 24 


88.6 (70-104) 7 


Wt (kg.) 


M 




6.0 




F 


4.6. 


4.8. 4.8. 5.5 


CIL 


M 


133.7 (125.5-142.3) 13 


124.2(119.8-132.9)7 




F 


130.7 (123.0-139.6) 22 


115.6(109.5-120.4) 13 


ZW 


M 


71.2(65.1-76.5) 12 


66.8(62.5-69.1) 11 




F 


70.2 (66.2-75.8) 24 


63.7 (59.8-69.1) 13 


RW 


M 


21.8(20.5-22.6) 13 


20.2(19.1-22.3) 12 




F 


21.6(19.3-22.5)23 


18.7(17.1-20.1) 13 


POW 


M 


21.0 (19.5-22.8) 12 


21.2 (19.7-23.4) 12 




F 


21.2(18.9-25.6)23 


21.8(19.8-23.2) 13 


BCW 


M 


45.6 (43.0-50.9) 12 


44.1 (42.7-45.4) 12 




F 


44.6 (42.8-46.5) 23 


43.0(41.6-55.3) 13 


NL 


M 


48.9 (38.4-56.7) 13 


44.5(41.5-49.6) 11 




F 


48.6 (45.3-52.6) 23 


41.8 (38.2-48.2) 13 


C-M» 


M 


61.0 (55.3-64.5) 13 


57.1 (55.0-62.1)9 




F 


59.7 (56.8-63.6) 22 


54.4 (51.4-57.0) 12 


M-M' 


M 


37.8 (35.0-39.9) 13 


35.4(33.1-36.9) 10 




F 


37.2 (35.5-40.6) 22 


34.2 (32.9-36.8) 10 


SH 


M 


47.8(44.0-49.1) 12 


45.8 (44.6-46.7) 8 




F 


46.8 (45.4-48.6) 10 


44.0 (43.2-45.4) 8 



Discussion of characters of domestic versus wild canids are in 
Gidley (1914), Lawrence (1967), and Lawrence and Bossert (1967). 

/Jemar^s. — References to wolves in literature on Egypt can be in- 
ferred as meaning jackals. The Arabic name, Deeb, is applied to 
both C. lupus and C. aureus. Embalmed remains of jackals and dogs 
have been reported from tombs near Asyut, ancient Egyptian city 
of Anubis, the "jackal" god, which was also known as Lycopolis, 
city of wolves, during the Ptolemaic period (Ebers, 1878; Murray, 



OSBORN&HELMY: MAMMALS OF EGYPT 367 

Table 46. — Means (and ranges) of measurements, ratios, and weight of adult 
male (M) and female (F) Vulpes r. rueppelli. 







Sinai Peninsula 


Eastern Desert 


Western Desert 


HBL 


M 





468.5 (428-519) 15 


463.2 (419-476) 13 




F 


505.0 (469-559) 3 


442.5(411-468)8 


449.0 (418-460) 5 


TL 


M 





343.6 (290-387) 15 


337.8 (305-380) 13 




F 


308.6 (281-363) 3 


326.4 (273-354) 8 


318.8 (292-345) 5 


TL/HBL% 


M 





73.3 (67.4-78.8) 15 


73.4 (67.6-83.8) 13 




F 


61.6 (50.4-74.5) 3 


73.8 (61.0-77.9) 8 


71.0 (68.4-75.0) 5 


FL 


M 





127.0(115-138) 16 


126.6(119-134)13 




F 


121.6(110-129)3 


121.5(112-131)8 


121.4(119-125)5 


EL 


M 





101.6(95-110) 14 


95.0 (89-107) 13 




F 


102.3 (96-108) 3 


97.5(93-110)8 


91.2 (88-94) 5 


Wt (kg.) 


M 





1.9 ( 1.4- 2.3)6 


1.7 ( 1.4- 1.9)7 




F 





1.5, 1.7 


1.7 ( 1.4- 1.8)4 


CIL 


M 


109.0 


106.4(98.2-113.9) 15 


106.1 (97.5-110.5) 13 




F 


105.9(110.2-108.8)4 


102.3 (99.0-105.8) 8 


101.2(91.0-103.8)5 


ZW 


M 


59.6 


60.0 (55.5-65.6) 15 


58.1 (55.6-62.4) 13 




F 


58.8 (56.7-60.3) 4 


57.3 (56.4-58.4) 8 


57.8 (55.7-59.8) 5 


RW 


M 


17.7 


16.8 (15.3-18.4) 15 


16.5(15.7-17.3)13 




F 


17.2 (16.8-17.4) 4 


16.8(15.9-17.5)8 


16.2(15.2-18.1)5 


POW 


M 


22.0 


20.0 (17.9-23.4) 15 


20.4 (18.6-22.9) 13 




F 


20.2 (18.3-22.9) 4 


20.6 (18.8-21.9) 8 


20.2 (19.2-22.4) 5 


BOW 


M 


42.0 


40.3 (38.8-42.0) 15 


40.2 (38.0-42.0) 13 




F 


39.3 (38.9-39.7) 4 


39.9 (38.0-41.6) 8 


39.3 (38.4-40.8) 5 


NL 


M 


38.5 


36.5 (31.8-41.1) 15 


38.2 (34.4-41.2) 13 




F 


36.6 (34.6-38.3) 4 


35.4 (33.0-38.4) 8 


36.9(36.1-38.1)5 


C-W 


M 





49.4 (46.4-52.4) 13 


50.0 (43.7-52.9) 13 




F 


48.2 (44.7-49.8) 4 


47.4 (44.6-49.6) 7 


48.6 (46.8-51.2) 5 


M'-M' 


M 





31.0 (29.5-33.5) 13 


31.2 (29.9-33.4) 12 




F 


31.7 (29.6-33.1) 4 


30.3(29.3-31.1)6 


31.4 (29.5-34.0) 5 


SH 


M 


42.9 


41.0 (39.6-42.4) 15 


41.4 (40.3-42.5) 13 




F 


41.0(40.6-41.3)4 


41.4 (40.7-42.4) 8 


41.2 (40.4-42.5) 5 



1891; Gaillard, 1927). In 1965, we searched many of the Gebel 
Durunka tombs, but found only fox (Vulpes vulpes) remains, along 
with mummified humans. We are inclined to believe that Anubis 
was a fox rather than a jackal, because all of the statuary and 
heiroglyphs of Anubis are of an animal with the tail of a fox. 

References to wolves in Sinai could possibly mean C. lupus. 
Palmer (1872) accounted the fighting of dogs with a large "wolf" in 
Wadi Aleyat, a tributary of Wadi Feiran near the Oasis. He also 



368 



FIELDIANA: ZOOLOGY 




JACKAL 




DOG 

Fig 111. Left mandibular teeth (pnij. ^; m,) of jackal and dog. Drawings are not to 
same scale. See explanation in text. 

found remains of a camel in Wadi Gharandel which, according to 
Bedouins, had been killed by a pack of wolves. Hart (1891) made no 
mention of wolves in Sinai, but several "wolves" were said to have 
been shot during an expedition to Sinai in 1927 (Bodenheimer and 
Theodor, 1929), and Howells (1956) reported wolves in Wadi Gaz- 
zah, northeastern Sinai. Wolves are still widespread in the Arabian 
Peninsula (Harrison, 1968), but are now considered to be intruders 
in Israel from the east and north (Bodenheimer, 1958). Wolves in 
Sinai could doubtlessly be considered as strays. 

Specimens examined,— Total 62. 



OSBORN&HELMY: MAMMALS OF EGYPT 369 

DAQAHLIYA: Faraskur (2). 

ALEXANDRIA: Ramleh (1). 

MINUFIYA: Mohammed Ali Barrage Park (1). 

QALYUBIYA: Qalyub (1); Sindbis (1); Sanafir. Ezbet Ibsan (1). 

GIZA: Giza (2); Imbaba (1); Abu Rawash (3): Abu Ghalib (1); Giza Pyramid area 
(1). 16 km. W (1); Badrshein (1); Kirdasa (1); El BaragU (1). 

EL FAIYUM: Faiyum (1); Kom O Shim (2), 1.6 km. W (1). 10 km. N (1); Kom O 
Shim Forest (10); Tamiya (3). 

BENI SUEF: Beni Suef (1). 

QENA: Qena-Safaga road km. 5 (1), Luxor (1). 

ASWAN: Wadi Allaqi (1). Wadi AUaqi 11.2 km. S of AUaqi Village (2|. Wadi el 
Targama (1). Khor Abusku S of Wadi Nogdeb (8). 

MATRUH: El Afritat 17 km. SW of El Hammam (1); Alam Shaltut 72 km. S of 
Bahig (1); between Nakhlet el Barraq and Qaret el Mashruka (1); Bir Shaqqa 
(1); Siwa Oasis 20 km. E of (1). Aghurmi (2). 

EL WADI EL GEDEED: Dakhla Oasis. Dakhla (2); El Gezira SE of Kharaga 
(1). 

Sight records of I. Helmy and D. Osbom.— 

CAIRO: Helwan. 

EL FAIYUM: El Mishigeiga. 

MATRUH: Nuweimesa, Wadi Labaq (tracks). 

ASWAN: Road between Idfu and Kom Ombo. 

Published records.— Records sire from Anderson (1902), Harding- 
King (1925), Flower (1932), Omer-Cooper (1947), Tregenza (1955), 
Howells (1956), and Setzer (1961b). 

SINAI: Wadi Gazzaht (probably C. a. syriacus). 
RED SEA: Wadi Midhais. 
DAQAHLIYA: Faraskur. 
QUALYUBIYA: Sindbis. 

GIZA: Giza, Abu Rawash, Abu Ghalib, El Baragil, Bahariya Oasis. 
EL FAIYUM: Faiyum, Kom O Shim 1 km. W, Lake Qarun. 
BENI SUEF: Beni Suef. 
QENA: Luxor, Thebes. 
ASWAN: El Dirr (sight record). 
BEHEIRA: Wadi el Natroun. 

MATRUH: Mersa Matruh, Redunkalil Tutuatee, Sitra, Salum (sight record). El 
Alamein (sight record). 

EL WADI EL GEDEED: Dakhla Oasis, Rashida, Mut, Kharga Oasis (sight 
record). 

Collection,— Shot at night using spotlight. Trapped with sardine 
bait. 



370 FIELDIANA: ZOOLOGY 

Habitats. — Nile Delta and Valley and adjacent desert; Western 
Mediterranean Coastal Desert; rocky area» in southern edge of 
Coastal Desert; oases. Frequently seen in isolated cliffs and rocky 
hillocks in semi-barren desert. 

Dens.— Tombs, natural caves, and crevices. 

//a6z7s.— Nocturnal, but often seen in late afternoon. 

Food— Jackals are known to eat dates, mulberries, apricots, and 
other fruits in season. North of El Faiyum, they supposedly lived on 
fish that were easily caught in shallow water {Anderson, 1902). They 
are attracted to carcasses (Dorst, 1970), and Manson-Bahr (1936) 
shot jackals beside a dead donkey near Cairo. 

Jackals living near the Nile Valley and Delta are reputed to feed 
on various cultivated crops and fruit and to prey upon domestic 
animals (Flower, 1932; Setzer, 1961b; Hoogstraal, 1964). Howells 
(1956) wrote that, in Wadi Gazzah, northeastern Sinai, jackals 
entered camps and villages to kill chickens, young goats, and sheep. 
Corn, watermelons, pumpkins, and grapes were also eaten, and 
jackals reportedly destroyed corn and melon patches. 

Kasim (1912) found evidence that a great number of desert snails 
{Eremica desertorum) were eaten by jackals near Bir el Malla on the 
Sidi Barrani-Siwa road. Sandford (1936) discovered a mud pan in 
northeastern Sudan where jackals had dug hundreds of Pila (Am- 
pullaria of authors) wernerei, an operculate fresh water snail, out of 
the cracks. 

Stomach contents of a jackal trapped near Bir Shaqqa contained 
shell fragments of the desert snail, gazelle remains, wings of a hawk 
moth {Acherontia atropos), a muscid fly, and a fly larva. In Wadi 
Allaqi, Osborn (1968a) shot a jackal and found its stomach to be full 
of fish scavenged from the waste of fishermen. 

In Iran, stomachs of jackals examined by Lay (1967, p. 20^) con- 
tained "grasshoppers, grapes, blackberries, grain seed, dates, 
freshwater crabs, carrion, and one Mus musculus." 

Although some authors are inclined to consider the jackal a 
scavenger (Setzer, 1952, 1961b; Hoogstraal, 1964; Dorst, 1970), 
evidence indicates that it is an opportunistic omnivore. 

Reproduction. — Flower (1932) recognized a definite breeding 
season. Nine wild litters were born in March, April, and May. Thirty 
litters were born in Giza Zoological Gardens between the second 



OSBORN&HELMY: MAMMALS OF EGYPT 371 

week in March and first week in May. The average litter size was 
4.5, maximum eight. 

Fo/^/ore. — According to Arab belief, the jackal is feared more 
than the fox by poultry, and if a jackal happens to pass under 
roosting chickens, they will all fall down from fear. A jackal's 
tongue left in a house will supposedly cause the inmates to argue 
and become hostile to one another. The flesh is considered to be 
useful in cases of madness and epilepsy. The right eye is said to pro- 
tect the wearer from the evil eye and the heart to protect him from 
attack by beasts of prey (Jayakar, 1906). 

Genus Vulpes Oken, 1816 

Reddish foxes. Tail relatively long, bushy, and club-shaped; tip 
white. Pupil of eye elongate vertically. Frontal region of skull not in- 
flated. Posterior end of nasals anterior to posterior level of fronto- 
maxillary suture. Cranium broadest on sides, narrower at base of 
zygomatic processes of temporals. 

Postorbital process concave dorsally, posterior margin ridged. 
Postorbital swelling nil. Vertical curvature of zygomatic arch high. 
Tip of zygomatic process of temporal relatively thick. Canines 
slender, elongate; point of upper canine reaching level of mental 
foramen when jaws are closed. Cheek teeth narrower and more tren- 
chant than in genus Canis. 

Key to Egyptian Species of Vulpes 

1. Size large, foot 100-150 mm., skull length 110-145 mm. Back of ear black. Belly 
blackish. Black mark on foreleg. Nasomaxillary contact narrow or lacking (fig. 
112) vulpes, p. 371. 

2. Size smaller, foot 90-115 mm., skull length 90-109 mm. Back of ear pale brown. 
Belly white. No black mark on foreleg. Nasomaxillary contact broad (fig. 112) 
rueppelli, p. 379. 

Vulpes vulpes (Linnaeus, 1758) 

Canis vulpes Linnaeus, 1758, Syst. Nat., 10th ed., p. 40. 

Type locality. —Sweden: Upsala. 

General distribution.— Eastern North America, British Isles, 
Europe, Asia (northern and southwestern), Saudi Arabia, Sinai 
Peninsula, Egypt and Sudan west to Morocco. 

Common names. — Red Fox, Nile Fox, Taaleb, Abu Hussein. 

Subspecies in Egypt.— 



372 



FIELDIANA: ZOOLOGY 





V.VULPES 



V.RUEPPELLI 



Fig 112. Comparison of nasomaxillary contact and shape of posterior margins of 
nasals in Vulpes vulpes and V. rueppelli. 

Vulpes vulpes aegyptiaca (Sonnini, 1816) 

Canis aegyptiacus Sonnini. 1816, Nouv. Diet. Sci. Nat.. Vol. VI, p. 524. 

Type locality.— Egypt. GIZA: Giza Pyramids. 

Distribution in Egypt— Figure 113. Sinai Peninsula, northern 
part of Eastern Desert, Nile Delta and Valley, Western Mediterra- 
nean Coastal Desert. 

Diagnosis.— harge fox. Ear relatively large, black posteriorly. 
Tail long, bushy, and club-shaped; tip white. Dorsum reddish to red- 
dish brown; side yellowish gray; venter brownish or blackish. 
Foreleg with prominent, elongate, black marking. 

Cranial ridges prominent. Superior edge of lambdoidal ridge ex- 
tending beyond level of occipital condyle. Postorbital process con- 
cave dorsally. Nasal bones taper gradually posteriorly. Nasomax- 
illary contact nil or very narrow. 

Adult head and body length average of male and female, respec- 
tively, 578, 551 mm.; tail 362, 341 mm., 62 per cent of head and body 
length; hind foot 143, 136 mm.; ear 96, 83 mm.; condyloincisive 
length 128, 123 mm. 



OSBORN & HELMY: MAMMALS OF EGYPT 



373 



25* 26' 27* 28* 2 9* 30* 31* 3 2* 3 3* 34* 35* 36* 37* 




Fig. 113. Collection localities of Vulpes vulpes aegyptiaca. 

External characters.— Dorsal stripe reddish to reddish brown, 50 
to 80 mm. wide, extending from eye to basal one-third of tail, 
broadest on shoulders (forming a "cross") and on pelvis, darkened 
between ear and shoulder by black and black-tipped guard hairs. 
Grizzling, due to long guard hairs with blackish tips and white 
subterminal bands, occurs over entire dorsum, is most pronounced 
on shoulder and hip, and occurs on cheek, throat, and chest. Side 
grizzled gray and yellowish. Throat and belly brownish or blackish. 
Muzzle buff dorsally to about level of eye, reddish laterally. Chin 
whitish. Cheek deep buff to reddish. Dark stripe from mystacial 
area to eye indistinct; prominent on juvenile pelage only. Hairs on 
side of neck buffy with black tips. A whitish or buffy stripe 
sometimes separates this marking from brown of throat. Axillary 
and groin patches whitish, buffy, or orangish. Ear whitish to cream 
on inner side with brush of long hairs on lower medial margin. Back 



374 FIELDIANA: ZOOLOGY 

of ear black, base brownish or color of back. Tail long, bushy, and 
club-shaped; reddish or brownish dorsally; paler ventrally, hairs 
buffy with dark tips; gland on upper base marked with blackish 
hairs; tip white. Foreleg with brownish and whitish markings and 
prominent black anterior stripe. Back of foreleg and foot brown to 
reddish brown. Hind limb similarly marked, black usually limited to 
upper foot. Hair of palm and sole not covering pads. 

Juvenile pelage brownish dorsally; side, throat, and belly grayish. 
Black stripe on foreleg prominent. 

Cranial characters. — Figure 114. Skull elongate. Frontoparietal, 
sagittal, and lambdoidal ridges prominent. Superior edge of the 
lambdoidal extends caudad beyond level of occipital condyle. Fron- 
tal region not inflated. Postorbital process concave dorsally, 
posterior ridge conspicuous and continuous with cranial ridge. 
Nasals taper gradually posteriorly, terminating anterior to 
posterior level of frontomaxillary suture. Nasomaxillary contact 
very narrow or absent due to contact of premaxillary and frontal 
processes (fig. 112). Paroccipital process large, prominent, and at- 
tached to bulla. Bulla large, rounded, and smooth. 

Baca/um.— Baculum triangular in cross-section, ventral channel 
extends entire length, surface somewhat rough, base not enlarged. 
There are two obscure distal tuberosities (Didier, 1946). Total 
lengths of two adult bacula, 46 and 47 mm. 

Teeth.— Teeth of Egyptian canids are all very similar. Foxes have 
slenderer teeth with lower crowns, smaller carnassials, and greater 
relative length of canines than jackals. 

Measurements.— Table 45. Male dimensions average considerably 
larger than female. 

Age determination.— Old adults have frontoparietal ridges fused 
posterior to frontoparietal suture and forming a sharp-edged sagit- 
tal ridge, cranial sutures closed, teeth well worn. 

Adults have frontoparietal ridges forming a narrow "V" or, if 
fused, sagittal ridge rounded and without sharp edges; cranial 
sutures closed; teeth slightly worn. 

Subadults have frontoparietal ridges lyre-shaped to broadly 
V-shaped, basioccipital-basisphenoid suture usually open, teeth 
sometimes slightly worn. 

Juveniles have cranium smooth or with inconspicuous lyre- 





E 
u 




Fig. 114. Skull of Vulpes vulpes aegyptiaca. 



375 



376 FIELDIANA: ZOOLOGY 

shaped ridges, basioccipital-basisphenoid suture open, teeth 
unworn. 

VariatioTu— Adult pelages are paler in summer than in winter; 
bellies are brownish in summer, blackish in winter. Two color 
phases, brownish red with little yellow and reddish yellow, occur in 
Nile Valley and Delta populations in frequencies of about 60 and 40 
per cent, respectively. Desert specimens are paler in either of above 
color phases. Desert specimens with cranial ridge development com- 
parable to Nile Valley and Delta specimens are considerably smaller 
in all dimensions (table 45), and teeth of subadults sometimes show 
as much wear as old adults of the latter. No old adults have been col- 
lected from desert localities. Setzer's (1961b) and Hoogstraal's 
(1963) statements that foxes from the Delta are smaller than those 
from elsewhere in Egypt are erroneous. 

Comparisons.— Vulpes vulpes differs from V. rueppelli in darker 
color, back of ear being black instead of pale brown, venter blackish 
instead of white, presence of black mark on foreleg, larger average 
dimensions (tables 45, 46), more prominent cranial ridges, less in- 
flated bulla (figs. 114, 117), more gradual posterior taper of nasals 
(fig. 112), lambdoidal ridge extending posteriorly beyond level of 
occipital condyle, and narrower nasomaxillary contact (fig. 112). 

Means (and ranges) of ratios of length of nasomaxillary contact to 
nasal length x 100 in 19 juvenile and subadult V. vulpes and 52 
adult V. rueppelli, respectively, are 12 (4 to 17) and 25 (15 to 38). 

Vulpes vulpes aegyptiaca is a larger and darker race than V. v. 
arabica and V. v. palaestina. 

Specimens examined.— Total 216. 

SUEZ: Suez (1); Gebel Sukhna, N of (2): Wadi Qiseib (1); Abu el Darag (11. 1 km. N 
(1). 

SHARQIYA: Bilbeis (6). 

BEHEIRA: El Khatatba (2). Hafs (3). Damanhour (1). 

MINUFIYA: Ashmun (1). Saqyet Abu Shara (5). 

QALYUBIYA: Sindbis (15). Abu Zabal (1). 

GIZA: El Qatta (1): Ausim (3). Bashtil (2), Tanash (3). Saft el Laban (1), Warraq el 
Arab (1). Wardan (1), Minshat el Bakkari (5). Kafr Hakim (9). El Mansuriya (5). Abu 
Rawash (7). El Kom el Akhdar (1). Beni Magdul (1). Abu Ghalib (2), Nahya (2). El 
Mitimdiya (2). Kafret Nassar (1). El Talbiya (1). Imbaba (3). Ezbet Moneib (1). Giz- 
zaya (2), Geziret Muhamed (9), Giza Pyramids (3), Giza (6). Sakkara (5). Dahshur 
Pyramid (1). Kafr Ammar (3). El Tabin (1). 

CAIRO: HeUopoUs (1). Cairo (2). 



OSBORN&HELMY: MAMMALS OF EGYPT 377 

EL FAIYUM: Faiyum (5). Kom O Shim (4). Tamiya (3). Fanus (4). Qasr Rashwan 
(4). Lake Qarun NW end (2). 

BENI SUEF: Maidum (1). 

QENA: Wadi Qena (1). 

ASWAN: Kom Ombo (2); Aswan (3), Koror area (1): Gebel Adda (3): Wadi el 
Targama (1); Wadi AUaqi (1). 

ALEXANDRIA: Mariut (1). 

EL TAHREER: Cairo-Alexandria desert road km. 164 (1). 

BEHEIRA: Bir Victoria (1); Wadi el Natroun (5), El Beida (1). 

MATRUH: Burg el Arab (3); Bahig (17), 4.8 km. S (1). 8 km. S (1), 15 km. S (1), 18 
km. S (1). 6 km. SW (1). 9 km. SW (4|; El Qarasat (1), 9 km. SW (2); El Afritat. 17 km. 
SW El Hammam (2); Abu Hagag (1); Salum 4.8 km. SE (1), Bir Wair (1). 

EL WADI EL GEDEED: Kharga Oasis (2); Dakhla Oasis, Mut (1), Asment (2), El 
Sheikh el Waly (2); Balat (1). 

Sudan. NORTHERN: Wadi Haifa (2). 

Published records.— Records are from Anderson (1902), Bonhote 
(1902), Flower (1932), Tregenza (1958), and Setzer (1952, 1961b). 

SINAI: Ayun Musa, various northern coastal locahties. 

SUEZ: Gebel Sukhna, N of. 

RED SEA: Gemsa {report from guide). 

SHARQIYA: Bilbeis, Faqus. 

DAQAHLIYA: SimbUlawein 8 km. W. 

BEHEIRA: Hafs. 

MINUFIYA: Ashmun. 

QALYUBIYA: Sindbis, Kanka, Abu Zabal. 

GIZA: Abu Ghalib, Imbaba, El Mitimdiya, Gizzaya, El Mansuriya, El Kom el 
Akhdar, Kafret Nassar, Nahya, Kafr Hakim, El Talbiya, Giza Pyramids area, 
Tanash, Bamha, Kafr Ammar, El Tabin, Abu Rawash, Ezbet Moneib, Bashtil, Min- 
shat el Bakkari, Geziret Muhamed, Warraq el Arab, El Baragil, Beni Magdul, Aiyut 
Barnasht, Dahshur Pyramid, Badrshein. 

CAIRO: Hehopolis. 

EL FAIYUM: Qasr Rashwan, Fanus, Lake Qarun W end 3 km. N, Tamiya. 

BENI SUEF: Maidum. 

BEHEIRA: Bir Victoria, between Bir Victoria and El Khatatba, Wadi el Natroun. 

MATRUH: Bahig 4.8 km. S, Burg el Arab. 

Sudan. NORTHERN: Wadi Haifa. 

Collection.— Shot at night under a spotlight; trapped near bur- 
rows or in foraging areas, usually with sardine bait. 

Habitats. —Sinai Peninsula: Northern coastal desert according to 
Flower (1932) and reported from vicinity of Ayun Musa (Anderson, 



378 FIELDIANA: ZOOLOGY 

1902). Later collectors failed to find this species in Sinai (Wassif, 
1954c; Wassif and Hoogstraal. 1954; Hoogstfaal. 1964). 

Eastern Desert: Northern part. Vegetated wadis from Abu el 
Darag northward. Also reported from Gemsa (Tregenza, 1958) and 
seen occasionally along Cairo-Suez road. 

Nile Valley and Delta: Inhabits date and fruit groves, cultivated 
areas, and suburban gardens. Dens in desert hills allow easy access 
to cultivated areas. 

Western Mediterranean Coastal Desert: Common throughout this 
area (figs. 8, 19), occasionally southward in semibarren and barren 
desert. 

Oases: Known to occur in Wadi el Natroun, El Faiyum, and 
Kharga and Dakhla Oases where habitats are similar to those of the 
Nile Valley. 

Habits. — Vulpes vulpes is not strictly nocturnal and is commonly 
seen during daylight hours. Foxes will often run out of dens when 
approached by a vehicle or a man on foot. 

Dens.— Dens usually have several openings and preferred sites 
are in hillsides under rocks. One den was under an old concrete floor 
in barren gravel north of El Beida, Wadi el Natroun. Clay hills or 
harms, excavations from Roman cisterns, south of Burg el Arab are 
well-known burrowing sites (Sandford and Arkell, 1939). Foxes also 
burrow in palm groves, fields, gardens, and beneath walls, stables, 
and houses. Other common den sites are ruins, tombs, and quarries. 

Food.— Stomachs of V. vulpes have contained green figs, various 
plant remains, insect remains, bread stolen from a Bedouin camp, 
remains of birds, Gerbillus sp. Dipodillus simonU Mus musculus, 
and sardine bait. A skull of Poecilictis libyca was found beside fox 
dens in hills west of Wadi el Natroun. Flower (1932) reported one 
with its stomach full of mole crickets. He fed captive foxes plums, 
fresh dates, and raw eggs. According to Tregenza's (1958) guides, 
foxes near Gemsa on the Gulf of Suez fed on crabs and dead fish 
thrown out by fisherman. Whether vegetable material forms a 
greater part of the diet than meat is not known. 

Wafer.— Availability of water may be a factor limiting the 
distribution of V. vulpes to the Nile Valley and Delta, the Western 
Mediterranean Coastal Desert, oases, and northern parts of the 
Eastern Desert and Sinai Peninsula. The southernmost locality of 



OSBORN&HELMY: MAMMALS OF EGYPT 379 

collection on the Gulf of Suez was near Abu el Darag. That far 
south, fresh water is available at all times a few kilometers from the 
seacoast. Tregenza's (1958) guides told him the Nile fox was found 
near Gemsa, but had to go into the mountains to drink. 

Reproduction.— No data are available on reproduction of V. 
vulpes in Egypt, except the recording of four pups in March 
(Hoogstraal et al., 1957b). 

Sex ratio.— A sample of 174 museum specimens contained equal 
numbers of males and females. 

Remarks.— The ranges of V. vulpes and V. rueppelli overlap only 
slightly. The latter is the more desert-adapted, probably due to its 
smaller size plus its ability to survive in waterless areas. 

Folklore.— The fox is considered by Arabs to be a cowardly beast 
of prey; weak, wily, deceitful, and cunning. It is said to feign death 
by filling its abdomen with air so as to appear bloated and then to lie 
on its side and raise two legs high in the air. Thus it awaits the 
approach of unsuspecting prey so it can capture them. Hunting 
dogs, supposedly, are not fooled by this ruse. 

Klunzinger (1878, p. 401) told the following as an example of 
numerous tales concerning the fox's reputed cunningness: 

"A fox wanted the chickens which he saw a man carrying to 
market in a basket. The fox ran ahead of the man and played dead in 
the middle of the road. The man passed with little more than a 
glance. The trick was repeated two more times. On seeing a third 
dead fox the man decided that three fox skins would be worth carry- 
ing to market to sell. He put down the basket of chickens and went 
back along the road to retrieve the first two foxes, but returned 
empty handed only to find that the third fox had vanished and his 
chickens as well." 

Vulpes rueppelli (Schinz, 1825) 

Corns rueppelli Schinz. 1825, Das Thierreich, Vol. 4, p. 508. 
Type locality.—Sudan. NORTHERN: Dongola. 

General distribution.— Afghanistan, Iran, Jordan, Saudi Arabia, 
Sinai Peninsula, Egypt, Sudan, Somaha, Asben, Libya, Algeria. 

Common names.— Rueppell's Sand Fox, Taaleb, Abu Hussein. 

Subspecies in Egypt— 



380 



FIELDIANA: ZOOLOGY 



,. 25* 26* 27* 28* 2 9* 30* 31* 32* 33* 34* 35* 36* 37 




Fig 115. Collection localities of Vulpes rueppelli rueppeUi and sight record (S). 



Vulpes rueppelli rueppelli (Schinz, 1825) 

Distribution in Egypt— Figyire 115. Sinai Peninsula, Eastern 
Desert, Western Desert. 

Diagnosis.—SmaW fox. Ear large, pale brown or rufous posterior- 
ly. Tail long, bushy, club-shaped; tip white. Dorsum grizzled red- 
dish, side buffy gray, and venter whitish. 

Frontoparietal ridge lyre- to V-shaped. Postorbital process con- 
cave dorsally. Superior edge of lambdoidal ridge not extending 
beyond posterior level of upper lip of foramen magnum. Nasal bones 
tapering abruptly posteriorly. Nasomaxillary contact broad, always 
separating premaxilla and frontal. Bulla proportionately large. 

Adult head and body length average 456 mm.; tail 281 mm., 72.8 



OSBORN & HELMY: MAMMALS OF EGYPT 



381 



per cent of head and body length; hind foot 124 mm.; ear 96 mm.; 
condyloincisive length 104 mm.; weight 1.7 kg. 

External characters.— Fig[ire 116. Fur long, soft, dense, par- 
ticularly in winter. Dorsum grizzled reddish from nape to base of 
tail. Guard hairs with short reddish tips, broad white distal subter- 
minal bands, fuscous to reddish proximal subterminal bands, and 








Fig 116. Live specimen of Vulpes rueppelli rueppelli. 



382 FIELDIANA: ZOOLOGY 

whitish bases. Under hairs fuscous: sometimes with pale or whitish 
bases. Side grayish buff to whitish with grizzling on upper part and 
scattered black-tipped hairs on lower. Grizzling extends over 
shoulders, entire dorsum, upper side, pelvis, and upper parts of fore 
and hind Umbs. Winter pelages often have a bluish tint over the 
grizzled areas. Under hairs of side have pale buffy tips, whitish to 
grayish bases. Side of neck is yellowish buff. Axillary area is 
orangish buff. Chin, cheeks, throat, belly, and inside of legs are 
whitish. Lower throat and chest are variable, with areas of brownish 
or grayish tipped hairs. Mammary area of adult females is reddish. 
Belly hairs, except as noted, are usually white to bases; occasionally 
with dark bases. Facial area is buff to orangish buff. Mystacial area 
brownish, continuous with brownish to orangish lacrimal stripe. 
Ear white to cream anteriorly and on margin, fuscous to pale red- 
dish brown posteriorly. Crown and nape are concolorous with back 
of ear. Tail bushy, club-shaped, buff to pale orangish buff with dor- 
sal stripe of brownish to blackish tipped hairs, and tip white. Dark 
hair and depression in fur over gland on upper tail base prominent. 
Anterior foreleg and outer side of hindleg pale reddish buff. Back of 
foreleg from elbow to palm and back of thigh and sole reddish 
brown. Hair of palm and sole partly covering pads. 

Cranial characters.— Figure 117. Skull elongate. Cranial ridges 
lyre- to V-shaped, much less prominent than in V. vulpes. Superior 
margin of lambdoidal ridge not extending beyond posterior level of 
upper lip of foramen magnum. Postorbital process concave dorsally, 
posterior ridge inconspicuous, continuous with cranial ridge. Nasals 
tapering abruptly posteriorly, terminating anterior to posterior 
level of frontomaxillary suture. Nasomaxillary contact broad (fig. 
112). 

Baculum.—TYie baculum is essentially a miniature of that of V. 
vulpes, except more sharply ridged dorsally. Mean (and ranges) of 
total length of five adult bacula are 38 (34 to 41) mm. 

Teeth.— Teeth smaller and slenderer than in V. vulpes. 

Measurements.— Table 46. Male dimensions average slightly 
larger than female. 

Age determination. — Adults have a short sagittal ridge; parietal 
ridges V-shaped, well developed; cranium rough; teeth worn. 

Subadults have parietal ridges lyre-shaped, poorly developed; 
cranium smooth to slightly rough; teeth slightly worn. 




Fig, 117. Skull of Vulpes rueppelli rueppelli. 
383 



384 FIELDIANA: ZOOLOGY 

Juveniles have a smooth cranium. 

Variation, —Summer pelages are more grayish than winter ones 
due to thinness of guard hairs. Some individuals have a strip of 
black-tipped hairs on side of foot. Specimens from southern parts of 
Western and Eastern Deserts have lacrimal stripes slightly darker 
and area of dark-tipped hair on throat larger than do individuals 
from northern localities. A few southern specimens have belly hairs 
with gray bases. Some Sinai specimens are brownish instead of 
reddish. 

An accessory cusp was present on pmg in four (50 per cent) of 
eight specimens from Sinai Peninsula, seven (28 per cent) of 25 from 
the Eastern Desert, and four (38 per cent) of 29 from the Western 
Desert. In two specimens only, an accessory cusp was also present 
on pmg. 

No subspecific differences could be found among Sinai, Eastern 
Desert, and Western Desert population samples. 

Comparisons.— The adult skull of V. rueppelli is a replica of that 
of a subadult V. vulpes, except for relatively larger bulla, broadness 
of nasomaxillary contact, and shape of posterior nasal margin (figs. 
112, 114, 117). Differences between the two species are under the 
latter. Dimensions can be compared in Tables 45 and 46. In com- 
parison with Fennecus zerda, V. rueppelli is larger, much darker, 
has a longer tail, smaller bulla, frontals not elevated, and white tail 
tip instead of black. Specimens in molt are sometimes mistaken for 
F. zerda. 

From V. pallida of Sudan, V. rueppelli is distinguishable by red- 
dish color, longer ears, lack of black mark on foreleg, and tail tip 
being white instead of black. 

From V. r. sabaea of Saudi Arabia, V. r. rueppelli differs in having 
darker color and slightly larger dimensions. 

Specimens examined,— Total 115. 

SINAI: Wadi Abu Zeitouna (3). Wadi el Sheikh 16 km. W of St. Catherine 
Monastery (4), Wadi el Raba (1). Tor (3). 

SUEZ: Wadi Iseili (3). Wadi Qiseib (1). 

CAIRO: near Cairo (1). 

RED SEA: Wadi Araba. Bir Zafarana (2), St. Anthony Monastery area (1). Bir 
Abu Shaar (2): Hurghada 14 km. S (2), 16 km. S (1): Wadi Umm Huweitat (1): 
Fawakhir Mine area 12); Mersa el Alam (1): Wadi Gemal (2). 

SUDAN ADMINISTRATIVE: Bir Shalatein (1). Wadi Ibib (2). Wadi Darawena 
(4). Bir Akwamtra area (1). 



OSBORN&HELMY: MAMMALS OF EGYPT 385 

GIZA: El Mansuriya (1): Sakkara (1); Bahariya Oasis (1); Bawiti (3); Ain el Guffara 
E of Bawiti (7); Ain el Beilda W of Bawiti (2); El Qasa. No. 2 NE of Bawiti (2), No. 3 
NW of Bawiti (1); Ain Marun (2): El Hara (3); El Ghaba el Qiblya (1); El Agouz (1); 
Wadi Ghorabi (1); Cairo-Bahariya Oasis track km. 208, acacia grove area 6 km. SE 
(1). 

EL FAIYUM: Qasr el Sagha (1). 20 km. N of cultivation (1). 

MINYA: Hatiyet el Sunt (1). 

ASWAN: Kurkur Oasis (9); Wadi Dihmit. Bir Umm Hibal area (1); Bir Umm 
Qareiyat area (4); Wadi Murra, Bir Murra area (2); Wadi AUaqi, n.2 km. SE of Allaqi 
Village (1). 

BEHEIRA: Wadi el Natroun. Deir Makaryus area (1). 

MATRUH: El Maghra (4); Bir Abd el Nabi (2); Siwa Oasis. El Maragi, W of (1); El 
Malfa 110 km. W of Siwa (1). 

EL WADI EL GEDEED: Farafara Oasis, Abu Minqar (1); Ain Gellaw (1); Dakhla 
Oasis. Mut (1); Kharga Oasis, Dush (1); Ain Amur (2), El Gizera (1); Bir Kiseiba (4); 
Bir Kurayim (3); Bir el Shab (3). 

Sudan. NORTHERN: Gebel Uweinat, Karkur Murr (4). 

Sight record of D. Osbom and I. Helmy.— 
RED SEA: Wadi MeUaha. 

Published records.— Records are from Anderson (1902), De Win- 
ton (1903), Flower (1932), Bagnold (1933), Hoogstraal et al. (1957b), 
Setzer (1961b), and Hoogstraal (1964). 

SINAI: Wadi el Sheikh; Wadi Abu Zeitouna; Wadi el Raba. 

RED SEA: St. Anthony Monastery area. 

QUALYUBIYA: Sindbis. 

GIZA: Imbaba. 

EL FAIYUM: 20 km. N of cultivation. 

BEHEIRA: Wadi el Natroun. Deir Makaryus. 

EL WADI EL GEDEED: Karkur Tahl (sight record). 

SUDAN ADMINISTRATIVE: Wadi Darawena. 

Collection.— Readily enters live traps baited with meat, sardines, 
or even bread. On the Red Sea shore, some were shot at night from 
distances of a few meters when they came to lick empty sardine and 
corned beef tins. 

Habitats.— Vulpes rueppelli is the most ubiquitous of Egyptian 
foxes, not restricted to sandy areas, and far more widely distributed 
than V. vulpes, particularly in waterless regions. 

Sinai Peninsula: Reported from northern and southern Sinai 
(Flower, 1932) in littoral semideserts and rocky wadis (Hoogstraal, 
1964). 



386 FIELDIANA: ZOOLOGY 

Eastern Desert: Figure 16. Ranges throughout the Eastern 
Desert in wadis and coastal areas. 

Western Desert: Figures 9, 18. Ranges throughout the Western 
Desert south of the Coastal Desert, particularly in vegetated areas, 
isolated acacia groves, tamarisc clumps, palm groves, and oases, 
where Rueppell's foxes hunt in patches of grass and mound-forming 
vegetation (fig. 17). They can always be found in the vicinity of 
springs and shallow wells. 

A few specimens have been collected on the borders of the Nile 
Valley and Delta, but none from the Delta or the Western Mediter- 
ranean Coastal Desert, probably because of the predominance of V. 
vulpes in those areas. 

Habits. — Vulpes rueppelli is sometimes seen during the day. One 
was flushed in mid-afternoon in the rocky canyon of Wadi Mellaha. 
Foxes were seen in late afternoon prior to being trapped in Karkur 
Murr, Gebel Uweinat, and near an acacia grove south of the Cairo- 
Bahariya Oasis track. 

Three males and one female were trapped beside palms at Bir 
Kiseiba within two hours after sundown. Hoogstraal (1964) ob- 
served Rueppell's foxes at dusk in the Gebel Elba area. 

On several occasions, when we were camped on the shores of the 
Red Sea, Rueppell's foxes came within a few meters to lick empty 
meat and sardine tins. Bagnold (1933) noted the indifference of this 
fox toward his camp in Karkur Tahl, Gebel Uweinat. 

Hungry sand foxes often become a nuisance by setting off rodent 
live traps in trying to reach the peanut butter bait. Rodents in traps 
were sometimes mangled by them. 

Burrows and dens.—De Winton (1903) dug a sand fox from a 
shallow burrow where the animal's nose was seen protruding. At Bir 
el Shab and Bir Kiseiba, we found dens in dead fronds under dense 
clumps of dom palms (Hyphaena thebaica), but nowhere else were 
we able to locate them. In rocky areas, V. rueppelli doubtlessly dens 
in crevices. 

Food. — Hoogstraal (1964) reported that, in Sinai, a camel carcass 
attracted Rueppell's sand foxes. He also said that Bisharin 
tribesmen in the Gebel Elba area maintained that this fox stole 
lambs. 

Stomachs have contained rodents, feathers of small birds, lizards 
{Euromastix aegyptius and Eremias sp.), remains of insects 



OSBORN & HELM Y: MAMMALS OF EGYPT 387 

(grasshoppers, mole crickets, and scarabid beetles), and dates. In 
the Bir el Shab area, we observed where sand foxes had gnawed the 
fibrous fruits of dom palm. Tregenza (1958) remarked on their 
ability to climb date palms. 

Water.— Captive V. rueppelli will drink water. The species is 
always found in the vicinity of wells and springs, and tracks have in- 
dicated that it drank water. Tracks have also been reported long 
distances from water (Hurst, 1910), and we have collected it in 
waterless areas. There is a possibility that this fox can utilize 
brackish water. 

An old Arab fable mentioned by Hurst (1910) is that this fox 
drinks from the wind by sleeping with its head into the breeze. 

Reproduction.— One female collected from Wadi Iseili in June had 
three small fetal scars. 

Sex ratio.— A sample of 67 museum specimens contained 39 (58 
per cent) males and 28 females. 

Economic importance.— As mentioned, this fox is suspected of 
preying on lambs. According to Hurst (1910), Arabs eat sand foxes. 

Genus Fennecus Desmarest, 1804 

Small, pale, buffy fox with narrow reddish dorsal stripe. Ear 
relatively large. Tail relatively short, brushy; tip black. Pupil of eye 
round. Frontal region of skull slightly inflated. Nasals ending 
anterior to posterior level of frontomaxillary suture. Cranium 
broadest on sides, narrower at base of zygomatic processes. Postor- 
bital process concave dorsally, posterior margin not ridged. Postor- 
bital swelling lacking. Vertical curvature of zygomatic arch low. Tip 
of zygomatic process of temporal thin. Canines slender, elongate; 
point of upper canine reaching level of mental foramen when jaws 
are closed. Tympanic bulla markedly inflated. Lower jaw very 
shallow and slender. Cheek teeth narrower than in Vulpes. 

Fennecus zerda (Zimmermann, 1780) 

Canis zerda Zimmermann, 1780, Geographische Geschichte des Menschen, Vol. 2, 
p. 247. 

Type locality.— "Sahara and other regions back of the Atlas 
Mountains and in Tripoli" (describer) and "sandy deserts of North 
Africa" (Flower, 1932, p. 401). 

General distribution. — Kuwait, northern Sinai Peninsula, Egypt, 



388 



FIELDIANA: ZOOLOGY 



and northern Sudan west of the Nile River; thence westward across 
the Sahara into Mauritania. ^ 

Common names.— Fennec Fox, Fennec. 

Distribution in Egypt — Figure 118. Northern Sinai Peninsula 
and Western Desert south of Mediterranean Coastal Desert. 

Diagnosis.— Small, buff colored; dorsal stripe narrow, reddish. 
Ear very large, triangular. Tail relatively short and brushy with a 
black tip. 

Skull rounded, ridges inconspicuous, frontal region slightly 
inflated. Rostrum short, narrow. Tympanic bulla greatly inflated. 

Adult head and body length average 367 mm.; tail 205 mm., 55 
per cent of head and body length; foot 103 mm.; ear 95 mm.; 
condyloincisive length 83.8 mm.; weight 1 kg. 



,. 2 5* 2 6* 2 7* 28* 2 9* 30* 31* 32* 3 3* 34* 35* 36* 37* 




Fig. 118. Collection localities of Fennecus zerda and sight record (S). 



OSBORN & HELMY: MAMMALS OF EGYPT 



389 



External characters.— Figure 119. Fur long, soft, silky. Dorsal 
stripe pale grizzled reddish and buff, 2.5 to 5.0 cm. wide from 
shoulder to base of tail; sometimes widening over hips. Grizzling 
due to guard hairs with narrow black tips and narrow rufous and 
white subterminal bands. Side and outer surface of legs pale 
yellowish buff. Chin, cheek, side of throat, belly, and inner side of 
legs white. The mammary area of adult females is rufous. Facial 
area buffy. Mystacial area and lacrimal stripe pale brownish. Ear 
white anteriorly and on margin, pale brownish posteriorly. Crown 
and nape pale brownish to rufous. Narrow line of black on neck and 
shoulders due to black-tipped hairs lacking white subterminal 
bands. Scattered guard hairs, in addition to color hairs mentioned 
above, are all black or with gray bases. Under hairs of dorsum have 
minute blackish tips, conspicuous rufous subterminal bands, and 
grayish or buffy bases. Juvenile pelages paler than adult, with little 
or no rufous, and facial marking inconspicuous. Tail brushy, not 
club-shaped as in Vulpes; dorsal hairs with blackish or reddish tips; 
color grading to pale reddish or whitish below. Tail tip and hair over 
gland on upper base of tail black. Hair of palm and sole buffy, long, 
completely concealing pads. 




Fui. 119. Young Fennecus zerda. 



390 FIELDIANA: ZOOLOGY 

Cranial characters. — Figure 1 20. Cranium rounded, frontal region 
slightly inflated, rostrum short, narrow. Cranial ridges, if present, 
lyre-shaped, inconspicuous. Lambdoidal ridge low, superior edge 
not exceeding posterior level of upper lip of foramen magnum. 
Postorbital process concave dorsally, posterior ridge lacking. 
Nasals tapering abruptly posteriorly, terminating anterior to 
posterior level of frontomaxillary suture. Nasomaxillary contact 
broad as in V. rueppelli. Tympanic bulla greatly inflated. Basioc- 
cipital and basisphenoid markedly constricted. External auditory 
meatus opening very large. Lower jaw very shallow and slender. 

Bacu/um. — Baculum relatively large and heavy compared with 
Vulpes sp. Ventral surface channeled almost entire length, edges 
irregular. Cross section triangular. Base rounded, bulging slightly; 
tip small, slender (Didier, 1946). Total lengths of two bacula were 31 
and 36 mm. 

Teeth.— Teeth smaller and slenderer than in V. rueppelli. 
Cingulum on anterior of upper cranassial (pm^) more strongly 
developed than in other foxes. An accessory cusp is occasionally 
present on pm^. 

Measurements.— Table 47. Male and female measurements are 
subequal. 

Age determination.— Adults have cranium roughened on sides; 

Table 47. — Means (and ranges) of measurements, ratios, and weight of adult 
Fennecus zerda. 





Desert Oases 


Desert-Delta 


HBL 


367.6(357-387) 11 


368.0 (337-387) 35 


TL 


203.7(187-226)11 


207.4 (186-230) 35 


TL/HBL% 


55.4 (51.4-60.8) 11 


56.2 (51.8-63.7) 35 


FL 


103.8(94-111) 11 


103.2(93-110)35 


EL 


94.2(88-104) 11 


96.3 (92-104) 35 


Wt (kg.) 


1.05 ( 0.8- 


1.15)9 


CIL 


83.9 (80.4-88.0) 12 


83.6 (80.6-87.6) 38 


ZW 


46.6 (45.4-47.9) 12 


46.2 (37.4-48.3) 29 


RW 


13.1 (12.5-13.6) 12 


12.7(11.9-13.6)37 


POW 


18.6(16.1-20.7) 12 


18.4(16.5-20.7)36 


BCW 


36.4(35.6-37.8) 12 


36.7(34.9-38.1)35 


NL 


28.3(25.3-30.3) 12 


27.3 (24.7-29.3) 36 


CM' 


36.2(34.5-37.8) 11 


35.9 (34.2-37.7) 35 


M-M' 


25.2 (24.5-26.9) 10 


24.9(23.1-26.2)32 


BL 


21.3 (20.0-22.4) 12 


21.4 (20.7-23.0) 37 


SH 


37.2(35.8-38.2) 11 


36.8 (35.5-38.2) 34 





Fig. 120. Skull of Fennecus zerda. 



391 



392 FIELDIANA: ZOOLOGY 

parietal ridges low, lyre-shaped; postorbital process pointed, slight- 
ly hooked; and basioccipital-basisphenoid sutlire fused. 

Subadults have cranium smooth, parietal ridges slightly 
developed, postorbital process blunt, and basioccipital- 
basisphenoid suture not completely fused. 

Juveniles have cranium smooth, postorbital process blunt, and 
sutures open. 

Variation.— Adult and subadult pelages have varying amounts of 
grizzling on dorsum. Summer pelages are thinner, but about same 
general color as winter pelages. Molting pelages appear grayish, 
yellowish, or rufescent dorsally. 

Dorsum under hairs in winter pelages have a broad, dark gray 
basal band fading to whitish proximally. Under hairs in summer 
pelages are buff to base. 

Winter pelages are divisible into pale and dark and narrow and 
broad dorsal stripes, which are about 2.5 and 5.0 cm. wide, respec- 
tively. In a sample of 16 males and 20 females, dorsal stripes were 
dark in 90 per cent of males and 75 per cent of females. Narrow and 
broad dorsal stripes occurred in equal numbers in both sexes. There 
was no indication of geographical variation in either character. 
Setzer (1961b, p. 118) recognized a "wide range of color. . . .from a 
pale sandy hue to rich brownish tone" and "no age or seasonal dif- 
ference in this range of color." 

Comparisons.— Small size, pale color, narrow reddish dorsal 
stripe, shorter tail, black tail tip, and proportionately larger ears 
distinguish F. zerda externally from other Egyptian foxes. Cranial- 
ly, F. zerda differs in having a shorter, narrower rostrum; slightly 
elevated frontals; smoother, more rounded braincase; large tym- 
panic bulla; and smaller, slenderer lower jaw (tables 45-47; figs. 114, 
117, 120). 

Specimens examined.— Total 114. 

BEHEIRA: El Khatatba (3). Wadi el Natroun (8). Wadi el Farigh (3). 

GIZA: El Qatta (15). Abu Ghalib (24). between Abu Ghalib and Cairo on Cairo- 
Alexandria desert road (37). Kirdasa (1). Giza Pyramids area (2). 

EL FA 1 YUM: Lake Quarun 10 km. SW (1); Wadi Muwellih (3). 

MATRUH: Qaret el Mashruka 8 km. S (1). Nakhlat el Barraq (1). El Maghra (10). 
Bir Mikheimin (1 drowned in well), Nuweimisa (1). 

EL WADI EL GEDEED: Kharga Oasis (1). 10 km. S (1). Dakhla Oasis. Mut (1). 



OSBORN&HELMY: MAMMALS OF EGYPT 393 

Sight record of I. Helmy.— 

EL WADI EL GEDEED: Farafara Oasis, El Qasr. 

Published records.— Records are from Anderson (1902), De Win- 
ton (1903). Bonhote (1912). Bagnold (1931), Flower (1932), Setzer 
(1961b), and Hoogstraal (1964). 

SINAI: 33 km. NE of Suez and 25 km. E of Suez Canal. 

BEHEIRA: El Khatatba, Wadi el Natroun. 

GIZA: Wardan, Abu Rawash, Abu Ghalib, between Abu Ghalib and Cairo on 
Cairo-Alexandria desert road. El Qatta, Kirdasa, Giza Pyramids area, Bahariya 
Oasis. 

EL FAIYUM: Lake Qarun 10 km. SW. 

EL WADI EL GEDEED: Dakhla Oasis. Mut; Kharga Oasis; Bir Abu Hussein; 
Sir Murr: Bir el Shab (sight record). 

Collection.— 'Dmq from burrows, shot at night using a spotlight, 
and trapped alive with sardine bait. 

//a6itats.— Burrows in sandy areas of desert (Anderson, 1902; 
Thomas, 1913), usually near vegetation. We have dug fennecs from 
burrows in hillocks under Nitraria retusa, Zygophyllum album, and 
Calligonum comosum and shot them at night in the same vegeta- 
tion. Stands of Artemisia monosperma and bunch grass (Panicum 
turgidum) are also frequented by fennecs (fig. 20). 

Flower (1932. p. 402) reported a specimen from "undulating coun- 
try with sand dunes" in Northwestern Sinai Peninsula. Most of 48 
specimens brought into Giza Zoological Gardens, he wrote, came 
from an area west of Giza north to Wadi el Natroun. Vegetated 
sandy areas between Giza Pyramids and El Khatatba have also pro- 
vided a large sample of fennecs (Hoogstraal, 1964). None have been 
collected or reported from the Western Mediterranean Coastal 
Desert. 

Activity. — Yoodm^ is almost strictly a nocturnal activity, 
although we have noted fennecs feeding on insect larvae during the 
day. Fennecs are often seen in afternoon, singly or in small groups, 
sitting on mounds or small hills (Hoogstraal, 1964). 

Voice. — Flower (1932) described the voice correctly as like the 
bark of a small dog. 

Burro m;s. — Burrows we have dug were simple tunnels 1 to 2.5 m. 
long and reaching a depth of about 1 m. Gauthier-Pilters (1967) 
reported a burrow in Algeria 10 m. long and 1 m. deep. Burrows may 



394 FIELDIANA: ZOOLOGY 

be in flat barren sand, low hills, vegetated hillocks, or according to 
Bruce (1790), among the dead fronds of date palms. We have col- 
lected V. rueppelli, but not F. zerda, in the last. 

Food. — "The stomach of a fennec shot in the vegetated area of 
Nuweimisa Oasis in March, 1964 contained only lepidopterous 
larvae [Heliothis (Chlorides); Noctuidae"] (Hoogstraal, 1964, p. 
214), which we found everywhere on the ground between 
Zygophyllum album at the time. Stomachs from other specimens 
contained feathers of small birds and remains of Jaculus jaculus. 

Insects, beetles in particular; small rodents (Gerbillinae), lizcirds, 
birds, and plant material have been listed as food of wild fennecs 
(Thomas, 1913; Buxton, 1923; Gauthier- Filters, 1967). Dekeyser 
(1955) included roots of orobanche (Cistanche phelipaea). 

Bruce (1790) fed a captive fennec dates, sweet fruits, bread with 
honey or sugar, and eggs of pigeons or small birds. He observed the 
animal did not know how to handle a hen's egg, but a broken egg 
was quickly eaten. Schmidt-Nielsen's (1964) fennecs ate a similar 
variety, including table scraps. Pet fennecs of Gauthier-Pilters 
(1967) were said to hide food like a red fox. 

One captive fennec had an affinity for sweets such as cake, 
chocolate, sweetened stewed fruits, whipped cream, and honey, but 
not for sour foods. It also ate various nuts and fruits (Vogel, 1962). 

Water.— The fennec api>ears to be the only desert carnivore that is 
entirely independent of drinking water, according to Schmidt- 
Nielson (1964) and Gauthier-Pilters (1967). Fennecs we have kept in 
captivity for a week or so did not drink water even though panting 
from heat. Fennecs kept as pets (Vogel, 1962) and in zoos drink 
water regularly. 

Reproduction. — Flov/er (1932) reported litters of three born in 
Giza Zoological Garden in July and April. Captive fennecs from 
Algeria bred in February, March, and April and gave birth in May 
and June after 50 to 52 days of gestation. Litter sizes were one to 
three (Saint-G irons, 1962). In Gauthier-Pilters' (1967) notes on 
mating in captivity, the rutting period was January and February, 
gestation 49 to 52 days, and birth period February and March. We 
dug two young, which were probably weaned, from a burrow in 
Wadi Muwellih in May (fig. 119). 

Sex ratio. — A sample of 71 from Giza Govemorate contained 34 
(48 per cent) males and 37 females. 



OSBORN&HELMY: MAMMALS OF EGYPT 395 

Predators.— According to Gauthier-Pilters (1967, p. 117), 
"enemies of the fennec are jackals, hyenas, vultures as well as dogs 
and men." 

Economic importance.— Young fennecs are fattened and eaten in 
the Western Sahara (Monod, 1958). 

Family 2. Mustelidae 

Small to medium size, slender carnivores. Pelage marked or 
unmarked. Muzzle short; ears small, rounded; legs short in propor- 
tion to body length. Feet plantigrade to digitigrade, functional toes 
5-5, claws nonretracile. Tail cylindrical or bushy, two-thirds of or 
less than length of head and body. Anal scent glands well developed. 
Males larger than females in some genera. 

Rostrum and nasals short. Upper tooth row length less than one- 
half skull length. Paroccipital process not prominent, fused com- 
pletely to tympanic bulla. Bulla moderately inflated, septum 
absent. Postglenoid process curved over glenoid fossa locking man- 
dible in place. Alisphenoid canal absent. Baculum well developed. 

Incisors unspecialized; canines elongate, sharp; premolars small, 
reduced in number, constriction commonly present between lateral 
and medial parts of m'. Dental formula: |, f, x. 2X 2 = 34 or 36. 

Key to Egyptian Genera of Mustelidae 

1. Dorsum striped black and white. Parapterygoid fused with tympanic bulla. 

a. Black middorsal stripes four or five. Palm and sole haired. Postorbital swelling 
inconspicuous. Meatal lip inflated Poecilictis, p. 395. 

b. Black middorsal stripes three. Palm and sole naked. Postorbital swelling very 
conspicuous. Meatal Up not inflated Ictonyx, p. 403. 

2. Dorsum brown. Parapterygoid not fused with tympanic bulla Mustela, p. 405. 

Genus Poecilictis Thomas and Hinton, 1920 

Shaggy, black- and white-striped, bushy-tailed weasels. Mid- 
dorsal longitudinal black stripes four or five. Palm and sole haired; 
toe pads concealed. Skull short, cranium outline triangular in dorsal 
view. Postorbital swelling moderate. Postorbital constriction slight- 
ly wider than rostrum. Tympanic and mastoid bullae inflated. 

Poecilictis libyca (Hemprich and Ehrenberg, 1833) 

Mustela libyca Hemprich and Ehrenberg, 1833, Symbolae Physicae Mamm., 
2, folio k. p. 6. 

Type locality.— Libya (Setzer, 1959c); Egypt, MATRUH: Libyan 
Desert between Siwa and Alexandria (Flower, 1932). 



396 



FIELDIANA: ZOOLOGY 



,. 25* 26* 27* 28* 2 9* 30* 31* 32* 3^ 34* 35* 36* 3 7* 




Fig 121. Collection localities of Poecilictis libyca libyca (circles) and records of 
tracks (T), Ictonyx striatus erythreae (triangle), Mustela nivalis subpalmata (dots) 
and sight records (S), and Genetta genetta senegalensis (squares). 

General distribution.— Mediterranean Coastal Desert from 
western margin of Nile Delta west to Morocco; northern Sudan, 
Chad, Niger, Mali, and Mauritania. 

Common mimes.— Striped Weasel, Abu Menten. 

Subspecies in Egypt— 

Poecilictis libyca libyca (Hemprich and Ehrenberg, 1833) 

Poecilictis libyca alexandrae Setzer. 1959, J. Egypt. Publ. Health Assn.. 33, 
No. 6, p. 201. 

Type locality.— Libya. 

Distribution in Egypt— Fig\xre 121. Western margin of Nile 



OSBORN & HELMY: MAMMALS OF EGYPT 



397 




Fig 122. Mustelidae. Left to right: dorsal and ventral views of Mustela nivalis 
subpalmata; two dorsal views and a ventral view of Poecilictis libyca libyca; dorsal 
and ventral views of Ictonyx striatus erythreae. 

Delta, Western Mediterranean Coastal Desert, Wadi el Natroun, El 
Maghra, and Gebel Uweinat. 

Diagnosis.— Dorsum with four or five black stripes alternating 
with white. White band encircling head. Outline of cranium broadly 
triangular in dorsal view. Postorbital swellings moderate. Mastoid 
and paroccipital processes obsolete. Tympanic and mastoid bullae 
and, lower lip of auditory meatus inflated. Adult head and body 
length average 256 mm.; tail 174.0 mm., 67.7 per cent of head and 
body length; foot 41.0 mm.; ear 22.2 mm.; condyloincisive length 
52.9 mm.; averaged weight of various individuals 200 gm. 

External characters. — Yigare 122. Pelage shaggy, markings black 
and white. Dorsum with three black stripes beginning behind ears. 
Middle stripe subdivided middorsally into two or three additional 
stripes that fuse on rump and base of tail. Black stripes are pro- 
duced by black under hairs and white stripes by guard hairs that are 
white to their bases. Pelage may appear spotted. Venter, legs, and 






Fu; 123. Skull of Poecilictis libyca libyca. 



398 



OSBORN&HELMY: MAMMALS OF EGYPT 399 

feet black. Palm, sole, and toes haired; toe pads concealed by hair. 
Snout black, mystacial area white. Head encircled by white band 
passing between eyes and ears and on underside of jaws. Ear tip 
sometimes white. Upper tail hairs with long, white tips, black 
subterminal bands, and white bases. Lower heiirs with black tips in- 
creasing in length from middle to end of tail. Underside of tail tip 
black. 

Cranial characters. — Figure 123. Skull short, broad; cranium 
relatively shallow, outline triangular in dorsal view. Sutures not 
visible, except in juveniles. Sagittal ridge broad and low in 
subadults, becoming slightly higher and narrower in adults. Lamb- 
doidal ridge fairly prominent, lateral portion extending posterior to 
supraoccipital, but not beyond level of posterior margin of mastoid 
bulla nor even approaching level of upper lip of foramen magnum. 
Postorbital process prominent. Postorbital inflation moderate. 
Postorbital constriction slightly wider than rostrum. Tympanic and 
mastoid bullae and lower lip of auditory meatus strongly inflated. 
Mastoid and paroccipital processes obsolete. Parapterygoid fused 
with tympanic bulla. Zygomatic arch not strongly curved upward. 
Coronoid process of lower jaw rounded. 

Teeth.— Outer upper incisor considerably larger than others. Up- 
per and lower premolars not crowded; anterior cusps triangular and 
larger than posterior cusps. Carnassials markedly sectorial. Upper 
carnassial with outer anterior cingulum cusp-like, inner cusps prom- 
inent, and constriction between anterior and posterior crowns deep. 
Width across crown of m' greater than width of carnassial (pm"*). 
Lower carnassial with anterior crown and inner cusp subequal; heel 
much larger than mg. M2 relatively l£U"ge, with three fairly promi- 
nent cusps. 

Baculum.-Baculum length about 32 mm. in adult; base swollen 
slightly, rugose; shaft smooth, curved slightly dorsad; tip teardrop 
shaped and oblique to shaft (Setzer, 1960a). 

Measurements.— Table 48. Male and female measurements sub- 
equal. Means (and ranges) of condyloincisive length (in millimeters) 
of 11 subadult males and eight subadult females are 49.9 (48.2 to 
54.9) and 49.8 (46.8 to 52.8), respectively. 

One adult female, one subadult female, and one juvenile of 
unknown sex weighed 181.0, 171.2, and 192.0 gm., respectively. 
Three males examined after death by Flower (1932) weighed 200, 
200, and 250 gm. 



400 FIELDIANA: ZOOLOGY 

Table 48. — Means (and ranges) of measurements and ratios of adult Poecilictis L 
libyca and adult male (M) and female (F) Ictonyx striatum erythreae. 





P. LI 


ybica 


/. s. erythreae 




Western Mediterranean 


Western border of 


Gebel Elba 




Coastal Desert 


Nile Delta 


M 


F 


HBL 


263.4 (254-279) 5 


231. 243 


369 


325 


TL 


178.2 (160-193) 4 


179. 162 


273 


288 


TL/HBL% 


67.0(61.1-72.2)5 


71.5. 66.6 


73.9 


88.6 


FL 


41.2 (38-46) 5 


40,41 


58 


53 


BL 


22.8 (21-25) 5 


22.20 


26 


24 


CIL 


54.5 (52.7-56.5) 5 


51.4 (48.8-54.3) 7 


61.0 


56.7 


ZW 


34.5 (32.8-35.4) 5 


32.3 (30.0-34.0) 7 


37.9 


33.8 


POW 


11.8(11.4-12.6)5 


11.2(11.0-11.7)6 


13.3 


13.2 


MW 


31.9 (29.9-32.7) 5 


29.9 (28.5-31.6) 6 


31.9 


29.8 


RW 


11.1 (10.6-11.8)5 


10.3 ( 9.9-10.8) 7 


14.1 


12.2 


SH 


23.1(21.8-23.7)5 


22.8 (22.0-23.4) 6 


23.3 


22.4 


M' 


6.5 ( 5.6- 7.2) 5 


6.2 ( 5.8- 6.5) 7 








P-P 


19.3 (18.5-19.8) 5 


18.4 (18.0-18.9) 7 


21.9 


21.2 


I-M' 


20.7(20.1-21.4)5 


19.8(19.1-20.7)7 


22.7 


21.3 


Post M' 


33.8 (32.1-35.2) 5 


31.6 (29.5-33.6) 7 








I-M'/CIL% 


38.0 (37.6-39.0) 5 


38.4 (37.4-39.8) 7 








Post M'/CIL% 


62.0 (60.9-62.6) 5 


61.4 (60.2-62.5) 7 









Age determinatiorL— Juveniles have a smooth cranium, some 
sutures not fused, frontal swellings prominent, and all permanent 
teeth not in position. Subadults have parallel parietal ridges form- 
ing a broad plateau about as wide as or wider than the occipital con- 
dyles, lambdoidal ridge not prominent, frontal swellings prominent, 
all sutures fused, and all permanent teeth in position. Adults have 
parietal ridges close together and forming a sagittal ridge which is 
narrower than the occipital condyles, lambdoidal ridge prominent, 
frontal swellings obsolete, and teeth worn. 

Vanation,— Specimens from areas adjacent to the Nile Delta and 
west to Bir Victoria average slightly smaller in most dimensions 
than those from the Western Mediterranean Coastal Desert (table 
48). 

Three variations in median lumbar stripe— complete, incomplete, 
and absent— appear to be clinal in distribution from west to east, 
but data are insufficient for definite conclusions. Amount of white 
on ear tip varies individually. Photographs of specimens from Libya 
(Zammarano, 1930) also indicate some variation in color pattern. 

Comparisons.— Poecilictis libyca differs from all other small 
Egyptian carnivores, except Ictonyx striatus, by its black and 



OSBORN & HELMY: MAMMALS OF EGYPT 401 

white dorsal striping. From the latter it differs in having the median 
dorsal stripe subdivided in the lumbar region. The black stripes are 
due to black under hairs, whereas in /. striatus black stripes are of 
black guard hairs. Cranially, P. libyca differs from the latter in 
having smaller postorbital swellings, bulla more inflated, meatal lip 
inflated, mastoid process obsolete, and smaller dimensions (table 
48). 

Poecilictis I libyca is intermediate in size between the subspecies 
vaillanti of Tunisia and Algeria and multivittata of Sudan (Thomas 
and Hinton, 1920). 

i?emarfes.— Following is an annotated list of characters which 
were considered to be diagnostic for P. I. alexandrae ssp. nov. Setzer 
(1959c) from the western border of the Nile Delta in comparison 
with P. I. libyca from the Western Mediterranean Coastal Desert. 

1. "Small body size." Setzer compared his type, a subadult, with 
two adult specimens of P. I. libyca. External measurements in Table 
48 are of four to five adult libyca and two adult ''alexandrae.'' Mean 
(and range) of head and body length (in millimeters) for two adult 
plus 12 subadult ''alexandrae'' are 230.4 (205 to 243); foot length 
37.2 (32 to 43). These data in conjunction with those in Table 48 
indicate that there is clinal reduction in size from west to east in P. I. 
libyca in Egypt. 

2. "Tail relatively long." Tail length is not relatively longer in the 
"alexandrae" sample and also averages slightly shorter than in 
libyca. Mean (and range) of tail length (in millimeters) of two adult 
plus 11 subadult "alexandrae" are 151.6 (124 to 179), whereas the 
mean (and range) of ratio of tail length to head and body length in 
per cent are 64.9 (58.2 to 71.5). 

3. "Skull small." Adult skulls in the "alexandrae" sample are 
slightly smaller and have less prominent ridges than older adult 
skulls of P. I. libyca (table 48). Mean condyloincisive length, plus or 
minus two standard errors, of P. I. libyca is 54.5 ± 3.16, and for the 
"alexandrae" sample, 51. 4± 1.396. Furthermore, statistical 
analysis indicates no significant difference between samples, where 
( = 1.79 with 10 degrees of freedom. 

4. "M and p"* small." M' averages slightly smaller in the "alexan- 
drae" sample, but shows more variation in libyca (table 48). Visual 
examination indicated no marked size difference in p^ between 
samples. 

5. "Upper premolars and molars small." Length of upper tooth 



402 FIELDIANA:ZCX)LOGY 

row (I-M') in the "alexandrae" sample averages 0.7 mm. less than 
libyca (table 48). The ratio of tooth row to condyloincisive length 
(I-M'/CIL) is essentially the same in both samples. The teeth of 
'"alexandrae'' are not small and weak in proportion to skull size as 
suggested elsewhere (Hoogstraal, 1964). 

6. "Audital portion of auditory bulla strongly inflated; mastoidal 
portion of auditory bulla but slightly inflated." Setzer's (1959c) 
type of "alexandrae"' is the only specimen that shows this condi- 
tion. Prominent inflation of the mastoid bulla is a characteristic of 
P. libyca. 

7. "Postpalatal length of skull relatively long." Ratios of 
postpalatal length to condyloincisive length (Post M'/CIL) are 
about the same in both samples (table 48). Postpalatal length 
averaged 2.2 mm. less in the eastern sample (table 48). 

8. "The posterior white rosette is not divided by the central dorsal 
black stripe." This character (median stripe absent) was present in 
25 per cent of the ''alexandrae'' sample and in one specimen of 
libyca from Bahig (table 49). The high frequencies of two additional 
variations, median stripe complete and median stripe incomplete, in 
Table 49 reduce further the value of "rosette all white" as a 
diagnostic character of Setzer's (1959c) proposed subspecies. 

The above notes indicate that the "a/ejcandrae " sample is not tax- 
onomically distinct from P. I. libyca. Data in Tables 48 and 49 
indicate further that dimensions and color characters are clinal in 
nature. The two populations under consideration are doubtlessly 
continuous. 

Specimens examined.— Total 45. 

MATRUH: SIDI Barrani 31 km. E (1); Mersa Matruh (3); Bahig (4); Giza 16 km. 
NW (1). 32 km. NW (1); Bir Nahid (1). 

BEHEIRA: Wadi el Natroun (2). hUls, W of (1 skuU beside fox den); El Beida (2); 
Bir Victoria (1). 

EL TAHREER: El Tahreer (1). 

GIZA: El Qatta (5). Abu Ghalib (10), between Abu Ghalib and Cairo-Alexandria 
desert road (1). Abu Rawash (2). Giza Pyramids area (3), Giza (2). 
CAIRO: Near Cairo (4). 

Sight record of D. Osborn.— Tracks photographed at El Maghra, 
AprU. 1977. 

Published records.— Records are from Anderson (1902), De Win- 



Western Mediterranean 

Coastal Desert 

No. % 


Western Border 

of Nile Delta 

No. % 


4 


44.4 


8 


28.5 


4 


44.4 


13 


46.4 


1 


11.1 


7 


25.0 


9 




28 





OSBORN&HELMY: MAMMALS OF EGYPT 403 

Table 49. — Variation in median lumbar stripe in samples of Poecilictis libyca. 
Terminology in parentheses is from Setzer (1959c). 

Median lumbar stripe 

Complete (white rosette divided) 

Incomplete (white rosette partially 
divided or with central black spot) 

Absent (rosette all white) 

Totals 

ton (1902), Flower (1932), Setzer (1959c), Hoogstraal (1964), and 
Missone (1970). 

MATRUH: Sidi Barrani 31 km. E, Mersa Matruh, Bahig. 

BEHEIRA: Wadi el Natroun. 

GIZA: El Qatta. Abu Rawash, Giza, Sakkara. 

Libya. CYRENAICA: Gebel Uweinat (tracks photographed). 

Collection.— Dug from shallow burrows in flat sand, hard ground, 
or in mounds (Hoogstraal, 1964). 

//a6itat.— Vegetated sandy desert bordering the Nile Delta. 
Discontinuous patches of vegetation west of the Delta, sandy areas 
of Wadi el Natroun, and Western Mediterranean Coastal Desert 
south of the coastal salt marshes (De Winton, 1903; Hoogstraal, 
1964). 

Behavior.— Reported by Hoogstraal (1964, p. 217) to "growl, 
bark, and hiss viciously at sudden sounds" and to raise the nape and 
spinal hair when annoyed. Flower (1932, p. 404) said that, in cap- 
tivity, "these animals become tame, do not smell and make 
interesting pets." 

Food.— Thought to live almost entirely on lizards (De Winton, 
1903). Animals in captivity have been fed young live mice. 

Reproduction. — Flower (1932, p. 404) recorded five litters of one 
to three born in Giza Zoological Gardens in January, February, and 
March. He described the newborn as "pink, hairless and helpless." 
Hoogstraal (1964) examined a female from Bahig in September, 
which contained advanced embryos. 

Genus Ictonyx Kaup, 1835 
Long-haired, black- and white-striped, bushy-tailed weasels. Mid- 



404 FIELDIANA: ZOOLOGY 

dorsal longitudinal black stripes three. Palm and sole naked. Skull 
elongate. Postorbital swelling prominent. Postorbital constriction 
about same width as rostrum. Tympanic bulla inflated. 

Ictonyx striatus (Perry, 1810) 

Bradypus striatus Perry, 1810, Arcana or the Mus. Nat. Hist., pt. II, pi. (41), text. 
Type locality. —South Africa: Cape of Good Hope. 

General distribution. —Sudan west through Chad, Niger, Mali, 
Mauritania, and Senegal; and south through Ethiopia, south- 
western Somalia, Kenya, Tanzania, and the remainder of Africa 
south of Zaire. 

Common names.— Zoril, Abu Afene. 

Subspecies in Egypt — 

Ictonyx striatus erythreae De Winton, 1898 

Ictonyx striatus erythreae De Winton, 1898 Ann. Mag. Nat. Hist., (ser. 7). 1, 
p. 248. 

Type locality.— Sudan, KASSALA: Suakin. 

Distribution in Egypt— Figure 121. Southeasternmost part of 
Eastern Desert. 

Diagnosis.— Dorsun\ with three prominent black stripes alter- 
nating with white. White band across forehead. Palm and sole bare. 
Skull elongate. Postorbital swelling prominent. Mastoid and paroc- 
cipital processes protruding. 

Head and body length average 347 mm.; tail 280 mm., 81 per cent 
of head and body length; foot 56 mm.; ear 25 mm.; and occipitonasal 
length 58.8 mm. 

External characters. — Figure 122. Pelage shaggy, markings black 
and white. Dorsum with three clearly defined black stripes begin- 
ning behind head, broadening and separating widely on middorsum, 
fused on rump. Black stripes of all black hairs, white markings of all 
white hairs. Belly, legs, and feet black; latter sometimes with scat- 
tering of white hairs. Palm and sole bare. Lips sometimes white. Ear 
tip white. A broken white band crosses the head between eyes and 
ears. Upper and lower basal one-third of tail black; remainder of tail 
hairs with white tips and black bases. 

Cranial characters.— Skull elongate; cranium high, rounded. No 
sutures visible in adults. Nasofrontal region markedly swollen. 



OSBORN&HELMY: MAMMALS OF EGYPT 405 

Sagittal ridge narrow and clearly defined. Lambdoidal ridge is 
prominent, superior margin extends beyond the posterior level of 
the occipital condyle. Postorbital process moderately developed, 
postorbital inflation prominent. Postorbital constriction about 
same width as rostrum. Tympanic bulla inflated, fused with 
parapterygoid. Mastoid bulla and lower lip of auditory meatus not 
inflated. Mastoidal and paroccipital processes prominent, pro- 
truding. Zygomatic arch strongly curved upward. Tip of coronoid 
process of lower jaw rounded. 

Baca/um.— Baculum length about 52 mm. Base enlarged; shaft 
tapering gradually to flared tip, curved dorsad (Didier, 1947). 

Teeth.— Outer upper incisor (I^) much larger than others. Upper 
premolars crowded. First premolars simple, second upper and lower 
with large triangular anterior cusps and small posterior cusps. Car- 
nassials are markedly sectorial. Upper carnassial with outer 
anterior cingulum cusp-like; inner cusps prominent, two-thirds 
height of anterior crown; and deep constriction between anterior 
and posterior crowns. Inner cusps and anterior crown of lower car- 
nassial subequal. Heel about the size of mg. Width across crown of 
m' greater than width of carnassial. Mg relatively large with three 
prominent cusps. 

Measurements.— Table 48. 

Comparisons.— Ictonyx striatus differs from all other small Egyp- 
tian carnivores, except Poecilictis libyca, in having black and white 
markings. Comparison with P. libyca is under the latter. Ictonyx 
striatus erythreae can be distinguished from subspecies sudanicus 
by reddish tone to black parts of pelage, black pigmented areas 
more extensive, and smaller dimensions (Setzer, 1956). 

Specimens examined.— Total two. 

SUDAN ADMINISTRATIVE: Wadi Darawena (2). 

Co//ection.— Trapped in parkland in traps set for fox in Wadi 
Darawena (Hoogstraal et al., 1957ab). 

Food— Reptiles, rodents, and bird eggs (Dorst, 1970). 

Reproduction. — Number of young two to three. Fur is short and 
marked like adults (Dorst, 1970). 

Genus Mustela Linnaeus, 1758 
Slender, short-haired, brown weasels. Legs short; tail short, cylin- 



406 FIELDIANA: ZOOLOGY 

drical. Rostrum short, broad. Cranium elongate, dorsoventrally flat- 
tened. Postorbital swelling absent. Postorbital constriction nar- 
rower than rostrum. Bulla moderately inflated and not fused with 
parapterygoid. 

Mustela nivalis Linnaeus, 1766 

Mustela nivalis Linnaeus. 1766. Syst. Nat.. 12th ed., p. 69. 

Type locality.-Sweden: VESTERBOTEN. 

General distribution.— Circumpolar in temperate, north 
temperate, and arctic regions. Also in Lebanon, Egypt, Morocco, 
and Algeria. 

Common names.— Weasel, Ersa. 

Subspecies in Egypt — 

Mustela nivalis subpalmata (Hemprich and Ehrenberg, 1833). 

Mustela subpalmata Hemprich and Ehrenberg. 1833, Symbolae Physical Manun., 
Vol. 3. foUo k. p. 2. 

Type locality.— Egypt. Houses of Cairo and Alexandria. 

Distribution in Egypt.— Figure 121. Lower Nile VaUey and Nile 
Delta. 

Diagnosis. — BrovfYi above, whitish below. Body slender, tail and 
legs short. Skull flattish, rostrum very short and broad. Hamular 
process of parapterygoid not fused with bulla. 

Head and body length of adult male and female, respectively, 
average 278, 242 mm.; tail 116, 99 mm., 41.6 per cent of head and 
body length; foot 50, 38 mm.; ear 21, 18 mm.; condyloincisive length 
50.0, 43.2 mm. 

External characters. — Figure 122. Dorsum and side dark brown. 
Venter whitish to cream. Demarcation between side and belly 
straight or irregular. Chin white, throat sometimes spotted. Tail 
unicolor, tip shghtly darker than rest of tail and body. Toes whitish. 
Hair of palm sometimes whitish, sole brown. 

Cranial characters. — Figure 124. Cranium elongate, shallow, 
sagittal and lambdoidal ridges prominent; the latter not exceeding 
posterior level of upper lip of foramen magnum. Rostrum markedly 
short and broad. Postorbital process relatively small. Postorbital 
swelling nil. Postorbital constriction narrower than rostrum (table 
50). Tympanic and mastoid bullae moderately inflated. Tympanic 
bulla not fused with parapterygoid. Mastoid process prominent. 






E 




Fig. 124. Sku]\ of Mustela nivalis subpalmata. 



407 



408 FIELDIANA: ZOOLOGY 

Paroccipital fused to bulla. Zygomatic arch not strongly curved 
upward. Tip of coronoid process of lower jaw. angular. 

Teeth.— Outer upper incisor slightly larger than others. Upper 
and lower premolars single cusped, triangular in lateral view, 
crowded, and crowns oblique to axis of tooth row (Miller. 1912, p. 
414). Carnassial trenchant. Outer anterior cingulum of upper car- 
nassial obsolete, inner anterior cusp reduced. Constriction between 
anterior and posterior crowns shallow. Inner cusp of lower car- 
nassial obsolete and heel slightly larger than m,,. Width across 
crown of m' less than width of carnassial (pm'). M;^ reduced and 
simple. 

Baculum. — Baculum is slender, grooved below, and hooked dis- 
tally. 

Measurements.— Table 50. Male dimensions average considerably 
larger than female. Flower (1932) listed the weights of three females 
as 200 gm. each. 

Age determination.— Adults have well-developed cranial ridges. 

Variation.— BeWy color varies from nearly pure white to creamy 
yellow. The area varies individually from a narrow, irregular, and 
occasionally broken midventral line with wider patches on chest and 
throat to a completely pale underside, sometimes extending onto 
side of throat and head. Bonhote (1909) mentioned variation in 
amount of white on the underparts. 

Table 50. — Means (and ranges) of measurements and ratios of adult male (M) and 
female (F) Mustela nivalis subpalmata. 



HBL 


M 


288.8 (252-301) 9 


RW 


M 


11.4(10.7-14.2)7 




F 


241.8(232-259)5 




F 


9.2 ( 8.7-10.2)6 


TL 


M 


116.8(109-129)9 


POW 


M 


8.5 ( 7.9- 8.8) 7 




F 


99.4(94-110)5 




F 


7.9 ( 7.4- 8.7) 6 


TL/HBL* 


70 M 


42.2 (39.2-45.9) 9 


MW 


M 


25.6 (25.0-26.8) 7 




F 


41.1 (37.8-46.4)5 




F 


21.1 (20.0-22.3) 6 


FL 


M 


50.2 (45-55) 9 


P-P 


M 


15.8(15.4-16.5)7 




F 


38.6 (34-42) 5 




F 


13.6(13.3-13.9)4 


EL 


M 


21.3 (20-23) 9 


C-M' 


M 


14.0(13.5-14.2)7 




F 


18.0(15-20)5 




F 


11.9(11.5-12.5)5 


CIL 


M 


50.0 (48.2-51.2) 7 


SH 


M 


16.7(16.2-17.4)7 




F 


43.2(41.8-43.9)5 




F 


14.8(14.4-15.2)5 


ZW 


M 

F 


28.4 (26.8-29.3) 6 

23.5 (22.2-24.9) 6 









OSBORN&HELMY: MAMMALS OF EGYPT 409 

Comparisons.— Mustela nivalis differs from other Egyptian 
Mustelidae in having brownish color; slender, short tail; absence of 
postorbital swelling; and bulla not fused with parapterygoid. 
Mustela nivalis subpalmata is larger than northern subspecies. 

Specimens examined— Total 38. 

QALYUBIYA: Shubra Shihab (1). Kafr el Shurafa (1). 

GIZA: Tanash (2), Mena (1), Giza Zoological Gardens (4). El Mansuriya (2), 
Sakkara (1). 

CAIRO: Cairo (15). Bulaq el Dakrur (2), Abassia (4), Sharabiya (1). 
EL FAIYUM: Shakshuk (2). Sella (2). 

Published records.— Records are from Taylor (1897), Anderson 
(1897, 1902), Flower (1932), Setzer (1952, 1959c), and Hoogstraal 
(1964). 

ALEXANDRIA: Alexandria. 
SHARQIYA: El Qanayat (sight record). 
QALYUBIYA: Sindbis. 
DAQAHLIYA: SimbiUawein 8 km. W. 

GIZA: Tanash, El Mansuriya. Kafr Teharmes, Kuneissa. Saqyet Meki, Abu 
Rawash, Sakkara. 

CAIRO: Cairo, Abassia, Old Cairo walls (sight record). 
EL FAIYUM: Shakshuk, SeUa. 

Habitat— Most specimens are from houses and public buildings; 
a few from cultivated fields and canal banks. According to 
Hoogstraal (1964), the species is no longer numerous in public 
places, such as clubs, restaurants, and theaters, as Flower (1932) 
observed earlier. 

Habits. — In Egypt, M. nivalis is almost completely commensal. It 
is mainly nocturnal, but individuals have been seen during the day 
(Taylor, 1897; Flower, 1932). 

Food— Anderson (1897) reported natives saying weasels killed 
rats and mice in houses. Stomach contents have contained 
cockroaches, tenebrionid beetles, red ants, a small bird, fish, and 
fish bait. 

Reproduction. — Flovfer (1932) noted a Utter of five born in 
December. 

Remarks.— RiippeW (1826) considered the Egyptian and European 
weasels to be conspecific, but thought that the species had been 
introduced into Egypt. 



410 FIELDIANA: ZOOLOGY 

Family 3. Viverridae 

Small to medium size carnivores. Pelage coarsely grizzled or spot- 
ted and stripyed. Muzzle long; ears relatively short, rounded; legs 
short in proportion to body length. Feet semi-plantigrade to 
digitigrade. Toes 5-5, poUex and hallux vestigial, claws semiretrac- 
tile in some genera. Tail somewhat bushy, longer than two-thirds 
length of head and body. Anal scent glands usually well developed. 

Rostrum relatively long, upper tooth row length less than one-half 
skull length. Paroccipital process fused completely to tympanic 
bulla. Bulla constricted externally, divided by septum. Alisphenoid 
canal absent. Baculum well developed. 

Middle lower incisor raised above level of other two; canines 
small, elongate; premolars small, pm, reduced or absent; upper car- 
nassial usually without anterior lobe, lower with well-developed 
talon; molars relatively large, first much larger than second. Dental 
formula: i I, i. ix 2=40, 



Key to Egyptian Genera of Viverridae 

1. Body spotted and striped, tail banded. Tail cylindrical. Ear longer than broad. 
Frontal swelling slight, postorbital bar lacking. Postpalatal margin slightly 
behind last molar. Posterior chamber of tympanic bulla not inflated below level of 
anterior chamber Genetta, p. 410. 

2. Body and tail grizzled. Tail tapering. Ear broader than long. Frontal and post- 
orbital swelling prominent. Postorbital bar complete in adults. Postpalatal 
margin at level of glenoid fossa. Posterior chamber of tympanic bulla inflated 
below level of anterior chamber Herpestes, p. 415. 

Genus Genetta Oken, 1816 

Cat-like, long-bodied, long-tailed, short-legged carnivores. Pelage 
striped and spotted. Tail with contrasting dark and light rings. 
Claws short, semi-retractile. 

Skull elongate, postorbital processes moderately developed. 
Palatal margin not extended posteriorly. Chambers of tympanic 
bulla about subequal. 

Genetta genetta (Linnaeus, 1758) 

Viverra genetta Linnaeus, 1758. Syst. Nat., 1 0th ed., p. 45. 

Type locality.— Spain. 

General distribution.— Southv/estern Europe, northwestern 
Africa, southeastern Egypt and Sudan, west to Senegal and south 



OSBORN&HELMY: MAMMALS OF EGYPT 411 

into Somalia, thence west and south into South Africa. In the 
Arabian Peninsula: Israel, Aden, and Yemen, 

Common name.— Common Genet. 

Subspecies in Egypt — 

Genetta genetta senegalensis (Fischer, 1829) 

Viverra senegalensis Fischer, 1829, Synopsis Mammalia, p. 170. 

Type locality.— Senegal. 

Distribution in Egypt. — Figure 121. Southeastern and 
southwestern parts of Eastern Desert. 

Diagnosis.— Body elongate, weasel-like. Pelage short, soft, 
grayish with black spots and stripes. Tail long, cylindrical, ringed 
with black and yellowish bands. Palm and sole haired. Ear long and 
narrow. 

Skull elongate. Cranium markedly constricted posteriorly. 
Nasofrontal region slightly inflated. Postpalatal margin slightly 
posterior to last molar. Postorbital process moderately developed. 
Cranium broadest at sides. 

Head and body length average 454 mm.; tail 382 mm., 85 per cent 
of head and body length; foot 78 mm.; condyloincisive length 79.8 
mm. 

External characters.— Figure 125. Dorsum with median black 
crest extending from shoulder to base of tail. Six thin stripes on 
neck and shoulders and rows of elongate spots on dorsum and sides. 
Stripes and spots blackish or brownish on yellowish gray 
background. Chest and belly grayish to buffy. Axilla and groin 
whitish. Lacrimal stripe black. Muzzle tip and suborbital area 
whitish. Frontal area pale gray with a dark median stripe. Crown 
and ear grayish. All hairs with gray bases. Tail with series of alter- 
nate blackish and pale rings. Blackish rings number 9-10 and are 
complete; pale rings are yellowish above, whitish below. Tail tip is 
usually whitish, sometimes black. Outer side of legs and upper part 
of feet are grayish. Palm and sole haired, black; toe pads not 
conceciled. 

Cranial characters.— Figure 126. Cranium elongate, narrowly con- 
stricted posteriorly; broadest on sides, narrower at bases of 
zygomatic processes of temporals. Nasofrontal region slightly 
swollen, postorbital swelling nil. Parietal ridges obscure, sagittal 




Fig. 125. Museum specimen 
of Genetta genetta senegalensis. 



412 




J.rt. Qror s. 



Fig. 126. Skull of Genetta genetta senegalensis. 



413 



414 FIELDIANA: ZOOLOGY 

ridge prominent posteriorly; lambdoidal ridge prominent, with 
superior edge slightly caudad of occipital condyle. Frontal process 
extends about one-half length of nasal, but does not contact 
premaxilla. Posterior margin of nasals anterior to frontomaxillary 
suture. Malar in contact with lacrimal. Postorbital process 
moderately developed. Malar not thickened vertically, postorbital 
process nil. Infraorbital foramen large, roundish. Incisive foramen 
elongate. Postpalatal foramen posterior in position, opposite 
anterior edge of pm"*. Postpalatal margin has a median spine and is 
slightly posterior to last molar. Hamular process of parapterygoid 
slender and pointed. Post-tympanic chamber larger than anterior, 
but not so large as in Herpestes. Meatal opening large, roundish. 
Coronoid process of mandible high and slender; angle slender and 
unmodified. 

5acu/um.— Baculum small, 6 to 7 mm. long; swollen at both ends, 
especially at base; and shaped somewhat like a phalanx (Didier, 
1948). 

Teef A.— Similar to, but smaller than, Herpestes, except for 
anterior premolars. Anterior face of upper incisor row slightly con- 
vex. Outer incisors larger than others. First premolars, upper and 
lower, larger than corresponding outer incisor. Pm simple; pmj 
with small posterior cusp; pm 2. 3. 4 with prominent posterior cusps. 
Upper carnassial has a prominent inner anterior lobe bearing low 
cusp and an obscure antero-lateral cusp-like cingulum. First upper 
molar width narrower than pm\ Lower carnassial with three promi- 
nent anterior cusps; posteriormost largest, inner smallest; heel area 
less than one-half that of crown and smaller than m.2. 

Measurements.— Tdhle 51. Male and female dimensions are 
subequal. 

Comparisons.— Genetta genetta differs from all other small Egyp- 
tian carnivores in having prominent stripes and rows of spots on 
body and rings or bands on tail. Cranially and dentally, it differs 
from the Mustelidae in the elongate cranium and rostrum and 
greater number of teeth. From Herpestes it differs cranially in lack 
of prominent frontal and postorbital swellings, poorly developed 
postorbital processes, and less prominent inflation of posterior tym- 
panic chamber. 

Remarks.— The trinomen G. g. senegalensis is tentatively re- 
tained for Egyptian specimens. 



OSBORN&HELMY: MAMMALS OF EGYPT 415 

Table 51. — Means (and ranges) of measurements and ratios of Genetta genetta 
and Herpestes ichneumon. 





Genetta g. 


Herpestes 




senegalensis 


i. ichneumon 


HBL 


453.8 (408-528) 4 


560.3 (546-608) 6 


TL 


381.8 (352-516) 4 


433.6 (363-460) 6 


TL/HBL% 


85.2 (66.6-98.8) 4 


77.4 (65.8-82.8) 6 


FL 


77.8 (74-86) 4 


104.8(101-113)6 


EL 


41.0 (37-46) 4 


35.8 (35-37) 6 


CIL 


79.8 (78.7-80.8) 4 


102.9 (99.8-109.4) 9 


ZW 


41.1 (38.9-43.7) 5 


51.2 (49.4-54.1) 9 


RW 


12.3(11.4-13.0)5 


19.2 (18.4-20.6) 9 


POW 


12.6(12.0-13.1)5 


18.4 (17.0-20.3) 9 


BOW 


28.6 (27.7-30.5) 5 


34.9 (33.6-35.5) 9 


NL 


18.0 (17.2-19.0) 5 





I-M' 


34.7 (34.1-35.6) 4 


42.6 (41.1-44.8) 8 


M'-M' 


24.1 (22.2-25.6) 5 


30.8 (29.7-31.9) 9 


SH 


30.1 (29.5-30.6) 4 


38.1 (36.7-39.3) 9 



Specimens examined.— Total six. 

SUDAN ADMINISTRATIVE: Bir Kansisrob (2). 
ASWAN: Gebel Adda (1). 

Sudan. UPPER NILE: Paloich 1.2 km. NE (1); Malakal 25 km. N (1), 13.3 km. N 
(1). 

Collection.— Trapped alive and shot at night under a spotlight 
(Hoogstraal et al., 1957ab; Hoogstraal, 1964). 

Habitat— Dry savanna, acacia parkland, and rocky slopes of 
desert mountains. 

//a6its.— Nocturnal. Dorst (1970) refers to genets as blood- 
thirsty, wasteful killers. 

Food— Various rodents, birds, reptiles, insects, and some 
vegetable material (Dorst, 1970). Hoogstraal (1964) suggested spiny 
mice (Acomys cahirinus) were a main food source in the Gebel Elba 
area. 

Genus Herpestes Illiger, 1811 

Weasel-like, long-bodied, long-tailed, short-legged carnivores. 
Pelage coarse, grizzled. Tail broad and flattened at base, distal one- 
half tapered. Claws nonretractile. 

Skull elongate, narrow, and deep. Postorbital processes well 
developed, fused in adults to form a postorbital bar. Postpalatal 



416 FIELDIANA: ZOOLOGY 

margin extended posteriorly. Posterior chamber of tympanic bulla 
much larger than anterior chamber. ^ 

Herpestes ichneumon (Linnaeus, 1758) 

Viverm ichneumon Linnaeus, 1758, Syst. Nat., 10th ed.. p. 43. 
Type locality.— Egypt: ad ripas Nili. 

General distribution.— Spain, Portugal, Dalmatia, Turkey, 
Lebanon, Israel, Jordan, Egypt, Libya, Morocco, Algeria, Sudan, 
Ethiopia, Kenya, Central Africa west through Nigeria, southern 
Mali etc., eastern equatorial Africa and southern Africa except for 
parts of southwestern and South Africa. 

Common names.— Egyptian Mongoose, Nims. 

Subspecies in Egypt — 

Herpestes ichneumon ichneumon (Linnaeus, 1758) 

Distribution in Egypt— Figure 127. Nile Delta, Nile Valley south 
to Asyut, El Faiyum, and Burg el Arab. 

Diagnosis.— Body elongate, weasel-like. Pelage long, coarse, 
grizzled blackish brown and cream. Tail long and tapering with 
black tip. Palm and sole naked. Ear short, broad and rounded. 

Skull elongate, nasofrontal and postorbital regions prominently 
inflated. Postpalatal margin at level of glenoid fossa. Postorbital 
bar complete or nearly so. Cranium broadest at base of zygomatic 
processes of temporals. 

Adult head and body length average 560 mm.; tail 434 mm., 77 
per cent of head and body length; foot 104 mm,; ear 41 mm.; con- 
dyloincisive length 102,9 mm. 

External characters.— Figure 128. Dorsum and side hairs grizzled. 
Guard hairs long, coarse, with eight alternating blackish brown and 
cream bands. Under hairs yellowish brown to orangish with 
brownish bases. Venter partly grizzled, clear yellowish brown or 
orangish medially. Muzzle blackish; frontal, cheek, and throat hairs 
short and grizzled. Mystacial area to orbit and circumorbital area 
sparsely haired or almost bare. Ear short, broad, and pale brownish. 
Tail color of dorsum, hair longest at base, and gradually shortening 
toward tip. Tip a tuft of long, black hairs. Feet black or brown. Palm 
and sole naked, pigmented. Juvenile pelage like adult but paler. 

Cranial characters. — Figure 129. Cranium elongate, deep and 




en 



.13 
tlO 



es 



& 
4 



o 
O 



417 




Fig. 128. Cadavers of Herpestes ichneumon ichneumon. 
418 






Fig. 129. Skuli of Herpestes ichneumon ichneumon. 



419 



420 FIELDIANA: ZOOLOGY 

narrow, constricted slightly posteriorly; broadest at base of 
zygomatic processes of temporals. Nasofrontal and postorbital 
regions prominently swollen. Postorbital constriction relatively 
broad. Ridges strongly developed. Superior edge of lambdoidal 
ridge extending beyond posterior level of upper lip of foramen 
magnum. Frontal process extending about one-third length of nasal 
and contacting premaxilla. Posterior margin of nasals level with 
frontomaxillary suture. Postorbital bar complete in adults. Malar 
thickened vertically behind postorbital process and not contacting 
lacrimal. Infraorbital foramen small, elongated dorsoventrally. In- 
cisive foramen ovoid. Postpalatal foramen small, forward in posi- 
tion, opposite anterior edge of pm^. Postpalatal margin truncate or 
serrate, never with median projection, and extending posteriorly 
almost to level of glenoid fossa. Hamular process of parapterygoid 
thickened terminally. Post-tympanic chamber swollen ventrally 
below level of anterior chamber and laterally beyond level of 
mastoid. Meatal opening small, elongate. Coronoid process of man- 
dible short, broad; angle expanded laterally. 

Baca/um.— Baculum relatively large, about 18 mm. long; anterior 
portion slender; posterior enlarged, hollow, and ladle-like ventrally, 
with saddle-like dorsal projection (Didier, 1948). 

Tee tA.— Anterior face of upper incisor row straight. Outer incisors 
much larger than others. First premolars, upper and lower, simple, 
smaller than corresponding outer incisors. Upper and lower second 
molars triangular in lateral view, pm^ ^ without secondary cusps; 
pm2. 3 with posterior basal cingular cusps; pm4 with well-developed 
posterior secondary cusp. Upper carnassial with large anteromedial 
lobe bearing prominent cusp; anterolateral cingulum prominent and 
bearing a distinct cusp. First upper molar width greater than pm\ 
Lower carnassial with three prominent anterior cusps; anterior 
crown and inner cusp subequal and about two-thirds height of 
posterior crown. Heel low, one-half area of m2. 

Measurements.— Table 51. Male and female dimensions are 
subequal. 

Age determination. — Adults have nasofrontal suture fused, 
median sagittal ridge well developed. 

Comparisons.— Herpestes ichneuman is distinguishable from all 
other Egyptian carnivores by its speckled coloring; long, tapering 
tail; short, broad ears; high, narrow skull; swollen frontal region; 



OSBORN&HELMY: MAMMALS OF EGYPT 421 

and elongate palate. Further differences between this species and 
Genetta genetta are under the latter. 

Ten subspecies in addition to H. i. ichneuman are listed from 
Africa (Allen, 1939). Differences among them appear to be trivial. 

Specimens examined.— Total 51. 

BEHEIRA: Dilingat (2). El Tarrana (1). 

TAHREER: Nubareia (1). 

SHARQIYA: Bilbeis (3). Zagzig (1). 

QALYUBIYA: Sindbis (1). 

MINUFIYA: Mohammed Ali Barrage park (1). 

GIZA: El Mitimdiya (1). Kirdasa (1), Abu Rawash (17), El Baragil (3), Zawyet Abu 
Musallam (1), Nahya (1), Minshat el Bakkari (3), Kafret Nassar (3), Cairo- Alexandria 
desert road km. 5 (1), El Harraniya (1), Beni Magdul (5), El Kom el Ahmar (1). 

CAIRO: Cairo (3). 

Sight record of I. Helmy.— 

MATRUH: Burg el Arab, 1976. 

Published records.— Records are from Anderson (1902), Flower 
(1932), Setzer (1952), and Hoogstraal (1964). 

BEHEIRA: Kom Hamada. 

DAMIETTA: Fariskur. 

QALYUBIYA: Sindbis. 

SHARQIYA: Bilbeis. 

DAQAHLIYA: Simbillawein 8 km. E. 

GIZA: Mena, Giza, Imbaba. 

CAIRO: Abassia Fever Hospital grounds. 

Distribution notes. — Flower (1932) reported having seen this 
species throughout El Faiyum and said it occurred "for certain" in 
the Upper Egyptian Governorates of Beni Suef, Minya, and Asyut. 
He said it was reported from as far south as Wadi Haifa, but had no 
evidence of its occurrence in Lower Nubia. "Reports of mongooses 
south of Asyut are unconfirmed" (Hoogstraal, 1964, p. 219). 

The specimen from Nubareia and the sighting at Burg el Arab 
represent recent expansion of range following completion of an 
irrigation canal into the desert. 

Co//ection.— Trapped or dug from burrows. 

//a6i(a(s.— Cultivated areas of Nile Valley and Delta, near water. 

//a6its.— Terrestrial, but readily enters water and swims well. 
Diurnal and crepuscular. Although appearing to be slow moving 



422 FIELDIANA: ZOOLOGY 

when seen crossing roads, mongooses are extremely alert and agile. 
When excited, the long hair is raised and back arched, nearly dou- 
bling the animal's bulk, a common trait among mongooses (Pocock, 
1941: Hinton and Dunn, 1967). 

Burrows. — Burrows are in cultivated areas and in canal banks. 

Food. — Rodents, birds, bird eggs, probably poultry, reptiles, 
frogs, fish, and various aquatic and terrestrial invertebrates. 
Reported to eat eggs of Nile crocodile (Anderson, 1902). 

/Reproduction.— Wild-born litters have been found in February, 
May, July, September, and October, which suggests there is no 
fixed breeding season (Flower, 1932). Litter sizes are two to four 
(Dorst, 1970). 

Remarks.— The mongoose, or Pharaoh's cat, was revered in 
ancient Egypt because of its taste for crocodile eggs and ability to 
kill poisonous snakes (Anderson, 1902). This author and others have 
also mentioned its popularity as a household pet (Russell, 1831; 
Flower, 1932; Hoogstraal, 1964). 

Family 4. Hyaenidae 

Large carnivores with body or legs striped or body spotted, and a 
dorsal mane. Muzzle relatively long; ears large, erect. Hind limbs 
shorter than fore. Feet digitigrade, toes 4,5-4; 4-4 functional. Claws 
short, blunt, nonretractile. Tail bushy, less than two-thirds length 
of head and body. 

Rostrum relatively long and broad. Alisphenoid canal absent. 
Upper tooth row slightly longer than one-half length of skull. Paroc- 
cipital in contact with bulla and projecting below it. Bulla moderate- 
ly inflated. 

Kky to Egyptian Gknkra of Hyaknidae 

1. Size large. Skull and jaws massive. Bulla undivided. Incisors unspecialized. outer 
much larger than inner; canines powerful: carnassials well developed; premolars 
large, crowns conical (function in crushing bone): molars large. . . Hyaena, p. 422. 

2. Size smaller. Skull and jaws weak. Bulla divided. Canines long and slender. Outer 
incisors slightly larger than inner. Premolars small, widely spaced; carnassials 
undeveloped: molars small Proteles, p. 432. 

Genus Hyaena Brisson, 1762 

Large-headed, dog-like carnivores. Shoulders markedly higher 
than rump. Body striped or stripes on legs only. Toes four on fore 



OSBORN & HELMY: MAMMALS OF EGYPT 423 

and hind feet. Skull very large, teeth massive. Dental formula: |, |, §, 
iX2=34. 

Hyaena hyaena (Linnaeus, 1758) 

Canis hyaena Linnaeus, 1758. Syst. Nat., 10th ed., p. 40. 

Type locality— Iran, LARISTAN: Benna Mts. (Thomas, 1911). 

General distribution. — India, Nepal, Afghanistan, Pakistan, Iran, 
Southern Russian Turkestan, Transcaucasia, Asian Turkey, Syria, 
Lebanon, Iraq, Saudi Arabia, Yemen, Jordan, Israel, Sinai Penin- 
sula, Egypt, Libya, Algeria, Morocco, Sudan, Asben, Ethiopia, 
Somalia, Kenya. 

Common names.— Striped Hyena, Dubbah, Dab. 

Subspecies in Egypt — 

Hyaena hyaena dubbah (Meyer, 1793) 

Hyaena dubbah Meyer, 1793, Uebersicht der Entdeckungen in Neu-Holland und 
Africa, p. 94. Based on Bruce 's travels to discover source of Nile. 

Type locality.-Sudan. NORTHERN: Atbara. 

Distribution in Egypt— Figure 130. Sinai Peninsula, Eastern and 
Western Deserts in part. 

Diagnosis.— Body and legs transversely striped black and 
grayish. Dorsal crest of long, black-tipped hairs. Tail relatively 
short, brush-like. Head disproportionately large. Ear large, 
blackish. 

Skull massive, frontal region inflated, cranium slightly wider than 
rostrum, sagittal ridge extremely high and prominent. Angular pro- 
cess of lower jaw prominent, spoon-shaped, and above level of tooth 
row. Teeth very large, especially carnassials. Protocone of p'* not 
sloping forward, but extending at right angle to main axis of tooth. 

Head and body length average 1,038 mm.; tail 308 mm., 30 per 
cent of head and body length; foot 210 mm.; ear 152 mm.; condyloin- 
cisive length 214 mm.; weight 18 to 20 kg. 

External characters. — Figure 131. Stripes brownish or blackish on 
whitish or pale buff ground color. Stripes on neck and side 
transverse, broken. Three broad diagonal stripes cross shoulder 
onto chest; two or three narrower stripjes cross hips diagonally onto 



424 FIELDIANA: ZOOLOGY 

, 25* 26* 27' 28* 2 9* 30* 31* 32* 33* 34* 35* 36* 37* 




Fig. 130. Collection localities of Hyaena hyaena dubbah, sight records IS), and 
tracks IT). Proteles cristata Icircle). 

hind limb. Markings on neck and belly faint. Throat blackish or 
brownish. Stripes on legs sharply defined, incomplete on inner side. 
Toes brownish above, buff between. Muzzle thinly haired, grizzled 
or grayish. Vibrissae sparse, very stiff; whitish, grayish, or black. 
Lips thick, blackish. Ground color of cheek, head, and neck darker 
than rest of body. Ear large, pointed; hair sparse, except on edges, 
and buffy. Skin of ear blackish on outer surface, paler with dark 
spots in inner. Dorsal crest or mane broad and extending from nape 
to base of tail. Hairs long (about 250 mm.), coarse with blackish ter- 
minal band (55 mm.), white subterminal band (45 mm.), four or five 
alternating bands of black or brown and white (80 mm.), and broad 
white basal band (65 mm.). Tail relatively short, brush-like, white 
with narrow dorsal stripe of black-tipped hairs and a black tip. 



OSBORN & HELMY: MAMMALS OF EGYPT 



425 




Fk; 131. Cadaver of Hyaena hyaena dubbah. 

Young have same markings as adult. Sexes easily recognized by 
external genitalia. Gland dorsal to anus prominent. 

Locomotion. — Hyenas running on level ground appear clumsy. 
The gait is a slow, laboring canter accentuated by the long forelimbs 
and up and down movement of head. In rough country, movement 
appears more rapid and graceful. 

Cranial characters. — Figure 132. Skull massive, cranium high and 
narrow. Sagittal ridge very high and prominent; supraoccipital pro- 
jection extending well beyond posterior level of occipital condyle. 
Zygomatic arch very thick, flaring widely posteriorly. Rostrum 
relatively short, broad; breadth nearly that of sides of cranium. 
Nasals deeply separated anteriorly, attenuated posteriorly. 
Nasomaxillary contact nil. Frontal slope steep and frontal region in- 
flated. Postorbital process very large, slightly concave dor sally, 
ridged posteriorly. Postorbital process of malar equally large. 
Postorbital swelling of cranium prominent. Greatest width of 
cranium is at base of zygomatic processes of temporal bones. 




Fici 132. Skull of Hyaena hyaena dubbah. 



426 



OSBORN & HELMY: MAMMALS OF EGYPT 427 

Posterior end of malar is at mid-section of zygomatic arch. Incisive 
foramen elongate. Palatine foramen at level of second premolars. 
Posterior margin of palate at level of posterior edge of m'. Occipital 
area narrow and triangular. Paroccipital process prominent, project- 
ing ventrally slightly below bulla. Tympanic bulla prominently 
swollen, surface smooth, undivided. External auditory meatus with 
tubular bony orifice. Lower jaw curved upward posteriorly. Angular 
process prominent, spoon-shaped, above level of tooth row. 

Teeth.— Outer upper incisors about twice size of inner. Canines 
very heavy and powerful. First upper premolar small, conical, 
sometimes absent in adults; pm^ and pm^ broad, not compressed, 
long axes oblique to jaw margin. Carnassial (pm^) large, powerful, 
with protocone extending at right angle to main axis of tooth, not 
sloping forward. M' large and three-rooted. Lower teeth comparable 
in development; m, with functional talonid. Carnassials function in 
slicing, crushing, and chopping (Ewer, 1954). 

Baculum. — Absent. 

Measurements.— Table 52. Male and female measurements are 
subequal. Dorst (1970) gives the weight as 50 kg. 

Variation.— Co\or of stripes and throat vary from blackish to 
brownish; ground color whitish to pale and grayish buff. 

Comparisons.— Hyaena h. dubbah of eastern Africa differs from 
H. h. barbara of northwestern Africa chiefly in the smaller size of 
skull and pm^ (Pocock, 1934). From H. h. syriaca, H. h. dubbah dif- 
fers in smaller cranial dimensions when compared with Harrison's 
(1968) data. Presence or absence of underwool in winter coat as a 
character appears to be of no taxonomic importance. 

Specimens examined.— TotaX 25. 
MINUFIYA:Minuf (1). 

Table 52. — Means (and ranges) of measurements, ratios, and weight of adult 
Hyaena h. dubbah. 

HBL 1038.0(1020-1075)4 RW 51.9(50.0-54.2)5 

TL 308.0(290-350)4 POW 37.2(30.3-42.3)5 

TL/HBL% 29.6(27.8-32.6)4 MW 80.2(76.1-84.1)5 

PL 210.0 (205-215) 4 NL 61.4 (59.2-64.2) 3 

EL 152.2(145-155)4 PM'-PM' 83.2(75.7-86.7)5 

Wt(kg.) 19.2(19-20)4 PM' 31.1(29.8-32.3)5 

CIL 213.8(209-220)5 I-PM* 107.1(105.8-109.4)5 

ZW 150.5 (146.2-157.1) 5 



428 FIELDIANA: ZOOLOGY 

GIZA: Giza Pyramids area, gallery 1 km. NW of Cheops Pyramid (one skull and 
two lower jaws). 

MATRUH: Bahig (one lower jaw from cave). 

ASWAN: Bir Murra (3). Bir Haimur (1), AUaqi VUlage9-10 km. S (4). El Dirr. E of 
(1). Wadi el Targama (1). Khor Abusku (9). 

Sudan. NORTHERN: Wadi Haifa (1). 

Sight records. — Records of D. Osborn, I. Helmy, H. Hoogstraal, 
and Frontier Corps Soldiers (last reported by I. Helmy). 

SUEZ: Wadi Dom. Wadi Abu Sanduq (tracks. 1956). Ras Abu el Darag. 
GIZA: Giza Pyramids area. 

ASWAN: Wadi Abu Subeira. Barqet Tokham (tracks. 1966). Wadi Kurkur. Qustul 
West. Gebel Adda (tracks. 1963). Umm Shilman Plains (tracks and dens. 1966). 
EL WADI EL GEDEED: Bir Karawein. Mut. 
QENA: Dandara. S of (tracks. 1966); Abydos. W of (tracks. 1968). 
MATRUH: Qara (1976). El Ghazalat near Bir Abd el Nabi (tracks. 1976). 

Published reports.— Hyenas in Sinai may represent either 
subspecies syriaca or sultana. Specimens from this area were not 
available. Reports are from Oliver (1804, cited by Anderson, 1902), 
Belzoni (1819, cited by Fagan, 1975), Palmer (1872), Murray (1891), 
Buxton et al., (1895), Anderson (1902), Barron (1907ab), Weigall 
(1909), Hurst (1910), Harding-King (1925), Murray (1935, 1967), 
Omer-Cooper (1947), Wassif (1953b), Wassif and Hoogstraal (1953), 
Howells (1956), and Hoogstraal (1964). 

SINAI: Wadi Kid (den). Wadi Gazzah; Maqdaba (tracks). Gebel Musa. Gebel 
Umm Rijlein. St. Catherine Monastery area. 

ISMAILIA: "Hyena Quarries" E of Lake Timsah. 

SUEZ: Gebel Iweibid (caves and old bone middens). 

ALEXANDRIA: Alexandria. 

GIZA: Bahariya Oasis. 

CAIRO: Wadi Hof 1933. 1956. 

EL FAIYUM: Qasr el Sagha and Qasr Qarun. 

RED SEA: Wadi Fatira. 

SOHAG: Gebel el Haridi (tracks). 

QENA: Karnak Temple. Luxor. 

ASWAN: Wadi Abu Subeira. 

MATRUH: Sitra. Siwa. and Maragi Oases. 

EL WADI EL GEDEED: Farafara Oasis. Bir Murr and Bir Karawein; Dakhla 
Oasis; Bir Nakheila. one day's journey N of (tracks). 
• Sudan. KASSALA: Bir Meisa. 

Remarks on distribution.— Judging from statements of early 
writers, hyenas were more numerous and widespread in Egypt 



OSBORN&HELMY: MAMMALS OF EGYPT 429 

previously than today. They commonly entered towns in search of 
food (Russell. 1831). 

Fitzinger (1860) included in the distribution the Western Mediter- 
ranean Coastal Desert and Nile Valley and Delta. Anderson (1902) 
listed desert margins of the Nile Valley from Cairo to Esna, desert 
surrounding El Faiyum, and Bahariya and Farafara Oases. Flower 
(1932) said that hyenas were not known to occur in the Nile Delta, 
but were on both sides of the Nile all the way to Nubia. He also men- 
tioned valleys inland from Salum and along the Mediterranean 
coast east to Alexandria and in Sinai. Omer-Cooper (1947, p. 26) 
reported that hyenas in the vicinity of Siwa and Maragi Oases had 
been "practically exterminated by poison." Native helpers told him 
that hyenas occurred at Sitra. Hyenas which occasionally range 
through the Eastern Desert are thought by natives to come from 
Sudan following rains (Tregenza, 1955). 

The Nile Valley south of Cairo to Nubia appears to be the area of 
highest concentration of H. hyaena in Egypt today. 

Collection.— HyenaiS were shot at night from a car using a 
spotlight (Osborn, 1968a). In Palestine, Bird (1946) shot hyenas at 
night from a blind when they came to a goat carcass he had staked 
out. 

Habitats.— \A]fie other large Egyptian carnivores, hyenas inhabit 
desert areas bordering the Nile Valley and oases and invade 
cultivated areas at night to feed and drink. Those in remote desert 
areas 15 to 25 km. from cultivation probably followed camel 
caravans in anticipation of a dead camel (Anderson, 1902). Osborn 
(1968a) collected hyenas along the ancient caravan road from Sudan 
to Daraw over which thousands of market camels pass each year. 
The Darb el Arbaein (Forty Days Road) and other routes in the 
Western Desert are now rarely used, and hyenas no longer live along 
them. 

In Sinai, hyenas frequent areas of human habitation and coastal 
regions. 

Dens.— Dens are in natural caves and cracks or among boulders or 
blocks of stone and are recognizable by accumulations of bones 
(Anderson, 1902; Barron, 1907b). Tombs and temple ruins are also 
known to have been occupied by hyenas (Murray, 1891). 

//a6its.— Striped hyenas are strictly nocturnal. They are shy, yet 
approach closely camps or dwellings in search of food. No 



430 FIELDI ANA: ZOOLOGY 

documented evidence is available that this species has attacked 
men, although folklore is full of such "reports." 

Osborn (1968a) was told by a Bishari that striped hyenas resorted 
to cannibalism at times of food shortage. Osborn killed one hyena 
that was carrying off the day-old carcass of another hyena used as a 
lure. Brown and spotted hyenas are reputed to be cannibalistic, but 
this was disputed by Hughes (1954). However, cannibalism in the 
Sfjotted hyena was witnessed by Kruuk (1968). 

Striped hyenas drag or carry carcasses to their dens, where con- 
spicuous bone middens accumulate (Anderson, 1902; Barron, 1907b; 
Reed, 1966; Reed, personal communication; and observations of D. 
Osborn). According to Hughes (1954), other species of hyenas do 
not. 

Drake- Brockman (1910) mentioned that old striped hyenas in 
Somalia became destructive and hunted sheep and goats by day and 
slaughtered them wantonly. Murray (1935) reported the mutilation 
of a herd of sheep by striped hyenas near Gebel Umm Rijlein in 
Sinai. 

The cowardliness of striped hyenas has been remarked upon by 
numerous authors. Anderson (1902, p. 200) was able to approach 
within a few yards of hyenas during the day. "When disturbed they 
show no fight, but only anxiety to make off with all possible haste." 
Tate- Regan (1946) said the striped hyena refrained from biting 
when attacked and had a suppressed instinct of self preservation. 
This aspect of hyena behavior figures strongly in folklore, and 
claims have been made that a man can creep into a den and capture 
a hyena alive without any resistance by throwing a cloak over it and 
tying a rope around its legs (Kitto, 1841; Jayakar, 1908). Authen- 
ticated observations of this feat occur in the literature (Wood, 1807; 
Murray, 1935; Kullman, 1965; Street, 1967; Hassinger, 1973). 

Food— Striped hyenas are notoriously eaters of carrion, and their 
powerful jaws and teeth enable them to crack large bones. Hyenas 
collected by Osborn (1968a) in Nubia subsisted chiefly on the re- 
mains of dead camels along the road from Sudan to Daraw. Stomach 
contents also included seeds of heglig {Balanites aegyptiaca). 
Specimens collected from the Nile area near Allaqi Village con- 
tained fish thrown out by fishermen and bones of gazelle and stork. 
There is ample evidence that hyenas will eat human carcasses if 
available (Doughty, 1888; Zeuner, 1963; Harrison, 1968; Osborn. 
1968a). Adams (1870) and Kitto (1841) reported that hungry hyenas 



OSBORN&HELMY: MAMMALS OF EGYPT 431 

are sometimes destructive to crops, especially Indian corn, and they 
are known to feed on dates in the Nile Valley (Murray, 1967). Cap- 
tive hyenas will eat ripe dates, bananas, tomatoes, plums, 
watermelons, etc., "in addition or almost in preference to meat" 
(Flower, 1932, p. 394). If carrion is unavailable, hyenas will move to 
the sea and break open shells (Klunzinger, 1878). In Iraq, hyenas 
were reported to have killed a horse and a donkey, and one person 
told of feeding desert tortoises to a pet hyena (Hatt, 1959). Earlier 
writers mention the hyena as a killer of asses and mules and, occa- 
sionally, cattle (Wilkinson, 1878; Bruce, 1790). The latter stated 
that hyenas had a fondness for dogs and would hunt them in his 
camp. Bird (1946) described hyena depredation in flocks of sheep 
and goats in Palestine. Further notes on the catholic food habits of 
the striped hyena are in Kruuk's (1976) studies in the Serengeti. 

An old hyena cave in the limestone cliff 1 km. NW of Cheops 
pyramid contained skulls and bones of domestic animals, including 
dogs, foxes, pig, and bones of a giant freshwater turtle. Hyena 
skulls and lower jaws were also retrieved from this cave. 

M^ater.— Hyenas drink periodically and are no doubt unable to 
survive without a source of water. 

Economic importance.— Ancient Egyptian peasants hunted 
hyenas for duty and amusement along with other animals that 
destroyed fields or flocks (Kitto, 1841; Wilkinson, 1878). Old 
Kingdom Egyptians force-fed hyenas to fatten for the table, as 
depicted in the tomb of Mereruka (Sixth Dynasty, 2300 B.C.) at Sak- 
kara. Brentjes (1966) has presented evidence that these animals 
were aardwolf rather than hyena. Hyenas were supposedly tamed 
and used in hunting. There is no evidence, however, that Ancient 
Egyptians considered them sacred. 

Numerous sources mentioned the eating of hyenas by modern 
Egyptian peasants. Others known to eat them are certain Arabian 
Bedouins (Doughty, 1888), Palestinian laborers (Zeuner, 1963), 
Sinai Bedouins (Murray, 1935), and Tuaregs (Ihote, 1946). 

Hyenas were once a valuable commodity. The flesh was sold in the 
markets, and Ulema, or religious leaders, were the chief buyers. 
Various parts were sold for charms and medicines (Klunzinger, 
1878). 

Reproduction.— The only records from Egypt are a nearly full- 
term female with one 504-gm. fetus and one resorption, and an old 



432 FIELDIANA: ZOOLOGY 

female with a resorption. Both were taken in March from tributaries 
of Wadi AUaqi. 

Fo/^Zore.— Although considered to be a cowardly animal, the 
striped hyena is feared by Egyptians; particularly farmers and 
Bedouins. Legends depict it as savage, dangerous, treacherous, 
cunning, and greedy. Nevertheless, it is commonly believed that to 
eat the heart will give one courage. Whiskers and eyeballs are 
believed to give protection from the evil eye. Many parts of the 
hyena are used to increase virility or to impart strength or bravery 
in men. Various ailments and afflictions are treated with specific 
organs. Further information on hyena folklore is available in Fitz- 
inger (1860), Klunzinger (1878), Jayakar (1908), Weigall (1909), and 
Osbom (1968a). 

Genus Proteles I. Geoffroy St.-Hilaire, 1824 

Hyena-like insectivore. Striped on body and legs. Toes 5-4; inner 
toe on forefoot vestigial. Teeth peg-like. Dental formula: |, |, i^, 
nx 2=28-32, usually 30. 

Proteles cristatus (Sparrmann, 1783) 

Viverra cristata Sparrmann. 1783, Resa Goda-Hopps-Udden, Vol. 1, p. 581. 

Type /oca/ity.— South Africa. Cape of Good Hope, Somerset East, 
near Little Fish River. 

General distribution.— SonthesiStem Egypt, Sudan, Ethiopia, 
Somalia, Kenya, Tanzania, Mozambique, Rhodesia, South Africa, 
South West Africa. 

Common names.— Aard Wolf, Deeb. 

Probable subspecies in Egypt — 

Proteles cristatus pallidior Cabrera, 1910 

Proteles cristatus pallidior Cabrera, 1910, Ann. Mag. Nat. Hist., (ser. 8), 6, p. 464. 

Distribution in Egypt — Knoy/r\ only from two specimens 
reportedly shot by Negumi (1949) in 1940 near Halaib in SUDAN 
ADMINISTRATIVE GOV. Hoogstraal et al. (1957b) gave the col- 
lection locality as Gebel Hamra Dom some 80 km. N of Halaib (fig. 
130). 

Diagnosis. — Hyena-like, but smaller and slenderer. Skull not 
massive. Teeth peg-like. 

External c/iaracfers. — Hyena-like in body form, forelimbs longer 



OSBORN&HELMY: MAMMALS OF EGYPT 433 

than hind, forefeet with five rather than four toes. Striping on body 
and legs as in Hyaena hyaena. Muzzle, chin, and part of area around 
eyes, nude and black. Dorsal crest prominent. Ear long, tip narrow- 
ly rounded. 

Cranial characters.— ^]sm\\ proportionately smaller and more 
lightly built than in Hyaena. Cranium low, broadest on sides, nar- 
rower at bases of zygomatic processes of temporals. Sagittal ridge 
low. Supraoccipital process extending slightly beyond posterior 
level of occipital condyle. Rostrum blunt, relatively long, sides 
parallel, and width almost equal to that of cranium. Nasals not as 
deeply separated anteriorly as in Hyaena, attentuated posteriorly. 
Nasomaxillary contact very broad. Frontal slope gradual, and fron- 
tal region only slightly swollen. Postorbital swelling nil. Postorbital 
process of frontal large, slightly concave dorsally. Postorbital pro- 
cess of malar equally large. Zygomatic arch flaring. Posterior end of 
malar extending to level of glenoid fossa. Incisive foramen ovoid. 
Posterior margin of palate extending to level of optic foramen. 
Paroccipital process adnate to bulla, not projecting. Alisphenoid 
canal lacking. Tympanic bulla prominent, posterior chamber much 
larger than anterior and extending beyond level of paroccipital pro- 
cess and below level of tooth row. Lower jaw constricted behind 
canines, curved upward posteriorly so that angular process is above 
level of tooth row. Angular process projects posterior to articular, 
but is not spoon-shaped as in Hyaena. 

Teeth. — Incisors and canines normal. Outer incisors slightly 
larger than inner. Canines long, slender, cheek teeth small, conical, 
widely spaced, and in parallel rows. 

Comparisons.— Proteles cristatus differs from H. hyaena in 
smaller size, five toes rather than four on forefoot, skull much 
smaller and lighter, and cheek teeth peg-like rather than sectorial. 

Specimens examined.— Total two. 

SUDAN ADMINISTRATIVE: Gebel Hamra Dom (1). 
Southern Rhodesia: Bulawayo 30 km. W (1). 

//a6i to (.—Plains and savanna. 

i/a6its.— Nocturnal and apparently shy and secretive. 

Food — Insects, primarily termites and larvae when available; 
probably carrion, eggs, and small vertebrates. 



434 FIELDIANA: ZOOLOGY 

Family 5. Felidae 

Medium to large carnivores. Pelage usually marked with stripes 
and/or spots. Muzzle conspicuously short; ears more or less 
triangular, sometimes tufted. Legs moderately long relative to body 
length. Feet digitigrade; toes 5-4; inner toe on forefoot vestigial. 
Claws sharp, strongly curved, retractile (semiretractile in 
Acinonyx). Tail cylindrical, length variable among species. Tongue 
covered with horny, curved papillae. 

Rostrum and nasals short; cranium short, rounded. Upper tooth 
row length less than one-half skull length. Paroccipital process flat- 
tened against bulla. Tympanic bulla conspicuously inflated, divided 
by septum. Postpalatal foramen on maxillopalatine suture, not on 
maxilla. Alisphenoid canal absent. Baculum absent or vestigial. 

Postorbital process of zygomatic arch prominent. Incisors small, 
chisel-like, and in tranverse line; canines elongate, sharply pointed; 
post canine diastema present, except in Acinonyx', premolars sharp; 
pm' reduced, often absent; carnassials large, well developed, lower 
smaller than upper; upper molar small, crown tranverse. Dental for- 
mula: |, }, |, j x 2=30. 

Key to Egyptian Genera of Felidae 

1. Small to medium size cats. Markings faint stripes and spots. Ear tuft, if present, 
very short. Nasal branch of premaxilla broad. Posterolateral margins of palate 
deeply notched Felis, p. 434. 

2. Medium size cats. Markings very faint to nil. Ear tuft very long. Facial stripe 
from nose through lacrimal onto forehead. Nasal branch of premaxilla long, thin, 
usually contacting frontal process. Posterolateral margins of palate shallowly 
notched Caracal, p. 447. 

3. Large cats. Markings prominent spots. Nasal branch of premaxilla narrow, not 
contacting frontal process. Posterolateral margins of palate without notches. 

a. Spots in rosettes. Black facial stripe absent. Skull elongate, profile flattish. 
Malar-maxillary suture below infraorbital foramen at lowest level. Infraorbital 
foramen large, ovoid Panthera, p. 451. 

b. Spots solid. Black facial stripe from mouth to eye. Claws nonretractile. SkuU 
short, profile highly domed. Malar-maxillary suture completely above infra- 
orbital foramen. Latter small, narrowed vertically Acinonyx, p. 455. 

Genus Felis Linnaeus, 1758 

Small to medium-size cats. Color pattern of indistinct stripes and 
spots in adults. Back of ear black and/or reddish. Claws completely 
retractile. Skull broad, rather evenly rounded or domed in lateral 
outline. Nasal branch of premaxilla broad opposite tip of nasal, then 
becoming abruptly pointed. Posterolateral margins of palate deeply 



OSBORN & HELMY: MAMMALS OF EGYPT 435 

notched. Postcanine diastema wide. First upper premolar usually 
present. Inner cusp of upper carnassial usually well developed, ex- 
cept in F. margarita. Anterior accessory cusp of second upper 
premolar small and not in line with main cusp. 

Key to Egyptian Species of Genus Felis 

1. Color dark. Ear small. Black elbow mark absent. Tympanic bulla moderately in- 
flated. Nasals ending about at level of frontomaxillary suture. 

a. Size large, tail relatively short (one-third head and body length), hind foot 
length over 140 mm. Ear tuft present, short. Cheek stripe absent. Skull large, 

condyloincisive length over 95 mm. Postorbital swelling present 

chaus, p. 435. 

b. Size small, tail relatively long (two-thirds head and body length), hind foot 
length under 140 mm. Ear tuft absent. Cheek stripe present. Skull small, con- 
dyloincisive length under 95 mm. Postorbital swelling absent 

sylvestris, p. 440. 

2. Color pale. Ear large, broad. Black elbow mark present. Tympanic bulla greatly 

inflated. Nasals long, ending posterior to level of frontomaxillary suture 

margarita, p. 444. 

Felis chaus Giildenstaedt, 1776 

Felis chaus Guldenstaedt, 1776, Nov. Com. Acad. Petrop., 20, p. 483. 
Type locality.— \J .S.S.R.: Terek River N of Caucasus. 

General distribution.— Yietnam, Thailand, Burma, Yunnan Pro- 
vince of Western China, Nepal, India, Ceylon, Chinese and Russian 
Turkestan, Afghanistan, Pakistan, Iran, Iraq, west shore of Cas- 
pian to Volga Delta, eastern Transcaucasia, southern Turkey, 
Israel, Jordan, Egypt. 

Common names.— Jungle Cat, Swamp Cat, Qut Barri (male), 
Qutta Barria (female). 

Subspecies in Egypt— 

Felis chaus nilotica De Winton, 1898 

Felis chaus nilotica De Winton, 1898, Ann. Mag. Nat. Hist., (ser. 7), 2, p. 292. 
Type locality.— Egypt. CAIRO: near Cairo. 

Distribution in Egypt— Figure 133. Nile Delta, Nile Valley south 
to Aswan, El Faiyum, Farafara and Dakhla Oases, Western 
Mediterranean Coastal Desert. 

Diagnosis.— Color dark, grizzled buff. Body markings indistinct. 
Lacrimal stripe dark brown, prominent. Cheek plain. Ear reddish 
brown with black tip and small tuft. Tail relatively short with 
several black disted rings and black tip. 



436 FIELDIANA: ZOOLOGY 

,. 2 5* 2 6* 2 7* 28* 2 9* 30* 31* 32* 3 3* 3 4* 35* 36* 37* 




Fig. 133. Collection localities of Felis chaus nilotica (squares). F. sylvestris Uhyca 
(dots), F. s. tristrami (circles), and F. margarita (triangle). 

Rostrum and cranium elongate. Postorbital width slightly more 
than rostral width. Anterior end of zygomatic process attenuate. 

Head and body length average 674 nrni.; tail 254 mm., 38 per cent 
of head and body length; foot 168 mm.; ear 71 mm.; condyloincisive 
length 112.8 nmi.; weight 9.0 kg. 

External characters.— Similar to F. sylvestris, but head and body 
markings less conspicuous, except for black ear tufts and prominent 
dark brown lacrimal stripe. Dorsal line blackish, reddish tinted 
posteriorly. Dorsum yellowish brown grizzled with black and 
yellow. Side and outer side of legs paler. Under hair fuscous. Chin 
pale buff; throat darker, grizzled; chest, belly, and inside of legs 
yellowish. Axilla and groin whitish. Faint, pale brownish spots on 
side and belly and faint stripes on upper legs. Nasal region pale buff, 
lacrimal stripe dark brown, supraorbital patch whitish, cheek plain. 



OSBORN & HELMY: MAMMALS OF EGYPT 



437 



Forehead, crown, and nape faintly striped. Ear reddish brown 
behind, base blackish, tip black with short tuft and inner side 
whitish. Tail relatively short, concolorate with back proximally and 
grayish distally with two narrow black bands and short black tip. 
Feet orangish to brownish yellow above, palm and sole blackish or 
brownish. Young have a more distinct pattern than adults. 

Cranial characters.— Figure 134. Rostrum and cranium somewhat 
elongate. Frontal and postorbital swelling prominent. Postorbital 
width slightly more than rostral width. Nasals tapering gradually 
posteriorly; posterior margin level with or slightly posterior to fron- 
tomaxillary suture. Cranial crests and ridges strongly developed. 
Anterior end of zygomatic process attenuate. Malar-maxillary 
suture below infraorbital foramen at its lowest level. Mastoid pro- 
cess large, protruding. 

TeetA.— Dentition similar to but much larger than F. sylvestris, 
especially anterior cusp of upper carnassial. 

Measurements.— Table 53. 

Comparisons.— Felis chaus differs from F, sylvestris in having 
body meu-kings less conspicuous, cheek stripe lacking, lacrimal 
stripe more prominent, black ear tufts, tail shorter, skull more 
elongate, postorbital swellings, anterior end of zygomatic process 

Table 53. — Means (and ranges) of measurements, ratios, and weight of Felis 
sylvestris and F. chaus. 





Fs. 


. tristrami 


F. s. libyca 


F. c. nilotica 


HBL 


498.5 (471-545) 4 


449.5 (373-483) 6 


674.5 (595-760) 12 


TL 


319.2 (282-390) 4 


291.2 (237-337) 6 


254.8 (210-280) 12 


TL/HBL% 


64.0 (57.8-78.3) 4 


64.8 (62.5-70.8) 6 


37.8 (33.8-44.5) 12 


FL 


121.0(115-131)4* 


124.8(110-136)6 


168.4 (145-178) 12 


EL 


56.5 (55-58) 4 


61.5 (56-70) 6 


71.4 (63-78) 12 


Wt (kg.) 







2.5, 3.8 


9.0 ( 7.0-11.2)5 


CIL 







83.1 (70.8-90.6) 11 


112.8(98.2-123.9) 18 


ZW 







64.6(53.9-72.5) 11 


79.8 (70.0-92.1) 16 


RW 







22.6(19.6-25.3) 11 


30.8 (26.6-34.0) 18 


POW 







32.6(26.5-35.0) 11 


36.2 (31.5-38.6) 17 


MB 







40.6 (37.2-42.9) 7 


48.0 (42.6-53.8) 15 


NL 







24.4 (22s5-26.3) 10 


36.6 (31.8-41.8) 17 


PM' 







10.9 ( 9.9-11.9)9 


14.9(13.5-16.5) 18 


PM'-PM' 







36.7 (34.5-39.4) 8 


48.6 (44.8-53.4) 15 


CM' 







26.3 (26.1-31.7) 12 


39.6 (34.9-41.9) 14 


SH 







45.4 (41.2-49.3) 9 


57.1 (48.8-61.6) 14 


♦Not including claw. 











Fl(i 134. Skull of Felis chaus nilotica. 



438 



OSBORN&HELMY: MAMMALS OF EGYPT 439 

attenuated instead of rounded, and larger dimensions. According to 
Pocock (1951), the only way some individuals of the two species can 
be distinguished externally is by relative tail lengths, 

Felis chaus differs from F. margarita in having darker color, less 
conspicuous markings, narrower ears, pads of feet not covered with 
hair, relatively shorter tail, relatively shorter nasals, and much 
smaller bullae. 

From F. chaus furax of the Eastern Mediterranean, F. c. nilotica 
differs in having darker color and smaller teeth. 

Specimens examined.— TotsX 38. 

ALEXANDRIA: Alexandria (1), Amiriya-Alexandria road, about 15 km. W of 
Alexandria (1), about 20 km. W of (1). 

QALYUBIYA: Tukh, El Ahmar (4); Sanafir, Esbet Ibsan (2); Qalyub (1). 

GIZA: Giza (1), El Baragil (1), Abu Rawash (2), Beni Yusef (1), Giza Pyramids (1), 
Sakkara (1), Kafr Hakim (1). 

CAIRO: Cairo (Type, 3). 

EL FAIYUM: Tamiya (2), Sinnuris (1), Kom O Shim (1), Fanus (1), no exact locali- 
ty (1). 

SOHAG: Akhmim (2). 

QENA: Farshout (1), Wadi Nassim (1). 

ASWAN: Aswan (1). 

MATRUH: Bahig (1), 14 km. S (1). 

EL WADI EL GEDEED: Farafara Oasis, Hatiyet el Sheikh Marzuk (one old 
poorly mounted skin); Dakhla Oasis; Mut (1), 4.8 km. N (1). 

Published records.— Records are from De Winton (1898), Ander- 
son (1902), Bonhote (1909), Flower (1932), Wassif (1960b), and 
Hoogstraal (1964). 

QALYUBIYA: Benha; Tukh. 
GIZA: Giza Zoological Gardens. 
EL FAIYUM: Various localities. 

QENA: Wadi Nassim; Nag Ayed, Tuftish Farshout (Farshout). 
ASWAN: Aswan. 

MATRUH: Bahig; Bahig 14 km. SW; Mersa Matruh, E and W of; Wadi el Rami 
(16.6 km. S of Mersa Matruh). 

EL WADI EL GEDEED: Dakhla Oasis. 

Co/fection.— Easily trapped or shot. 

Habitat— how cultivated or marshy ground, reed beds, fields of 
sugar cane, bean fields, or any similar thick cover (Anderson, 1902; 
Flower, 1932). Reference by Anderson to cornfields between Alexan- 
dria and Siwa must mean Bedouin barley fields west of Alexan- 



440 FIELDIANA: ZOOLOGY 

dria. We have taken specimens near Alexandria in reed (Phragmites 
aus trails) swamp and in low vegetation of the Western Mediterra- 
nean Coastal Desert (fig. 19). Flower (1932) reported this cat from 
cliffs east and west of Mersa Matruh. 

Behavior.— Felis chaus shows "remarkably little fear of man" 
(Hoogstraal. 1964, p. 222). One kiUed with a .22-caliber pistol at 
night near Alexandria was hit on the third shot. It showed no in- 
dication of fright (I. Helmy, personal communication). 

Food— According to Flower (1932), F. chaus eats snakes of genera 
Coluber and Psammophis and dead fish. One collected from near 
Alexandria had its stomach full of fish which it had either stolen 
from fishermen or scavenged. Bonhote (1909) remarked that this cat 
did considerable damage each year to animals and birds in the Giza 
Zoological Gardens. Reference was also made to predation on sheep. 

Reproduction.— lAtter sizes are two to three, rarely four to six, 
and young are born from January to April (Flower, 1932). 

Remarks.— fiiiicient Egyptians mummified F. chaus, but whether 
it was domesticated at the time is debatable (Morrison-Scott, 1952; 
Monaiery, 1965). 

Felis sylvestris Schreber, 1777 

FeUs {Catus) sylvestris Schreber, 1777, Die Saugeth., 3, p. 397. 
Type locality.— Germany. 

General distribution.— British Isles, Western Europe, Balkans, 
Turkey, Ukraine, Transcaucasia, Russian and Chinese Turkestan, 
Kazakstan, India, Afghanistan, Iran, Iraq, Arabian Peninsula, 
Syria, Lebanon, Israel, Jordan, Sinai Peninsula, Egypt, Sudan 
across North Africa to Morocco, south of the Sahara into North 
Nigeria, Asben, Ethiopia and Somalia, and southward into South 
Africa. 

Common names.— Wild Cat, Qut Gebeli. 

Distribution of subspecies in Egypt— Figure 133. Felis sylvestris 
libyca: margins of Nile Valley and Delta, oases, and Western 
Mediterranean Coastal Desert; Felis sylvestris tristrami: Sinai 
Peninsula. 

Diagnosis.— Slender, similar to house cat. Color grizzled buff with 
indistinct stripes and spots. Lacrimal stripe pale brown. Cheek 



OSBORN&HELMY: MAMMALS OF EGYPT 441 

striped. Ear reddish brown, tuft nil. Tail relatively long with several 
distal rings and black tip. 

Rostrum and cranium short and broad. Postorbital width about 
one and one-half times rostral width. Anterior end of zygomatic pro- 
cess broad and rounded. 

Head and body length average 450 mm.; tail 291 mm., 64 per cent 
of head and body length; foot 124 mm.; ear 62 mm.; condyloincisive 
length 83.1 mm.; weight 3 kg. 

External characters.— SimiXax to domestic cat, but legs and tail 
longer. Dorsal line blackish. Dorsum buff grizzled with black and 
white or yellow, side and outer side of legs paler. Under hair grayish. 
Chin and throat whitish; chest, belly, and inside of legs whitish to 
buff. Axilla grayish or white, groin white. Pale brownish spots on 
belly and side change to faint vertical lines on shoulder and flank. 
Upper legs with broad brownish bands. Broken stripes on feet. 
Nasal region orangish, lacrimal stripe pale brown, and supraorbital 
patch whitish. Faint stripes on cheek, crown, nape, and dorsum. Ear 
reddish brown behind, margin blackish, tip without tuft, inner hairs 
whitish or cream. Tail relatively long, blackish above at base, with 
three black distal rings, and black tip. Feet yellowish above, palm 
and sole black. 

Cranial characters.— Figure 135. Rostrum and cranium short and 
broad. Frontal and postorbital swelling nil. Postorbital width about 
one and one-half times rostral width. Nasals tapering abruptly 
posteriorly, with posterior margin anterior to or level with fronto- 
maxillary suture. Cranial ridges not strongly developed. Anterior 
end of zygomatic process rounded. Malar-maxillary suture with ven- 
tral margin level with lower edge of infraorbital foramen. Mastoid 
process small and appressed. 

Bacu/um.— Baculum about 5 mm. in length; cross-shaped due to 
lateral projections near base (Didier, 1949). 

Measurements.— Table 53. 

Comparisons.— Comparison of F. sylvestris and F. chaus are 
under the latter. Felis sylvestris differs from F. margarita in having 
less conspicuous markings, smaller ears, relatively shorter nasals, 
and smaller bullae. 

Remarks.— Some authors (Smithers in Meester and Setzer, 1971) 
consider F. libyca as a separate species. Mummified cats described 



442 



FIELDIANA: ZOOLOGY 




Fkj. 135. Skull of Felts sylvestris libyca, 

as F. lihyca bubastis Hemprich and Ehrenberg (1833) are considered 
to be a variety of domestic cat (F. catus) by Morrison-Scott (1952) 
and Haltenorth (1953a). Monaiery (1965) proposed that domestic 
cats originated in Egypt from wild stock. 

Habitats.— Dry situations in rocky or wooded districts (Ander- 
son, 1902). Hoogstraal (1964) reported F. sylvestris from the 
Western Mediterranean Coastal Desert in a barley field in flat, 
vegetated desert, and a desert valley. One was collected near Abu 
Mena in habitat shown in Figure 8. 



OSBORN&HELMY: MAMMALS OF EGYPT 443 

//a6its. —Nocturnal. 

Food— Hares, rodents, reptiles, birds; probably young of gazelles. 
Said to eat insects and fruits (Dorst, 1970). 

Reproduction.— No data from Egypt. Dorst (1970) lists two to five 
as number in litters. He also remarked that F. sylvestris interbred 
with domestic cats. This we heard also from Bedouins in Egypt. 

Subspecies in Egypt— 

Fells sylvestris libyca (Forster, 1780) 

Felis libyca Forster. 1780, in Buff on. Nat. Vierf. Thiere. Vol. 6. p. 613. 
Type /oca/ity.— Tunisia: Gafsa. 

Distribution in Egypt— Figure 133. Margins of Nile Valley and 
Delta, Western Mediterranean Coastal Desert. 

External characters.— In comparison with F. s. tristrami, F. s. 
libyca is paler and more buffy, markings are less prominent, back of 
ears is paler, and there is shghtly more black on the feet. 

Cranial characters. —See species description. 

Measurements.— Table 53. 

Specimens examined.— Total 18. 

GIZA: Bahariya Oasis. Mandisha (1). EI Aguz (1). 

MATRUH: Burg el Arab (1); Bahig (5). 5 km. S (1), 16 km. S (1). 48 km. S (1); Abu 
Mena 5 km. SE (1); El Qarasat (1). El Maghra (1). 
EL WADI EL GEDEED: Dakhla Oasis. Mut (1). 
BENI SUEF: Maidum (1). 
ASWAN: Aswan (1). Wadi el Targama (1). 

Published records.— Records are from Anderson (1902), Flower 
(1932), and Hoogstraal (1964). 

MATRUH: Bahig SSW 16 to 50 km. 
CAIRO: Wadi Hof (skull found in cave). 
BENI SUEF: Maidum. 

Remarks.— A specimen from Bir Victoria was described as having 
yellowish body color and reddish tail (De Winton, 1903). 

Felis sylvestris tristrami (Pocock, 1944) 

Felis libyca tristrami Pocock, 1944, Ann. Mag. Nat. Hist., (ser. 11). 11, p. 125. 
Type locality.— Jordan, MOAB: Ghor Seisaban. 



444 FIELDIANA: ZOOLOGY 

Distribution in Egypt— Figure 133. Sinai Peninsula. 

External characters. — In comparison with F. s. libyca, F. s. 
tristrami is darker and more grayish, markings are slightly more 
prominent, backs of ears are darker, and there is less black on the 
feet. 

Cranial characters.—See under species description. 

Measurements.— Table 53. No external measurements available 
from Sinai specimens. Cranial measurements of one Sinai specimen 
are: CIL 89.9, ZW 72.2. POW 35.2, RW 26.0. NL 27.3, PM'-PM* 
38.9, SH 47.8. (See Appendix 1 for explanations of abbreviations.) 

Specimens examined.— Total one. 
SINAI: No exact locality (1). 

Published records.— Records are from Flower (1932) and Harrison 
(1968). 

SINAI: Awlad Ali in Wadi el Arish (sight record), Abu Durda Mines between Tor 
and Has Jehan (specimen). 

Felis margarita Loche, 1858 

Felis margarita Loche. 1858, Rev. Mag. Zool., 10, No. 2.. p. 49, pi. 1. 

Type locality.— Algeria. SAHARAN OASES: Ngoussi (Negousa, 
Negonca). 

General distribution.— Southern Russian Turkestan, Iran, Ara- 
bian Peninsula, Algeria, Egypt, Libya, Morocco, and Asben. 

Common name.—Sand cat. 

Probable subspecies in Egypt— 

Felis margarita margarita Loche, 1858 

Type locality. —See above under species. 

Distribution in Egypt— Not known, but reportedly "found in 
very sandy tracts of desert only" by Flower (1932, p. 390), who 
observed two in the 2^1ogical Garden at Zagazig, 1912-1914. 
Listing of Sinai by Ellerman and Morrison-Scott (1951) was without 
documentation. One specimen was collected by L Helmy in 1975 in 
the southwestern part of the Eastern Desert (fig. 133). 

Diagnosis.— Color pale grizzled buff. Body markings indistinct. 
Foreleg markings prominent. Ear very broad; distal one-fourth 
black behind, base rufous. Tail slightly more than one-half head and 



OSBORN&HELMY: MAMMALS OF EGYPT 445 

body length. Hair of palm and sole covering pads. Ears set low 
giving broad, flat appearance to head. 

Rostrum and cranium short, broad. Postorbital width one and 
one-fourth times rostral width. Anterior end of zygomatic process 
gradually tapering. Bulla greatly inflated. Basioccipital constricted. 

Head and body length average 461 mm.; tail 225 mm., 55 per cent 
of head and body length; foot 110 mm.; ear 66 mm.; condyloincisive 
length 85.4 mm. 

External characters.— Fur longer and silkier than in other species 
of Felis. Dorsal line blackish. Dorsum pale buff grizzled with black 
and white. Side and outer side of legs paler. Under hair grayish. 
Venter white except for orangish throat. Indistinct longitudinal 
streaks on head, nape, and shoulders. Vertical grayish stripes on 
side and flank. Two black stripes encircle foreleg and fuse with 
patch inside elbow. Thigh with five fairly distinct stripes. Face 
whitish. Upper muzzle, lacrimal, and postorbital stripes orangish 
buff. Ear broad, inner side white, margin pale buff, distal one-fourth 
of back black, base rufous. Latter color extends onto nape and side 
of neck. Tail grayish above, whitish to buff below, with four in- 
distinct stripes dorsally near tip and conspicuous black tip. Feet 
whitish to buff above; palm and sole brownish; hair long, curly, and 
completely covering pads. 

Cranial characters.— Figure 136. Rostrum and cranium short and 
broad. Zygomatic arches relatively wide. Skull arched in lateral 
outline. Frontal and postorbital swellings prominent. Nasals con- 
stricted slightly medially, converging abruptly posteriorly, and 
ending well behind level of frontomaxillary suture. Lambdoidal 
ridge well developed, sagittal crest prominent posteriorly, frontal 
ridges continuous with postorbital processes. Maxillary pro- 
tuberance dorsolateral to infraorbital foramen obsolete. Anterior 
end of zygomatic process attenuate. Mastoid process prominent. 
Paroccipital process adnate to bulla. Tympanic bulla greatly in- 
flated. Basioccipital noticeably constricted and narrower than 
mesopterygoid fossa. Malar-maxillary suture sloping and below 
level of infraorbital foramen posteriorly. 

Teet/i. — Inner lobe of upper carnassial reduced, though 
protoconid is distinct. 

Measurements.— External and cranial measurements of one im- 
mature Egyptian specimen are: HBL 438, TL 248, HF 117, Ear 71, 



446 



FIELDIANA: ZOOLOGY 







Flu 136. Skull of Felis margarita. 

CIL 77.9, ZW (approx.) 61.8, POW 35.4, RW 18.3, NL 24.4, PM'- 
PM' 32.0, SH 44.6. (See Appendix 1.) 

Comparisons.— Felis margarita is easily identified from other 
si)ecies of Felis by its paler color; prominent facial and foreleg mark- 
ings; large, broad ears lacking tufts; rounded, shortened cranium 
and rostrum; relatively large tympanic bulla; and constricted 
basioccipital. Hemmer et al. (1976) recognized F. m. margarita as 
the smallest of four races. 



OSBORN&HELMY: MAMMALS OF EGYPT 447 

Specimens examined.— Total three. 

Saudi Arabia: El Rub el Khali (1). 

Aden: Beihan (1). 

Egypt. ASWAN: Wadi el Targama (1). 

Sight record of I. Helmy.— 
ASWAN: Wadi el Targama (3) 1974. 

//a6i(a(.— Reported to occur in sandy areas of desert, to which the 
species appears to be adapted due to the long fur covering the feet, 
large ears, and greatly inflated tympanic bullae. 

Be/iayior.— Nocturnal. Short legs and low-set, broad ears allow 
the sand cat to present a low profile; an advantage to a predator in 
sparsely vegetated areas. Further notes are in Hemmer (1974) and 
Hemmer et al. (1976). 

Food— Reptiles, birds, and rodents. 

/Reproduction.— Known to give birth to four and five young. 

Genus Caracal Gray, 1843 

Medium size, long-limbed, short-tailed cat with long ear tufts. 
Pelage without pattern except striping on side and spotting on 
venter. Nasal branch of premaxilla long, attenuate; sometimes con- 
tacting frontal process, which is also elongate. Notch in postero- 
lateral edge of palate shallow. Postcanine diastema short. First 
upper premolar usually absent. Inner cusp of upper carnassial 
reduced. Inferior edge of lower jaw strtiight. 

Caracal caracal (Schreber, 1776) 

Felis caracal Schreber. 1776, Die Saugeth.. pi. 110. text 3. pp. 413. 587. 

Type locality.— Sonth Africa: Cape of Good Hope, Table 
Mountain. 

General distribution.— Central India, Afghanistan, Russian 
Turkestan, Iran, Asian Turkey, Syria, Jordan, Israel, Saudi Arabia, 
Sinai Peninsula, Egypt, Libya, Algeria, Morocco, Sudan west to 
Mauritania. Ethiopia, Somalia, Kenya south and southwest into 
Mozambique, and South Africa. 

Common names.— Caracal, Umm Rishat. 

Probable subspecies in Egypt.— 



448 



FIELDIANA: ZOOLOGY 



,.2 5* 2 6* 2 7* 28* 2 9* 30* 31* 3 2* 3 3* 34* 35* 36* 37 




Fig. 137. Collection localities of Caraco/ caraca/ scAmttei. 



Caracal caracal schmitzi (Matschie, 1912) 

Felis {Caracal) caracal schmitzi Matschie, 1912, S.B.Ges. Nat. Fr. Berlin, p. 64. 
Type locality.— Jordan: Ain ed Dachubeijir. 

Distribution in Egypt— Figyire 137. Sinai Peninsula, northern 
part of Eastern Desert, 

Diagnosis.— Light reddish brown on dorsum and side, venter 
white. Tail short, color of dorsum, tip black. Ear with long black 
tuft. 

Cranium relatively long, rostrum short. Postorbital and rostral 
widths subequal. Anterior end of zygomatic process gradually 
tapered, tip rounded. 

External characters.— Dorsum light reddish brown grizzled with 
white. Side paler and with scattering of buffy spots or faint striping. 



OSBORN & HELMY: MAMMALS OF EGYPT 



449 




Fk; 138. Skull of Caracal caracal schmitzi. 



Chin and gular regions whitish; throat and chest rufous; belly and 
inside of legs whitish with scattering of spots. Faint striping on 
outer side of legs. Face reddish brown, paler above eye. Blackish 
stripe continuous from nose through lacrimal onto forehead. 
Mystacial aiesi blackish. Ear elongate, blackish behind, tip with 
long black tuft (40 to 60 mm.); inner side and margins whitish. Tail 
relatively short, indistinctly bicolored. Feet faintly marked with 
brown, palm and sole light brown, pads partly concealed. 



450 FIELDIANA: ZOOLOGY 

Cranial characters.— Figure 138. Rostrum short, cranium relative- 
ly long. Skull outline rounded in lateral view. Cranial ridges 
moderately developed. Nasal branch of premaxilla long, attenuate; 
sometimes contacting elongate frontal process. Nasals narrowed 
gradually posteriorly, posterior margin rounded or truncate and 
level with or slightly anterior to frontomaxillary suture. Postorbital 
swelling nil. Postorbital processes relatively short. Postorbital and 
rostral widths subequal. Malar-maxillary suture sloping and below 
level of infraorbital foramen posteriorly. Anterior end of zygomatic 
process gradually tapered, tip rounded. Mastoid process pro- 
truding. Paroccipital adnate to bulla. Bulla inflation as in Felis 
chaus. Posterolateral margin of palate with slight notch, medial 
margin further posterior than in Felis. Lower jaw with inferior edge 
straight and angular process not curving upward. 

Bacu/um.— Baculum length about 6 mm.; base broadened, 
triangular in outline (Didier, 1949). 

Teef A.— Dentition similar to Felis, except the inner cusp of the 
upper carnassial is more reduced. 

Comparisons.— Caracal caracal differs externally from other 
Egyptian Felidae by having less conspicuous m£u*kings, reddish 
color, long ear tufts, and shorter tail. Cranially, C. caracal differs 
from Felis species in having a long, attenuate nasal process of the 
premaxilla, less developed postorbital processes, and a straight 
inferior margin on the lower jaw. From Panthera and Acinonyx^ 
Caracal differs in lacking spots, having ear tufts, small postero- 
palatal notches, and contact between premaxilla and frontal 
processes. 

Specimens examined.— Total four. 

SINAI: El Arish (1); Tor. N of (1). 

CAIRO: Foum el KhaUg (Old Cairo) (1). Gebel MokatUm (1). 

Published records and reports.— The following are listed by date, 
where possible, from Anderson (1902), Flower (1932), Russell 
(1949ab), Hoogstraal et al. (1957b) and Hoogstraal (1964). 

SINAI: El Arish (19201. Tor N of (1902). 

SUEZ: Between Helwan and Gulf of Suez (1902). 

CAIRO: Foum el Khalig (Old Cairo) (1904). Gebel MokatUm (1939). 

RED SEA: Bir Abraq (1940). Wadis of Eastern Desert (1940's). 

//a6i (ats.— Savanna, rocky and hilly desert. 



OSBORN&HELMY: MAMMALS OF EGYPT 451 

Food— Gazelles, hares, rodents, birds, reptiles. 

Remarks.— T\us cat was known to the ancient Egyptians as in- 
dicated by drawings on tomb walls at Beni Hassan (Anderson, 
1902). 

Genus Panthera Oken, 1816 

Large, long- tailed cats. Color plain or with stripes or broken 
spots. 

Skull elongate, nasals broad. Profile of cranium flattish or broadly 
convex. Nasal branch of premaxilla narrow. Posterolateral margins 
of palate without notches. Tympanic bulla relatively small. Anterior 
cusp of second upper premolar internal and relatively small. 

Panthera pardus (Linnaeus, 1758) 

Felis pardus Linnaeus, 1758, Syst. Nat., 10th ed., p. 41. 

Type locality.— WaRey of the Nile (describer and HoUister, 1918), 
Egypt (Thomas, 1911), Egypt or Sudan (Flower, 1932). 

General distribution.— Eastern Siberia, Manchuria, China, 
Malaysia, Vietnam, Java and Kangean Islands, Burma, Nepal, 
Kashmir, Tibet, India, Sri Lanka, Pakistan, Baluchistan, 
southwestern Turkestan, Iran, Transcaucasia, Asian Turkey, Saudi 
Arabia, Yemen, Syria, Lebanon, Israel, Jordan, Sinai, Egypt, 
Algeria, Morocco, Sudan, Ethiopia, Somalia, thence westward to 
Senegal and the remainder of Africa southward. 

Common names.— Leopard. Memoura, Nimr. 

Probable subspecies and distribution in Egypt— Fig[ire 139. Pan- 
thera pardus jarvisi: Sinai Peninsula; Panthera pardus pardus: 
Eastern and Western Deserts. 

Diagnosis.— harge, long-tailed cat. Color yellowish dorsally with 
pattern of brown to black spots forming rosettes. Venter white. Ear 
black behind with white spot. 

Cranium relatively flat in lateral profile. Postpalatal margin 
posterior to anterior end of presphenoid. Protocone on inner lobe of 
upper carnassial. 

Head and body length average 95-100 cm.; tail 60-95 cm. 

External characters.— harge cat marked with black spots forming 
open rosettes. Dorsal color buff to pale yellow fading to white on 
venter. Spots less dense on underparts and inner sides of legs, 



452 FIELDIANA: ZOOLOGY 

,♦2 5* 2 6* 27* 28* 2 9* 30* 31* 3 2* 3 3* 34* 35* 36* 37* 




Fig. 139. Collection localities of Panthera pardus pardus (dot) and P. p. jarvisi 
(circles). 

broken on chest. Facial markings nil. Ear black behind with white 
spot, whitish or cream on inner side. Tail spotted as body; spots 
becoming solid and appearing as bands distally. 

Cranial characters.— Figure 140. Skull massive, strongly ridged, 
elongate. Nasal process of premaxilla narrow, attenuate. Nasals 
broad, convex, and tapering gradually posteriorly to rounded 
margin about level with frontomaxillary suture. Premaxillary pro- 
cesses of nasal short. Postorbital swelling prominent. Postorbital 
width considerably less than rostral width. Malar very deep. Malar- 
maxillary suture straight, sloping, and ending below level of infra- 
orbital foramen. Latter very large and ovoid with maxillary pro- 
tuberance present and situated above posterior level of upper 
second premolar. Anterior end of zygomatic process acute. Incisive 
foramen ovoid. Palatal foramen opposite inner lobe of carnassial. 







FlU. 140. SkuU oi Panthera pardus. 



463 



454 FIELDIANA: ZOOLOGY 

Lateral postpalatal margins without notches; posterior margin well 
behind anterior end of presphenoid. Mesoptefygoid space narrower 
than basioccipital and with edges curved inward. Mastoid and 
paroccipital processes prominent. Mandible very strong, inferior 
edge curved. 

Tee t A.— Canines relatively long and powerful. Postcanine 
diastemas wide. Cheek teeth similar to Felis, except second upper 
premolar less compressed and with small anterior secondary inter- 
nal cusp. There is a protocone on the inner lobe of the upper 
camassial. 

Comparisons.— Panthera pardus differs from most other Egyp- 
tian feUds by Isu-ger size, pattern of broken spots, flatness of skull, 
largeness of infraorbital foramen, posterior postion of latter, 
posterior position of postpalatal margin, and narrowness of 
mesopterygoid space. 

Habitats.— RocVy mountains, cliffs, and wadis. Known to have in- 
habited Western Mediterranean Coastal Desert and Qattara 
Depression around oases-like areas. 

Food.— According to Murray (1930), the Sinai leopard feeds main- 
ly on hyraxes and ibex. Instances of predation on camels and 
donkeys in Sinai have been reported (Hume, 1906; Murray, 1935). 

iiemar/js.— Leopards are portrayed on tomb walls at Beni Hassan 
(Anderson, 1902). 

Probable subspecies of Panthera in Egypt.— 

Panthera pardus jarvisi Pocock, 1932 

Panthera pardus jarvisi Pocock, 1932, Abstr. Proc. Zool. Soc., London, 1932. No. 
347. p. 33. 

Type locality.— SIN Ah southwestern Sinai, no exact locality. 

Distribution in Egypt— Fignre 139. Sinai Peninsula. 

External characters.— Dorsum creamy buff, flank grayish, venter 
white. Rosettes with centers slightly darkened. 

Specimen examined— SINAI: no exact locality (TVpe). 

Published records and reports.— The following are listed by date, 
where possible, from Palmer (1872), Hart (1891), Buxton et al. 
(1895), Hume (1906), Barron (1907a), Pocock (1932), Negumi (1949), 
Hardy (1949), Wassif and Hoogstraal (1954). and Murray (1935, 
1967). \ 



OSBORN&HELMY: MAMMALS OF EGYPT 455 

SINAI: Wadi Sigilliyeh (tracks. 1872); Ain el Taba (tracks, 1891); Gebel Shomer 
and Gebel Serbal (Bedouin rejwrt, 1891); Wadi Nasli and Wadi er Rimm (tracks, 
1894); Wadi Isla, Wadi Aad, and Gebel Ferani (tracks. 1906): Wadi Threya and Wadi 
Nasb (camels killed by leopards, 1906); Haid Merzega Pass (Bedouin leopard trap, 
1906): Wadi Geba (tracks, 1907); Wadi Sheqer (camels killed by leopards, 1907); 
Moiyet Luliya (Pearl's Spring) below Gebel Yithmid (one seen, 1929): Wadi Hebron 
(11 donkeys killed by leopards, 1929); mountains of Sinai (male and female shot. 
1939-1940): Gebel Serbal (skin, 1942); no exact locality (type and skin. 1900; one 
shot. 1945; mounted skin, early 1950's). 

Panthera pardus pardus (Linnaeus, 1758) 

Type locality.— See under species. 

Distribution in Egypt— Figure 139. Northern part of Western 
Desert (previously) and possibly Gebel Elba area. 

External characters.— Large leopard, variable in size and color. 
More ochraceous buff colored and with spots smaller and darker 
than in P. p. jarvisi. 

Specimens examined.— Total one. 

Sudan: No exact locality (1). 

Published records and reports. —Sources are Barron and Hume 
(1902), Flower (1932), Fahmy (1936), Tregenza (1955), Hoogstraal et 
al. (1957b). 

RED SEA: Said to be absent in Eastern Desert. Guides have mentioned that 
leopards were present in the Eastern Desert in times of more vegetation. 

SUDAN ADMINISTRATIVE: Occasionally found in the Elba Mountains and 
legendary among the Bisharin of that area. 

MATRUH: Skin seen in 1913 from El Maghra 10 km. SW, and said to occur be- 
tween Mariut and Siwa. 

Genus Acinonyx Brookes, 1828 

Monotypic genus of long-limbed, panther-like cats. Pelage spot- 
ted. Ears small. Claws not retractile. 

Skull short, broad, conspicuously domed. Frontal region excep- 
tionally broad, swollen. Nasal branch of premaxilla attenuate. 
Posterior margin of palate without lateral notches. Postcanine 
diastemas nil. Anterior accessory cusp of second upper premolar in 
line with others. Inner cusp of upper carnassial obsolete; lower car- 
nassial with posterior talonid-like cusp. 

Acinonyx jubatus (Schreber, 1776) 

Felisjubata Schreber. 1776. Die Saugeth., 3, pi. 105. 

Type locality.— South. Africa: Cape of Good Hope. 




CO 



JS 
00 



o 

o 



456 



OSBORN&HELMY: MAMMALS OF EGYPT 457 

General distribution.— Parts of India (previously), Baluchistan, 
Afghanistan, southern Russian Turkmenia, Iran, Iraq, Syria, Jor- 
dan, Saudi Arabia, Sinai Peninsula, Egypt, Libya (?), Morocco, Rio 
di Oro, Sudan, Ethiopia, Somalia, Chad, northern Nigeria south into 
southern and southwestern Africa. 

Common names.— Cheetah, Hunting Leopard, Fahd. 

Distribution in Egypt— Figure 141. Northern Sinai Peninsula 
and northern half of Western Desert. 

Diagnosis.— harge, long-tailed cat. Grayish to yellowish dorsally, 
with pattern of solid black spots; venter white. Ear small, tip 
yellowish, base black. 

Cranium conspicuously domed. Infraorbital foramen small, nar- 
rowed vertically. Postpalatal margin almost level with anterior end 
of presphenoid. Upper carnassial with inner lobe reduced and lack- 
ing protocone. 

External characters.— Body size and tail length about as in Pan- 
thera. Legs long, slender. Ear small, short. Paws small, compact. 
Mane on nape and shoulders. Latter covers entire dorsum in cubs. 
Dorsum, side, and outer legs yellowish to pale buff or grayish white 
covered with close-set, solid blackish spots. Venter white. Tail 
marked like body proximally, distal one-third ringed, tip white. 
Prominent, black, curved stripe between mouth and eye. Ear tip 
yellowish, base of ear black. 

Cranial characters. — Figure 142. Cranium thin boned, 
lightweight, relatively short, broad and markedly domed with peak 
anterior to postorbital process. Premaxillary process thin, elongate. 
Nasals very broad, flat, tapering gradually to an abruptly rounded 
posterior margin, and ending at level of frontomaxillary suture. 
Dorsofrontal region very broad and conspicuously swollen. Postor- 
bital processes short. Postorbital swelling prominent. Postorbital 
width slightly more than rostral width. Anterior of zygomatic pro- 
cess broadly rounded. Sagittal crest developed posteriorly only. 
Lambdoidal ridge prominent. Infraorbital foramen opposite middle 
cusp of upper carnassial; small, narrowed vertically, and lacking 
maxillary protuberance. Malar relatively shallow. Malar-maxillary 
suture angular in outline, not extending below level of infraorbital 
foramen. Incisive foramen rounded. Palatal foramen opposite 
posterior cusp of carnassial. Postpalatal margin well behind m^ 
Anterior lip of glenoid slightly developed. Mesopterygoid wider 
than basioccipital, edges curved inward. Mastoid and paroccipital 






Fig 142. SkuU of Acinonyx jubatus. 



I 



458 



OSBORN&HELMY: MAMMALS OF EGYPT 459 

processes protruding and prominent. Exoccipital relatively low and 
broad. Mandible relatively weak, inferior edge almost straight. 

Teeth. — Canines relatively short. Postcanine diastema nil. 
Anterior cusp of second upper premolar well developed and in line 
with main cusp. Upper carnassial length subequal with length of 
canine. The inner lobe is vestigial, and an anterior cingular cusp is 
present. The lower carnassial has a talonid-like cusp. 

Specimens examined.— Total four. 

MATRUH: Cairo-Alexandria desert road km. 125, 15 km. N (1); Qur el Hilab 45 
km. ENE of El Maghra. killed in 1974 by Bedouins (2). 
Kenya: Athi River (1). 

Sight records of I. Helmy.— 

MATRUH: Sitra Oasis (six, 1975), El Zeitun (seven, 1975). 

Published records and reports.— The following are listed by date, 
where possible, from Flower (1932), Hardy (1949), Russell (1951), 
Murray (1935, 1967), Hoogstraal (1964), and Hoogstraal et al. 
(1968). 

SINAI: Sinai Desert (two seen, early 1946). 

MATRUH: Alexandria 66 km. W (tracks of two, 1909); El Maghra 8 km. N (one 
shot, 1910); common around El Maghra (early 1930's); Giza Pyramids 166 km. W 
(tracks, late 1920's); present prior to World War II in low-lying parts of Western 
Desert, such as El Maghra; Salum area (three cubs, 1927; one shot, 1934; skin seen, 
1937; two verbal reports. 1934); Sitra (one seen, 1964); Sidi Barrani (one shot, 1964); 
Qattara Depression cliffs (few pairs reported, late 1920's); Tal el Fawakhir (one seen, 
1965); Hatiyet Labaq (tracks, 1967); Cairo- Alexandria desert road km. 125, 15 km. N 
(one shot, 1967). 

In addition to the above, Omer-Cooper (1947, p. 21) reported that 
hunting leopards occurred "in the coastal belt (a few miles in width) 
of the Western Desert." Bedouins reported cheetahs in Qattara 
Depression from El Maghra to Tal el Fawakhir (Hoogstraal et al., 
1968). 

Habitat— Savanna and semi-desert. Known from Western 
Mediterranean Coastal Desert and sparsely vegetated areas of Qat- 
tara Depression, acacia groves, and oasis-like depressions. 

//a6its.— Usually approaches prey then runs it down, rather than 
stalking like the leopard. Western Desert Bedouins claim "that it 
perches in acacia branches to attack gazelles." (Hoogstraal et al., 
1968, p. 65). 

Food— Hares, gazelles (Russell, 1951; Murray, 1967), domestic 
sheep (Hoogstraal et al., 1968), and probably rodents and birds. 



ORDER HYRACOIDEA 
Family Procaviidae 

Genus Procavia Storr, 1780 

Rock hyraxes. Grayish to yellowish or orangish brown dorsally, 
venter buffy. Frontals of skull flat, triangular, Postorbital process 
of frontal and malar not fused. Tempoparietal ridges fused 
posteriorly in adults. Parietals extend slightly posterior to inter- 
parietal. Cheek teeth brachydont, four-rooted. Upper premolars of 
adults shorter than molars. Dental formula: |, 5, 1, | x 2=34. 

Procavia capensis (Pallas, 1766) 

Cavia capensis Pallas, 1766, Miscellanea 2jOo1., p. 30 pis. 3, 4. 
Type locality.— Africa: Cape of Good Hope. 

General distribution.— Syria, Lebanon, Israel, southern Arabian 
Peninsula, Egypt, Algeria, Libya, Tibesti, Azbine, Sudan, Ethiopia, 
and Somalia southwards to South Africa and westwards to Senegal. 

Common names. — Hyrax, Coney, Dassie, Wabar, Buar, Kalidob. 

Distribution of subspecies in Egypt— Figure 143. Procavia capen- 
sis syriaca: Sinai Peninsula; Procavia capensis ruficeps: Eastern 
Desert. 

Diagnosis.— Rodent or hare-like. Grayish to yellowish brown; 
tailless; ear short, rounded. Palm and sole with large, firm pads. 
Toes with nails. 

Skull strongly built, angular; nasals broadest posteriorly; frontal 
flat, triangular in outline with prominent postorbital process. 
Zygomatic process of temporal begins at level of nuchal ridge. 
Lower jaw deep, ramus and angle greatly expanded. Upper incisors 
tusk-like, lower incisors pectinate. Lophs of upper cheek teeth 
U-shaped; lower, W-shaped. 

Height at shoulders about 25 cm., head and body length 40 cm., 
foot 6.6 cm., ear 3.3 cm., weight 2 to 4 kg. 



460 



OSBORN&HELMY: MAMMALS OF EGYPT 461 

, 25* 26* 27* 28* 2 9* 30* 3l' 32° 3 3' 34* 35* 36* 37* 




Fk; 143. Collection localities of Procauia capensis syriaca (circles), P. c. ruficeps 
(dots), and sight records (S). 

External characters.— Fur relatively short, soft. Dorsum and side 
grayish to yellowish brown; throat, belly, and inside of legs pale 
brownish or buff. Middorsal scent gland marked, in some in- 
dividuals, with yellowish or orangish hairs. Dorsum and side hairs 
with brownish tips, yellowish to orangish subterminal bands, and 
brown bases. All other hairs also have brown bases. Long, blackish 
tactile hairs scattered over dorsum and side. Mystacial, supra- 
orbital, and postauricular areas paler than body. Gular area with 
large vibrissae. Snout short, pointed. Ears short, rounded. Tail lack- 
ing. Feet plantigrade. Palm and sole with firm, naked pads. Forefoot 
with four short toes bearing blunt nails. Hind foot with two long, 
webbed toes bearing nails and an inner grasping toe with sharp, 
curved claw. 

Cranial characters.— Figures 144, 145. Skull strongly built and 




Fu; 144. Skull of Procavia capensis. 



462 





Fig 145. Comparison of position of anterior end of malar relative to lacrimal in 
Procavia capensis ruficeps (A) and P. c. syriacus (B). 



463 



464 FIELDIANA: ZOOLOGY 

angular. Rostrum relatively short, compressed laterally, and nasals 
very broad posteriorly. Interorbital region broad, flattened dorsally; 
frontals triangular with prominent postorbital processes. Lacrimal 
with prominent peg-like projection. Tempoparietal ridges fused 
posteriorly in adults forming a high, median sagittal crest. Inter- 
parietal small, triangular to pentagonal in outline. Parietal extends 
posteriorly slightly beyond level of interparietal. Nuchal ridge well 
developed in adults. Zygomatic process of temporal begins almost 
at the nuchal ridge. Zygomatic arch does not flare laterally. Malar 
very long, extending from level of lacrimal posteriorly to form outer 
part of glenoid cavity; middle broadened vertically and has a long 
postorbital process. Supraoccipital vertical. Paroccipital process 
elongate, extending below levels of occipital condyle and auditory 
bulla. Incisive foramen small, round; posterior palatine foramen 
minute. Posterior margin of palate anterior to last molar. Para- 
pterygoid fossa prominent. Mandible deep; angular region rounded 
and greatly enlarged. 

Tee tA.— Upper incisors two only and continuously growing, wide- 
ly spaced, tusk-like, triangular in cross section, pointed tips. Lower 
incisors broad, pectinate. Cheek teeth brachydont with deep lophs; 
upper with U-shaped lophs, lower, W-shaped. 

Measurements.— Table 54. 

Age determination.— Adult specimens have median sagittal crest 
on posterior cranium and majority of sutures closed. 

Comparisons.— Procavia c. syriaca is darker and more orangish 
than P. c. ruficeps, has hairs of dorsal spot concolor rather than 
banded, and anterior end of malar not contacting lacrimal. 

Vanaf ion.— Pelage of young is markedly grayer than adult. In- 
dividual variation in adult color was noted by Gray (1868), Ander- 
son (1902), and Flower (1932). 

Table 54. — Means (and ranges) of measurements of Procavia capensis. 

P. c. syriaca* P. c. ruficeps 

HBL 473.8 (345-574) 17 464.8 (458-478) 4 

Fl, 72.2 (56-81) 17 68.8 (67-72) 4 

EL 34.8 (26-40) 17 33.8 (30-36) 4 

CIL 88.0(72.7-99.8)21 88.9(86.2-92.1)3 

ZW 50.6(41.2-61.9)21 52.0(51.0-54.0)3 

TRL 36.6(26.2-41.7)21 35.8(36.2-36.2)4 
*Data sumnuirized from Harrison (1968). 



OSBORN&HELMY: MAMMALS OF EGYPT 465 

Collection.— Kyraxes have been caught by hand and by dogs 
when well away from the vicinity of rocks (Hoogstraal et al., 
1957ab; Bedan, 1928). Steel traps set in trails between cliffs and 
food trees will sometimes catch them (Osborn, 1968a). Shooting is 
satisfactory, providing animals are on exposed rocks away from 
dens, otherwise, if an animal does not die immediately, it can 
escape. Gunfire and other noises usually cause hyraxes to disappear 
for half an hour or more (Hoogstraal et al., 1957ab). 

//a6itat.— Figure 146. Rock falls and crevices in cliffs in the 
vicinity of acacia trees. Dens are always in deep crevices too narrow 
for human passage. Remaining walls of Roman buildings provided a 
suitable den site in Wadi Semna. 

Behavior.— Hyraxes live in colonies, are nocturnal and diurnal in 
activity and may be seen sunning on exposed rocks. Feeding is 
mainly at night. They are very agile and can move rapidly and 
gracefully in trees and rocks. Climbing ability is due to glandular, 
cohesive foot pads and almost opposable inner hind toe. A high- 
pitched alarm call is given by sentinels. Other sounds have been 
noted. Feces and urine are deposited habitually in specific places. 
White, bleached urine streaks on rocks are a sign of hyrax dens (fig. 
146), although they remain visible for years after dens have been 
deserted. 

Popu/ations.— Information on populations is sketchy. According 
to Anderson (1902), hyraxes were so abundant in upper reaches of 
Wadi Sheitun that Schweinfurth called it "VeJle di Hyrax." Barron 
and Hume (1902) wrote that hyraxes were once numerous in the 
Eastern Desert as far north as Bir Inglizi (Beida) on the Qena- 
Quseir road, but that, due to promiscuous cutting of acacia trees, 
only a small population remained near the Sudan frontier. Flower 
(1932) reported the disappearance of many colonies between 1914 
and 1920, a period of critical food and fuel shortage in Egypt. 

Small colonies still exist in southern Sinai Peninsula and the 
Eastern Desert (fig. 143) far north of the limits decided by Barron 
and Hume (1902). Floyer (1887) and Tregenza (1955) reported 
hyraxes from Wadi Qattar, and we observed them there and in Wadi 
Umm Yassar in 1966. Cutting of acacia trees for making charcoal, a 
practice that continues over much of the Eastern Desert, has prob- 
ably reduced or eliminated hyraxes in many areas, e.g., parts of 
Wadi Sukari and Wadi Ghadir and probably wadis in the Maaza 




Fig 14H. Habitat of Procavia capensis in Wadi Nagib. Note white urine streaks 
on rocks. Tree is Acacia ehrenbergiana. 



466 



OSBORN&HELMY: MAMMALS OF EGYPT 467 

Plateau. Bedouins with guns are also a factor to be considered 
(Hoogstraal, 1964). 

Hoogstraal et al. (1957ab) observed many old dens marked with 
white urine streaks and suggested that this indicated either periodic 
movements or that populations were smaller than before. Their 
estimates of numbers of hyraxes in the Bir Kansisrob and Bir 
Abraq areas were 100 and 12 to 20, respectively. Osborn (1968a) 
reported seeing 11, including two young specimens in Wadi Nagib. 

Reproduction.— A colony from Sinai Peninsula stock in Giza 
Zoological Gardens produced 14 litters, averaging 2.5 young per lit- 
ter, five being the largest (Flower, 1932). Roche (1960) reported lit- 
ter size is usually two to three, maximum three in captive P. c. 
syriaca in France. Captive P. c. syriaca in Israel usually begin 
mating in August and bear young from mid-March into May. The 
usual number of young is three to four. Young are fully active soon 
after birth (Mendelssohn, 1965). 

Economic importance.— Russell (1949b) mentioned that hyrax 
guano is sometimes used for fertilizer. Bedouins kill them for food. 

Food— Over much of the Eastern Desert, acacia leaves emd pods 
are staple food of hyraxes. In Wadi Aqwamtra, Gebel Elba, they fed 
selectively on Rumex vesicarius. Reports indicate they can eat 
almost any plant, even those toxic to other animals such as 
Solanum sp.. Euphorbia sp., and Ficus sp. Halophytic plants are 
probably eaten, and listings also include grasses, herbs, berries, 
small fruits, bark, lichens, leaves of chmbable shrubs and trees 
(Mendelssohn, 1965; Sale, 1965; Dorst, 1970). Captive hyraxes eat 
almost anything except meat. They have been fed barley, bread, 
bread and milk, sorghum grain, cooked rice, clover, alfalfa, alfalfa 
hay, carrots, beets, eggplant, other vegetables, dates, grapes, other 
fruits, mealworms, insects, and small vertebrates (Bruce, 1790; 
Mendelssohn, 1965; Flower, 1932; Hauser, 1951). 

VViater.- Rock hyraxes drink very little water, but according to 
Dorst (1970), may travel distances of 640 m. to obtain it. Most col- 
onies in the Eastern Desert live in waterless areas. 

Fo/A/ore.— According to Sinai Bedouins of earlier times, the Con- 
ey is ". . .man's brother. The pecuhar conformation of its feet are 
proof that it is the descendent of a human being transformed; they 
will not eat its flesh and declare that if a man were to do so, he would 
never look upon his father and mother again" (Palmer, 1872, p. 89). 



468 FIELDI ANA: ZOOLOGY 

Apparently, modem Bedouins have forgotten this taboo, for they 
wantonly slaughter the animal. 

Key to Egyptian Subspecies of Pmcavia capensis 

1. Dorsum yeUowish to orangish brown. Hairs of dorsal spot concolor. Anterior end 
of malar not contacting lacrimal (fig. 145). (Sinai Peninsula) syriaca, p. 468. 

2. Dorsum grayish to yellowish brown. Hairs of dorsal spot with dark base and 
generally dark tip. Anterior end of malar usually contacting lacrimal (fig. 145). 
(Eastern Desert) ruficeps, p. 468. 

Procavia capensis syriaca (Schreber, 1784) 

Hyrax syriacus Schreber. 1784. Die Saugeth.. Vol. 4, pi. 240B (1792). p. 923. 

Type locality.— Lebanon: Mt. Lebanon. 

Distribution in Egypt— Figure 143. Sinai Peninsula. 

External characters.— Dorsum orangish brown, venter buff to 
pale brownish. Hairs of dorsal spot around gland concolor, at least 
inner margin, yellowish to orangish. 

Cranial characters. —See species description and Figures 144, 145. 
Malar not contacting lacrimal. 

Measurements.— Table 54. 

Comparisons.— Procavia c. syriaca differs from P. c. ruficeps in 
darker, more orangish color; dorsal spot more clearly meu-ked, and 
malar not contacting lacrimal. 

Specimens examined.— Total 17. 

SINAI: Wadi Abu Zeitouna (3). Gebel Musa (1). Wadi Taba (1). Wadi Ergein (4). 
Gebel Hammami (I), St. Catherine Monastery area (I), "Sinai" (6). 

Published records.— Records are from Hart (1891), Anderson 
(1902), Brauer (1917), Flower (1932), Hahn (1934), and Wassif and 
Hoogstraal (1953). 

SINAI: Wadi Ergein, Gebel Musa, St. Catherine Monastery area. Gebel Ham- 
mami, El Tor, Wadi Adani, Has Muhammed, and Dahab (reported by Bedouins). 

Procavia capensis ruficeps (Hemprich and Ehrenberg, 1832) 

Hyrax ruficeps Hemprich and Ehrenberg. 1832. Symbolae Physicae Mamm., 

Dec.l, folio h. pi. 2. upper fig. 
Hyrax burtonii Gray. 1868. Ann. Mag. Nat. Hist. (ser. 4). 1, p. 43. 

Type /oca/ity.- Sudan. NORTHERN: Dongola 

Distribution in Egypt— Figure 143. Eastern Desert. 

External characters.— Dorsum grayish to yellowish brown, venter 



OSBORN&HELMY: MAMMALS OF EGYPT 469 

buffy. Hairs of dorsal spot around gland usually with dark tips, pale 
yellowish subterminal bands, and brownish bases. 

Cranial characters. —See species description and Figures 144 and 
145. Malar usually contacting lacrimal. 

Comparisons.— Procavia c. ruficeps differs from P. c. syriaca in 
paler more yellowish and sometimes grayish pelage; dorsal spot 
hairs not clearly separable from dorsum; and malar usually contact- 
ing lacrimal. 

Remarks.— Procavia c. ruficeps, on basis of priority, is considered 
synonymous with P. c. burtoni described from an individual variant. 

Specimens examined.— Total 21. 

Egypt: No exact localities (3). 

RED SEA: Wadi Abu Kaleja (4). Wadi Habib (1). Bir Abraq (4), Gebel Abraq (1), 
Wadi Hodein (2). Wadi Atallah. 6.6 km. inside (old skull). 
ASWAN: Wadi Nagib (2). 
SUDAN ADMINISTRATIVE: Bir Kansisrob (3). 

Sight records {individuals, fresh tracks, fresh feces) of D. Oshorn 
and I. Helmy. — 

RED SEA: Wadi Umm Yassar. Wadi Qattar. Gebel Umm Disi. Wadi Semna, Wadi 
Ghadir. Wadi Sukari. 

ASWAN: Wadi Magal GabrU. 

SUDAN ADMINISTRATIVE: Wadi Aqwamtra. Wadi Kansisrob. 

Published records.— Records are from Floyer (1887), Anderson 
(1902), Barron and Hume (1902), Bonhote (1909, 1912), Bedan 
(1928), Negumi (1952), Tregenza (1955), Hoogstraal et al. (1957ab), 
and Osborn (1968a). 

RED SEA: Wadi Qattar. Wadi Umm Delfa, Wadi Habib. Wadi Abu Kaleja (Abu 
Khalifa). Wadi Fertili. Wadi Sheitun, Bir Inglizi (Beida), Ras Gurdi, Bir Abraq, 
Gebel Abraq, Wadi Hodein. 

ASWAN: Wadi Nagib. 

SUDAN ADMINISTRATIVE: Wadi Kansisrob. 



ORDER PERISSODACTYLA 
Family Equidae 

Genus Equus Linnaeus, 1758 

Genus of large-headed, slenderly built horse-like mammals, with 
or without stripes. Tail long, tufted. Short mane on neck. Bare 
callosity on inner side of foreleg. Skull with cranium short, but 
large; rostrum elongate; nasals long, narrow, projecting freely; 
orbits small, enclosed. Ethmoidal fissure absent. Canines in 
diastema, usually only in males. Cheek teeth extremely hypsodont, 
crowns squarish with complex foldings of enamel, dentine, and 
cement. Anterior pillar of upper cheek teeth united by narrow 
enamel bordered neck to dentine of main body of tooth. Dental 

3 q± lA 3, 
3» 0-l» 3 • §■ 



formula: i ^. ¥. 5 x 2=36-42 



Equus asinus Linnaeus, 1758 

Equus asinus Linnaeus, 1758, Syst. Nat., ed. 10, p. 73. 
Type locality.—''. . .in Oriente." 

General distribution. —Southeastern Egypt, eastern Sudan, 
Eritrea, Somalia, northern Chad. 

Common names.— Wild Ass, Hamar el Wadi, Homard BerL 

Subspecies in Egypt — 

Equus asinus africanus (Fitzinger, 1857) 

Asinus africanus Fitzinger, 1857. Naturgesch. Saugeth., Vol. 3, p. 667. 
Asinus taeniopus Heuglin, 1861, Nova Acta Acad. Caes. Leop.-Carol., Jena, 
Vol. 28. No. 10. pi. 1. p. 1. 

Type locality.— Nubia. 

Distribution in Egypt— Figure 147. Southeastern Egypt. 

Diagnosis.— Dorsum and side pale gray with reddish sheen; legs 
paler than body; head, neck, and legs without stripes; shoulder cross 
present, variable. Eye ring, inside of ear, muzzle, underside of lower 



470 



OSBORN & HELMY: MAMMALS OF EGYPT 



471 



2 5* 2 6* 2 7* 28* 2 9* 30* 31* 3 2* 3 3* 3 4* 35* 36* 37* 




Fig 147. Localities of observations of Equus asinus africanus in 1800's (circles) 
and from 1950 to date (dots); paleolithic and prehistoric remains (X). 

jaw, and venter white. Short mane on neck, forelock absent. Ear 
very large. Tail long, tufted. 

Shoulder height 115 to 125 cm. 

Comparisons.— Equus a. africanus differs from £^. a. somaliensis 
in darker color, lack of leg stripes, presence of white eye ring in all 
individuals, more strongly developed dorsal stripe and shoulder 
cross, and a dark spot on outer side of the fetlock. From the 
domestic ass, E. a. africanus differs in paler color, lack of leg stripes, 
legs paler than body, lack of dark patch at ear base and tip, and 
presence of dark spot on outer side of fetlock. 

//a6itat.— According to Ziccardi (1970), wild asses prefer arid, 
remote wadis, and plains with xerophytic vegetation. 



472 FIELDIANA: ZOOLOGY 

Remarks.— Equus cl taeniopus is considered obsolete on bases of 
the description having been of a Uving animal with no definite locali- 
ty (Sclater, 1884; Harper, 1940), and the drawing to be of either a 
domestic donkey (Neumann, 1935) or an inaccurate representation 
of a wild one (Antonius, 1937). Lydekker (1916, p. 38) favored reten- 
tion of the subspecies and listed "typical locality Ha wash district of 
Abyssinia." Hoogstraal et al. (1957b) listed it tentatively from 
southeastern Egypt. Ziccardi (1970) called it the Eritrean wild ass. 

Historical notes.— hate Paleolithic shoreline deposits in El 
Faiyum contain Equus fragments, and wild ass occurred in the 
prehistoric Kom Ombo Plain fauna (Butzer and Hansen, 1968). 
Prehistoric rock drawings of wild ass are in the Wadi Muktil (Mu- 
qtil)-Wadi Natash area (Floyer, 1893) and further north in Wadi el 
Hammamat. 

Domestication in the Nile Valley was estimated to have been be- 
tween 3,500 and 3,300 B.C. A predynastic period slate depicts asses 
being traded with Libya. Ramses III hunted donkeys, and they 
were well known as wild animals in Egypt until about 1,100 B.C. 
(Talbot, 1960; Zeuner, 1963). 

Originally, Equus asinus probably ranged from Somalia through 
the Libyan Desert to Morocco (Zeuner, 1963). The Nubian wild ass 
(E. a. africanus) is thought to have inhabited most of the Eastern 
Desert and parts of the Western Desert along the Sudan border 
(Talbot, 1960). Harding- King (1925) referred to Uweinat as a place 
where wild asses fed when there was pasture. 

References to wild asses in literature on Egypt are scarce. Ander- 
son (1898, 1902) wrote that wild asses had been seen near Wadi Kit- 
tar (Qattar) and Ayd near Old Keneh (Qena) in the 1820's by James 
Burton (MSS in British Museum). Flower (1932) maintained that 
Anderson's quotations from Burton's MSS were in error, because 
they referred to Nubia. Burton's localities are validated, however, 
by the fact that he traveled with Wilkinson (1832) in 1823 in the 
Eastern Desert northeast of Qena. 

Burton wrote that the Bedouins let their female donkeys loose to 
be bred by wild males and that white offspring, called ''Homar 
Wahsh," always resulted from these matings. The animal seen near 
Old Qena was described as white with a dark dorsal line. 

Anderson (1898, p. XXV) also wrote of Linant de Bellefonds' ex- 
plorations in the mid 1800's in southeastern Egypt: "Having 



OSBORN&HELMY: MAMMALS OF EGYPT 473 

cleared the defiles of Wadi Daffeti; Linant de Bellefonds visited the 
mountain and valley of Beint el Fegue. Hereabouts many wild asses 
occurred, extremely shy, and scenting man from a great distance. 
They were trapped by the Bisharin Arabs who used their flesh as 
food." 

A pair of semi-wild donkeys with one young were seen in the Qat- 
tar area by Floyer (1887) who met with the owner 32 km. further 
north. These donkeys, wrote Floyer (1887, p. 671), "leapt from rock 
to rock with the agility of goats." Later he (Floyer, 1893) remarked 
that wild ass were still found in the area south of a line from 
Berenice to the Nile. 

According to Flower (1932, p. 432), "there appear to be no certain 
records of genuine wild ass having occurred in Egypt during the 
nineteenth century." Conflicting reports continued to be published 
in the 20th century. Barron and Hume (1902) said wild asses were 
fairly common south of the Qena-Quseir road in the Eastern Desert, 
but according to Ball (1912), wild asses had disappeared from 
southeastern Egypt. Bedouins often pointed out "wild donkeys" to 
Murray (1935), which he assumed were feral. One sight record which 
he considered authentic was near Gebel Shindeib (Shendib). 

The last wild ass of Nubian origin was supposedly shot in 1925 
near Gebel Rababa on the Sudan-Eritrea border (Antonius, 1937) 
and "considered to be extinct in its former range" (Setzer, 1956, p. 
569). However, Zeuner (1963) published a photo of a Nubian wild ass 
captured near Abu Hamad (19° 32' N lat., 33° 19' E long.) in 
northern Sudan. Hoogstraal (1964) received authentic reports of 
wild asses in the Red Sea area of Kassala Province of Sudan. 
Negumi (1952) concluded that E. asinus was spottily distributed in 
the Eastern Desert. 

After years of uncertainty as to the status of this animal in 
Egypt, several herds of 2 to 10 wild asses were seen in the spring of 
1954 (Hoogstraal et al., 1957ab; Hoogstraal, 1964) on the coastal 
plains north of Gebel Elba. Talbot (1960) also reported scattered 
herds in southeastern Egypt and northeastern Sudan. In October 
1964, we saw two donkeys in Wadi Sukari which, our guide told us 
later, were wild asses. In spring of 1967, we inquired about wild 
asses in the Gebel Elba area and were told by our Bishari guide that 
they were south of us in Sudan. Apparently, the herds move about 
over an area of considerable size. In Talbot's (1960, p. 272) opinion. 



474 FIELDIANA: ZOOLOGY 

the present herds may be partly feral "as they are considered prop- 
erty of the local Bedouin." 

In 1974, one wild ass was observed by I. Helmy in Wadi Allaqi. 

Remains of E. a, africanus have recently been reported from Late 
Paleolithic sites near Kom Ombo (Reed and TurnbuU, 1969; 
Churcher, 1972). 



ORDER ARTIODACTYLA 
Family 1. Suidae 

Genus Sus Linnaeus, 1758 
Description under species. 

Sus scrofa Linnaeus, 1758 

Sus scrofa Linnaeus, 1758, Syst. Nat., ed. 10, p. 49. 

Type locality.— Germany . 

General distribution.— Continental Europe, southwestern 
U.S.S.R., Turkey; Transcaucasia eastward and northward into 
Siberia, China, Japan, Vietnam and Malay; parts of Syria, northern 
Arabian Peninsula, Israel, Jordan, Libya, Tunisia, Algeria, Moroc- 
co, Spanish Sahara, and Sudan. Extinct in Egypt. 

Common names.— Wild Boar, Hanzir. 

Previous distribution in Egypt— Figure 148. Nile Delta, Wadi el 
Natroun, El Faiyum, El Maghra (possibly), and probably most of 
the Nile Valley in prehistoric and historic times. 

Diagnosis.— Large, sturdily built with hump over shoulders. 
Pelage bristly and shaggy. Color dull brownish black to yellowish 
gray. Young striped brown and yellow. Ear moderately large, erect, 
pointed, and densely pilose. Dorsal crest present. Tail relatively 
short. Lateral digits sfnall, but functional. Muzzle elongate, ending 
in a flat, disk-shaped, naked pad. Skull narrow, elongate; braincase 
small; supraoccipital high; paroccipital extremely long. Incisors 
proodont. Canines tusk-like and continuously growing. Upper 
canine turning outward, forward, and upward; wearing against 
lower and producing sharp, flat edges. Molariform teeth brachydont 
and bunodont, increasing in size and complexity posteriorly. Pm, 
small, transient, and in diastema between canine and pm2. Dental 
formula:!, 1, 1,1, X 2=44. 

Historical notes.— The last Egyptian wild boar is supposedly 



475 



476 



FIELDIANA: ZOOLOGY 



,. 2 5* 2 6* 2 7* 26* 2 9* 30* 31* 32* 33* 34* 35* 36* 37 




Fig. 148. Previous distribution of Sus scrofa. 

represented by British Museum specimen No. 2450 which died 
December 20, 1912, in Giza Zoological Gardens, where it had Uved 
for more than 14, possibly 18, years (Flower, 1931, p. 224). Wild 
boar became extinct in Egypt about 1900. Strekalovsky (1949) said 
the last wild boar was shot in Wadi el Natroun in 1894. Russell 
(1951, p. 19) commented, "when I came to Egypt in 1902 the last 
wild boar had just been killed in the reed beds and swamps of Wadi 
Natrun and Moghra oasis." It was common previously in swamps of 
the Nile Delta, Giza, El Faiyum, and Wadi el Natroun (Anderson, 
1902; De Winton, 1903; Flower, 1932). According to the last. Prince 
Kemal el Din restocked Wadi el Natroun in 1907 with wild boar 
from Hungary, but they were soon shot by poachers. Negumi (1952) 
wrote that, 50 to 70 years before, wild boar attacked Bedouin sheep 
near the Giza swamps. Persistent hunting and burning of habitat 
exterminated the species. 



OSBORN&HELMY: MAMMALS OF EGYPT 477 

Butzer and Hansen (1968) listed Sus remains from Late 
Paleolithic deposits in El Faiyum. 

Ancient Egyptians reportedly domesticated pigs from Neolithic 
at least into Dynastic times and later in the 18th Dynasty (Zeuner, 
1963). During Roman times, boar hunts in El Faiyum were fairly 
common (Lindsay, 1965). 

Family 2. Hippopotamidae 

Genus Hippopotamus Linnaeus, 1758 
Description under species. 

Hippopotamus amphibius Linnaeus, 1758 

Hippopotamus amphibius Linnaeus, 1758, Syst. Nat., ed. 10, p. 74. 

Type locality.— Egypt: Nile River. 

General distribution.— River systems of Africa. 

Common names.— Hippopotamus, Barnik. 

Previous distribution in Egypt-Figure 149. Nile River, except- 
ing Rosetta branch. 

Diagnosis.— Very large, heavily built, and amphibious. Skin 
almost hairless, except for bristles on muzzle and tail. Mouth enor- 
mous; snout broad; nostrils dorsal, widely separated, and pro- 
truding. Ears relatively small and erect. Tail short. Legs short and 
heavy. Feet short, broad, four-toed, with rounded hoofs. Skull 
massive and extremely broad across the canines. 

Incisors and canines tusk-like and continuously growing. Cheek 
teeth bunodont with trefoil-shaped dentine islands. Dental formula: 
rl.U. 1x2=38 or 42. 

Historical notes.— Sporadic occurrences of hippopotamus along 
the Nile were documented by Flower (1932). He found no indication 
of it having lived in the Rosetta branch, but gathered evidence from 
early writers and dredged bones that this beast had lived along the 
Damietta branch, the papyrus swamps east of there, and the ancient 
Pelusian branch of the Nile. Kock (1970a) has shown that H. am- 
phibius occurred originally in two disjunct areas (fig. 149). 

The last two hippos were reportedly killed near Damietta in 1600 
(quoted from Buffon's Natural History, 1764, by Anderson, 1902). 
Sonnini (1807) wrote that the last one killed in Egypt was in 1685. 
However, Burckhardt (1819, p. 67) reported one killed in 1816 near 



478 



FIELDIANA: ZOOLOGY 



,. 2 5* 2 6* 2 7' 28' 2 9* 30* 31* 32* 3 3* 34* 35* 36* 37* 




Fk; 149. Previous distribution of Hippopotamus amphibius. 



Wadi Haifa and a second traveling to Derau (Daraw) 36 km. N of 
Aswan. Flower (1932) recorded the latter as 1816 or 1818. Butzer 
(1959) gives 1658 as the date of the last hippopotamus killed in the 
Delta and 1850 for Upper Egypt. Additional records listed by Kock 
(1970a) for Egypt are: Damietta (1600, 1815). near Cairo (1580), Abu 
Girgeh (1658), and Wadi Haifa (1812). 

The ancient Egyptians called the hippopotamus "Nile Horse." It 
damaged their fishing nets and ate their crops and was therefore 
considered an incarnation of an evil force and was hunted and har- 
pooned. Harpooning of hippos was also a sport of the Ptolemaics 
(Lindsay, 1965). 

Hippo remains from Late Pleistocene middens were found at Kom 
Ombo (Reed and Turnbull, 1969). 



OSBORN&HELMY: MAMMALS OF EGYPT 479 

Fo/ife /ore.— Around the hippo, as with many other large mammals, 
there has developed a repertory of legends and medicinal properties. 

Burned hippopotamus skin mixed with flower of pulse and ap- 
plied to cancer is supposed to cure it in three days. The gall bladder 
soaked for 30 days in water then pounded and mixed with honey 
that has not been over fire is said to cure a black eye in two or three 
days. Rubbing with a hippopotamus tooth is a "cure" for stomach 
ache. Burning pieces of skin in the middle of a town is a 
"protection" from calamities. Application of burned skin is sup- 
posed to remove swelling and inflammation. A hippopotamus in a 
dream "indicates" a lie and an affair that will not be completed 
(Jayakar, 1908). 

Family 3. Bovidae 

Small to large ungulates with horns present on both sexes. Legs 
long relative to body. Feet unguligrade, toes four with hoofs, and 
lateral digits usually vestigial. Anterior portion of skull elongate, 
braincase relatively large, postorbital bar present, paroccipital pro- 
cess prominent, but not extending much below level of tympanic 
bulla. 

Upper incisors and canines absent. Cheek teeth usually h5T)so- 
dont. Posterior premolars like molars. Dental formula: % j, |, 
ix2=32. 

Key to Egyptian Genera of Bovidae 

1. Horns arising from a frontal pedicel or buttress. Facial and other markings ab- 
sent. Muffle naked. Skull with large lacrymal or infraorbital fossa 

Alcelaphus, p. 484. 

2. Horns not arising from a pedicel. Muffle haired. 

a. Facisd markings present. 

i. Size small, height at shoulder less than 1 m. Tail short, reaching less than 
one-half way to heel. Horns short, arising above orbits. Skull with large in- 
fraorbital or lacrymal fossa Gazella, p. 486. 

ii. Size large, height at shoulder more than 1 m. Tail long, reaching heel. Horns 
long, arising behind orbits. Infraorbital fossa lacking. 

(a) Horns straight or curved Oryx, p. 480. 

(b) Horns spiralled Addax, p. 482. 

b. Facial markings absent. 

i. Horns scimitar-shaped, curving dorsally and posteriorly, anterior surface 
broadest with numerous large transverse knobs. Beard present. Black and 
white leg markings Capra, p. 514. 

ii. Horns not as above, curving dorsally, laterally, and inwardly; transversely 



480 FIELDIANA: ZOOLOGY 

ridged from base to tip. Beard absent. Long hair from sides of jaws to upper 
forelegs. No leg markings Ammotragus. p. 52L 

Genus Oryx Blainville. 1816 
Description under species. 

Oryx dammah (Cretzschmar, 1826) 

Antilope dammah Cretzschmar. 1826, in Riippell, Atlas zu der Reise im nord- 
lichen Afrika von Rupi>ell, pt.l, Saugeth., p. 22. 

Type locality. -Sudan. KORDOFAN: Gebel Haraza. 

General distribution. — Desert regions of North Africa from north 
of Khartoum west to southern Tunisia and as far south as Sene- 
gambia. 

Common names.— Scimitar Horned Oryx, White Oryx, Meha, 
Abu Herab. 

Previous distribution in Egypt— Figure 150. Western Desert 
until about middle of 19th century (Flower, 1932; Kock, 1970b). 

Diagnosis.— Large, pale, long-horned antelope. Pelage yellowish 
or reddish white; neck, shoulders, dorsal stripe, upper base of tail, 
and tail tuft brownish; head and muzzle whitish with large grayish 
or brownish patches on nose and forehead which may or may not 
connect with stripe through eye from base of ear and horn; ear 
whitish; legs whitish, forepart brownish; flank stripe faint. Tail tuft 
long and nearly reaching dew claws. White rump patch absent. 
Knee tufts absent. 

Skull lacking infraorbital fossa. Ethmoidal fissure small. Pre- 
maxilla not contacting nasal. 

Horns very long, cylindrical, straight or scimitar-like; basal half 
ridged; postorbital in origin with base on plane of frontals. 

Molars large; internal cusps between inner lobes of upper molars 
and external cusps between outer lobes of lower molars. 

Historical notes. — Until the first half of the 19th century, oryx 
occurred over much of the Western Desert, according to records 
accumulated by Kock (1970b). LocaHties where oryx were observed 
are as follows: Western Desert (1859); Siwa Oasis, Kharga Oasis, 
and south of El Faiyum (1869); El Faiyum area (1800, 1835); 
western Giza Governorate and Wadi el Natroun (until 1800). Prob- 
ably, Schweinfurth's (1874) "I'antilope Dama" of Kharga and 
Dakhla Oases was, correctly, Antilope dammah Cretzschmar, 1826. 



OSBORN & HELMY: MAMMALS OF EGYPT 



481 



,. 25* 26* 27* 28* 2 9* 30* 31* 3 2* 3 3* 3 4* 35* 36* 37* 




Fig 150. Previous distribution of Oryx damah and recent sight record (S). 



The only recent record from Egypt is a single individual seen near 
the Siwa road, 130 km. S of Mersa Matruh by I. Helmy in 1975. 

Prehistoric rock paintings of oryx exist at Gebel Uweinat 
(Almasy, 1936), indicating the former range included northwestern 
Sudan, southwestern Egypt, and adjacent parts of Libya (Osborn 
and Krombein, 1969). 

Oryx leucoryx was reported to have existed in northern Sinai and 
lower Israel until about 1800 (Talbot, 1960). 

Ancient Egyptians captured oryxes, kept them in semi- 
domestication, and sacrificed them for their gods. In Butzer (1959), 
the oryx is one of the commonest of animals depicted in temples and 
tombs of dynastic Egypt. 

During the reign of Ptolemy II (283 to 246 B.C.), oryx and 



482 FIELDIANA: ZOOLOGY 

hartebeests were among the animals displayed in Alexandria 
(Jennison, 1937). 

Genus Addax Rafinesque, 1815 
Description under species. 

Addax nasomaculatus (Blainville, 1816) 

Cerophorus {Gazetla) or Antilope nasomaculata Blainville, 1816. Bull. Sci. Soc. 
Philo.. Paris, p. 75. 

Type /oca/jfy. — Probably Senegambia, West Africa (Ellerman and 
Morrison-Scott, 1951). 

General distribution.— Desert areas from Dongola west of Nile 
River to Senegal. 

Common names.— Addax, Meha, Akash, Begra el Ouash. 

Previous distribution in Egypt. — Figure 151. Probably ranged 
over most of the Western Desert. 

Diagnosis.— Large pale antelope. Horns long (65 cm. or more), 
slender, and twisted. Hoofs exceedingly broad and shallow. 
Shoulder height 97 to 108 cm. Color brownish gray in winter, pale 
reddish in summer. Head darker than body with white X-like facial 
patch. Muffle haired. Patches of long hair below eyes and on side of 
neck. Forehead tuft, dorsal crest, and tail tuft short, blackish. Post- 
auricular area, hindquarters, tail (except tuft), venter, and legs 
white. Tail tip reaching below heel. Facial and inguinal glands lack- 
ing. Interdigital glands present. Skull without supraorbital pit or 
infraorbital fossa. Ethmoidal fissure small. Horns postorbital, long, 
twisted, and with prominent annulations. 

Upper molars squarish and with accessory columns on either side. 

Historical notes. — Information on previous distribution of the 
addax in Egypt is very limited. It probably inhabited most of the 
Western Desert periodically, depending upon availability of 
grazing. 

According to HeugUn (quoted by Sclater and Thomas, 1894-1900), 
in the mid-1800's, the addax ranged northwards into the Libyan 
Desert of Egypt to the oases and El Faiyum. "EarHer than the two 
wars, but within the memory of living man, occasional herds of 
Addax followed the grazing into Egypt from further west" (Russell, 
1951, p. 19). It is now one of the rarest mammals in Libya (Setzer, 
1957c), although it had been recorded from hinterlands of Bengazi 



OSBORN & HELMY: MAMMALS OF EGYPT 



483 




Fig. 151. Previous distribution of Addax nasomaculatus. 

(de Beaux, 1932), Giarabub, and Cufra (Toschi, 1954) in Cyrenaica. 
Other localities of observation listed by Kock (1970b) are: Libyan 
Desert, no exact locality (1855, 1877); Western Mediterranean 
Coastal Desert, no exact locality (1877); north of El Faiyum (1869); 
oases of Middle and Upper Egypt (1869); Surarieh south of Beni 
Suef (1863); and Herwer (Antinoe) (1863). A skeleton, date of death 
unknown, was found in 1938 in Wadi Aqaba in the southern Gilf el 
Kebir (Kock, 1970b). An old weathered horn was found at Bir 
Terfawi in 1972 by Ali Abu Resha, official desert guide, Kharga 
Oasis (personal communication). 

In 1927, Murray (1967, p. 167) was told by a Bedouin that, 30 
years before, "four wild cows" (addax antelope) had been seen near 
Minqar Abu Dweiss in Qattara Depression. Flower (1932) quoted a 
personal communication from T. W. Russell of the Egyptian police 



484 FIELDIANA: ZOOLOGY 

that the last known specimen of addax was shot by a Bedouin in 
1900 in the Mariut District about 65 km. W of Alexandria. In a foot- 
note, Murray (1967, p. 167) wrote: "Abu Fidel killed the last addax 
in Egypt by running it down with his car near Sheb while on 
Clayton's 1931 expedition." Negumi (1952) listed two locaUties: 
near Bir Dibbis and south of Uweinat. Of the two areas, Uweinat is 
presently the most likely to be visited by addax grazing northward 
(Osborn and Krombein, 1969). 

This species, like other Egyptian antelopes, was hunted and kept 
captive by ancient Egyptians. It is depicted in many tombs and 
temples (Butzer, 1959). 

Genus Alcelaphus Blaineville, 1816 
Description under species. 

Alcelaphus buselaphus (Pallas, 1766) 

Antilope buselaphus Pallas, 1766, Misc. 2kx)l., p. 7. 

Type locality.— Probably Morocco (Lydekker, 1916), 

General distribution.—Soma]ia, Ethiopia, Uganda, Sudan, Chad, 
Central African Republic, and Cameroons. Previous distribution 
included Egypt, Libya, Tunisia, Algeria, and Morocco. 

Common names.— Bubal Hartebeest, Begra el Ouash. 

Previous distribution in Egypt— Figure 152. Oases and oasis-like 
depressions of Western Desert; Western Mediterranean Coastal 
Desert. 

Diagnosis.— Large reddish, ungainly antelope with abnormally 
long face. Horns doubly curved. Shoulder height about 110 cm., 
markedly higher than rump. Color uniformly pale reddish or 
yellowish with no contrasting markings on head and limbs and 
rump patch paler than body. Faint blackish areas on forehead, 
muzzle, shoulder, and thigh. Whitish patch on hip. Muffle bare. Tail 
reaching heel; tuft thin and brownish. Lacrymal glands prominent. 
Skull with large infraorbital fossa; ethmoidal fissure and supra- 
orbital pit lacking. Rostrum narrow and elongate. Frontal region 
abnormally elongate. Premaxillary not contacting nasal. Fronto- 
parietal suture on lower side of skull. 

Horns U-shaped in frontal view, diverging posteriorly in line with 
face, then rising almost vertically, and bending again sharply 
posteriorly in the middle. Ridges prominent to second curvature, 



OSBORN&HELMY: MAMMALS OF EGYPT 485 

25* 26* 27* 28* 2 9* 30* 31* 32* 33* 34* 35* 36* 37* 




Fig. 152. Previous distribution of Alcelaphus buselaphus. 

tips smooth. Cores angulated; united at base on a single elevated 
frontal pedicel which projects posteriorly in line with face and 
beyond level of the occiput. 

Molars hypsodont, upper molars narrow, and inner accessory 
columns absent. 

Historical notes.— The hartebeest is probably the animal of the 
Western Mediterranean Coastal Desert referred to by Sonnini (1807) 
who saw herds of 8 to 10, said the Arabs hunted them, and had seen 
them domesticated by Bedouins of Abu Qir and Alexandria, who 
had captured them when young. Cailliaud (1826, p. 134) described a 
ruminant, "baquar ou boeuf sauvage," which was as large as a calf 
and seen in El Bahrein. Anderson (1902) wrote that Dr. R. C. 
Mitchell had been told by Bedouins of El Faiyum that hartebeest 
were to be found two days journey by camel due west of Lake Kurun 



486 FIELDIANA: ZOOLOGY 

(Qarun), a distance of roughly 80 to 100 km. On the basis of descrip- 
tions by Siwa residents of an animal "the .size of a donkey, of a 
yellowish brown colour, with two horns like a cow's," Belgrave 
(1923, p. 202) searched the uninhabited oasis of Gagub (Qeiqab) near 
Giarabub without finding any trace of the hartebeest. In 1965, a 
Bedouin in Salum mentioned having seen a "wild cow" about 30 
years before in Siwa Oasis. Jennison (1937) assumed that hartebeest 
must have once been common all along the North African coast 
because of the traffic in these animals in Roman times. They were 
among the animals displayed in Alexandria during the reign of 
Ptolemy II (283 to 246 B.C.). 

Indefinite as these reports appear, plus the fact that the name 
Begra el Ouash (also spelled Begra el Ouach, Bakkar Wahash, and 
Bekker el Wash or Wahash), meaning wild cow or ox, has been used 
indiscriminately by Arabs for both Addax and Alcelaphus, some 
were certainly of the latter. Within historical time, the hartebeest 
no doubt existed in vegetated areas of the Western Desert. 

There are no specimens from Egypt except mummified remains 
from El Abadiyah, El Faiyum, and Sakkara (Anderson, 1902; Loret 
and Gaillard, 1908; Blaine, 1914), and Pleistocene fossil fragments 
from Kom Ombo and Wadi Haifa, Sudan (Reed and Turnbull. 1969). 
Tombs and temples of ancient Egypt were adorned with figures of 
hartebeests (Butzer, 1959). 

Genus Gazella Blainville, 1816 

Small, slender antelopes with horn bases curving dorsally and 
posteriorly and tips recurving. Color buff to brown with contrasting 
facial, side, hip, and gluteal markings and a white rump patch. Tail 
short, blackish, and tufted. Muffle completely haired. Carpal tufts 
present. Inguinal, interdigital, and lacrimal (facial) glands present. 

Braincase expanded, bulla enlarged; infraorbital (lacrimal) fossa 
and ethmoidal fissure large and prominent. Premaxilla contacting 
nasal. Horn core supraorbital and extending dorsally from base. 

Ki-;y to Egyptian Speciks of Gazella 

1. Color buffy, facial markings indistinct, lacking blackish hairs. Male horns slight- 
ly recurved, spreading apically (fig. 157). Skull with fenestra in infraorbital fossa, 
posterior nasal margin and interparietal suture round (fig. 156). (Western Desert) 
leptoceros, p. 487. 

2. Color brownish, facial markings distinct, with blackish hairs. Male horns slightly 
to strongly recurved, curving forward and inward apically (fig. 157). SkuU lacking 
fenestra in infraorbital fossa, interparietal suture angular (fig. 156). 



OSBORN&HELMY: MAMMALS OF EGYPT 487 

a. Horns of adult male long and straight to semi-lyrate. Horns of adult female 
long and slender. Posterior nasal margin triangular. Parietal ridges present. 
(Sinai Peninsula, Eastern and Western Deserts) dorcas, p. 501. 

b. Horns of adult male short and semi-lyrate. Horns of adult female short and 
slender. Posterior nasal margin round. Parietal ridges absent. (Sinai Peninsula) 
gazeUa, p. 513. 

Gazella leptoceros (F. Cuvier, 1842) 

Antilope leptoceros F. Cuvier, 1842, in E. Geoffrey St. Hilaire and F. Cuvier. Hist. 
Nat. Manmi., Vol. 4, pt. 72, pis. 424, 425. 

Type locality.— Egypt. MATRUH: "Probably desert between 
Giza and Wadi Natroun, Lower Egypt, as the type specimen was 
brought to Paris by James Burton circa 1833 though in 1842 Cuvier 
wrote 'Senaar'" (Flower, 1932, p. 438). 

General distribution.— Western Desert of Egypt, Libya, Tunisia, 
Algeria, and probably northwestern Sudan. 

Common names.— Slender-Horned Gazelle, White Gazelle, Reem, 
Ghazal Abyad. 

Subspecies in Egypt— 

Gazella leptoceros leptoceros (F. Cuvier, 1842) 

Type locality.— Given above under species. 

Distribution in Egypt.— Figure 153. Northern part of Western 
Desert south of Mediterranean Coastal Desert vegetation. Possibly 
in the vicinity of Gebel Uweinat. 

Diagnosis.— Fale yellowish to buff with indistinct facial and body 
markings. Horns in male not lyrate in frontal view; slightly re- 
curved and spreading at tips. Skull with elongate fenestra in infra- 
orbital fossa and posterior margin of nasals round. Interparietal 
suture outline semicircular. 

Height at shoulders in adult male about 63 cm., head and body 
length average 94 cm., tail length 15 cm., hind foot 30 cm., ear 14 
cm., condylobasal length 17.4 cm., weight approximately 15 kg. 

External characters.— Pale, yellowish to buffy brown dorsally 
with pale and dark side stripes, triangular hip patch, and pygal 
border indistinct. Color of side not extending down outside of legs. 
Circumoral region, throat, chest, belly, rump, and legs white. Facial 
markings indistinct, paler than body markings and lacking 
brownish or reddish hairs. Ear pale buff on outer surface, cream or 



488 



FIELDIANA: ZOOLOGY 



,. 25* 26* 27* 28* 2 9* 30* 31* 32* 3 3* 34* 35* 36* 37 




Fk; 153. Collection localities of Gazella leptoceros leptoceros; sight records (S): 
skulls and horns, date of death unknown (X). 

whitish on inner surface. Tail, knee tufts and hair between dew 
claws and hoofs, brownish to blackish. 

Hoofs. — Hoofs of specimens from sandy areas are reported to be 
much longer and more diverging than of those from hard and stony 
desert (Thomas, 1894b; Sclater and Thomas, 1897-1900) and con- 
sidered to be an adaptation for easier movement on sand (Loder, 
1894; Schomber and Kock, 1960). Thomas (1894a), Bramley (1895), 
and Hoogstraal (1963) listed longer hoofs as a taxonomic character 
separating G. leptoceros from G. dorcas. 

In both species of Egyptian gazelles, those living on sand tend to 
have longer hoofs, probably because of lack of wear as in the case of 
hoofed mammals in zoos on soft substrates. Hoof measurements of 
specimens from hard, stony desert together with a few from sandy 



OSBORN&HELMY: MAMMALS OF EGYPT 489 

Table 55. — Hind hoof length (and ranges) in subadult and adult gazelles from 
hard and sandy desert. 





G. dorcas 


G. leptoceros 


Hard desert: 
Males 
Females 


40.6 (36-46) 10 
43.0 (34-48) 9 


50.5 (45-55) 4 
42, 44 


Sandy desert: 
Males 


52,67 


60 



desert in Table 55 indicate that hoofs are longer in leptoceros than 
in dorcas, but that abnormalities occur in both species when living 
on sand. Measurements of hind hoofs are given because they are less 
variable individually than measurements of fore hoofs. Thomas 
(1894b) gave the hind hoof length of 56 mm. for G. /. loderi from 
sand dunes in Algeria. 

Cranial characters.— Y'lgyires 154, 155. Skull rather elongate, nar- 
rowing posteriorly, without parietal ridges, supraoccipital crest pro- 
nounced and extending posteriorly beyond level of occipital condyle. 
Posterior margin of nasals tapering abruptly (rounded) from a point 
posterior to the ethmoidal fissure and the beginning of the nasofron- 
tal suture. Interparietal suture semicircular (fig. 156). Infraorbital 
fossa with elongate fenestra perforating the nasolacrimal canal. 
One-half of the infraorbital fossa is formed by encroachment of the 
jugal onto the lacrimal bone. Jugomaxillary suture is angular in 
outline (fig. 155). Length of premaxillary contact with the nasal, a 
character stressed by Gentry (1964) in comparing African gazelles, 
is quite variable, but usually greater than length of the maxillary 
contact (fig. 155). Auditory bulla with broken ridge on ventral sur- 
face. Basioccipital broad, sides parallel, and tuberosities slightly 
developed. Additional characters are listed in Table 56. 

Horns and horn cores.— Male horns are long, slightly recurved, 
and diverging gradually from the base (fig. 157). The tips curve 
slightly anteriorly and outwardly. Annulations are conspicuous, 
close together, and complete, except for distal three or four below 
the smooth, curving tips. Smooth tips are 25 to 50 per cent of total 
horn length, depending on age (fig. 157). Female horns are 
straighter and slenderer, annulations shallower and less con- 
spicuous than in males. 

Horn cores are conspicuously pitted and grooved on all surfaces, 
with intermittent grooves running from base to tip (fig. 154). Like 



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FIELDIANA: ZOOLOGY 




Fig 154. Skull of Gazella leptoceros leptoceros. 

the horns, the cores are relatively straight and not diverging. The 
base of the male horn core is ovoid in cross section and broader 
anteriorly than posteriorly. The female core is much smaller in 
diameter and round in cross section (fig. 158). 

Teeth.— Figure 156. Lingual ridges are prominent in lower molars 
(nxj, mg), particularly in subadults. Large posterior labial folds are 
also present on these teeth. 

Measurements.— Table 57. Lack of female specimens limits 
knowledge of sexual difference in dimensions in this species. 

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manent teeth have emerged. Subadults have m^ emerging. 



OSBORN & HELMY: MAMMALS OF EGYPT 



493 



Table 57. — Means (and ranges) of measurements and weight of adult male (M) 
and female (F) Gcuella dorcas and G. leptoceros. 







G. d. dorcas 


G. d. Uttoralis 


G. I. leptoceros 


HBL 


M 


902.0 (871-958) 8 


893.2 (860-930) 6 


937.0 (885-999) 4 




F 


869.3 (830-924) 6 


938.1 (909-991) 7 


955 


TL 


M 


151.8(120-180)8 


142.6(135-157)6 


162.0 (155-166) 4 




F 


137.8(120-179)6 


129.0 (102-145) 7 


125 


FL 


M 


309.0 (289-320) 8 


320.0 (309-330) 6 


320.0 (307-335) 4 




F 


291.2 (261-308) 6 


308.6 (247-332) 7 


300 


EL 


M 


124.8 (120-130) 8 


135.3 (129-149) 6 


140.0 (132-145) 4 




F 


125.0 (120-129) 6 


138.1 (130-148) 7 


130 


Wt (kg.) 




15.4 (14.0-18.0) 5 


15.0 


CBL 


M 


165.3(160.5-170.7) 11 


171.0(167.4-176.0)5 


175.1 (171.8-180.4) 3 




F 


156.3(151.0-161.5)5 


165.0(156.6-171.1)8 


169.3 


TRL 


M 


51.4 (46.5-53.7) 9 


54.2 (51.2-58.3) 5 


61.8 (57.5-69.7) 4 




F 


52.1 (50.1-54.2) 4 


52.4 (50.9-54.3) 8 


54.8 


OW 


M 


78.2(75.0-81.9)11 


78.4 (77.0-81.0) 5 


82.8 (80.4-86.6) 5 




F 


73.7 (70.8-78.2) 6 


74.4 (71.9-77.8) 8 


80.0 


POW 


M 


51.7(48.4-54.4) 11 


51.2 (49.6-52.9) 5 


52.7 (51.5-54.5) 5 




F 


50.3 (49.0-52.8) 5 


50.6 (48.9-52.6) 8 


52.4 


BCW 


M 


54.6(51.7-57.0) 11 


56.3 (55.5-57.1) 4 


55.3 (53.2-58.1) 5 




F 


54.0 (51.0-55.5) 5 


54.9 (53.6-55.3) 8 


52.6 


NL 


M 


45.4(39.0-48.5) 11 


42.4 (35.7-45.1) 5 


49.2 (46.0-53.5) 5 




F 


40.8(36.5-46.1)6 


44.3 (37.1-47.0) 8 


50.7 


MW 


M 


44.6 (42.5-47.3) 9 


47.1 (44.1-51.2) 4 


49.2 (42.9-51.6) 6 




F 


44.9 (44.3-45.3) 3 


44.0 (43.0-45.5) 7 


48.6 


BOW 


M 


12.9(11.0-14.5)9 


14.1 (13.7-14.8)4 


17.6 (16.3-19.2) 3 




F 





13.4 (12.5-14.5) 7 





SH 


M 


69.6(66.1-74.7) 11 


69.5 (67.2-73.5) 5 


71.8 (69.6-75.2) 4 




F 


65.2 (62.5-67.5) 6 


66.4 (62.3-70.4) 8 


69.7 


BL 


M 


26.9 (23.8-29.9) 9 


26.0 (23.8-28.8) 4 


26.4 (25.4-27.9) 3 




F 





24.4 (23.3-25.8) 6 


23.9 


HCW 


M 


55.6 (52.3-57.9) 8 


68.2 (62.6-73.5) 7 


58.8 (57.7-62.2) 4 




F 


51.8(49.2-52.6)7 


50.9 (48.7-53.4) 5 


45.9 


HL 


M 


233.1 (215-275) 7 


326.6 (294-366) 3 



Sexual dimorphism.— DiUerences between sexes in horns and 
horn cores were discussed above and shown in Figures 157, 158. 

Comparisons.— Gazella I. leptoceros is somewhat smaller and 
darker than G. I. loderi. Compared with G. dorcas, G. leptoceros is 
considerably paler, but young female specimens cannot always be 



~-^ 





G. DORCAS 



Fig 155. Lateral views of anterior part of skulls of Gazella leptoceros (opposite) 
and G. dorcas (above) showing variations in lengths of premaxillary and maxillary 
contacts with nasal bone, differences in position of the jugo-maxillary suture, and 
the fenestra in the suborbital fossa of G. leptoceros. 



494 





G. LEPTOCEROS 



496 









I cm 



G.DORCAS 



G.LEPTOCEROS 



Fk; 156. Nasal bones, interparietals, and lower second and third molars of 
Gazella leptoceros and (i. dorcas. Anterior ends of nasals and interparietals toward 
top of figure. Lingual side of teeth upward, anterior to right. Scale refers to molars 
only. 



496 



OSBORN&HELMY: MAMMALS OF EGYPT 497 

identified by color alone. Facial markings, however, are consistently 
paler in leptoceros. The two species differ in numerous other ways 
listed in Table 56 and shown in Figures 154-158, 161. The auditory 
bulla is less inflated in leptoceros. The basioccipital has straight 
sides compared with the narrower constricted basioccipital of G. 
dorcas. The latter, however, has larger, more prominent tuberosities 
on the basioccipital for muscle attachment. 

A line drawn from the ventral surface of the occipital condyle to 
the posterior margin of m^ will not pass through the bulla, except in 
young individuals of leptoceros. Such a line will pass through the 
bulla in dorcas, indicating that basioccipital and basisphenoid meet 
at a sharper angle in G. leptoceros than in G. dorcas. Differences in 
horns and horn cores are illustrated in Figures 157 and 158 and 
listed in Table 56. Horn cores of male leptoceros are conspicuously 
pitted on all surfaces, with intermittent grooves running from base 
to tip (fig. 154), whereas in dorcas, grooves extend to the tip only on 
the posterolateral surface. Differences in dentition are most obvious 
in m2, mg. Enamel is thinner, dentine thicker, islets larger, labial sur- 
faces more folded, and posterolateral folds much more pronounced 
in G. leptoceros (fig. 156). 

Gazella leptoceros differs externally from G. gazella about as from 
G. dorcas in longer, straighter, and outward turning horns; paler 
coloration; and fainter markings. The posterior nasal margins of 
leptoceros and gazella are similar, but interparietal shapes are dif- 
ferent, and the latter rarely has a fenestra in the infraorbital fossa 
(table 56). 

Remarks.— The fenestra in the infraorbital fossa is also present in 
species beyond the boundaries of Egypt — G. subgutturosa of 
southwestern Asia, G. marica of Arabia, and occasionally G. g. 
arabica. 

Specimens examined.— Skins and/or skulls, 17. (Year of collection 
follows number of specimens.) 

BEHEIRA: Wadi el Natroun (1) 1896. 

GIZA: Abu Rawash (1) 1921; Giza, W of (1) 1930. 

FAIYUM: Wadi Rayan (2) 1951. Wadi Mishigeiga (1) 1966, Wadi Muwellih (3) 
1966. 

MATRUH: Cairo, 48 to 64 km. W of (1) 1895; El Maghra, 95 km. SSE. between 
Misaada and El Rammak Dunes (3) 1964; Nuweimisa (2) 1964; Bahrein (1) 1935; 
desert near Siwa (1) 1935. 

Horns or skulls, date of death unknown: 






G.DORCAS 




G.LEPTOCEROS 

Fio 157. Frontal views of horns of Gazella dorcas (left to right: adult male, adult 
female, and subadult female) and G. leptoceros (left to right: adult male, subadult 
male, and adult female). 



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500 FIELDIANA: ZOOLOGY 

MATRUH: Acacia grove 120 km. S of El Maghra (3). Acacia grove 48 km. SW of 
south end of Abu Sennan Ehine (1); Acacia groves, QatCara Depression (1). 

Two "recent" skulls were recorded from north and south of Gebel Uweinat by 
Missone (1969). 

Sight record of I. Helmy.— One large male on limestone plateau 
between El Naqb el Abyad and El Naqb el Ahmar on Siwa-Qara 
track about 10 km. E of Qara (May 1965). 

Published records.— Records are from Bramley (1895), Anderson 
(1902), Shaw (1933), and Churcher (1972). 

BEHEIRA: Wadi el Natroun. 

MATRUH: Cairo 48 to 64 km. W. Western Desert, wadis of Gebel Uweinat in 
1932. 

ASWAN: Kom Ombo (horn core from Late Paleolithic deposits). 

Habitats.— Areas between dunes with scattered Comulaca 
monocantha shrubs south of El Maghra, depressions with stands of 
Acacia raddiana (fig. 18), and sandy areas bordering oases support- 
ing Nitraria retusa (fig. 17) are frequented by this gazelle. One in- 
dividual has been observed on hard, barren desert between Qara and 
Siwa. The general opinion on habitat of G. leptoceros, from observa- 
tions in Algeria (Loder, 1894; Thomas, 1894ab; Pease, 1896), 
Tunisia (Schomber and Kock, 1960), and Egypt (Hoogstraal, 1964) 
is that it lives almost exclusively in dunes or sandy areas. Bramley 
(1895) described a hunt for white gazelle 48 to 64 km. W of Cairo on 
hard desert interspersed with patches of sand supporting perma- 
nent and ephemeral vegetation. 

Although not strictly a sand gazelle in Egypt, G. leptoceros is 
referred to as "Gazelle des Sables" (Pease, 1896) in Algeria. The fact 
that it has never been reported from the Mediterranean Coastal 
Desert, where G. dorcas still exists, suggests that it is more of a 
desert inhabiting species than G. dorcas. Both species, however, fre- 
quent sandy areas because of the browse plant, Nitraria retusa (fig. 
17). 

Flower's (1932, p. 438) statement that G. leptoceros "existed but 
was not numerous in the desert west of Giza which stretches from 
the Wadi Natroun southward to the Faiyum," was verified by con- 
temporary collections from the area (fig. 153). There is no evidence 
however, that G. leptoceros now occupies this area. 

Behavior.— Gazella leptoceros is active during cooler periods of 
the day and jjossibly at night. During the hot part of the day. 



OSBORN&HELMY: MAMMALS OF EGYPT 501 

gazelles lie in the shade of acacia trees or scrape a pit in the sand 
mound under a shrub to get into the small amount of shade that is 
cast. 

Bramley (1895) observed in Egypt that this gazelle would stand 
motionless when danger approached. Both species, we have ob- 
served, run out of the protection of Nitraria bushes in soft sand onto 
hard desert or out of an acacia grove into barren desert when 
frightened by an approaching vehicle. 

Water— Nothing is known of the water economy of the white 
gazelle. We have observed it coming to drink from a spring in Wadi 
Muwellih. 

Food.— Nitraria retusa (fig. 17), a shrub of saline sandy areas 
bordering oases, is a staple food of gazelles in the northern part of 
the Western Desert and often heavily browsed. Specimens of G. lep- 
toceros with stomachs full of spiny Cornulaca monocantha (fig. 108) 
were collected between Misaada and El Rammak Dunes, 95 km. 
SSE of El Maghra. A flower head of Launaea capitata was found in 
one gazelle from Wadi Muwellih. Tracks and clippings around 
Calligonum comosum in that area (fig. 9) indicated that this plant 
was also browsed. 

Gazella leptoceros probably feeds on acacia and other plants 
which have been recorded from G. dorcas where the ranges of the 
species overlap. 

Desert guides have remarked that the celery-flavored Pituranthos 
tortuosus (fig. 10) is a favorite food of white gazelle. Bramley (1895) 
was informed that a plant resembling cranesbill (Geranium sp. or 
Erodium sp.) was also relished. 

Associates.— Gazella leptoceros has been found together with G. 
dorcas in Wadi Muwellih and Nuweimisa and flushed from the same 
acacia groves as the latter, but at different times. 

Reproduction.— In Giza Zoological Gardens, the gestation period 
was five and one-half months, or less than 167 days (Flower, 1932). 
In Algeria, female G. leptoceros often have two young and G. dorcas 
only one (Pease, 1896). 

Gazella dorcas (Linnaeus, 1758) 

Capra dorcas Linnaeus, 1758, Syst. Nat., ed. 10, p. 69. 

Type locality.— "Lov/er Egypt" (Blaine, 1913, p. 292). 

General distribution.—Syria, Palestine, Arabia, Sinai Peninsula, 



502 FIELDIANA: ZOOLOGY 

Egypt, Libya, Tunisia, Algeria, Morocco, Sudan, northern Ethiopia, 
and Chad. ^_^,^ ,. 

Common names.— Dorcas Gazelle, Afri, Ghazal. 

Distribution of subspecies in Egypt— Figure 160. Gazella dorcas 
saudiycu Sinai Peninsula; Gazella dorcas littoralis: Eastern Desert; 
Gazella dorcas dorcas: Western Desert. 

Diagnosis.— Brownish red with distinct facial and body markings. 
Horns in male straight to semilyrate in frontal view. Male horns are 
either almost straight and slightly spreading with tips not hooked, 
or strongly curved and semilyrate with tips hooked forward and 
inward. Fenestra lacking in infraorbital fossa, posterior margin of 
nasals triangular, interparietal suture outline angular. 

Height at shoulder of adult about 55 cm., head and body length 
average 90 cm., tail length 15 cm., hind foot 30 cm., ear 13 cm., con- 
dylobasal length 16.4 cm., weight 15 kg. 

External characters. — Reddish brown dorsally with pale side 
stripe and triangular hip patch distinct, dark side stripe and pygal 
border about the same shade or darker than dorsum. Color of side 
extending down outside of legs. Circumoral region grayish. Throat, 
chest, belly, rump, and inside of legs white to varying shades of 
buff. Facial markings distinct. Central facial band from base of 
horns reddish, either becoming paler toward nostrils or with 
blackish nose spot. Light lateral facial stripe from base of horns to 
muzzle, whitish. Dark lateral facial stripe from lacrimal gland to 
mouth, brownish to blackish. Ear grayish on outer surface, buffy on 
inner surface with pale border. Tail, carpal tufts, and hair between 
dewclaws and hoofs black. 

Hoofs. — Hoofs of specimens from sandy areas tend to be longer 
than those from hard and stony desert (table 55 and discussion 
under G. leptoceros). 

Cranial characters.— Figures 156, 161. Skull broad posteriorly 
with parietal ridges present in all ages and supraoccipital crest not 
pronounced nor extending beyond level of occipital condyle. 
Posterior margin of nasals tapering gradually (angular) beginning 
at level of ethmoidal fissure and anterior to nasofrontal suture. In- 
terparietal suture outUne angular (fig. 156). Infraorbital fossa lack- 
ing fenestra. One-third of the infraorbital fossa is formed by 
encroachment of the jugal onto the lacrimal bone. 

Jugomaxillary suture straight or slightly curved (fig. 155). 



OSBORN&HELMY: MAMMALS OF EGYPT 503 

Length of premaxilla contact with nasal variable, but usually less 
than length of maxilla contact (fig. 155). Auditory bulla with 
smooth ventral surface. Basioccipital constricted due to medial 
swelling of bullae. Tuberosities on basioccipital strongly developed. 
These and additional characters are listed in Table 56. 

Horns and horn cores.— Figures 157, 158. Male horns are of two 
types. In subspecies dorcas and littoralis, horns are semilyrate in 
frontal view with tips curving strongly anteriorly and inwardly, 
strongly curved backward and forward in lateral view. In saudiya, 
horns are not lyrate, but almost straight and spreading very slight- 
ly, with tips slightly curved inwardly in frontal view, and nearly 
straight in lateral view. Annulations are conspicuous, not as close 
as in G. leptoceros, and complete except for distal 10 to 12 below the 
short tip. Smooth tips are about 25 per cent of total horn length (fig. 
157). Female horns are slenderer and straighter than male, with 
thin, poorly developed annulations (fig. 157). 

Horn cores of males diverge widely from the base in dorcas and lit- 
toraUs, and like the horns, are strongly curved (fig. 161). Proximal 
grooves extend to tips only on the posterolatersd surface. Horn 
cores of females are less diverging and smoother. Base of male horn 
core in cross section is ovoid and narrower anteriorly than posterior- 
ly. The female horn core is slightly ovoid and much smaller in 
diameter (fig. 158). 

TeetA.— Figure 156. Lingual ridges not prominent on lower molars 
(m2, mg). Posterior labial folds absent. Enamel is thick, dentine thin, 
and islets small. 

Measurements.— Table 57. Males are larger than females, with ex- 
ception of a few dimensions in littoralis. Tail length, although quite 
variable, averages shorter in females. 

Age determination. —Specimens are considered adult when all per- 
manent teeth have emerged. 

Sexual dimorphism.— Differences between sexes in horns, horn 
cores, and measurements are discussed above and illustrated in 
Figures 157, 158. 

Viariation.— Coloration varies from pale to dark reddish brown, 
with dark side stripe the same shade as the back in Western Desert 
dorcas, slightly darker than the back in the Eastern Desert lit- 
toralis, and paler again in saudiya of Sinai and the Arabian Penin- 
sula. The last two usually have a dark or blackish spot on the nose. 



504 FIELDIANA: ZOOLOGY 

and dorcas does not. In saudiya, the belly is marked with reddish 
buff, but other subspecies have pure white bellies. 

Horns are often blacker in the Eastern Desert form, as discerned 
by Blaine (1913). Most littoralis have a V-shaped postpalatal 
margin, but in dorcas, it is U-shap)ed. 

Ear, foot, and a few other measurements average slightly larger in 
littoralis than in dorcas (table 57). 

Co mpansows.— Comparison with G. leptoceros is under that 
species and in Table 56. Gazella dorcas can be distinguished from 
the more similar G. gazella by longer and straighter male horns and 
longer female horns, posterior margin of the nasals triangular 
rather than rounded. Where the ranges overlap, G. dorcas has red- 
dish buff markings on the belly, and the belly is white in G. gazella. 
Species characters are compared in Table 56. 

Habitats.— Eastern Desert: Accessible wadis and canyons of 
North and South Galala Plateaus, wadis of Red Sea Hills, vegetated 
beaches on Gulf of Suez and Red Sea coasts, and coastal plains of 
the southeast. "The proximity of gazelles to the sea coast in the 
Elba area is in strong contrast to their complete absence from the 
relatively rich Mediterranean littoral of northern Egypt, where 
larger human populations and easy accessibility by jeep have driven 
them miles inland to more barren desert" (Hoogstraal et al., 1957b, 
p. 61). 

Western Desert: Littoral desert inland from the coast, sandy and 
hard desert, margins of oases, wadis and canyons in cliff and moun- 
tain areas. 

Be/iayior.— Activity generally appears to be similar in G. dorcas 
and G. leptoceros. The seeking of shade is characteristic of both, and 
in the Eastern Desert, G. dorcas takes advantage of overhanging 
cliffs. 

The alarm note of dorcas, according to Flower (1932, p. 440), 
'*. . .is a single, short bark uttered by one animal. It is an instant 
signal to the whole herd." 

Gazelles are very nervous and wild, but if raised from a young 
age, they become tame and docile. Gazelles become comparatively 
tame in oases in summer when they come to drink at the springs 
(Anderson, 1902). Buxton et al. (1895) and Osborn and Krombein 
(1969) experienced the habit of G. dorcas of keeping in advance of 



OSBORN&HELMY: MAMMALS OF EGYPT 505 

pursuers in a wadi until it was nearly at an end or became steep and 
rocky, then turning and attempting to run through its followers. 

Vyater.— Observations made on G. dorcas in the vicinity of Khar- 
toum, Sudan, indicate that gazelles must drink, even in winter, and 
appear to inhabit areas "where some water, fresh or saline, or dew 
and succulent food are available, even if considerable distances have 
to be travelled in order to obtain them" (Cloudsley-Thompson and 
Ghobrial, 1965, p. 1313). Captive gazelles deprived of water were 
able to survive up to 12 days in winter and five days in summer in 
Khartoum (Ghobrial and Cloudsley-Thompson, 1966), and 
withstood a weight loss of 17 to 20 per cent (Ghobrial, 1976). During 
the dry season in Sudan, dorcas gazelles ate leaves only of Acacia 
tortilis, which contained about 60 per cent moisture, and not ends of 
twigs or bark. Grasses (Panicum turgidum, Aristida sp.) and green 
shrubs {Capparis decidua, Leptadenia pyrotechnica, Boscia 
senegalensis, and Cassia senna) were ignored (Carlisle and Ghobrial, 
1968). 

The demand for water provoked gazelles to race through Wilkin- 
son's (1832) camp in a narrow wadi in order to drink from a spring 
further inside the valley. Tracks of gazelles along the banks of the 
Nile River in Nubia are common, and occasionally, the animals 
themselves are seen drinking. 

In the interior of the Egyptian deserts, gazelle tracks are always 
present around shallow, saline wells (personal observations of the 
authors). Meinertzhagen (1954) observed gazelles drinking sea 
water on an island off the Somalia coast and on the Red Sea 
Littoral. However, Ghobrial's (1976, p. 490) experimental results 
showed "that gazelles do not voluntarily ingest sea water to any 
great extent, even when deprived of fresh water." 

Food — In the Eastern Desert, Acacia ehrenbergiana and A. rad- 
diana are staple food of G. dorcas. Leaves, thorns, and flowers are 
stripped from accessible branches (fig. 159), and pods are eaten 
green or dry. The one-and-a-half-inch long thorns have been found in 
stomach contents. Nitraria retusa (fig. 17) is also browsed along the 
Gulf of Suez and northern part of the Red Sea coast. 

Additional plant species which have been found in the stomachs 
of gazelles are: pieces of the midrib of a date palm leaf (Phoenix dac- 
tylifera) and flower head of Cuscuta sp. near Bir Abbad; leaflets of 
Psoralea plicata, Wadi Allaqi 11 km. inland from AUaqi Village; 



506 



FIELDIANA: ZOOLOGY 




Fig. 159. Acacia ehrenbergiana browsed by GazeLla durcas. Twigs are stripped of 
leaves, flowers, bark, and thorns. 



pods of Astragalus vogelii, Wadi Umm Karayiet and Wadi Allaqi 
junction; leaf fragments of P. plicata and possibly Cleome sp., Wadi 
Ibib; leaves of P. plicata and fruits of Crotalaria sp., Wadi Eteigan; 
leaves, flowers, and fruits of Hippocrepis cons trie ta, pods of 
Crotalaria aegyptiaca (?), fruits of Euphorbia granulata, pods of 
Lotus glinoides, and leaf fragments of Aizoon canariense in Wadi 
Voider, near Gebel Nesla. 

In the northern sector of the Western Desert, Nitraria retusa, 
mentioned under G. leptoceros (fig. 17), is also a staple food of G. 
dorcas. Pods oi Acacia raddiana were found in a specimen from Tahl 
el Fawakhir near Qara, and doubtlessly Acacia sp. are eaten 
whenever available. Fragments of Moltkiopsis ciliata. Anabasis 



OSBORN&HELMY: MAMMALS OF EGYPT 507 

articulata, and Suaeda sp. were found in a gazelle from the vicinity 
of Qaret el Mashnika. Various plants mentioned under G. leptoceros 
are probably eaten by G. dorcas and vice versa. 

In wadis of Gebel Uweinat, browse plants are: Crotalaria 
thebaica, Argyrolobium saharae, Trichodesma africanum, Farsettia 
ramosissima, and bitter green fruits of Citrullus colocynthis 
(=Colocynthis vulgaris) (Osborn and Krombein, 1969). Continuous 
browsing of gazelles and periodic cropping by camels prevent some 
species such as Convolvulus lanatus and Astragulus spinosus from 
becoming more than a small, dense cushion (fig. 12). We observed 
that gazelle browse camel thorn, Alhagi mannifera (=A. maurorum) 
in vicinities of Bahariya, Dakhla, and Kharga Oases. Careful 
examination of a gazelle feeding area, an association of A. mannifera 
and Imperata cylindrica, revealed no evidence that the grass was 
eaten by gazelles. Anderson (1902) wrote that gazelles occasionally 
fed at night on crops in the oases. 

Reproduction.— The gestation period of five and a half months is 
the same for G. dorcas as G. leptoceros. Single embryos, fetuses, 
and newly born young taken from seven females in December, 
March, and April indicate that one young is usual in G. dorcas. 

Associates.— Gazella dorcas is found together with G. leptoceros, 
as mentioned under the latter, and occasionally with camel herds in 
the Mediterranean Coastal Desert. In Tunisia, it apparently joins 
herds of camels to play with them (Schomber and Kock, 1960). 

Predators.— Jackal, wild cats, and cheetah are presumably 
natural predators of gazelle. The most efficient predator is man, 
with snares and wheel traps (Anderson, 1902), guns, and 
automobiles. Flower (1932) mentioned the decrease of gazelles due 
to man in the previous 40 years. 

Wanton slaughter of gazelles under the guise of "sport" so that a 
car may be draped with dead bodies for a photograph, together with 
the traditional belief that "there are always plenty of gazelles," is 
rapidly reducing these animals to extinction. 

Remarks.— Remains of G. dorcas found in Late Paleolithic sites 
near Kom Ombo have been reported by Reed and Turnbull (1969) 
and Churcher (1972). 

Key to Egyptian Subspecies of Gazella dorcas 
1. Horns of male spreading from base, curved, tips hooked. Venter white. 

a. Dark side strif>e usually darker than back. Black nose spot present. (Eastern 
Desert) littoralis, p. 508. 



508 FIELDIANA: ZOOLOGY 

,. 25* 26* 27* 28* 2 9* 30* 31* 32* 33* 34* 35* 36* 37* 




Fig. 160. Collection localities of Gazella dorcas dorcas (dots), G. d. Uttoralis 
(circles), 0. d. saudiya (square), and 0. gazella ara6ica(triangle); sight records (S); 
and skulls and horns, date of death unknown (X). 

b. Dark side stripe usually paler than back. Black nose spot absent. (Western 

Desert) dorcas, p. 509. 

2. Horns of male not spreading from base, almost straight, tips not hooked. Dark 
side stripe usually paler than back. Black nose spot variable. Venter marked with 
reddish buff. (Sinai Peninsula) saudiya, p. 512. 

Gazella dorcas littoralis (Blaine, 1913) 

Gazella littoralis Blaine. 1913, Ann. Mag. Nat. Hist., (ser. 18). 11. p. 295. 
Type locality. —Sudan: Khor Asot, Nubian Desert. 
Distribution in Egypt— Figure 160. Eastern Desert. 

External characters. — Reddish brown, with dark side stripe 
darker than back. Blackish nose spot usually present. Belly and 
underparts white. 



OSBORN & HELM Y: MAMMALS OF EGYPT 509 

Cranial characters.— Figure 161. See species description. 
Postpalatal margin usually V-shaped. 

if oms.— Figure 157. See species description. Horns generally 
blacker than in other subspecies. Length and shape about as in G. d. 
dorcas, but shorter and more curved than in G. d. saudiya. Com- 
parison is given under the latter subspecies. 

Measurements.— TBhle 57. Gazella d. littoralis averages slightly 
larger in most dimensions, particularly ear length and horn core 
width, than G. d dorcas. From data given by Groves and Harrison 
(1967) and Harrison (1968), littoralis is generally larger than 
saudiya. Some discussion of measurements is given under the latter 
subspecies. 

Specimens examined.— Skins and/or skulls, 22. (Year of collection 
follows the number of specimens.) 

RED SEA: Wadi Umm Huweitat (1) 1965, Wadi Saqi (1) 1966, Wadi Sukari (1) 
1965, Bir Abbad (1) 1964, Wadi Umm Had (1) 1975. 

QENA: Wadi Qena mouth (2) 1966. 

SUDAN ADMINISTRATIVE: Wadi Ibib (1) 1967, Wadi Yoider near Gebel Nesla 
(2) 1967, Wadi Eteigan (1) 1967; Wadi Abeib 3.6 km. N of Bir Kansisrob (3) 1954, 6.4 
km. N of Bir Kansisrob (3) 1954. 

ASWAN: Wadi AUaqi 11.2 km. SE of AUaqi Village (2) 1966, Wadi Umm Karayiet- 
Wadi Allaqi junction (1) 1966, Bir Murra (2) 1966. 

Sight records.— Personal observations of I. Helmy and D. Osborn. 

SUEZ: Wadi el Gindali (smaU herd) and Wadi IseiU (2-3) 1964, Wadi el Ghuweibba 
(1) 1966, Wadi Abu Seyala (5) 1961. 

RED SEA: Wadi Mellaha (1) 1965, Wadi Atrash (1) 1965, Wadi Fatira el Zarka (1) 
1965, Wadi Abu Sheeh (4) 1969, Wadi Semna (10) 1966, Wadi Atallah (1) 1969, Dissht 
el Daba (1) 1965, Wadi Abu Ziran (1) 1966, Wadi Umm Had (1) 1975. 

ASWAN: AUaqi Village 20 km. S (1) 1966. 

SUDAN ADMINISTRATIVE: Wadi Eteigan (2) and Wadi Dub (smaU herds) 
1967. 

Published records.— Records are from Hoogstraal et al. (1957b). 

SUDAN ADMINISTRATIVE: Wadi Adeib, 3.6 km. N of Bir Kansisrob. 6.4 km. 
N of Bir Kansisrob, and coastal plain near Gebel Elba (herds of 2 to 20) 1954. 

Gazella dorcas dorcas (Linnaeus, 1758) 
Type locality.— Lower Egypt. 

Distribution in Egypt— Figure 160. Western Desert. 
External characters.— Pale reddish brown, dark side stripe usual- 




FlO. 161. Skull of Gtuella donas. 



510 



OSBORN&HELMY: MAMMALS OF EGYPT 511 

ly slightly paler than back. Blackish nose spot absent. Belly and 
underparts white. 

Cranial characters.— Figyire 161. See species description. Post- 
palatal margin usually U-shaped. 

Horns.— Figure 157. Similar in outline to G. d. littoralis. 

Measurements.— Table 57. Gazella d. dorcas averages slightly 
smaller in most dimensions, particularly ear length and horn core 
width, than G. d. littoralis. Further comparisons are under the 
species and G. d. saudiya. 

Specimens examined. —Skins, skulls, and horns, 40. (Year of col- 
lection follows number of specimens.) 

FAIYUM: Wadi Rayan (2) 1950. (1) 1951; Wadi Mishigeiga (1) 1966. 

QENA: Isna. Wadi Nassim (3) 1953. 

ASWAN: Kurkur Oasis (3) 1963. 

BEHEIRA: Wadi el Natroun (2) 1923. 

MATRUH: Qasr el Qatagi (2) 1962. (1) 1965; Nakhlet el Barraq (1) 1962, (1) 1963; 
Qaret el Mashruka (2) 1962; El Fureinat (2) 1974; El Maghra E of (1). 20 km. E of (1) 
1974; Acacia grove 120 km. S of El Maghra (1) 1965 and horns (date of death 
unknown); Raqabet el Rala (1) 1964; Wadi Umm Shedak (1) 1962; Qara, Tahl el 
Fawakhir (1) 1965; Bahrein (1) 1964; Nuweimisa (2) 1964; Siwa Oasis 10 km. N (1) 
1974, 48 km. N (1) 1951; Salum 24 km. SW (2) 1953; Bir Sidi Omar (1) 1964; Acacia 
groves, Qattara Depression (two cadavers, date of death unknown). 

Sudan. NORTHERN: Gebel Uweinat, Karkur Murr 12 km. W (2) 1967. 

Sight records.— Personal observations of I. Helmy and D. Osbom. 

GIZA: Bahariya Oasis N of (2) 1966. 

BEHEIRA: Bir Victoria (2) 1969. 

MATRUH: Wadi el Farigh (2) 1969; Abu Mena 10 km. E (2-3) 1963; El Afritat (2) 
1964. 1970; Wadi Labaq (1) 1966; Halfaya (small herd) 1965; Qur el Hilab (10) 1965; 
El Maghra (small herd) 1958, 1960, 1963, 1965, 1967, none seen in 1977; Bir 
Mikheimin (1), Hatiyet Labbaq (one seen and tracks of several), Samaket Gaballa (2), 
acacia groves in Qattara Depression (tracks of a few individuals) 1977. 

EL WADI EL GEDEED: Farafara Oasis, Hatiyet el Sheikh Marzuk (1), Karawein 
(4-5). El Gau (1), Wadi Hennis (2) 1969; Dakhla Oasis. Mut (1) 1969; Dakhla Oasis 60 
km. E (small herd) 1966; Bir el Shab (1) 1967. 

ASWAN: Kurkur Oasis (2) 1962. 

Published records.— Records are from Hoogstraal (1964). 

MATRUH: Siwa 48 km. N. 

GIZA: Giza pyramids a few miles W of. 

FAIYUM: Wadi Rayan. 

QENA: Wadi Nassim. 



512 FIELDIANA: ZOOLOGY 

Gazella dorcas saudiya (Camithers and Schwarz, 1935) 

Gazella gazella saudiya Carruthers and Schwarz, 1935, Proc. Zooi. Soc., London, 
1935. p. 155. / 

Type locality.— SanAi Arabia: Dhalm 240 km. NE of Mecca. 

Distribution in Egypt— Figare 160. Sinai Peninsula. 

External characters. — Pale reddish brown, dark side stripe usual- 
ly paler than back as in G. d. dorcas. Blackish nose spot variable. 
Differs from other subspecies in not having the belly pure white, but 
variably marked with reddish buff, especially below the side stripe 
(after Harrison, 1968). 

Cranial characters.— Differs from other G. dorcas subspecies in 
having a generally longer nasopremaxilla contact (Harrison, 1968, 
p. 357). 

Horns.— Horns, in contrast with other subspecies of G. dorcas, are 
longer, straighter, and not spreading. Harrison's (1968) 
measurements of horn length, in straight line from anterior base to 
tip, of adult males from Arabia range from 244.5 to 304 mm. 
Measurements taken on the front curve of G. dorcas from Egypt 
(table 56) range from 215 to 275 mm., which would be considerably 
less if taken in a straight line. 

Measurements.— Ear length, according to Groves and Harrison 
(1967), is very long in G. d. saudiya, but they were comparing the 
shorter-eared G. gazella arabica. No external measurements are 
available from saudiya. 

Cranial measurements from Arabian specimens given by Groves 
and Harrison (1967) and Harrison (1968), when compared with those 
in Table 57, indicate that littoralis is slightly larger than saudiycu 
Horn core width is considerably less in G. d. saudiya from Arabia 
than in Egyptian subspecies (table 57). The above authors list a 
mean horn core width of 37.6 ±1.8 mm. and maximum of less than 
60 mm., respectively, for saudiya. 

Remarks.— No specimens are available from Sinai Peninsula, and 
there are no examples of intergradation between the subspecies 
littoralis and saudiya. 

Sclater and Thomas (1894-1900) assumed G. arabica reported by 
Hemprich and Ehrenberg (1833) on the Sinai coast between Suez 
and Tor to be G. dorcas. 



OSBORN&HELMY: MAMMALS OF EGYPT 513 

Specimens examined.— Total three. 

Saudi Arabia: Dhalm (3). 

Published records.— Record is from Anderson (1902). 

SINAI: Ayun Musa. 

Gazella gazella (Pallas, 1766) 

Antilope gazella Pallas, 1766, Misc. Zool., p. 7. 

Type locality. —Syria. 

General distribution.— Syria, Palestine, Arabian Peninsula, Sinai 
Peninsula. 

Common names.— Mountain Gazelle, Arabian Gazella, Idmi. 

Subspecies in Egypt — 

Gazella gazella arabica (Lichtenstein, 1827) 

Antilope arabica Lichtenstein, 1827, Darstellung Saugeth., pi. 6. 

Type locality.— Saudi Arabia: Farsan Island, eastern Red Sea 
coast. 

Distribution in Egypt— Figare 160. Northeastern part of Sinai 
Peninsula. 

Diagnosis.— Pale reddish brown with distinct facial markings and 
a dusky nose spot. Body markings variable. Horns short and 
strongly curved. Fenestra usually lacking in infraorbital fossa. 
Posterior margin of nasals round, interparietal suture angular. 

Height at shoulder in adult male about 58 cm., condylobasal 
length about 18 cm. 

External characters.— Pale reddish brown dorsally with distinct 
gazelline markings. Dark side stripe grayish black. Belly and under- 
parts white. Very similar to G. d. dorcas. 

Cranial characters.— Braincase expanded posteriorly. Parietal 
ridges absent. Supraoccipital crest about level with posterior 
margin of occipital condyle. Posterior margin of nasals round, inter- 
parietal suture angular. Infraorbital fossa lacking fenestra, with few 
exceptions. Premaxilla usually not in contact with nasal. Auditory 
bulla more swollen than in G. dorcas (Gentry, 1964). 

Horns.— Male horns are shorter than in other gazelles, spreading 
from the base, strongly recurved, with tips turning forward and 
inward. Female horns are straighter, slenderer, and markedly 



514 FIELDIANA: ZOOLOGY 

shorter than in the male of the species or females of other species 
(Harrison, 1968). Straight line measurements of male horns from 
Harrison range from 150 to 254 mm. and from Morrison-Scott 
(1939). 205 to 256 mm. 

Teeth,—Simi\ar to G. dorcas in Figure 156. 

Measurements. — Means (and ranges) of measurements (in 
millimeters) of males listed by Harrison (1968) that are comparable 
with those in Table 57 are: Condylobasal length 183.1 (174.4 to 
191.0) 8. braincase width 54.6 (52.3 to 56.8) 11, postorbital width 
52.3 (49.1 to 55.8) 11, and upper tooth row length 57.0 (54.0 to 60.0) 
11. These data indicate that G. gazella is larger than G. dorcas. Ear 
lengths of 109, 119, and 120 mm. given by Harrison are shorter than 
measurements of G. dorcas in Table 57. Horn core width of adult 
males from Groves and Harrison (1967) is 64.5 ± 2.3 mm. and from 
Morrison-Scott (1939), 65.9 mm. (60.2 to 69.0) 15, somewhat less 
than G. d. littoralis of the Eastern Desert. 

Co mpansons.— According to Harrison (1968), G. gazella is larger 
and longer limbed than G. dorcas and has much shorter ears. The 
two species are similar in color and have comparable variations. 
Gazella gazella can be distinguished from G. dorcas and G. lep- 
toceros by its shorter horns in both sexes, shorter ears, and cranial 
combination of round posterior nasal margin, angular interparietal, 
and shortness of or absence of premaxillary-nasal contact. Various 
characters of gazelles are compared in Table 56. 

Specimens examined. — Four from Saudi Arabia. 

Published records. —Said to have been seen in Sinai, including 
Wadi el Arish (Flower, 1932, p. 438). 

Habitats.— Coastal plains, foothills, and mountains of the Ara- 
bian Peninsula; absent from interior steppe and desert (Harrison, 
1968). 

Genus Capra Linnaeus, 1758 

Large, goat-like bovid with scimitar-shaped horns curving dorsal- 
ly and posteriorly. Male horn with large, anterior knobs. Color 
brownish, leg markings black and white. Muffle completely haired. 
Beard present in male. Tail shorter than ear. Interdigital glands pre- 
sent on forefoot, subcaudal glands present in male. Lacrimal and in- 
guinal glands absent. 

Braincase not expanded. Orbital region noticeably broad. Fron- 



OSBORN&HELMY: MAMMALS OF EGYPT 515 

tonasal region concave. Infraorbital fossa lacking, ethmoidal fissure 
small. Premaxilla contacting nasal. Horn core supraorbital and ex- 
tending dorsally from base. 

Capra ibex Linnaeus, 1758 

Capra ibex Linnaeus, 1758, Syst. Nat., ed. 10, p. 68. 
Type locality.— VeXais, Switzerland. 

General distribution.— yiountains of southern and eastern 
Europe, Transcaucasia, Russian and Chinese Turkestan, Mongolia, 
Afghanistan, India, Saudi Arabia, Syria, Israel, Egypt, Sudan, 
Ethiopia. 

Common names. — Ihex., Taytal, Beden. 

Subspecies in Egypt — 

Capra ibex nubiana (F. Cuvier, 1825) 

Capra nubiana F. Cuvier, 1825, in I. Geoffrey St. Hilaire and F. Cuvier, Hist. Nat. 
Mamm., Vol. 3. pt. 50, pi. 397, p. 2. 

Type locaiity.—Egypt. Nubia or Upper Egypt. 

Distribution in Egypt— Figare 162. Sinai Peninsula and Eastern 
Desert. 

Diagnosis.— Upper parts brownish with contrasting black spinal 
crest and black and white leg markings. Beard present in male. Tail 
shorter than ear. Male horn long, scimitar-shaped, anterior surface 
knobbed. Skull broad at orbits, nasofrontal region concave. 
Shoulder height 84 to 87 cm. 

External characters.— General color brownish grizzled with 
whitish. Muzzle, chin, beard, chest, spinal crest, flank, side of tail, 
and front part of legs (except knees and pasterns) black. Belly, inner 
side of thigh, inner and back side of legs, knees, and a band above 
each hoof, whitish. Faint mark between eye and mouth. Outer side 
of ear brownish, inner blackish with white border. Muffle complete- 
ly haired. 

Cranial characters.— Figare 163. Orbital region broad, prominent. 
Frontonasal contour concave. Braincase constricted laterally. Oc- 
cipital condyle protrudes well beyond level of supraoccipital in 
adults. Frontoparietal suture almost straight. Interparietal suture 
curved. Premaxillary long, contacting nasal. Nasofrontal suture 
posterior to anterior margin of orbit. Infraorbital fossa lacking, 



516 FIELDIANA: ZOOLOGY 

, 25* 26* 27* 28* 2 9* 30* 31* 32* 3 3* 34* 35* 36* 37* 




Fig. 162. Collection localities of Capra ibex nudtano, undated and prior to 1932 
(circles) and 1948 to date (dotsi; horns or skulls, date of death unknown (X): and 
sight records (S). 

ethmoidal fissure small. Postpalatal margin has a narrow V-shai)ed 
cleft and apex of cleft reaches level of m^ 

Horns and horn cores.— Figure 163. Horns present in both sexes; 
smaller and relatively smooth in female. Male horn scimitar-shaped; 
curving dorsally and posteriorly, medially at tip; reaching outside 
curve length of 1 m. or more; flattened laterally, anterior surface 
broadest and with large, regularly arranged transverse knobs 
almost to tip. Pronounced keel between knobs from base to tip. Base 
of horn supraorbital in position. Core grooved and pitted on all sur- 
faces from base to tip. Cross-section ovoid, broadest anteriorly. 
Basal part hollow, without trabeculae. 

Teeth.— Figare 163. Upper cheek teeth, except pm' with well 



OSBORN & HELMY: MAMMALS OF EGYPT 



517 




Fk; 163. Skull of Capra ibex nubiana. 



developed labieil ridges. M^ without posterior accessory lingual 
ridge. 

Measurements.— Table 58. 

Sexual dimorphism.— Males are larger than females. Male horns 
in comparison with female are much longer, heavier, and have knobs 
on the anterior surface. Males and very old females have beards. 

Variation. — Knobs on horns are reported to be narrower and less 
regular in Sinai than in Eastern Desert specimens (Lydekker, 1916). 

Comparisons.— Capra i. nubiana, in comparison with the Abyssi- 
nian ibex, (C. i. walie), is smaller, paler, has a slightly longer beard, 
and longer and less massive horns (Dorst, 1970). 

Specimens examined.— Skins and skulls, four. (Year of collection 
follows number of specimens.) 



r 

518 FIELDIANA: ZOOLOGY 

Table 58. — Measurements of two specimens of Capra ibex nubiana. 

Immature Adult 

Male Female 

HBIj 999 

TL 82 

PL 260 

EL 114 

CBL 197 208 

OW 109.8 108.6 

BCW 68.9 68.1 

NL 70.3 73.6 

TRL 62.7 65.0 

HL 630 320 



SINAI: Gebel el Rabbah (2) 1905. 

RED SEA: Wadi Gozah (immature skin) 1964. Wadi Rawd Ayiad (1) 1965. 

Horns or skulls, date of death unknown: 

SUEZ: Wadi Qiseib (3). 

RED SEA: Gebel Abu Harba (1). 

ASWAN: Bir Umm Hibal (1). 

Sight records of D. Osborn and I. Helmy.— Bir Qiseib (small herd 
and individuals seen on several occasions) 1964 - 1967, Wadi Abu 
Sanduq (tracks, 1964). Bir Mellaha (1) 1965, Gebel Katamiya (drop- 
pings and reports of animals seen by observatory personnel) 1964. 

Published records and reports.— The following are listings by 
date, where possible from Palmer (1872), Wilkinson (1832), Hart 
(1891), Floyer (1893), Buxton et al. (1895), Buxton (1898), Anderson 
(1902), Hume (1906), Barron (1907a), Weldon (1909), Stuart (1910), 
Murray (1912. 1930, 1967), Flower (1932), Russell (1949a), Tregenza 
(1955, 1958), Couturier (1962), Wassif and Hoogstraal (1954), 
Hoogstraal et al. (1957ab), Hoogstraal (1964). and Kock (1971). 

SINAI: Wadi Isla (Isleh) (1923. 1893): near Tor (1823); Wadi Hebran (1823. 1851. 
1932); Gebel Horeb (1831): Wadi Feiran (1826); Gebel Serbal (1831, 1918. 1949. 1964); 
Mt. Sinai (1832); Wadi Salafe (1851); Wadi el Kharig (Rhatakit) (1851); Wadi Gharan- 
dal and Wadi Sigilliyeh (tracks, 1872); Wadi Hanjurat el Qattar (herd. 1872); Gebel 
Umm Shomer and Wadi Saal (herd, 1872; reported 1968); Wadi Araba and Gebel Hor 
(1891); Gebel Umm Alawi and Wadi Nasb (1893); Wadi Nesle (23 seen in mts. prior to 
1895); Gebel Baba. Wadi Shellal. Wadi Aleyat. Gebel Hamra. Nakb Baraq (1898); 
Gebel Genawi. Wadi Kid (Kyd). Wadi Ethmiemat, Tellat Gimal, and Fersh Sheikh el 
Arab (1906); Gebel el Raba (Rhaba. Rabbah) (12 seen. 1909; reported 1968); Gebel 
Sinn (good place for ibex. 1909); Gebel Catherine (no date); Wadi Satakh and Wadi 
Geba (1907): Gebel Tarbush (1932. 1949); St. Catherine Monastery area (1954); Ayun 
Musa (1956); Gebel Umm Afruth and Feiran Oasis (1968). 



OSBORN&HELMY: MAMMALS OF EGYPT 519 

ISMAILIYA: Gebel Shubrawit (one seen several times in 1967; personal com- 
munication of an Egyptian military officer). 

SUEZ: Gebel Ataqa (no date), mountains near Suez (1866), Gebel Naqud (1881). 

CAIRO: Toura cliffs (prior to 1893). El Saff Plateau (prior to 1932). 

RED SEA: Gebel Tenassib and Wadi Qattar (two shot. 1823). near Quseir (1878), 
Bir Hindusi and Gebel Abu Tiyur (1865. 1878), St. Paul Monastery area (1876, 1878), 
Wadi el Abyad and Wadi Naqud (1881), Gebel Gharib (1886), Gebel Qattar (Kitar) 
(1892. 1893). Gebel Abu Dokhan (1892. 1893. numerous in 1910. 1951). Gebel 
Zabarra and Wadi Lahama (Lehema) (two shot. 1893), Bir Sheitun (100 killed about 
1918; one shot in 1927), Wadi Asyuti and Wadi Habeeb (one killed, one seen in 1927), 
Wadi Umm Balad (1947), Gebel Umm Gidri and Gebel Shayeb (always present, 
1949), Wadi Rishrash (30 to 40 photographed in 1932; a few left in 1960). Wadi 
Markh. 40 to 45 km. NW of Qena (old horn. 1951). Wadi Mitgal, Gebel Abu Harba, 
and Wadi el Atrash (1951). 

ASWAN: Near Aswan, El Diwan near El Derr (1813). 

SUDAN ADMINISTRATIVE: Gebel SheUal (many seen. 1926). Gebel Elba area 
(horns seen in local huts, natives reported killings in dry seasons, 1954). 

Sudan. KASSALA: Gebel Asoteriba (about 50 seen on summit, 1926). 

Habitats.— Rocky wadis, cliffs, and mountains. One specimen was 
shot on the barren rocky, gravelly plain of Wadi Rawd Ayiad near 
Qusur el Banat. 

Behavior. — Ibex are alert and shy. Their agility enables them to 
ascend steep cliffs rapidly. A climbing ibex may suddenly become 
motionless and impossible to see. The alarm note is a sharp whistle. 

Water.-lhex cannot survive without water and apparently travel 
long distances in order to drink. Their vulnerability at watering 
places is mentioned below. 

Food.— Acacia raddiana was browsed by ibex in Wadi Abu San- 
duq. Woody shrubs browsed in Wadi Qiseib were: Lindenbergia 
sinaica; Lycium shawii; Capparis spinosa, particularly the flower 
buds (Osbom, 1968b); and Ficus pseudosycomorus. Phragmites 
australis, Imperata cylindrica, J uncus rigidus,. and Alhagi man- 
nifera were also eaten. Funnel-shaped pits in wadi gravel seen by 
Tregenza (1958) were supposedly made by ibex that twisted out the 
roots of Lotus arabicus. 

Aside from food plants, an Abadi told us that the male ibex rub 
their horns in the stiff, pungent foliage of Cleome droserifolia. 

Associates. — In times past, ibex and Barbary sheep probably 
inhabited much the same areas. 

Predators.— Buxton et al. (1895) reported killing of ibex by 
leopards in Sinai. 



520 FIELDIANA: ZOOLOGY 

Historical notes. — Ibex are depicted on rocks throughout the 
Eastern Desert. Particularly good Stone Age carvings are in the 
Nubian sandstones of Wadi Hammamat (personal observations of 
authors). 

Prehistoric man hunted ibex by bringing them to bay with dogs, 
then killing them with stones or arrows; or by lying in ambush in 
little stone blinds built near watering places where they could be 
stoned or shot. Hunting methods today are essentially the same, 
and the stone blinds remain as they were built centuries ago. 

During Dynastic times, temples and tombs the length of Egypt 
were adorned with figures of ibex. The early Egyptians hunted ibex 
for sport, kept them as pets, and offered them in sacrifice to their 
gods (Buxton et al., 1895). 

Capturing or killing animals for pleasure as well as profit is an 
ancient profession which has changed little with time. Northern 
Sinai tribes once had an ibex business with Suez (Weldon, 1909). 
Bedouins of the Hamada section of southwestern Sinai sometimes 
had tame ibexes with their sheep and goats (Murray, 1912). 

In the early 1900's according to Russell (1949ab, 1951), hunters 
were few and their weapons primitive. During World War I, Egyp- 
tians in the Nile Valley towns of Ekhmim, Ebnub, Badari, and 
Minya exchanged their flintlocks for modern military rifles. Armed 
with efficient weapons and rope snares, they mercilessly hunted the 
ibex for profit in the sale of meat. Bir Sheitun, sometimes being the 
only available water for hundreds of miles, was a favorite shooting 
and snaring spot. A steep-sided rain pool on Gebel Umm Boanik 
was said to be a natural trap for thirsty ibex (Murray, 1912). Russell 
(1951) estimated a kill of about 100 at Bir Sheitun during one 
summer. 

Travelers of the past century in Egypt reported ibex from the 
bluffs overlooking the Nile eastward to all the plateaus and peaks of 
the Eastern Desert and Sinai Peninsula (fig. 162). Flower (1932) 
noted that, before completion of the railway between Luxor and 
Aswan, ibex came to the Nile for water. 

About the year 1900, Prince Kemal el Din established an ibex 
sanctuary with food and water in Wadi Rishrash (Halton, 1935). It 
was maintained for about 40 years. Russell (1949a) photographed 30 
or 40 ibex there in 1932. Latest reports are that a few ibex may still 
be seen in the area (Talbot, 1960). This reserve probably saved the 
ibex from annihilation in this part of the Eastern Desert. 



OSBORN&HELMY: MAMMALS OF EGYPT 521 

We know that ibex still exist in the more remote mountain peaks, 
but the necessity for water makes them vulnerable to human preda- 
tion throughout much of the year. Were it not for the security of the 
Wadis Qiseib and Abu Sanduq enforced by the Frontier Patrol 
stations there, the small herds inhabiting these areas would prob- 
ably have been eliminated long ago. 

Today, crudely made swords fitted with an ibex horn sheath are 
popular tourist items in the bazaars of Aswan. The horns are said to 
be brought from Gebel Elba by Bisharin tribesmen. 

Genus Ammotragus Blyth, 1840 

Monotypic genus of large sheep-like bovid with horns curving out- 
ward, backward, downward, and inward. Color nearly uniform red- 
dish or chestnut. Beard absent. Hair from jaw to upper foreleg long, 
mane-like. Tail long, nearly reaching hock. Lacrimal, tarsal, and 
subcaudal glands absent. 

Braincase not expanded. Frontonasal region flat. Infraorbital 
fossa lacking, ethmoidal fissure very small. Premaxilla contacting 
nasal. Horn core postorbital and extending laterally and posteriorly 
from base on same plane as frontals. 

Ammotragus lervia (Pallas, 1777) 

Antilope lervia Pallas, 1777, Spicilegia, Zool.. Vol. 12, p. 12. 

Type locality. — "V/estern Algeria, Department of Oran" (Harper, 
1940, p. 327). 

General distribution.— Egypt, southeastern Libya, Sudan, 
Tunisia, Morocco, Algeria, Mauretania. 

Common names.— Barhary Sheep, Maned Sheep, Aoudad, Kebsh 
el Gebel, Wadden, Ami. 

Subspecies in Egypt — 
Ammotragus lervia ornatus (I. Geoffroy St. Hilaire, 1827) 

Ouis ornata I. Geoffroy St. Hilaire, 1827, Diet. Class. Hist. Nat., Vol. 11, p. 264. 

Type locality.— Egypt. CAIRO: Near Cairo. 

Distribution in Egypt— Figure 164. Central part of Eastern 
Desert, central and southwestern parts of Western Desert. 

Diagnosis. — Reddish color with long mane from jaw to upper 
foreleg. Beard absent. Tail considerably longer than ear. Fronto- 
nasal region flat. Infraorbital fossa lacking. Premaxilla contacting 



522 



FIELDIANA: ZOOLOGY 



25* 26* 27* 28* 2 9* 30* 31* 32* 3 J* 34* 35* 36* 37 




Fk; 164. Collection localities of Ammotragus lervia ornatus. undated and prior to 
1932 (circles). 1932-1948 (dots), and 1949 to date (squares); and horns and skulls, 
date of death unknown (X): and sight records 1932 to date (S). 

nasal. Horns heavy, postorbital and curving outward, backward, 
downward, and inward. Shoulder height about 1 m. 

External characters.— General color reddish or chestnut, fore and 
outer part of legs brownish. Inside of ear, chin, venter, upper inside 
of legs, and upper part of foot whitish. Long mane on side of jaw, 
neck, chest, and upper foreleg darker than body and becoming skirt- 
like on forelegs. Beard absent. Muffle completely haired. Tail con- 
siderably longer than ear, reaching nearly to hock, with long hair on 
distal half. 

Cranial characters.— Skull triangular in lateral outline. Fronto- 
nasal contour flat. Braincase constricted laterally. Occipital con- 
dyles not protruding beyond level of supraoccipital. Frontoparietal 



OSBORN&HELMY: MAMMALS OF EGYPT 523 

suture broadly V-shaped. Interparietal suture broadly shield- 
shaped. Infraorbital fossa lacking, and ethmoidal fissure very small. 
Premaxilla contacting nasal. Nasofrontal suture forward of anterior 
margin of orbit. Postpalatal margin with narrow U-shaped cleft and 
apex of cleft reaching level of m . 

Horns and horn cores.— Horns heavy, with ventral keel; con- 
spicuous annulations from base to tip and curving outward, 
backward, downward, and inward; larger and more strongly 
annulated in male. Cores close together at base, almost pedicellate, 
postorbital, and on same plane as flat frontonasal region of skull. 
Base deeply perforated, shallow grooves and pits from base to tip. 
Cross-section eliptical. Interior hollow to tip and irregularly sub- 
divided by thick trabeculae. 

Teeth.— Labial ridges of upper cheek teeth, except pm\ well 
developed. M^ with accessory posterior vertical lingual ridge. 

Comparisons.— Ammotragus I. ornatus differs from other 
subspecies in having slightly darker color and absence of white 
subauricular patch and dark median facial marking. 

Specimens examined.— Total two. 

EL WADI EL GEDEED: Bir el Obeiyid NW of Farafara Oasis (severed head 
examined and photographed by L Helmy, February 1972), Ain Amur NW of Kharga 
Oasis (old weathered skull, date of death unknown). 

Published records.— The following is a listing of kills and observa- 
tions by date, wherever possible, from Wilkinson (1832), Sclater 
(1895), Anderson (1898, 1902), Buxton (1898), Barron and Hume 
(1902), Bedan (1928). Flower (1932), Shaw (1933), Mason (1936), 
Negumi (1952), Tregenza (1955, 1958), Murray (1967), Missone 
(1969. 1970), and Kock (1971). 

CAIRO: Cairo, hills east of (late 1700's); near Cairo (type). 

FAIYUM: Birket el Qarun, W of (1875); near EI Faiyum (1902). 

MINYA: Near El Minya (1861. 1868). rocky hills near El Minya (1902). 

QENA: Near Qena (1893). Thebes (1861. 1868). 

ASYUT: Manfalut (1827). 

RED SEA: Gebel Abraq (no date); Gebel Qattar (1823, 1892, 1893); Wadi Medisa 
(horns. 1893); Gemsa area (1893. 1910); Wadi Sceitun (1893); Ain Yassar (old horn, 
1893); Wadi el Gosa (Gossal) (1893, 1902); Wadi Seqel and Wadi Esserba (prior to 
1898); Ras Banas. Ras Gharib. and Qena-Quseir road (1902); Bir Abu Shaar (1910); 
Bir Abu Laseifa (1912); Wadi Tarfa (prior to 1920); Gebel Aradia (droppings. 1920); 
Bir Sheitun (1927); Wadi Asyuti (1927. 1934); Wadi Badia and Wadi Umm Sidri (old 
horns. 1949); plateau E of Asyut (reported by guides. 1949); Wadi Qena plateau 
(1951). 



524 FIELDIANA: ZOOLOGY 

ASWAN: El Derr (Diwan) and Aswan (1813). Wadi Sibaa (1860), Wadi Hor (no 
date). Koro8ko(1861). 

MATRUH: Qattara Depression near Minqar Abu Dweiss (old horns. 1927). 

EL WADI EL GEDEED: GUf el Kebir. Wadi Hamra (1935): Wadi el Malik (1933. 
1934). Gebel Uweinat (1923. 1925. 1932. 1933. 1934. 1935. 1969). Karkur Tahl (1934). 

Libya. SYRENAICA: Gebel Uweinat. Ain Dua (1933. 1943). Ain Zueia (two seen. 
1962). 

Sudan. NORTHERN: Semneh (herd of about 13 in 1890). Abu Hamed (1861. 
1913). 

Unpublished sight records.— 

RED SEA: Wadi Mellaha. 1963 or 1964 (personal observation of Dr. Hassan Sabr. 
former director of Giza Zoological Gardens); Wadi Asyuti tributary, March 1969 
(personal observation of Dr. Hani Zeny. Director of Nag Hamadi sugar company). 

EL WADI EL GEDEED: Near Ain DaUa on Guss Abu Said Plateau NW of 
Farafara Oasis. 1969 (personal observation of I. Helmy). and Ain Umm Dabadib 
(personal observation of Abd el Magid el Doghal. Governor of El Wadi el Gedeed). 

Habitats.— Rocky desert mountain and cliff areas. Descend into 
wadis and plains to feed. 

Behavior.— hike ibex, Barbary sheep are alert, shy, and extremely 
agile in rocky terrain. 

Food and water.— Ammotragus feeds on a variety of desert 
plants. Bedan (1928) observed one browsing on Tamarix sp. in Wadi 
Habeeb. Mason (1936) said it thrived on the bitter Colocynthis 
vulgaris (=Citrullus colocynthis) gourds in Wadi Hamra. Dorst 
(1970) mentioned Acacia sp. and Calotropis sp., and said they can 
obtain moisture from plants such as Rumex sp., but drink if water is 
available. Russell (1949b, p. 7) commented that "unlike ibex, sheep 
are not snared at waterholes because they do not drink." 

Historical notes.— Within historical time, Barbary sheep prob- 
ably inhabited most of the Eastern Desert and areas of rugged ter- 
rain in the Western Desert (fig. 164). In the Eastern Desert, inciden- 
tally, a well, a wadi, and a mountain are called Umm Kibash (mother 
of wild sheep). The type oiA. I. ornatus was shot "outside the gates 
of Cairo" (Rothschild, 1913, p. 459), and Barbary sheep were 
reported to have existed in the hills east of Cairo in the late 1700's 
(Anderson, 1898). Russell (1831) commented that sheep lived in the 
rocky deserts bordering the Nile, but did not occur habitually in the 
vicinity of Cairo. Numerous explorers since have observed Barbary 
sheep and their remains and published these, together with reports 
from guides. Many of these references pertain to Wadi Qena, Wadi 
Asyuti, and adjacent drainages in the Maaza Plateau. Flower (1932, 



OSBORN&HELMY: MAMMALS OF EGYPT 525 

p. 435) stated that, although the Barbary sheep was said to occur on 
both sides of the Nile in Upper Egypt during 1900-1909, by 1910, it 
had become "really scarce." Bedan (1928) killed a Barbary sheep in 
Wadi Asyuti in February 1927. He commented on the hunting 
pressure during World War I in the Wadi Asyuti area and said that 
a 1920 expedition had found no game. Some sheep, he thought, took 
refuge in an inaccessible cliff east of Wadi Asyuti on the west side of 
Wadi Qena. Russell (1949ab, 1951) recounted the decimation of Bar- 
bary sheep and ibex in the Wadi Qena-Wadi Asyuti country by com- 
mercial hunters, particularly during the war years when meat was 
scarce and expensive. He concluded that wild sheep no longer 
existed north of Gebel Elba. 

Of interest is the comment by Hoogstraal (1964, p. 237) that 
"Legends of wild sheep on Gebel Elba are rife among Bishareen, but 
we obtained no specimens. ' ' We do not know if sheep ever existed in 
the Elba mountains, although they were known to occur on Gebel 
Hisse (Isse or Is) 100 km. SW of Elba (Sclater, 1895). 

Recent observations of Barbary sheep in Wadi Asyuti and Wadi 
Mellaha in the Eastern Desert and Ain Dalla and Gebel Uweinat in 
the Western Desert (see above) indicate that small populations sur- 
vive in isolated areas. The most recent record is a specimen killed by 
a hunter in 1972 near Bir el Obeiyid NW of Farafara Oasis. Further 
indication of the former extent of distribution is the horns found in 
1927 in Qattara Depression near Minqar Abu Dweiss (Murray, 
1967). 

According to Zeuner (1963), Barbary sheep, unlike other native 
bovids, were never "domesticated." They were hunted and 
presented as offerings by the ancient Egyptians and are fairly com- 
mon in tomb and temple reliefs (Butzer, 1959), 



APPENDIX 1 

Explanation of abbreviations.— AbhTeviations used in text and 
tables are: N, north; S, south; E, east; W, west; mm,, miUimeter(s); 
cm., centimeter(s); m., meter(s); km., kilometer(s); wt., weight; gm., 
gram(s); kg., kilogram(s); C, centigrade; F., Fahrenheit; R.H., 
relative humidity. 

In tables, all measurements are given in mm. and all weights in 
gm., unless stated otherwise. Numbers after ranges are number of 
specimens. 

Abbreviations for measurements are: 

AL Alveolar length of upper tooth row (molar row). 

BCW Braincase width. 

BL Bullar length. The greatest horizontal distance from the anterior- 

most surface to the point of contact with the paroccipital process on 
the posterior-most surface of the right auditory bulla. 

BOW Basioccipital width. Least width of basioccipital bone between the 

auditory bullae. 

CBL Condylobasal length. Greatest distance between the anterior-most 

surface of premaxilla to posterior-most surface of occipital condyles. 

CIL Condyloincisive length. Greatest length between posterior margin of 

occipital condyles to anterior-most surface of incisors. 

CNL Condylonasal length. 

1 2 

C-M . C-M Distance from anterior-most surface of canine to posterior of first or 
second molar. 

EL Ear length from notch to tip. 

FL Foot length. Length of the hind foot including claw, unless stated 

otherwise. 

HBL Head and body length. Total length minus tail length. Not to be con- 

fused with term "body size" of Ranck (1968) and others who mean 
total length. 

HCW Horn core width. Greatest distance across the outside margins of the 

horn cores. 

HL Horn length. Taken along anterior surface of horn, unless stated 

otherwise. 

IFL Greatest length of the right incisive foramen. 

lOW Least interorbital width. 

526 



OSBORN&HELMY: MAMMALS OF EGYPT 527 

4 2 

I-PM , I-M Distance from anterior-most surface of incisors to posterior of fourth 

upper premolar or second upper molar. 

M - M Width across first upper molars. 

MW Mastoid width. 

NL Nasal length. Greatest length of nasal bones, unless stated otherwise. 

ONL Occipitonasal length (see SL). 

OW Orbital width. Greatest width across orbital bones. 

4 4 

P - P Width across fourth upper premolars. 

PAW Paroccipital width. 

PL Palatal length. Greatest distance from the anterior-most surface of 

the premaxilla to the posterior palatal margin. 

POW Postorbital width. 

PM Length of fourth upper premolar. 

PPF Greatest length of right posterior palatine foramen. 

PW Least pterygoid width. 

RW Rostral width. In carnivores, it is greatest width across alveoli of up- 

per canines; in rodents, it is greatest width anterior to the zygomatic 
plate. 

SH Skull height. Taken from highest point of skull to underside of a plate 

of known thickness upon which the skull rests with incisors or 
canines and bulla. Thickness of plate is then subtracted to give the 
measurement. 

SL Skull length. Greatest horizontal length of skull, including mastoid 

bullae in some cases. 

TL Tail length. Dorsal length of tail vertebrae from articulation with 

sacrum to tip of last tail vertebra. 

TL/HBL% Tail length divided by head and body length. Tail, head, and body 
length ratio in per cent. 

TRL Upper tooth row length (crown length). 

Wt Weight in grams, unless stated otherwise. 

ZW Greatest zygomatic width. 



APPENDIX 2 

The auditory bulla.— Terminology of chambers and related struc- 
tures of the middle ear (Dr. D. M. Lay, personal communication) 
used in taxonomic sections of the text are illustrated in Figure 165. 



Opposite: 

Fig. 165. — Auditory bulla of Dipodillus campestris. A, Posterolateral view of ex- 
terior showing chambers and associated structures. B, Same view enlarged, with 
walls partially removed to show partitions and relationships of chambers and 
semicircular canals. Arrows indicate communication between chambers. Numbered 
parts are: 

1 . Posterior arm of tympanic bone. 

2. Tympanic chamber. 

3. Manubrium of malleus. 

4. External auditory meatus. 

5. Lip of external auditory meatus. 

6. Anterior arm of tympanic bone. 

7. Incisura tympanicum. 

8. Hamular process of temporal bone. 

9. Suprameatal triangle. 

10. Supraoccipital. 

11. Anterior mastoid chamber. 

12. Subarcuate fossa. 

13. Lateral superior posterior mastoid chamber. 

14. Medial inferior posterior mastoid chamber. 

15. Occipital condyle. 

16. Lateral inferior posterior mastoid chamber. 

17. Paroccipital process. 

18. Anterior semicircular canal. 

19. Posterior semicircular canal. 

20. Lateral semicircular canal. 

528 



8 ' V " '2,3 




1 



Fig 165. Auditory bullae of Dipodillus campestris. 
529 



APPENDIX 3 

Tooth terminology. — Figure 166 shows the terminology used in 
the text for rodent molars. 



Opposite: 

Fk; 166. Terminology used in describing molars of Gerbillinae. 

530 



1^ 



E 



.<o 



o 




a 
a 







E 



E 



col 
E 



531 



APPENDIX 4 
The Govemorates of Egypt 

Figure 167 shows the govemorates of Egypt at the time the 
manuscript went to press. 



,. 25* 26* 27* 28* 2 9* 30* 31* 3 2* 3 3* 34* 35* 36* 37* 




Fui 167. Govemorates of Egypt. 



532 



APPENDIX 5 

GAZETTEER OF LOCALITIES MENTIONED IN THE TEXT 

Where possible, coordinates are from the United States Board on 
Geographic Names, 1959, Egypt and the Gaza Strip, Gazetteer No. 
45. 



Locality 


Governorate 


NLat. 


E Long. 






(°) 


(') 


(°) 


(') 


Abar el Dafa 


MATRUH 


31 


19 


26 


53 


Abassia 


CAIRO 


30 


04 


31 


17 


Abd el Mawla (Gebel) 


MATRUH 


30 


33 


29 


13 


Abnub (Ebnub) 


ASYUT 


27 


16 


31 


09 


Abu Aweigila (Augeila) 


SINAI 


30 


50 


34 


07 


Abu Darag 


SUEZ 


29 


29 


32 


27 


Abu Durba (Darba) Mine 


SINAI 


28 


29 


33 


20 


Abu el Matamir 


BEHEIRA 


30 


55 


30 


11 


Abu Gandir (Jandir) 


EL FAIYUM 


29 


14 


30 


41 


Abu Ghalib 


GIZA 


30 


16 


30 


54 


Abu Girgeh 


SINAI 




not found 




Abu Haggag 


MATRUH 


31 


08 


27 


50 


Abu Hammad 


SHARQIYA 


30 


32 


31 


40 


Abu Hommos (Hummus) 


BEHEIRA 


31 


06 


30 


19 


Abu Kharif Mine 


RED SEA 


26 


48 


33 


25 


Abu Kir (Abu Qir) 


ALEXANDRIA 


31 


19 


30 


04 


Abu Makkar Monastery 


see Deir Makaryus 


— 


— 


— 


— 


Abu Mena (Mina) 


MATRUH 


30 


51 


29 


40 


Abu Minqar 


EL WADI EL GEDEED 


26 


30 


27 


38 


Abu Ramad 


SUDAN ADMIN. 


22 


21 


36 


27 


Abu Rawash (Ruawash, Roash) 


GIZA 


30 


02 


31 


06 


Abu Sennan Dune 


MATRUH 


29 


33 


28 


53 


Abu Shuruf 


see Ain Abu Shuruf 


— 


— 


— 


— 


Abu Shusha 


QENA 


26 


10 


32 


01 


Abu Simbil (Simbel) 


ASWAN 


22 


22 


31 


38 


Abu Simbil West 


see Abu Simbil 


— 


— 


— 


— 


Abu Sir 


GIZA 


29 


53 


31 


13 


Abu Sir 


MATRUH 


30 


57 


29 


31 


Abu Sultan 


ISMAILIA 


30 


25 


32 


19 


Abu Zabal (Zaabal) 


QALYUBIYA 


30 


15 


31 


21 


Abu Zenima (Zeneima) 


SINAI 


29 


03 


33 


06 


Abydos 


QENA 


26 


11 


31 


55 



533 



534 



FIELDIANA: ZOOLOGY 



Locality 


Governorate 


NLat. 


E Long. 




>^ 


(°) 





C) 


(') 


Acacia grove 


GIZA 


28 


55 


29 


31 


Acacia grove 


MATRUH 


29 


13 


29 


05 


Acacia gfroves, Qattara Depression MATRUH 


29 


37 


27 


32 


Adindan 


ASWAN 


22 


12 


31 


30 


Aga Minshat el Ikhwa 


see Minshat el Ikhwa 


— 


— 


— 


— 


Ageeba 


MATRUH 


28 km. 


Wof 






Mersa Matruh 


Aghurmi 


MATRUH 


29 


15 


25 


20 


Aguz 


see El Aguz 


— 


— 


— 


— 


Ain Abu Nateigina (Nateiqina) 


SINAI 


29 


15 


33 


30 


Ain Abu Shuruf 


MATRUH 


29 


11 


25 


45 


Ain Amur 


EL WADI EL GEDEED 


25 


39 


30 


00 


Ain Beshai (Ibshai) 


EL WADI EL GEDEED 


27 


02 


27 


57 


Ain Dalla 


EL WADI EL GEDEED 


27 


19 


27 


20 


Ain Eede 


EL WADI EL GEDEED 




not found 




Ain el Baqar 


MATRUH 


29 


13 


25 


37 


Ain el Beilda 


GIZA 


5 km. W of Bawiti 


Ain el Dakrur 


MATRUH 




not found 




Ain el Furtaga 


SINAI 


29 


03 


34 


33 


Ain el Gedeirat (Ain Gudairat) 


SINAI 


30 


39 


34 


26 


Ain el Qht 


GIZA ] 


10-15 km. E of Bawiti 


Ain el Senned 


SINAI 




not found 




Ain el Tinnin 


EL WADI EL GEDEED 


26 


52 


27 


58 


Ain el Wadi 


EL WADI EL GEDEED 


28 


20 


29 


04 


Ain Gellaw 


EL WADI EL GEDEED 


15-20 km 


1. SE of 








EIQ 


asr 




Ain Ghabah 


GIZA 


2-3 km. N of El Aguz 


Ain Guffara 


GIZA 


28 


18 


28 


56 


Ain Marun 


GIZA 


28 


25 


28 


54 


Ain Musib Nabbut 


EL WADI EL GEDEED 


24 


31 


30 


39 


Ain Qureishit 


see El Zeitun 


— 


— 


— 


— 


Ain Ruweishid 


RED SEA 


26 


58 


33 


23 


Ain Shams 


CAIRO 


30 


08 


31 


19 


Ain Sudr 


SINAI 


29 


49 


33 


06 


Ain Sukhna (Sokhna) 


SUEZ 


29 


35 


32 


20 


Ain Taba 


SINAI 


29 


30 


34 


53 


Ain Umm Dabadib 


EL WADI EL GEDEED 


25 


46 


30 


25 


Ain Zeitun 


MATRUH 


29 


10 


25 


47 


Ain Zueia 


Libya. CYRENAICA 


21 


53 


24 


50 


Aiyut Barnasht 


GIZA 


29 


41 


31 


15 


Akhmim (Ekhmim) 


SOHAG 


26 


34 


31 


44 


Alam Shaltut 


MATRUH 


30 


46 


29 


50 


Alexandria llskanderiya, 


ALEXANDRIA 


31 


12 


29 


54 


Eskenderiya) 












Allaqi Village 


ASWAN 


23 


07 


32 


45 


Amada Temple 


ASWAN 


22 


43 


32 


15 


Ambogma 


see Umm Bugma 


— 


— 


— 


— 


Ambukol (Ambikol. Ambikul) 


Sudan. NORTHERN 


21 


19 


30 


53 



OSBORN & HELMY: MAMMALS OF EGYPT 



535 



Locality ' 


Governorate 


NLat. 


E Long. 






(°) 


(') 


(°) 


{') 


Antinoe (Antinopolis) 


see El Sheikh Ibada 


— 


— 


— 


— 


Armena (Armina) 


ASWAN 


22 


27 


31 


51 


Armina Temple 


ASWAN 


22 


25 


31 


47 


Asaa 


see El Qasr 


— 


— 


— 


— 


Ashmun el Ghunamiya 


MINUFIYA 


31 


18 


31 


44 


Asment 


EL WADI EL GEDEED 


25 


55 


29 


25 


Aswan (Assouan) 


ASWAN 


24 


05 


32 


53 


Aswan Dam Hospital 


see Aswan 


— 


— 


— 


— 


Aswan West 


see Aswan 


— 


— 


— 


— 


Asyut (Assiut, Siout) 


ASYUT 


27 


11 


31 


11 


Atf 


see El Atf 


— 


— 


— 


— 


Atfih 


GIZA 


29 


24 


31 


15 


Ausim (Awsim) 


GIZA 


30 


07 


31 


08 


Awlad Ali 


SINAI 


30 


52 


34 


04 


Awlad Hamza 


SOHAG 


26 


24 


31 


49 


Awlad Hawra 


GIZA 




not found 




Ayun Musa 


SINAI 


29 


52 


32 


39 


Bab el Sharia (Bab el Shaariya) 


see Cairo 


— 


— 


— 


— 


Badari 


see El Badari 


— 


— 


— 


— 


Bahariya Oasis 


see Bawiti 


— 


— 


— 


— 


Bahig 


MATRUH 


30 


56 


29 


35 


Bahrein 


MATRUH 


28 


40 


26 


32 


Bahr el Tubat 


Libya. CYRENAICA 


29 


36 


24 


53 


Balat 


EL WADI EL GEDEED 


25 


33 


29 


16 


Ballana 


ASWAN 


22 


16 


31 


34 


Baltim 


KAFR EL SHEIKH 


31 


33 


31 


05 


Bardia 


Libya. CYRENAICA 


31 


46 


25 


06 


Baris (Berys) 


EL WADI EL GEDEED 


24 


40 


30 


36 


Barqet Tokham 


ASWAN 


23 


35 


33 


25 


Bashtil 


GIZA 


30 


05 


31 


11 


Basus 


QALYUBIYA 


30 


08 


31 


13 


Batras 


EL WADI EL GEDEED 


5 km. W of Qasr el 








Farafara 




Bawiti 


GIZA 


28 


21 


28 


52 


Bayadeia 


see PORT SAID 


— 


— 


— 


— 


Beheira Nakhla 


BEHEIRA 




not found 




Beit el Wall Temple 


see Kalabsha 


— 


— 


— 


— 


Benha 


QALYUBIYA 


30 


28 


31 


11 


Beni Adi 


ASYUT 


27 


15 


30 


35 


Beni Magdul 


GIZA 


30 


02 


31 


07 


Beni Mazar 


MINYA 


28 


30 


30 


48 


Beni Salami (Ezbet Bint Salami) 


BEHEIRA 


30 


20 


30 


25 


Beni Salami (Salama) 


GIZA 


30 


19 


30 


51 


Beni Suef 


BENI SUEF 


29 


05 


31 


05 


Beni Yusef 


GIZA 




not found 




Bik 'at Hayareach 


see Has el Naqb 










Bilbeis 


SHARQIYA 


30 


25 


31 


34 


Bir Abbad 


RED SEA 


25 


02 


33 


04 



536 



FIELDIANA: ZOOLOGY 



Locality 

Bir Abd el Nabi 

Bir Abraq (Bir Abrag) 

Bir Abu Hussein 

Bir Abu Kharif 

Bir Abu Laseifa 

Bir Abu Sanduq 

Bir Abu Seyala (Bir Sayal, 

Ma Sweillim) 
Bir Abu Shaar 
Bir Abu Zawal 
Bir Akwamtra 
Bir Beida (Inglizi) 
Bir Bosslanga 
Bir Dakaar 
Bir Dibbis (Dibis) 
Bir el Aradj 
Bir el Hammamat 
Bir el Iseila 

Bir el Maghara 
Bir el Malla 
Bir el Nokta 

Bir el Obeiyid 

Bir el Qattara 

Bir el Qryeia 

Bir el Shab (El Sheb) 

Bir el Suweir (Suweira) 

Bir Fatira (Abu Kharif) 

Bir Ghadir 

Bir Gindali 

Bir Gumbiet 

Bir Haimur (Haimur wells) 

Bir Hassana 

Bir Hindusi 

Bir Hooker 

Bir Inglizi 

Bir Kansisrob 

Bir Karawein 

Birket el Sabh 

Birket el Qarun 

Bir Kibash 

Bir Kiseiba 

Bir Kurayim (Kreyim) 

Bir Kussaima 

Bir Lehfan 

Bir Meisa 

Bir Mellaha 



Governorate 


N Lat. 


E Long. 


MATRUH 


29 


59 


\ 1 
26 


58 


RED SEA 


23 


25 


34 


48 


EL WAD! ELGEDEED 


22 


53 


29 


55 


see Bir Fatira 


— 


— 


— 


— 


RED SEA 


26 


54 


32 


27 


SUEZ 


29 


25 


32 


31 


SUEZ 


29 


32 


32 


22 


RED SEA 


27 


22 


33 


37 


RED SEA 


26 


40 


33 


14 


SUDAN ADMIN. 


22 


13 


36 


18 


RED SEA 


26 


05 


34 


07 


see Bir Wair 


— 


— 


— 


— 


see Bir Samweil 


— 


— 


— 


— 


EL WADI EL GEDEED 


22 


09 


29 


27 


see El Areg 


— 


— 


— 


— 


RED SEA 


25 


58 


33 


33 


GIZA 


10-15 km 


. SWof 






\in Guffara 


SINAI 


30 


42 


33 


23 


MATRUH 


31 


10 


26 


01 


EL WADI EL GEDEED 


5 km 


1. W of Gharb el 






Mawhoub 




EL WADI EL GEDEED 


27 


19 


27 


40 


MATRUH 


31 


35 


25 


10 


RED SEA 


26 


22 


33 


01 


EL WADI EL GEDEED 


22 


19 


29 


46 


SINAI 


29 


15 


34 


43 


RED SEA 


26 


50 


33 


30 


RED SEA 


24 


48 


34 


47 


SUEZ 


29 


55 


31 


40 


RED SEA 


23 


21 


34 


47 


ASWAN 


22 


43 


33 


47 


SINAI 


30 


28 


33 


47 


RED SEA 


25 


49 


34 


11 


BEHEIRA 


30 


23 


30 


20 


see Bir Beida 


— 


— 


— 


— 


SUDAN ADMIN. 


22 


15 


36 


22 


EL WADI EL GEDEED 


27 


06 


28 


32 


MINUFIYA 


30 


38 


31 


05 


see Lake Qarun 


— 


— 


— 


— 


see Bir Umm Kibash 


— 


— 


— 


— 


EL WADI EL GEDEED 


22 


41 


29 


55 


EL WADI EL GEDEED 


22 


24 


29 


43 


see El Qoseima 


— 


— 


— 


— 


SINAI 


31 


01 


33 


52 


Sudan. KASSALA 


21 


21 


35 


35 


RED SEA 


27 


34 


33 


27 



OSBORN & HELMY: MAMMALS OF EGYPT 



537 



Locality 


Governorate 


NLat. 


E Long. 






n 


(') 


(°) (') 


Bir Mikheimin (Nahda and 


MATRUH 


30 


13 


28 52 


Nahid on topog. maps) 










Bir Murr 


EL WAD! EL GEDEED 


23 


21 


30 05 


Bir Murr (Farafara) 


EL WADI EL GEDEED 


27 


06 


28 32 


Bir Murra 


ASWAN 


22 


32 


33 54 


Bir Nagib (Nakeyb) 


ASWAN 


22 


50 


33 44 


Bir Nahed (Nahid) 


MATRUH 


4.5 km. E of 








El Maghra 


Bir Nakheila 


EL WADI EL GEDEED 


24 


01 


30 52 


Bir Number two 


EL WADI EL GEDEED 


4 km. N of Mut 


Birqash 


GIZA 


30 


10 


31 02 


Bir Qasr, Nos. 1, 2. 3 


GIZA see El Qasr 


— 


— 


— — 


Bir Qattar 


MATRUH 


31 


35 


25 10 


Bir Qattar (Kittar, Guttar) 


RED SEA 


27 


05 


33 17 


Bir Qiseib 


SUEZ 


29 


24 


32 29 


Bir Qokshira 


EL WADI EL GEDEED 




not found 


Bir Safsaf 


EL WADI EL GEDEED 


22 


44 


29 18 


Bir Samweil 


EL FAIYUM 


28 


53 


30 30 


Bir Sarrara 


SUDAN ADMIN. 


22 


16 


36 30 


Bir Semna 


RED SEA 


26 


27 


33 35 


Bir Seyala 


RED SEA 


26 


07 


33 56 


Bir Shafarzin 


MATRUH 


31 


19 


24 53 


Bir Shalatein 


SUDAN ADMIN. 


23 


08 


35 36 


Bir Shaqqa 


MATRUH 


30 


52 


24 59 


Bir Sheitun 


RED SEA 


26 


48 


32 07 


Bir Sidi Omar 


MATRUH 


31 


24 


24 52 


Bir Terfawi (Tarfawi) 


EL WADI EL GEDEED 


22 


55 


28 53 


Bir Thai 


SINAI 


29 


10 


33 04 


Bir Umm Delfa 


RED SEA 


27 


00 


33 34 


Bir Umm Dud 


RED SEA 


26 


58 


31 44 


Bir Umm Hibal 


ASWAN 


23 


42 


33 14 


Bir Umm Kibash 


RED SEA 


26 


55 


33 38 


Bir Umm Qareiyat 


ASWAN 


22 


33 


33 22 


Bir Victoria (El Qaraya) 


BEHEIRA 


30 


24 


30 37 


Bir Wair (Bosslanga) 


MATRUH 


31 


33 


25 05 


Bir Wigaba 


MATRUH 




not found 


Bir Zafarana 


RED SEA 


29 


07 


32 33 


Biyala 


KAFR EL SHEIKH 


31 


10 


31 13 


Bubastis 


SHARQIYA 


30 


34 


31 31 


Bulaq 


see Cairo 


— 


— 


— — 


Bulaq 


EL WADI EL GEDEED 


25 


12 


30 32 


Bulaq el Dakrur 


CAIRO 


30 


02 


31 11 


Buq Buq 


MATRUH 


31 


31 


25 34 


Burg el Arab 


MATRUH 


30 


55 


29 32 


Burullus District 


KAFR EL SHEIKH 


31 


35 


31 05 


Busili (Buseili) 


BEHEIRA 


31 


20 


30 24 


Cairo 


CAIRO 


30 


03 


31 15 


Cairo Citadel 


see Cairo 


— 


— 


— — 



538 



FIELDIANA: ZOOLOGY 



Locality 


Governorate 


NLat. 

(Ok f\ 


E Long. 

(°l CI 
30 17 


Camel Pass Dune 


MATRUH 


\ 1 

29 


50 


(Ghard el Qattaniya) 










Dahab (Dhahab) 


SINAI 


28 


29 


34 32 


Dahshur Pyramids 


GIZA 


29 


48 


31 12 


Dakhla Oasis 


see Mut 


— 


— 


— — 


Dakka 


ASWAN 


23 


12 


32 45 


Damanhour 


BEHEIRA 


31 


02 


30 28 


Damietta (Dumiat) 


DAMIETTA 


31 


25 


31 48 


Dandara (Dendera) 


QENA 


26 


10 


32 39 


Daraw (Derau) 


ASWAN 


24 


25 


32 56 


Deir Makaryus 


BEHEIRA 


30 


18 


30 29 


Dikheila Airfield 


see Alexandria 


— 


— 


— — 


Dilingat 


BEHEIRA 


30 


50 


30 30 


Dinshawi 


MINUFIYA 


30 


36 


30 51 


Dishna 


QENA 


26 


07 


32 28 


Disshet el Dabba (Dissht el Daba) RED SEA 


27 


04 


33 53 


Dist el Ashraf 


BEHEIRA 


30 


43 


30 39 


Durunka 


see Gebel Drunka 


— 


— 


— — 


Dush 


EL WADI EL GEDEED 


24 


34 


30 42 


El Abadiyah 


GIZA 


31 


22 


31 07 


El Abbasa 


SHARQIYA 


30 


32 


31 42 


El Afritat 


MATRUH 


17 km. 


SWof 






El Hammam 


El Aguz 


GIZA 


5 km 


1. SSE of Bawiti 


El Ahiah 


see Hurghada 


— 


— 


— — 


El Ahmar 


QALYUBIYA 




not found 


El Aiyat 


GIZA 


29 


37 


31 15 


El Alamein 


MATRUH 


30 


49 


28 57 


El Amiriya 


ALEXANDRIA 


31 


35 


31 01 


El Arbaein Monastery 


see Wadi el Arbaein 


— 


— 


— — 


El Areg (Arig. Arej. Aradj. 


Arag) MATRUH 


28 


56 


26 24 


El Arish 


SINAI 


31 


08 


33 48 


El Atarien 


see Alexandria 


— 


— 


— — 


El Atf 


GIZA 


29 


39 


31 16 


El Auberge 


see Shaksuk 


— 


— 


— — 


El Badari 


ASYUT 


26 


59 


31 25 


El Badrshein (Badrashein) 


GIZA 


29 


51 


31 16 


El Bahnasa (Behnessa) 


MINYA 


28 


32 


30 39 


El Bahrein 


see Bahrein 


— 


— 


— — 


El Bakhanis 


KAFR EL SHEIKH 


31 


11 


30 53 


El Ballah 


ISMAILIA 


30 


46 


32 19 


El Baradiah 


QALYUBIYA 


30 


14 


31 09 


El Baragil 


GIZA 


30 


04 


31 09 


El Barqil 


ELFAIYUM 




not found 


El Beida IBir Beida) 


BEHEIRA 


30 


27 


30 15 


El Birigat 


BEHEIRA 


30 


30 


30 50 


El Biyara 


ASWAN 




not found 


El Burg 


KAFR EL SHEIKH 


31 


35 


30 59 



OSBORN & HELMY: MAMMALS OF EGYPT 



539 



Locality 

El Daba (Dabah) 

El Dar el Bayda (Qasr Abbas I) 

El Deir 

El Deir el Beida 

El Dirr Temple 

El Faiyum 

El Ferden (Firdan) 

El Ferinat (Fureinat) 

ElGamil 

ElGau 

El Gezira 

El Ghaba el Qiblya 

El Ghazalat 

El Ghunamiya 

El Hamda (Hamtha) 

El Hammam 

El Hamra 

El Hamra 

El Hamul 

El Hara 

El Harraniya 

El Hawa 

ElHeiz 

EI Imayid 

El Kagug Cave 

El Kanayat 

El Kanayis 

El Khanafis 

El Khanka 

El Kharga 

El Khatatba 

El Kom el Ahmar 

El Koror 

El Kossaima 

El Kubri 

El Kunaiyisa (Kuneisa) 

El Kuntila 

El Kuraimat 

El Labban 

El Lisht Pyramid 

El Magedla 

El Maghra (Moghra) 

El Mahariq 

El Malfa (Ain Melfa) 



Governorate 

MATRUH 

SUEZ 

QENA 

see El Dar el Bayda 

ASWAN 

see Faiyum 

ISMAILIA 

MATRUH 

PORT SAID 

EL WADI EL GEDEED 

EL WADI EL GEDEED 
see El Aguz 
MATRUH 

MINUFIYA 

SINAI 

MATRUH 

BEHEIRA 

KAFR EL SHEIKH 

KAFR EL SHEIKH 

see Bawiti 

GIZA 

see Ilwat Hawa 

GIZA 

MATRUH 

ASWAN 

see El Qanayat 

RED SEA 

EL WADI EL GEDEED 

QALYUBIYA 

EL WADI EL GEDEED 

BEHEIRA 

GIZA 

see Aswan 

see El Quseima 

SUEZ 

GIZA 

SINAI 

GIZA 

see Alexandria 

GIZA 

see Beni Magdul 

MATRUH 

EL WADI EL GEDEED 

MATRUH 



N Lat. E Long. 

(°) (') (°) (') 

31 02 28 26 

30 08 31 51 

26 03 32 45 

22 44 32 15 

30 41 32 20 

30 14 29 15 

10 km. W of 

Port Said 

15 km. SW of 

Qasr el Farafara 

not found 

20 km. SW of 

Bir Abd el Nabi 

30 13 31 01 

30 55 33 52 

30 50 29 23 
not found 

31 10 30 52 
31 19 31 10 

29 58 31 10 

28 02 28 39 

30 47 29 12 

24 40 32 57 

25 00 33 19 
15-18 km. E of 

Qasr el Farafara 
30 13 31 21 
25 26 30 33 
30 23 30 50 
30 17 31 16 



30 02 32 33 

29 59 31 11 

30 00 34 41 
29 18 31 13 



29 34 



31 14 



30 15 28 55 
25 37 30 39 
29 45 24 50 



540 



FIELDIANA: ZOOLOGY 



Locality 



Mandara 

Manshiya 

Mansuriya 

Maragi (Maraqi) 

Maraziq 

Marg 

Mawhoub 

Mehalla el Kubra 

Mellaha 

Miharraqa 

Minya (Minia) 

Mishigeiga 

Mitimdiya 

Muhsib 

Muntazah 

Naqb el Abyad 

Naqb el Ahmar 

Nassariya 

Qanayat 

Qantara 

Qantara 

Qarasat 

Qaraya 

Qasr 

Qatta 

Quseima (Kossaima) 

Quweirat el Sud 

Raba (El Rabba) 

Rammak Dune 

Ras el Ahmar 

Saboua Temple (El Sibu) 

Saff 

Salhiya 

Saqiya (Saghee station) 

Shallufa 

Sheb 

Sheikh Ibada 

Sheikh el Waly 

Sibu 

Tabin 

Tahreer (Tahrir) 

Talbia (Talbiya) 

Taramsa 

Tarrana 

Wilidiya 

Zeitun 



Governorate 

ALEXANDRIA 

ALEXANDRIA 

GIZA 

MATRUH 

GIZA 

QALYUBIYA 

see Gharb el Mawhoub 

see Mehalla el Kubra 

RED SEA 

ASWAN 

MINYA 

EL FAIYUM 

GIZA 

see Ain Musib Nabbut 

ALEXANDRIA 

MATRUH 

MATRUH 

EL FAIYUM 

SHARQIYA 

SINAI 

ISMAILIA 

MATRUH 

see Bir Victoria 

GIZA 

GIZA 

SINAI 

MATRUH 

see Gebel El Rabbah 

MATRUH 

SINAI 

ASWAN 

GIZA 

SHARQIYA 

RED SEA 

SUEZ 

see Bir el Shab 

MINYA 

EL WADI EL GEDEED 

see El Saboua Temple 

GIZA 

TAHREER 

GIZA 

QENA 

BEHEIRA 

ASYUT 

MATRUH 



N Lat. E Long. 

(°) (') (°) (') 

31 13 30 41 

31 15 30 01 

30 08 31 05 

29 14 25 19 

29 49 31 16 

30 09 31 20 



28 12 33 10 

23 03 32 44 

28 06 30 45 

29 07 30 27 

30 03 31 10 

31 17 30 01 
29 27 26 20 
29 29 26 25 

29 21 30 41 

30 37 31 28 
30 51 32 19 
30 52 32 18 

5 km. SE of 
Abu Mena 

28 21 28 51 
30 13 30 58 
30 40 34 22 

20 km. SE of 

Qaret el Mashruka 

29 34 29 17 

30 59 33 48 
22 45 32 34 
29 34 31 18 

29 26 31 14 

26 44 32 53 

30 07 32 34 

27 48 30 52 
5 km. E of Mut 

29 47 31 18 

30 40 30 15 
30 46 30 52 

26 08 32 42 
30 26 30 50 

27 12 31 10 
29 10 25 47 



OSBORN & HELMY: MAMMALS OF EGYPT 



541 



Locality 


Governorate 


NLat. 


E Long. 






n 


(') 


n 


(') 


Ezbet Abu Zeid 


EL FAIYUM 




not found 




Ezbet Afifi Pasha 


GIZA 


30 


08 


31 


02 


Ezbet Ayub Ali 


EL FAIYUM 




not found 




Ezbet Beni Salami 


see Beni Salami 


— 


— 


— 


— 


Ezbet el Asfar 


EL FAIYUM 




not found 




Ezbet Ibhsan 


QALYUBIYA 




not found 




Ezbet Moneib 


see Giza 


— 


— 


— 


— 


Ezbet Muhsib 


see Ain Muhsib Nabbut 


— 


— 


— 


— 


Ezzeit (Marsa ez Zeitiya) 


RED SEA 


27 


50 


33 


35 


Faiyum (Feyum) 


EL FAIYUM 


29 


19 


30 


48 


Fani 


SUEZ 




not found 




Fanus 


EL FAIYUM 


29 


32 


30 


58 


Faqus 


SHARQIYA 


30 


44 


31 


48 


Farafara Oasis 


see Qasr el Farafara 


— 


— 


— 


— 


Faras 


Sudan. NORTHERN 


22 


10 


31 


27 


Faraskur (Fariskur) 


DAMIETTA 


31 


20 


31 


43 


Farshout (Farshut) 


QENA 


26 


03 


32 


09 


Fassulet Misaada 


MATRUH 


29 


27 


29 


12 


Fawakhir Mine 


RED SEA 


26 


01 


33 


36 


Fayid 


ISMAILIA 


30 


20 


32 


18 


Feiran Oasis 


SINAI 


28 


42 


33 


38 


Fersh Sheikh el Arab 


SINAI 




not found 




Fort Capuzzo 


Lybia. CYRENAICA 


31 


33 


25 


04 


Foum el Khalig (Old Cairo) 


see Cairo 


— 


— 


— 


— 


French Camp No. 2 


MATRUH 


29 


47 


27 


23 


Fuwa 


BEHEIRA 


31 


12 


30 


33 


Gabub (Qeigab) 


MATRUH 


29 


35 


24 


56 


Ganah 


EL WADI EL GEDEED 


25 


20 


30 


31 


Gara 


see Qara 


— 


— 


— 


— 


Gardaga 


see Hurgadah 


— 


— 


— 


— 


Gattah 


EL FAIYUM 




not found 




Gebel Abraq 


RED SEA 


23 


23 


34 


45 


Gebel Abu Dokhan (Dukhan) 


RED SEA 


27 


13 


33 


16 


Gebel Abu Harba 


RED SEA 


27 


17 


33 


13 


Gebel Abu Tiyur 


RED SEA 


25 


43 


34 


16 


Gebel Adda 


ASWAN 


22 


16 


31 


36 


Gebel Ain 


ASWAN 




not found 




Gebel Akheider 


RED SEA 


29 


44 


32 


11 


Gebel Ambish 


BENI SUEF 




not found 




Gebel Aradia 


RED SEA 


26 


20 


33 


29 


Gebel Asoteriba (Asotriba) 


Sudan. KASSALA 


21 


51 


36 


30 


Gebel Ataqa 


SUEZ 


29 


55 


32 


20 


Gebel Baba (el Babar) 


SINAI 


29 


16 


33 


43 


Gebel Catherine (Katrina) 


SINAI 


28 


31 


33 


57 


Gebel Deshesha (Dishasha) 


BENI SUEF 


28 


59 


30 


51 


Gebel Dhalfa (Dalfah) 


SINAI 


30 


45 


34 


12 


Gebel Dhulal (Dhalal, Dalai) 


SINAI 


28 


54 


33 


54 


Gebel Drunka (Durunka) 


ASYUT 


27 


07 


31 


10 



542 



FIELDIANA: ZOOLOGY 



Locality 


Governorate 


N Lat. 


E Long. 

<0| l'\ 


Gebel Egma 


SINAI 


29 


12 


\ 1 

34 


02 


Gebel el Ahmar 


CAIRO 


30 


03 


31 


18 


Gebel Elba 


SUDAN ADMIN. 


22 


11 


36 


21 


Gebel el Bruk (Buruk) 


SINAI 


30 


11 


33 


42 


Gebel el Ghigiga 


GIZA 


30 


01 


31 


02 


Gebel el Haridi 


SOHAG 


26 


47 


31 


55 


Gebel el Katamiya (Kutamiyal 


SUEZ 


29 


56 


31 


49 


Gebel el Rabbah 


SINAI 


30 


01 


33 


11 


Gebel el Silsila 


ASWAN 


24 


38 


32 


56 


Gebel el Themed (Yithmid) 


SINAI 


29 


42 


34 


23 


Gebel Faraid 


RED SEA 


23 


31 


35 


20 


Gebel Ferani 


SINAI 




not found 




Gebel Genawi 


SINAI 




not found 




Gebel Gharib 


RED SEA 


28 


07 


32 


54 


Gebel Gurdi 


see Ras Abu Gurdi 


— 


— 


- 


— 


Gebel Hamata 


RED SEA 


24 


12 


35 


00 


Gebel Hammami 


SINAI 


29 


12 


32 


58 


Gebel Hamra 


SINAI 


28 


35 


34 


30 


Gebel Hamra Dom 


SUDAN ADMIN. 


22 


39 


35 


39 


Gebel Hebron 


SINAI 


28 


33 


33 


37 


Gebel Hindus! 


RED SEA 


25 


51 


34 


14 


Gebel Hisse (Isse. Is) 


Sudan. KASSALA 


21 


55 


35 


29 


Gebel Hor (Har) 


SINAI 


29 


53 


32 


56 


Gebel Horeb 


SINAI 


W of Gebel Musa 


Gebel Hormadjan 


SINAI 




not found 




Gebel Iweibid (Oweibid, Awabed) 


SUEZ 


30 


06 


32 


09 


Gebel Katamiya (Kutamiya) 


SUEZ 


29 


56 


31 


49 


Gebel Lehfan (Lahfan) 


SINAI 


31 


01 


33 


52 


Gebel Magal Gabril 


ASWAN 


22 


53 


33 


36 


Gebel Maghara 


SINAI 


30 


42 


33 


23 


Gebel Mokattam 


CAIRO 


30 


02 


31 


17 


Gebel Muluk 


BEHEIRA 


31 


21 


30 


18 


Gebel Muqsim 


SUDAN ADMIN. 


22 


10 


34 


01 


Gebel Musa (Moosa) (Mount SinaijSINAI 


28 


32 


33 


59 


Gebel Naqud 


SUEZ 




not found 




Gebel Nesla 


SUDAN ADMIN. 


22 


15 


36 


16 


Gebel Nugrus 


RED SEA 


24 


49 


34 


36 


Gebel QatUr 


RED SEA 


27 


05 


33 


22 


Gebel Serbal 


SINAI 


28 


39 


33 


39 


Gebel Shallal (Shellal) 


SUDAN ADMIN. 


22 


01 


36 


31 


Gebel Shayeb (Shayeb el Banat) 


RED SEA 


26 


59 


33 


29 


Gebel Shindeib (Shendib) 


SUDAN ADMIN. 


22 


01 


36 


17 


Gebel Shubrawit 


ISMAILIA 


30 


17 


32 


17 


Gebel Sukhna 


see Khashm el Galala 


— 


— 


— 


— 


Gebel Tarbush 


SINAI 


28 


36 


33 


50 


Gebel Umm Afruth 


SINAI 


29 


10 


34 


15 


Gebel Umm Alawi 


SINAI 


28 


34 


34 


02 



OSBORN & HELMY: MAMMALS OF EGYPT 



543 



Locality 


Governorate 


NLat. 


E Long. 

IO\ l'\ 


Gebel Umm Boanik 


SINAK?) 


\ 1 


\ 1 \ 1 
not found 


\ 1 


Gebel Umm Disi 


RED SEA 


27 


02 


33 


15 


Gebel Umm Gidri 


RED SEA 


26 


58 


33 


36 


Gebel Umm Harba 


RED SEA 


23 


37 


34 


31 


Gebel Umm Kibash 


see Bir Umm Kibash 


— 


— 


— 


— 


Gebel Umm Rijlein 


SINAI 




not found 




Gebel Umm Shomer 


SINAI 


28 


22 


33 


55 


Gebel Umm Tenassib 


RED SEA 


28 


30 


32 


34 


Gebel Uweinat 


EL WADI EL GEDEED 


21 


54 


24 


58 


Gebel YeUeq (YiaUaq) 


SINAI 


30 


22 


33 


31 


Gebel Yithmid 


see Gebel el Themed 


— 


— 


— 


— 


Gebel Zabara 


SINAI 


24 


45 


34 


42 


Gemsa 


RED SEA 


27 


38 


33 


35 


Gezira Seud 


SHARQIYA 


30 


50 


32 


16 


Geziret Muhamed 


GIZA 


30 


07 


31 


12 


Gharah 


EL FAIYUM 


29 


08 


30 


42 


Gharb el Mawhoub 


EL WADI EL GEDEED 


25 


40 


28 


45 


Gharb el Qattaniya 


see Camel Pass Dune 


— 


— 


— 


— 


Ghardaqa 


see Hurghada 


— 


— 


— 


— 


Giarabub (Garabub) 


Libya. CYRENAICA 


29 


45 


24 


33 


GUf el Kebir 


EL WADI EL GEDEED 


23 


27 


26 


00 


Giza 


GIZA 


30 


01 


31 


13 


Giza Pyramids 


GIZA 


29 


59 


31 


08 


Giza Zoological Gardens 


see Giza 


— 


— 


— 


— 


Gizzaya 


see Giza 


— 


— 


— 


— 


Gobala 


GIZA 


13 km. E of Ba 


witi 


Guttar 


see Bir or Gebel Qattar 


— 


— 


— 


— 


Habou City Temple 


see Luxor 


— 


— 


— 


— 


Hafs 


BEHEIRA 


31 


00 


30 


20 


Haid Merzega Pass 


SINAI 




not found 




Haimur wells 


see Bir Haimur 


— 


— 


— 


— 


Halaib 


SUDAN ADMIN. 


22 


13 


36 


38 


Halfaya 


MATRUH 


31 


30 


25 


11 


Hammam Musa 


SINAI 


28 


17 


33 


55 


Hatiyet el Sheikh Marzuk 


EL WADI EL GEDEED 


26 


49 


27 


51 


Hatiyet el Sunt (Acacia grove) 


MINYA 


28 


26 


29 


42 


Hatiyet Labbaq 


MATRUH 


30 


20 


28 


36 


Hatiyet Tabany 


GIZA 




not found 




Hawamdiya (Hawamidiya) 


GIZA 


29 


54 


31 


15 


Heliopolis 


CAIRO 


30 


06 


31 


20 


Helwan 


CAIRO 


29 


51 


31 


20 


Herwer 


see El Sheikh Ibada 


— 


— 


— 


— 


Hibis Temple 


see El Kharga 


— 


— 


— 


— 


Hod Subeira (Abu Sobeiral 


ASWAN 


24 


13 


32 


53 


Hurghada (Ghardaqa) 


RED SEA 


27 


14 


33 


50 


Idfina 


BEHEIRA 


31 


18 


30 


31 


Idku 


BEHEIRA 


31 


18 


30 


18 


Idwa (Idwah) 


EL FAIYUM 


29 


19 


30 


52 



544 



FIELDIANA: ZOOLOGY 



Locality 


Govemorate 


N Lat. 


E Long. 




\^ 


D 


(') 


n 


(') 


Ilwat Hawa 


MATRUH 


30 


41 


29 


17 


Imbaba 


GIZA 


30 


04 


31 


13 


Ismailia 


ISMAILIA 


30 


35 


32 


16 


Isna lEsna) 


QENA 


25 


18 


32 


33 


Kafr Abu Sir 


QALYUBIYA 




not found 




Kafr Ammar 


GIZA 


29 


30 


31 


14 


Kafr Daoud (Dawud) 


BEHEIRA 


30 


28 


30 


49 


Kafr el Battikh 


DAM I ETTA 


31 


24 


31 


44 


Kafr el Dawar 


BEHEIRA 


31 


10 


30 


09 


Kafr el Sheikh 


KAFR EL SHEIKH 


31 


07 


30 


56 


Kafr el Shobak 


QALYUBIYA 


30 


17 


31 


19 


Kafr el Shurafa 


QALYUBIYA 


30 


22 


31 


20 


Kafret el Gebel 


GIZA 


29 


58 


31 


09 


Kafret Nassar 


GIZA 


30 


00 


31 


08 


Kafr Hakim 


GIZA 


30 


05 


31 


07 


Kafr Teharmes 


GIZA 


30 


01 


31 


11 


Kaiman el Matana 


QENA 


25 


49 


32 


43 


Kalabsha 


ASWAN 


23 


33 


32 


52 


Kalamsha 


EL FAIYUM 




not found 




Karawein 


EL WADI EL GEDEED 


27 


06 


28 


32 


Karkur Murr 


Sudan. NORTHERN 


21 


53 


25 


06 


Karkur Tahl 


EL WADI EL GEDEED 


22 


02 


25 


08 


Karmouz 


see Alexandria 


— 


— 


— 


— 


Karnak Temple 


see Luxor 


— 


— 


— 


— 


Kasr Rashwan 


see Qasr Rashwan 


— 


— 


— 


— 


Kasr Saghig 


see Zaghig 


— 


— 


— 


— 


Katamiya Canyon 


see Wadi el Katamiya 


— 


— 


— 


— 


Keneh 


see Qena 


— 


— 


— 


— 


Khabra Abu Guzour (Gazour) 


SINAI 


31 


00 


34 


20 


Khanka 


see El Khanka 


— 


— 


— 


— 


Kharga Oasis 


see El Kharga 


— 


— 


— 


— 


Khartoum 


Sudan. KHARTOUM 


15 


40 


32 


35 


Khashm el Galala 


SUEZ 


29 


34 


32 


20 


Khor Abusku 


ASWAN 


23 


13 


32 


52 


Khor Asot 


Sudan. KASSALA 


18 


18 


36 


10 


Khor el Madiq 


see Madiq 


— 


— 


— 


— 


Khor el Sil 


ASWAN 




not found 




Khor Musa Pasha 


Sudan. NORTHERN 


21 


51 


31 


16 


Khor Rhama el Bahari 


ASWAN 


23 


33 


32 


53 


Kirdasa 


GIZA 


30 


02 


31 


07 


Kittar 


see Bir or Gebel Qattar 


— 


— 


— 


— 


Kom Ashmun 


EL FAIYUM 


29 


18 


30 


58 


Kom Aushim 


see Kom Shim 


— 


— 


— 


- 


Kom el Hanash 


BEHEIRA 


30 


59 


30 


03 


Kom Hamada 


BEHEIRA 


30 


46 


30 


42 


Kom Ombo 


ASWAN 


24 


28 


32 


57 


Kom Ombo Temple 


see Kom Ombo 


— 


— 


— 


— 


Kom Shim (Kom Aushim) 


EL FAIYUM 


29 


34 


30 


55 



OSBORN & HELMY: MAMMALS OF EGYPT 



545 



Locality 


Governorate 


NLat. 


E Long. 


Kom Shim Forest 


see Kom Oshim 


\ 1 


\ 1 


\ 1 


\ ( 


Korosko 


ASWAN 


22 


36 


32 


20 


Kosseima 


see El Kosseima 


— 


— 


— 


— 


Kosseir 


see Quseir 


— 


— 


— 


— 


Kubra Abu Guzoar 


see Khabra Abu Guzour 


— 


— 


— 


— 


Kuneissa 


see El Kunaiyisa 


— 


— 


— 


— 


Kuntila 


SINAI 


30 


00 


34 


41 


Kurkur Oasis 


ASWAN 


23 


54 


32 


19 


Lake Burullus 


KAFR EL SHEIKH 


31 


30 


30 


50 


Lake Idku 


see Idku 


— 


— 


— 


— 


Lake Qarun (Kurun) 












(Birket Qarun) 


EL FAIYUM 


29 


28 


30 


40 


Lake Timsah 


ISMAILIA-SINAI 


30 


34 


32 


17 


Luxor 


QENA 


25 


41 


32 


39 


Maadi 


CAIRO 


29 


58 


31 


15 


Maatin el Garawla 


MATRUH 


31 


14 


27 


24 


Madiq 


ASWAN 


22 


44 


32 


09 


Maghagha 


MINYA 


28 


39 


30 


50 


Maidum 


BENI SUEF 


29 


22 


31 


10 


Mallawi (Mellawi) 


MINYA 


27 


44 


30 


50 


Mandisha 


MATRUH 


28 


21 


28 


55 


Manfalut (Monfalut) 


ASYUT 


27 


19 


30 


58 


Manshiyet Radwan 


GIZA 


30 


09 


31 


02 


Maqdabah (Magdaba) 


SINAI 


30 


53 


34 


02 


Maragi (Maraqi) 


MATRUH 


29 


14 


25 


19 


Marsaba 


MATRUH 




not found 




Maryiut (Mariut) 


ALEXANDRIA 


31 


01 


29 


48 


Mazraet el Gebel el Asfar 


QALYUBIYA 




not found 




Mehalla el Kubra 


GHARBIYA 


30 


58 


31 


10 


Mena (Mina) 


GIZA 


29 


58 


31 


08 


Mersa el Alem (Alam) 


RED SEA 


25 


04 


34 


54 


Mersa Matruh 


MATRUH 


31 


21 


27 


14 


Max 


ALEXANDRIA 


31 


09 


29 


51 


Mina el Basal 


see Alexandria 


— 


— 


— 


— 


Mina el Qamh 


SHARQIYA 


30 


30 


31 


15 


Minqar Abu Dweiss 


MATRUH 


30 


24 


28 


32 


Minqat Tukh 


see Tukh 


— 


— 


— 


— 


Minshat Beni Osman 


EL FAIYUM 




not found 




Minshat el Amir 












(Mohamed Ali Pasha) 


EL FAIYUM 




not found 




Minshat el Bakkari 


GIZA 


30 


01 


31 


08 


Minshat el Ikhwa 


DAQAHLIYA 


30 


56 


31 


21 


Minshat Tantawi 


EL FAIYUM 




not found 




Minuet el Sultan 












(Minyet el Sultan) 


GIZA 




not found 




Minuf 


MINUFIYA 


30 


28 


30 


56 


Misaada Dune 


MATRUH 


29 


25 


29 


05 


Mit Faris 


MINUFIYA 


30 


37 


31 


03 



546 



FIELDIANA: ZOOLOGY 



Locality 


Governorate 


N Ut. 


E Long. 

n (') 

31 16 


Mit Ghamr 


DAQAHLIYA 


1 \ 
30 


\ 1 
43 


Mit Riheina 


GIZA 


29 


51 


31 15 


Mohammed Ali Barrage Park 


MINUFIYA 


30 


12 


31 07 


Moiyet Luliya (Pearl's Spring) 


SINAI 




not found 


Mokattam Hills 


see Gebel Mokattam 


— 


— 


— — 


Monfalut (Monafalut) 


see Manfalut 


— 


— 


— — 


Mt. Sinai 


see Gebel Musa 


— 


— 


— — 


Mudrit el Tahreer 


see El Tahreer 


— 


— 


— — 


Muneiha 


ASWAN 


24 


29 


32 53 


Mut 


EL WADI EL GEDEED 


25 


29 


28 59 


Nadir 


MINUFIYA 


30 


33 


30 51 


Nag Ayed 


see Farshout 


— 


— 


— — 


Nag Farqanda West 


ASWAN 


22 


23 


31 45 


Nag Misaw 


ASWAN 


22 


22 


31 42 


Nahya 


GIZA 


30 


03 


31 07 


Naikhala 


ASWAN 




not found 


Nakhl (Nekhl) 


SINAI 


29 


55 


33 45 


Nakhlat el Barraq 


MATRUH 


30 


27 


29 32 


Naqb Abu Dweis 


MATRUH 


30 


27 


28 34 


Naqb Baraq (Barach) 


SINAI 




not found 


Nasser Village (Nasier) 


EL WADI EL GEDEED 


20 km. S of Kharga 


Nawa 


QALYUBIYA 


30 


14 


31 16 


Nefisha (Nafisha) 


ISMAILIA 


30 


34 


32 15 


Nohel 


SINAI 




not found 


Nubareia 


TAHREER 


30 


43 


30 46 


Nuweibah (Nuheibeh) 


SINAI 


28 


58 


34 39 


Nuweimisa 


MATRUH 


28 


42 


26 44 


Old Qena 


RED SEA 


ruins 20 km. S of 








El Saqiya 


Paris 


see Baris 


— 


— 


— — 


Port Said 


PORT SAID 


31 


16 


32 18 


Port Tawfik (Taufiq, Tewfiq) 


SUEZ 


29 


57 


32 34 


Qalama 


QALYUBIYA 


30 


13 


31 12 


Qalamsha(h) 


FAIYUM 


29 


10 


30 50 


Qalt Umm Disi 


RED SEA 


27 


00 


33 15 


Qalyub 


QALYUBIYA 


31 


11 


31 13 


Qanatir 


see Shibin el Qanatir 


— 


— 


— — 


Qara (Gara) 


MATRUH 


29 


37 


26 31 


Qaret el Ided (Idad) 


MATRUH 


29 


55 


28 54 


Qaret el Mashruka 


MATRUH 


30 


16 


29 32 


Qaret Sumara 


MATRUH 


30 


21 


29 05 


Qasr el Farafara 


EL WADI EL GEDEED 


27 


03 


27 58 


Qasr el Gebali 


EL FAIYUM 


29 


20 


30 38 


Qasr el Qatagi 


MATRUH 


30 


32 


29 39 


Qasr el Sagha 


EL FAIYUM 


29 


36 


30 40 


Qasr Qarun 


EL FAIYUM 


29 


25 


30 25 


Qasr Rashwan 


EL FAIYUM 


29 


30 


30 55 


Qasr Saghig 


see Zaghig 


- 


- 


- - 



OSBORN & HELMY: MAMMALS OF EGYPT 



547 



Locality 


Governorate 


NLat. 

f Ok 1 l\ 


E Long. 

1 o\ i f\ 


Qeigab 


see Gabug 


\ I 


\ ; 


\ 1 


( / 


Qena (Kenah, Kina, Qina) 


QENA 


26 


10 


32 


43 


Qokshira 


see Qasr el Farafara 


— 


— 


— 


— 


Quo Monastery 


SINAI 


28 


25 


34 


00 


Qur el Hilab 


MATRUH 


30 


21 


29 


17 


Quseir (Kosseir) 


RED SEA 


26 


06 


34 


17 


Qustul and Qustul west 


ASWAN 


22 


14 


31 


39 


Qusur el Banat 


RED SEA 


25 


55 


33 


16 


Quweisna (Quesna) 


MINUFIYA 


30 


34 


31 


09 


Rafa 


SINAI 


31 


17 


34 


14 


Ramleh 


ALEXANDRIA 


31 


14 


29 


58 


Raqabet el Halif 


MATRUH 


30 


44 


29 


40 


Raqabet el Rala 


MATRUH 


30 


21 


28 


52 


Ras Abu el Darag 


SUEZ 


29 


23 


32 


33 


Ras Abu Gurdi (Gebel Gurdi) 


RED SEA 


24 


00 


35 


05 


Ras Abu Rudeis 












(Rudeis-Sidri OUfield) 


SINAI 


28 


59 


33 


10 


Ras Banas 


RED SEA 


23 


54 


35 


48 


Ras el Bar 


DAM I ETTA 


31 


32 


31 


50 


Ras el Hekma (Hikma) 


MATRUH 


31 


20 


27 


50 


Ras el Ish (Esh) 


PORT SAID 


31 


08 


32 


18 


Ras el Kanayis 


see Ras el Hekma 


— 


— 


— 


— 


Ras el Naqb 


SINAI 


29 


36 


34 


51 


Ras el Sudar 


SINAI 


29 


36 


32 


40 


Ras Gharib 


RED SEA 


28 


21 


33 


06 


Ras Gurdi 


RED SEA 




not found 




Rashid 


see Rosetta 


— 


— 


— 


— 


Ras Jehan (Gihan) 


see Abu Durba Mine 


— 


— 


— 


— 


Ras Muhammed (Mohammad) 


SINAI 


27 


44 


34 


15 


Ras Zafarana (Zaafarana) 


RED SEA 


29 


07 


32 


39 


Redunkalil Tutuatee 


MATRUH 


29 


11 


25 


34 


Risan Aneiza (Risan Eineiza) 


SINAI 


30 


54 


33 


44 


Rishon el Zion 


ISRAEL 


31 


57 


34 


48 


Romani (Rumana) 


SINAI 


31 


00 


32 


40 


Rosetta (Rashid) 


BEHEIRA 


31 


24 


30 


25 


Royal Shooting Club 


see Kom Shim 


— 


— 


— 


— 


Safaga 


RED SEA 


26 


44 


33 


56 


Saft el Laban 


GIZA 


30 


02 


31 


10 


Saghee Station 


see El Sagiya 


— 


— 


— 


— 


Sakkara (Saqqara) 


GIZA 


29 


51 


31 


13 


Salum (Solium) 


MATRUH 


31 


34 


25 


09 


Samaket Gaballa 


MATRUH 


30 


28 


29 


05 


Samalut 


MINYA 


28 


18 


30 


42 


San el Haggar 


SHARQIYA 


30 


58 


31 


52 


Saqyet Abu Shara 


MINUFIYA 


30 


19 


31 


05 


Seket Meki (Saqyat Makki) 


GIZA 


30 


00 


31 


13 


Seila 


EL FAIYUM 


29 


21 


30 


58 


Seiyala (Seyala and Seiyala West) ASWAN 


22 


59 


32 


40 



548 



FIELDIANA: ZOOLOGY 



Locality 


Governorate 


N Lat. 


E Long. 


Shadwan Island 


RED SEA 


\ } 
27 


\ 1 
30 


\ 1 
33 


59 


Shakshuk 


EL FAIYUM 


29 


28 


30 


42 


Sharabiya 


see Cairo 


— 


— 


— 


— 


Shata 


DAM I ETTA 


31 


25 


31 


52 


Shatt el Mel 


DAM I ETTA 




not found 




Shalt Gheit el Nasara 


DAM I ETTA 


31 


24 


31 


49 


Sherbin (Shirbin) 


GHARBIYA 


31 


11 


31 


32 


Shibin el Qanatir 


QALYUBIYA 


30 


19 


31 


19 


Shubra Shihab 


QALYUBIYA 


30 


17 


31 


07 


Sidi Abd el Rahman 


MATRUH 


30 


58 


28 


44 


Sidi Barrani 


MATRUH 


31 


36 


25 


55 


Simbillawein 


DAQAHLIYA 


30 


53 


31 


32 


Sinnuris 


EL FAIYUM 


29 


25 


30 


52 


Sitra 


MATRUH 


28 


42 


26 


54 


Siwa and Siwa Oasis 


MATRUH 


29 


12 


25 


31 


Sohag 


SOHAG 


26 


33 


31 


42 


St. Anthony Monastery 


RED SEA 


28 


56 


32 


21 


St. Catherine Monastery 


SINAI 


28 


31 


33 


57 


St. Paul Monastery 


RED SEA 


28 


51 


32 


33 


Suez 


SUEZ 


29 


58 


32 


33 


Surarieh 


BENI SUEF 




not found 




Suweira 


SINAI 


29 


15 


34 


43 


Tahreer Forest 


EL FAIYUM 




not found 




Talbia 


GIZA 


30 


00 


31 


11 


Talha Station 


see Kom Hamada 


— 


— 


— 


— 


Talh el Fawakhir (Acacia Grove) 


MATRUH 


29 


45 


26 


38 


Tamiya 


EL FAIYUM 


29 


29 


30 


58 


Tanash 


GIZA 


30 


08 


31 


11 


Tanta 


GHARBIYA 


30 


47 


31 


00 


Tel Abu Ekaim (Akim) 


SHARQIYA 


30 


51 


32 


06 


Tel BasU 


SHARQIYA 


30 


34 


31 


31 


Tel el Amarna 


MINYA 


27 


39 


30 


58 


Tel el Kebir 


SHARQIYA 


30 


33 


31 


47 


Tellat Gimal 


SINAI 




not found 




Tel Khamis or Saad Khamis 


KAFR EL SHEIKH 




not 


found 




Thebes 


see Luxor 


— 


— 


— 


— 


Tor (El Tur) 


SINAI 


28 


14 


33 


37 


Toura (Tura) Cliffs 


CAIRO 


29 


56 


31 


19 


Tukh (Mingat Tukh) 


QALYUBIYA 


30 


50 


31 


06 


Umm Bugma 












(Umm Bogma. Ambogma) 


SINAI 


28 


59 


33 


21 


Umm Shilman Plains 


ASWAN 


22 


40 


33 


45 


Uyun Tablimun 


GIZA 


28 


02 


28 


44 


Valley of the Kings 


QENA 


25 


44 


32 


36 


Wadi Aad 


SINAI 




not found 




Wadi Abbad 


see Bir Abbad 


— 


— 


— 


— 


Wadi Abu Aweigila 


SINAI 


29 


20 


31 


29 


Wadi Abu Hor 


ASWAN 


23 


29 


32 


57 



OSBORN & HELMY: MAMMALS OF EGYPT 



549 



Locality 


Governorate 


N Lat. 


E Long. 






n 


(') 


(°) 


(') 


Wadi Abu Kaleja 












(KhaUfa. Haleifa) 


RED SEA 


26 


55 


31 


45 


Wadi Abu Quraiya 


RED SEA 


25 


16 


33 


59 


Wadi Abu Sanduq 


see Bir Abu Sanduq 


— 


— 


— 


— 


Wadi Abu Seyala 


see Bir Abu Seyala 


— 


— 


— 


— 


Wadi Abu Shaar 


see Bir Abu Shaar 


— 


— 


— 


— 


Wadi Abu Sheeh 


RED SEA 


26 


42 


33 


36 


Wadi Abusku 


see Khor Abusku 


— 


— 


— 


— 


Wadi Abu Subeira 


see Hod Subeira 


— 


— 


— 


— 


Wadi Abu Zawal 


see Bir Abu Zawal 


— 


— 


— 


— 


Wadi Abu Zeitouna (Zaitouna) 


SINAI 


28 


23 


33 


58 


Wadi Abu Ziran (Zeiran) 


RED SEA 


26 


09 


33 


58 


Wadi Adani 


SINAI 




not found 




Wadi Adeib 


SUDAN ADMIN. 


22 


15 


36 


26 


Wadi Ain el Gefeef 


SINAI 




not found 




Wadi Akwamtra 


see Bir Akwamtra 


— 


— 


— 


— 


Wadi Aleyat 


SINAI 


28 


41 


33 


41 


Wadi AUaqi 


see Allaqi Village 


— 


— 


— 


— 


Wadi Ambagi 


RED SEA 




not found 




Wadi Aqaba 


EL WADI EL GEDEED 


23 


28 


25 


48 


Wadi Araba 


see Bir Zafarana 


— 


— 


— 


— 


Wadi Araba 


SINAI 


28 


19 


33 


31 


Wadi Aruba 


SINAI 


28 


58 


34 


14 


Wadi Atallah 


RED SEA 


26 


03 


33 


36 


Wadi Bada 


SUEZ 


29 


43 


32 


16 


Wadi Bali 


RED SEA 


27 


27 


33 


39 


Wadi Bir el Abd 


SUEZ 


29 


32 


32 


22 


Wadi Daffeti (Defeit) 


SUDAN ADMIN. 


22 


13 


34 


11 


Wadi Dalma 


SINAI 




not found 




Wadi Darawena 


SUDAN ADMIN. 


22 


11 


36 


22 


Wadi Digla 


CAIRO 


29 


58 


31 


18 


Wadi Dihmit 


see Bir Umm Hibal 


— 


— 


— 


— 


Wadi Diib (Dib. Kiraf) 


SUDAN ADMIN. 


22 


28 


36 


06 


Wadi Dom (Doam, Doum) 


SUEZ 


29 


26 


32 


20 


Wadi el Abyad 


RED SEA 




not found 




Wadi el Arbaein (Arbain) 


SINAI 


28 


32 


33 


57 


Wadi el Arish 


see El Arish 


— 


— 


— 


— 


Wadi el Asyuti (Assiuti) 


ASYUT 


27 


10 


31 


16 


Wadi el Atrash 


RED SEA 


26 


39 


32 


46 


Wadi el Baharri (Bahharah) 


SUEZ 


30 


01 


32 


26 


Wadi el Deir 


RED SEA 


28 


54 


32 


40 


Wadi el Farigh (Faregh) 


BEHEIRA 


30 


13 


30 


20 


Wadi el Gafra 


SUEZ 


30 


24 


31 


36 


Wadi el Ghazal 


MATRUH 


south of Sidi Barrani 


Wadi el Gindali 


see Bir Gindali 


— 


— 


— 


— 


Wadi el Gosa (Gossal) 


RED SEA 


27 


10 


33 


05 


Wadi el Hammamat 


see Bir el Hammamat 


— 


— 


— 


— 



550 



FIELDIANA: ZOOLOGY 



Locality 

Wadi el Katamiya 

Wadi el Kharig (RhaUkit) 

Wadi el Laqeita 

Wadi el Melik (Abd el Malik) 

Wadi el Nasouri 

Wadi el Natroun (Natron, Natnin) 

Wadi el Nil 

Wadi el Qreiya (Qurrayyah) 

Wadi el Raba (Raha) 

Wadi el Ray an (Raiyan) 

Wadi el Rokham 

Wadi el Sheikh 

Wadi el Sheikh Isa 

Wadi el Targama (Targamy) 

Wadi Ergein (Ergayn) 

Wadi er Rimm 

Wadi Esserba (Heisurba) 

Wadi Eteigan 

Wadi Ethmiemat 

Wadi Fatira (Fatiri) 

Wadi Fatira el Zarka 

Wadi Feiran 

Wadi Fertili 

Wadi Figo 

(Fegar, Beint el Fegue) 
Wadi Gabgaba 
Wadi Garawi (Gerrawi) 
Wadi Gazzah (Ghazzah) 
Wadi Geba 
Wadi Gedeiret 
Wadi Gemal 
Wadi Ghadir 

Wadi Gharandal (Shurandel) 
Wadi Ghorabi 
Wadi Ghozah 

Wadi Ghuweibba (Ghweibba) 
Wadi Gindali 
Wadi Gossal 
Wadi Gray gar 
Wadi Gumbiet 
Wadi Gurdi 
Wadi Habib (Habeeb) 
Wadi Hagul 
Wadi Haimur Mine 
Wadi Haifa 
Wadi Hammad 
Wadi Hamra 
Wadi Hanjurat el Gattar 



Governorate 


N Lat. 


E Long. 


»^ 


n 


(') 


n 


CI 


SUEZ 


29 


58 


31 


49 


SINAI 


29 


03 


33 


22 


QENA 


25 


52 


33 


07 


EL WADI EL GEDEED 


23 


50 


25 


18 


SUEZ 


30 


10 


31 


29 


BEHEIRA 


30 


25 


30 


13 


RED SEA 


29 


20 


32 


35 


RED SEA 


26 


20 


32 


46 


SINAI 


29 


55 


32 


50 


EL FAIYUM 


29 


05 


30 


20 


SUEZ 


25 


22 


32 


21 


SINAI 


28 


44 


33 


50 


QENA 


26 


12 


32 


39 


ASWAN 


22 


55 


33 


10 


SINAI 




not found 




SINAI 




not found 




RED SEA 




not found 




SUDAN ADMIN. 


22 


07 


36 


03 


SINAI 




not found 




see Bir Fatira 


— 


— 


— 


— 


RED SEA 


26 


50 


33 


20 


SINAI 


28 


45 


33 


25 


RED SEA 


27 


08 


31 


40 


SUDAN ADMIN. 


22 


15 


35 


10 


ASWAN 


22 


37 


33 


17 


CAIRO 


29 


47 


31 


19 


SINAI 


31 


25 


34 


25 


SINAI 




not found 




see Ain Gedeirat 


— 


— 


— 


— 


RED SEA 


24 


40 


35 


06 


see Bir Ghadir 


— 


— 


— 


— 


SINAI 


29 


20 


33 


00 


GIZA 


28 


29 


29 


02 


RED SEA 


26 


59 


33 


12 


SUEZ 


29 


36 


32 


20 


see Bir Gindali 


— 


— 


— 


— 


see Wadi el Gosa 


— 


— 


— 


— 


RED SEA 




not found 




see Bir Gumbiet 


— 


— 


— 


— 


RED SEA 


26 


40 


32 


40 


RED SEA 


27 


11 


31 


46 


SUEZ .. 


29 


42 


32 


22 


ASWAN 


22 


36 


33 


17 


Sudan. NORTHERN 


21 


56 


31 


20 


RED SEA 


26 


48 


32 


46 


EL WADI EL GEDEED 


23 


50 


25 


26 


Israel 


30 


47 


34 


58 



OSBORN & HELM Y: MAMMALS OF EGYPT 



551 



Locality 

Wadi Hareidin (Hereidin) 

Wadi Hebron (Hibron) 

Wadi Hennis 

Wadi Hodein 

Wadi Hof (Hoaf) 

Wadi Hor (Or) 

Wadi I bib 

Wadi Iseili 

Wadi Isla (Isleh, IsUh) 

Wadi Kansisrob 

Wadi Kharit 

Wadi Kid (Kyd) 

Wadi Kiraf 

Wadi Kurkur 

Wadi Labaq 

Wadi Lahami (Lehema) 

Wadi Magal Gabril 

Wadi Markh (Merkh) 

Wadi Medisa 

Wadi Mellaha 

Wadi Midhais 

Wadi Mishigeiga 

Wadi Mitgal 

Wadi Muktil (Muqtil) 

Wadi Muwellih 

Wadi Naam 

Wadi Nagib 

Wadi Nakl (Nakhl) 

Wadi Naqud 

Wadi Nasb 

Wadi Nasli (Nesle) 

Wadi Nasouri 

Wadi Nasr 

Wadi Nassim 

Wadi Nekla 

Wadi Nogdeb (Nujdayb) 

Wadi Onib (Onibe) 

Wadi Qasab (Gusab) 

Wadi Qattar 

Wadi Qena 

Wadi Qiseib 

Wadi Quleib 

Wadi Rad Ayia 

Wadi Raha 

Wadi Rahaba 

Wadi Rished (Reshid) 

Wadi Rishrash (Rashrash) 

Wadi Saal 



Governorate 

SINAI 

SINAI 

EL WADI EL GEDEED 

RED SEA 

CAIRO 

ASWAN 

SUDAN ADMIN. 

SUEZ 

SINAI 

see Bir Kansisrob 

RED SEA 

SINAI 

see Wadi Diib 

ASWAN 

MATRUH 

RED SEA 

ASWAN 

RED SEA 

RED SEA 

see Bir Mellaha 

RED SEA 

EL FAIYUM 

RED SEA 

RED SEA 

see Bir Samweil 

RED SEA 

see Bir Nagib 

SUEZ 

RED SEA 

SINAI 

SINAI 

see Wadi el Nasouri 

SUEZ 

QENA 

MATRUH 

ASWAN 

Sudan. KASSALA 

RED SEA 

see Bir Qattar 

see Qena 

see Bir Qiseib 

ASWAN 

see Qusur el Banat 

SINAI 

SINAI 

SINAI 

RED SEA 

SINAI 



N Lat. E Long. 

(°) (') n {') 

30 59 33 53 

28 31 33 42 

27 24 28 16 
23 04 35 30 

29 53 31 18 
22 15 31 50 
22 50 35 46 

30 04 31 55 

28 08 33 43 



24 


26 


33 


03 


28 


07 


34 


30 


23 


56 


32 


35 


30 


20 


28 


32 


24 


13 


35 


25 


22 


53 


33 


36 


26 


21 


33 


03 


26 


55 


33 


10 



not found 

29 07 30 27 

not found 

24 25 34 00 

23 18 34 59 

29 32 32 22 
not found 

28 28 34 08 
not found 

29 31 32 23 

25 15 32 27 
not found 

23 15 33 07 

21 31 35 56 

26 19 32 02 



22 47 



33 12 



28 34 33 57 

28 25 34 00 
not found 

29 29 31 16 
28 45 34 27 



552 



FIELDIANA: ZOOLOGY 



Locality 



Wad 


1 Salafe (Solaf) 


Wad 


1 Saqi 


Wad 


1 Sceitun 


Wad 


iSaUkh 


Wad 


I Semna 


Wad 


iSeqel 


Wad 


1 Serimtai 


Wad 


1 Shait 


Wad 


Shawak 


Wad 


Sheitun (Scietun) 


Wad 


Sheger 


Wad 


Shellal 


Wad 


Sibaa 


Wad 


Sidr 


Wad 


Sigilliyeh 


Wad 


Sikait 


Wad 


Sukari 


Wad 


Taba 


Wad 


Tarfa 


Wad 


Threya 


Wad 


Timar (Thiman) 


Wad 


Tumilat 


Wad 


Umm Balad 


Wad 


Umm Delfa (Delfi) 


Wad 


Umm Dud 


Wad 


Umm el Seniyat 


Wad 


Umm Had 


Wad 


Umm Huweitat 


Wad 


Umm Qareiyat 


Wadi 


Umm Seleimat 


(Su 


laymat) 


Wad 


Umm Shedak 


Wad 


Umm Sidri 


Wad 


Umm Yassar 


Wad 


Wardan 


Wad 


Yesein 


Wad 


Yoider 


Wad 


Zeidun (Zaidun) 


Wad 


Zeidun 


Ware 


ian 


Warr 


aq el Arab 


Zaga 


zig 


Zagh 


ig (Qasr Saghig) 


Zahr 


el Rubin 


Zaiy< 


in Temple 


Zawj 


>^et Abu Musallam 


Zawj 


i^et el Mithniyan 


2^itii 


n 



Governorate 


N Lat. E Long 


SINAI 


\ 1 

28 


38 33 


47 


RED SEA 


26 


21 33 


52 


see Bir Sheitun 


— 


— — 


— 


SINAI 




not found 




see Bir Semna 


— 


— — 


— 


RED SEA 




not found 




SUDAN ADMIN. 


22 


12 36 


28 


RED SEA 


24 


33 33 


01 


RED SEA 




not found 




see Bir Sheitun 


— 


— — 


— 


SINAI 




not found 




SINAI 


28 


56 33 


18 


ASWAN 


22 


45 32 


34 


SINAI 


29 


39 32 


41 


SINAI 




not found 




RED SEA 


24 


40 34 


48 


RED SEA 


25 


03 34 


49 


SINAI 


29 


32 34 


52 


ASYUT 


28 


25 30 


50 


SINAI 




not found 




SINAI 


28 


15 33 


45 


SHARQIYA 


30 


31 31 


43 


RED SEA 




not found 




see Bir Umm Delfa 


— 


— — 


— 


see Bir Umm Dud 


— 


— — 


— 


RED SEA 




not found 




RED SEA 


26 


20 33 


23 


RED SEA 


26 


35 33 


57 


see Bir Umm Qareiyat 


— 


— — 


— 


RED SEA 


26 


16 32 


45 


MATRUH 




not found 




RED SEA 


27 


54 32 


33 


RED SEA 




not found 




SINAI 


29 


30 32 


43 


SUEZ 


29 


27 32 


28 


SUDAN ADMIN. 


22 


17 36 


18 


RED SEA 


25 


33 33 


04 


SINAI 




not found 




GIZA 


30 


19 30 


54 


GIZA 


30 


06 31 


12 


SHARQIYA 


30 


35 31 


31 


see Gebel Muluk 


— 


— — 


— 


SINAI 




not found 




EL WADI EL GEDEED 


25 


12 30 


32 


GIZA 


29 


56 31 


10 


MATRUH 


31 


30 26 


15 


see El Zeitun 


— 


— — 


— 



APPENDIX 6 

Definitions of terms used in the text — 

y4m.— Spring (plural Ayun). Sometimes applied to wells and cisterns. 

fiir.— Well. Sometimes applied to springs. 

£>ar6.— Camel road. 

Gait or Qa/t— Natural rock basin (plural Qulut) or hollow in rocks carved by water, 

together with gravel and stone. 
Gebel or JebeL— A hill or mountain or the desert. 

//atiyet. — Patches of vegetation in otherwise barren desert, with or without a well. 
Minqar.—A promontory or outstanding part of a cliff. 

Naqb.—A deep pass between cliffs from a plateau to low lands and vice versa. 
Oasis.— Vegetated area with natural occurring water, with or without cultivation. 
Qaret—A small hill, pile of boulders, or isolated rock formation. 
Qasr.— Literally, a palace. Used in reference to ruins. 
Qur.—A conical hill or rock pile. 

TaAt— Similar in meaning to Hatiyet, but the area would have larger trees. 
Wadi, Karkur, or Khor.—A gully, canyon, or valley; typically, a dry stream bed. 



553 



REFERENCES 

Abi) fl Rahman. A. A. and K. H. Batanouy 
1959. The phenology of the desert in relation to environment. Bull. Inst. Desert 

Egypte. 9. pp. 11-19. 
1966. Microclimatic conditions in Wadi Hoff. Bull. Soc. Geogr. Egypte, 39, pp. 
137-153. 
Abd kl Rahman. A. A. and M. N. el Hadidy 
1959. Some observations on the effect of wind on the desert vegetation along Suez 
road. Bull. Soc. G6ogr. Egypte, 32, pp. 207-216. 
Abu Al izz. M. A. 

1971. Landforms of Egypt. American University, Cairo, xv+281 pp. 
Adams. A. L. 
1870. Notes of a naturalist in the Nile Valley and Malta. Edmonston and Douglas, 
Edinburgh, xvi + 295 pp. 
Adolph. E. F. and associates 
1947. Physiology of man in the desert. Interscience Publ. Inc., New York, xiii-l-357 
pp. 
Aharoni. B. 

1932. Die Muriden von Palestina und Syrien. Z. Saugetierk.. 7, pp. 166-240. 
All F. M. 
1952. Outstanding variations in rainfall on the north coast of Egypt. Bull. Inst. 
Fouad 1 Desert Egypte, 2, pp. 5-6. 
Allen. G. M. 
1915. Mammals obtained by the Phillips Palestine Expedition. Bull. Mus. Comp. 

Zool., Harvard. 59. pp. 1-14. 
1939. A checklist of African mammals. Bull. Mus. Comp. Zool., Harvard, 83, pp. 
1-763. 
Almasy. L. E. de 
1936. R^entes exploration dans le Desert Libique (1932-1936). Publications 
Speciales de la Societe Royal de Geographie d 'Egypte. 97 pp. 
Alpinl p. 
1735. Historiae Aegypti naturalis. Vol. 1. Lungundi Batavonim, Apud Gerardium 
Potvliet. xvi-f 260 pp. 
Anderson. J. 
1892. Remarks on the occurrence of Spalax typhtus in Africa. Proc. Zool. Soc., 
London, 1892. pp. 472-476. 

1897. Exhibition of a coloured drawing of, and remarks upon, the Egyptian 
weasel. (Mustela subpalmata). Proc. Zool. Soc., London, 1897. pp. 600-601. 

1898. Zoology of Egypt. Vol. I. Reptilia and Batrachia. Bernard Quaritch. 
London, lxv-t-371 pp. 



564 



OSBORN&HELMY: MAMMALS OF EGYPT 555 

1902. Zoology of Egypt: Mammalia. (Revised and completed by W. E. De Winton). 
Hugh Rees Ltd., London, xvii+374 pp. 
Antonius. O. 
1937. On the geographical distribution, in former times and today, of the Recent 
Equidae. Proc. Zool. Soc., London, (ser. B) 107, pp. 557-564. 
ASCHERSON, P. F. A. 

1874. Exploration of the Libyan Desert. Card. Chron., (n.s.) 2, pp. 646, 647, 743. 
Atallah. S. I. 
1967. A new species of spiny mouse {Acomys) from Jordan. J. Mammal., 48, pp. 
258-261. 

AUDOUIN. V. 

1829. Description et sommaire des Mammiferes carnassiers. In Vol. II 
description de L'fegypte. De L'Imprimerie Imperiale, Paris, pp. 744-750. 
Bagnold. R. a. 

1931. Journeys in the Libyan Desert, 1929 and 1930. Geogr. J., 78, pp. 13-39, 
524-535. 

1933. A further journey through the Libyan Desert. Geogr. J., 82, pp. 103-129, 
211-235. 

1935. Libyan sands: Travel in a dead world. Hodder and Stoughton, London. 288 
pp. 

1936. The Libyan Desert. J. R. Afr. Soc, 35, pp. 294-305. 

1942. The principles of blown sand and desert dunes. Morrow, New York, xx+265 

pp. 
1954. The physical aspects of dry deserts. In Cloudsley-Thompson, J. L., ed., 

Biology of Deserts. Institute of Biology, London, pp. 7-12. 

Bahmanyar. M. and D. M. Lay 
1975. First record of GerbiUus henleyi De Winton, 1903, and records of other 
rodents from northern Yemen. Mammalia, 39, pp. 322-325. 
Ball. J. 
1912. The geography and geology of southeastern Egypt, Ministry of Finance, 

Survey Dept., Gov. Press, Cairo, xii-l-394 pp. 
1927. Problems of the Libyan Desert. Geogr. J., 70, pp. 21-38, 105-128, 209-224, 

512. 
1933. The Qattara Depression of the Libyan Desert and the possibility of its 

utilization for power production. Geogr. J., 82, pp. 289-314. 
1939. Contributions to the geography of Egypt. Ministry of Finance, Survey 
Dept., Cairo. 300 pp. 
Barron. T. 
1907a. The topography and geology of the Peninsula of Sinai (western portion). 

Nat. Print. Dept., Cairo. 241 pp. 
1907b. The topography and geology of the district between Cairo and Suez. Nat. 
Print. Dept., Cairo. 133 pp. 
Barron. T. and W. F. Hume 
1902. Topography and geology of the Eastern Desert of Egypt (central portion). 
Nat. Print. Dept., Cairo. 331 pp. 
Bate. D. M. A. 
1945. Notes on small mammals from the Lebanon mountains, Syria. Ann. Mag. 
Nat. Hist., (ser. 11), 12, pp. 141-158. 



556 FIELDI ANA: ZOOLOGY 

Bauer. K. 

1963. Ergebnisse der Zoologischen Nubien-Expedition 1962. Teil XIX, Saugetiere. 
Ann. Naturhistor. Mus. Wein, 66, pp. 495-506. 
Beadnkll. H. J. L. 

1910. The sand-dunes of the Libyan Desert. Geogr. J., 35, pp. 379-395. 

1931. Zerzura. Geogr. J.. 77, pp. 245-250. 
Bedan. -. 

1928. A desert expedition. Blackwood's Mag.. 223 (1350), pp. 519-548. 
Beecher. W. J. 

1969. Possible motion detection in the nuddle ear. Bull. Chicago Acad. Sci., 11, 
pp. 155-210. 
Belgrave. C. D. 
1923. Siwa, the oasis of Jupiter Ammon. John Lane the Bodley Head Ltd., 
London, xxix-l-275 pp. 
Bird. F. W. 

1946. The hyaena in Palestine. Bull. Jerusalem Nat. Club. 21, p. 1. 
Blaine. G. 

1913. On the relationship of Gazella Isabella to Gazella dorcas, with a description 
of a new species and subspecies. Ann. Mag. Nat. Hist., (ser. 8), 11, pp. 291-296. 

1914. An extinct hartebeeste from Egypt. Ann. Mag. Nat. Hist., (ser. 8), 13, pp. 
335-337. 

Bodenheimer. F. S. 

1935. Animal life in Palestine. L. Mayer, Jerusalem, xiii-l-506 pp. 

1958. The present taxonomic status of the terrestrial mammals of Palestine. Bull. 
Res. Council Israel, 7B (3-4), pp. 165-190. 
Bodenheimer. F. S. and O. Theodor 

1929. Ergebnisse der Sinai-expedition 1927. J. C. HinrichSche, Leipzig, viii+143 
pp. 

BONHOTE. J. L. 

1909. On a small collection of mammals from Egypt. Proc. Zool. Soc.. London. 
1909, pp. 788-798. 

1910. On the varieties of Mus rattus in Egypt. Proc. Zool. Soc., London, 1910, 
pp. 651-665. 

1912. On a further collection of mammals from Egypt and Sinai. Proc. Zool. Soc., 

London, 1912. pp. 224-231. 
BouLOS. L. 
1966a. A natural history study of Kurkur Oasis. Libyan Desert. Egypt. IV. The 

vegetation. Postilla, No. 100, pp. 1-22. 
1966b. Flora of the Nile region in Egyptian Nubia. Feddes Repert.. 73, pp. 184-215. 

1967. On the weed flora of Aswan, Egypt. Bot. Not., 120, pp. 368-372. 

1968. The discovery of Medemia Palm in the Nubian Desert of Egypt. Bot. Not., 
121, pp. 117-120. 

Bramley. W. E. J. 
1895. On Loder's gazelle in Egypt and the mode of capture. Proc. Zool. Soc., 
London, 1895, pp. 863-865. 

Brauer. a. 

1917. Neue Procaviiden. Sber. Ges. Naturf. Freunde Berlin. 1917, pp. 294-303. 
Brentjes. B. von 

1966. Einige Bemerkungen zur Darstellung der Hyanen. Erdwolfe und 
Hydnenhunde in den Kulturen des Alten orients. Z. Saugetierk.. 31, pp. 308-314. 



OSBORN&HELMY: MAMMALS OF EGYPT 557 

Briscoe. M. S. 
1956. Kinds and distribution of wUd rodents and their ectoparasites in Egypt. 
Amer. Midi. Nat., 55 (2). pp. 393-408. 
Brodoff. B. N. and G. Zeballos 

1970. Further studies on the effect of hypothalamic lesions in the sand rat 
iPsammomys obesus). Diabetologia, 6, pp. 366-370. 

Brodoff. B. N.. A. Kagan. B. Slotnik, and J. Hagedoorn 

1971. The effect of hypothalamic lesions in the sand rat maintained on a high fat 
diet. Unpublished manuscript. 

Brown. G. W. ed. 

1968. Desert biology. Academic Press. New York, xvii+635 pp. 
Bruce. J. 

1790. Select specimens of natural history collected in travels to discover the 
source of the Nile in Egypt, Arabia, Abyssinia, and Nubia, Vol. 6. Jackson, 
DubUn. xxiv + 286 pp. 
Brunk. R. and H. Strasser 

1967. Hamatologische Standardwerte bei der Sandratte (Psammomys obesus). 
Kleintierpraxis, 12, pp. 236-240. 

1969. Unterschiede im weissen Blutbild und in der Blutglukosekonzentration 
gefiitterter und gefasteter Stachelmailse {Acomys cahirinus, Desmarest, 1819). 
Berl. Munch. Tierarztl. Wochenschr., 82. pp. 93-96. 

Brunner. E. 

1969. Animals of ancient Egypt. Animals, 11, pp. 559-562. 
Budge. E. A. W. 
1926. The dwellers on the Nile. The Religious Tract Society, London, xxxii+326 
pp. 
Burckhardt. J. L. 

1819. Travels in Nubia. John Murray, London, cii+543 pp. 
Burns. T. W. 
1956. Endocrine factors in the water metaboUsm of the desert mammal G. 
gerbillus. Endocrinology. 58 (2). pp. 243-254. 
BUTZER. K. W. 

1959. Studien zum vor- und friihgeschichtlichen Landschaftswandel der Sahara. 
III. Die Naturlandschaft Agyptens wahrend der Vorgeschichte und der 
Dynatischen Zeit. Akad. Wiss. Literatur. Math. Naturw., No. 2. pp. 1-122. 

1965. Desert landforms at the Kurkur Oasis, Egypt. Ann. Assn. Amer. Geogr., 55, 
pp. 578-591. 

BuTZER. K. W. and C. L. Hansen 

1968. Desert and river in Nubia. Univ. Wise. Press, Madison, xxi + 562 pp. 
BuTZER. K. W. and C. R. Twidale 

1966. Deserts in the past. In Hills, E. S. ed.. Arid Lands, Methuen & Co., Ltd.. 
London, pp. 127-144. 

Buxton. E. N. 

1898. Short stalks. Edward Stanford, London. Second series, xi+226 pp. 
Buxton. H. M. B.. C. E. B. Buxton, and T. B. Buxton 

1895. On either side of the Red Sea. Edward Stanford. London, viii+163 pp. 
Buxton. P. A. 

1923. Animal life in deserts. Edward Arnold Ltd.. London. xv+ 176 pp. 



668 FIELDIANA: ZOOLOGY 

Cailliaud. M. F. 

1826. Voyage k M^ro^ et au Fleuve Blanc, Vol. 1. A L'Imprimerie Royal, Paris. 
XV + 429 pp. 
Carlislk. D. B. and L. 1. Ghobriai- 

1968. Food and water requirements of dorcas gazelle in the Sudan. Mammalia, 32, 
pp. 570-576. 
Caslick. J. W. 

1956. Color phases of the roof rat. Rattus rattus. J. Mammal., 32. pp. 255-257. 
Caton Thompson. G. and E. W. Gardnkr 

1932. The prehistoric geography of Kharga Oasis. Geogr. J., 80, pp. 369-409. 
Chavvorth Musters. J. L. and J. R. Ellerman 

1947. A revision of the genus Meriones. Proc. Zool. Soc., London. 117, pp. 478-504. 
Churcher. C. S. 

1972. Late Pleistocene vertebrates from archaeological sites in the plain of Kom 
Ombo. Upper Egypt. Life Sci. Contr., R. Ontario Mus., No. 82, 172 pp. 

Cloudsley-Thompson. J. L., ed. 

1954. Biology of deserts. Institute of Biology, London. 224 p. 
Cloudsley Thompson. J. L. and M. J. Chadwick 

1964. Life in deserts. G. T. Foulis & Co. Ltd.. London, xvi+218 pp. 
Cloudsley-Thompson. J. L. and L. Ghobrial 

1965. Water economy of the dorcas gazelle. Nature. 207, p. 1313. 
Cockrum. E. L.. T. C. Vaughan. and P. J. Vaughan 

1976a. A review of North African short-tailed gerbils (Dipodillus) with description 

of a new taxon from Tunisia. Mammalia. 40. pp. 313-326. 
1976b. Gerbillus andersoni De Winton. a species new to Tunisia. Mammalia, 40. 

pp. 467-473. 
Corbet. G. B. 

1966. The terrestrial manunals of Western Europe, Dufour. Philadelphia, xi+264 
pp. 

Couturier. M. A. M. 

1962. Le Bouqetin des Alpes. Couturier. Grenoble. xi+ 1.564 pp. 
Daly. M. and S. Daly 

1973. On the feeding ecology of Psammomys obesus (Rodentia: Gerbillidae) in 
Wadi Saoura. Algeria. Mammalia, 37, pp. 545-591. 

1974. Spatial distribution of a leaf-eating Saharan gerbil {Psammomys obesus) in 
relation to its food. Mammalia, 38, pp. 591-603. 

1975a. Socio-ecology of Saharan gerbils, especially Meriones libycus. Mammalia, 

39, pp. 289-311. 
1975b. Behavior of Psammomys obesus (Rodentia: Gerbillinae) in the Algerian 
Sahara. Z. Tierpsychol., 37, pp. 298-321. 
Davis. P. H. 

1953. The vegetation of the deserts near Cairo. J. Ecol., 41 (1), pp. 157173. 
Dawson. W. R. 
1924. The mouse in Egyptian and later medicine. J. Egypt. Archaeol.. 10, pp. 
83-86. 
De Beaux. O. 
1928. Risultati zoologici della missione inviata dalla R. Societa Geographica 
Italiana per I'esplorazione dell' oasis di Giarabub. (1926-1927). Mammiferi. Ann. 
Mus. Civ. Stor. Nat. Genova, 53, pp. 39-76. 



OSBORN&HELMY: MAMMALS OF EGYPT 559 

1932. Spedizione scientifica all' oasis di Cufra. Ann. Mus. Civ. Stor. Nat. Geneva, 
55. pp. 374-395. 
De Blase. A. 
1972. Rhinolophus euryale and R.,mehelyi (Chiroptera, Rhinolophidae) in Egypt 
and southwest Asia. Israel J. Zool., 21, pp. 1-12. 
De Blase. A. and R. E. Martin 

1974. A manual of mammalogy. Wm. C. Brown, Dubuque, xv + 329 pp. 
De Fronzo. R., E. Mikl and J. Steinke 
1967. Diabetic syndrome in sand rats. III. Observations on adipose tissue and 
liver in the non-diabetic stage. Diabetologia, 3, pp. 140-142. 
Dekeyser. p. L. 

1955. Les Mammif6res de L'Afrique Noire Fran^aise. Institut Fran^ais D'Afrique 
Noire, Dakar. 426 pp. 

De Winton. W. E. 

1898. Felis chaus and its allies, with descriptions of new subspecies. Ann. Mag. 

Nat. Hist., (ser. 7). 2. pp. 291-294. 
1902a. Descriptions of two new gerbils from Egypt. Ann. Mag. Nat. Hist., (ser. 7), 

9, pp. 45-47. 
1902b. Descriptions of two new hares from Egypt. Nov. Zool, 9, pp. 444-445. 
1903. List of mammals obtained by the Hon. N. Charles Rothschild and the Hon. 

Francis R. Henley in the Natroun Valley. Egypt. Nov. Zool., 10, pp. 279-285. 

DiDIER. R. 

1946. Etude systematique de 16s penien des mammif^res. Mammalia, 10, pp. 78-91. 

1947. Etude systematique de 16s penien des mammif^res. Mammalia, 11, pp. 
139-152. 

1948. Etude systematique de 16s penien des mammif^res. Mammalia. 12, pp. 67-93. 

1949. Etude systematique de 16s penien des mammif^res. Mammalia, 13, pp. 17-37. 

1953. Etude systematique de 16s penien des mammif^res. Mammalia, 17, pp. 
260-269. 

1954. Etude systematique de 16s penien des mammif^res. Rongeurs. Famille des 
Murides. Mammalia. 18, pp. 237-256. 

1956. Etude systematique de 16s penien des mammif^res. Mammalia, 20, pp. 
238-247. 

Dor. M. 
1947. Observations sur les micromammif^res trouv6s dans les pelotes de la 
Chouette effraye (Tyto alba) en Palestine. Mammalia, 11, pp. 50-54. 
DORST, J. 
1970. A field guide to the larger mammals of Africa. Hougton Mifflin, Boston. 
287 pp. 
Doughty. C. M. 

1888. Travels in Arabia Deserta. Jonathon Cape. London. XIX +690 pp. 
Drake Brockman. R. E. 
1910. The mammals of Somaliland. Hurst and Blackett. Ltd.. London, xvii+201 
pp. 
Drar. M. 

1955. Egypt, Eritrea. Libya and the Sudan. Arid Zone Research VI. Plant 
Ecology. UNESCO. Paris, pp. 151-194. 

DUMREICHER. A. VON 

1931. Trackers and smugglers in the deserts of Egypt. Methuen and Co. Ltd.. 
London, xii+248 pp. 



560 FIELDIANA: ZOOLOGY 

Dunbar. J. H. 
1941. The rock pictures of Lower Nubia. Service (^es Antiquit^s de L'Egypte, 
Cairo. x+lOO pp. 
Ebers. G. 
1878. Egypt: Descriptive, historical, and picturesque, Vol. 2. Cassell, Peter, 
Galpin and Co.. London, Paris. New York, xxii + 388 pp. 
Edney. E. B. 
1966. Animals of the desert. In Hills, E. S., ed., Arid Lands. Methuen and Co. 
Ltd.. London, pp. 181-215. 

ElSENTRAUT. M. 

1969. Das Gaumenfaltenmuster bei westafrikanischen Muriden. Zool. Jahrb. 
Syst. Bd.. 96 (478). pp. 478-490. 
ElDanasori. G. 
1957. The weather of Egypt. In Studies in Geography of Egypt (In Arabic), 
section 5. Misr Book Shop. Cairo, pp. 142-187. 
El Hadidi. M. N. and S. Ghabbour 
1968. A floristic study of the Nile Valley at Aswan. Rev. Zool. Bot. Afr.. 78, pp. 
394-407. 
Ellerman. J. R. 

1940. The families and genera of living rodents. Vol. \. British Mus. (Nat. Hist.). 
London, xxvi+689 pp. 

1941. The families and genera of living rodents. Vol. II. Brit. Mus. (Nat. Hist.). 
London, xii+690 pp. 

1948. Rodents of south-west Asia. Proc. Zool. Soc.. London. 118, pp. 765-816. 

1949. The families and genera of living rodents. Vol. III. pt. 1. Brit. Mus. (Nat. 
Hist.). London. 210 pp. 

Ellerman. J. R. and T. C. S. Morrison Scott 
1951. Checklist of Palaearctic and Indian mammals 1758 to 1946. Brit. Mus. 
(Nat. Hist.). London. 810 pp. 
Ellerman. J. R.. T. C. S. Morrison-Scott, and R. W. Hayman 

1953. Southern African mammals. Brit. Mus. (Nat. Hist.). London. 363 pp. 
Ewer. R. F. 

1954. Some adaptive features in the dentition of hyenas. Ann. Mag. Nat. Hist., 
(ser. 12). 7, pp. 188-194. 

Pagan. B. M. 

1975. The rape of the Nile. Charles Scribner's Sons. New York, xiv + 399 pp. 
Fahmy. I. R. 

1936. Report on Gebel Elba. Egypt. Univ. Press, Cairo. 58 pp. 
Fiedler. U. 
1973. Beobachtungen zur Biologie einiger Gerbillinen, insbesondere GerbiUus 
(Dipodillus) dasyurus (Myomorpha, Rodentia) in Gefangenschaft. Z. Saugetierk., 
38. pp. 321-340. 

FiTZINGER. L. J. 

1860. Naturgeschichte der Saugethiere. Vol. I. Aus der kaiserlichkbniglichen 

Hof- und Staatsdruckeri. Vienna. 487 pp. 
FiTZINGER. L. J. and Th Heuglin von 
1866. Systematische ubersicht der Saugethiere Nordost-Afrika's mit Einschluss 

der arabische Kiiste, des rothen Meeres, der Somal- und der Nilquellen-Lander. 

sudwarts bis zum vierten Grade nordlicher Breite. Sber. Akad. Wiss. Wien, 

54, pp. 537-611. 



OSBORN&HELMY: MAMMALS OF EGYPT 561 

Flower. S. S. 

1931. Contributions to our knowledge of the duration of life in vertebrate 
animals. Proc. Zool. Soc., London. 1931, pp. 145-234. 

1932. Notes on the recent mammals of Egypt, with a list of the species recorded 
from that kingdom. Proc. Zool. Soc., London, 1932, pp. 368-450. 

Floyer. E. a. 
1887. Notes on a sketch map of two routes in the Eastern Desert of Egypt. Proc. 

R. Geogr. Soc., London, 9, pp. 659-681. 
1893. Further routes in the Eastern Desert of Egypt. Geogr. J., 1 (5), pp. 408-431. 
Forskal, p. 

1775. Descriptions animalium. Molleri, Hauniae. xxxiv+164 pp. 
Gabr, M. F. a. and A. A. Shalaby 
1962. Glucose content of the blood of the desert mammal Jaculus jaculus in 

relation to its body water balance. Bull. 2k)ol. Soc. Egypt, 17, pp. 15-20. 
1964. Studies on the concentration of the total electrolytes and some organic ions 
in the urine of the desert mammal Jaculus jaculus, in relation to its water 
economy. Bull. Zool. Soc. Egypt, 19, pp. 45-59. 
Gaillard. C. and G. Daressy 
1905. La faune momifiee de I'antique Egypte. De L'Institut Francais 
D'Archeologie Orientale, Le Caire. 159 pp. 
Gaillard. M. C. 
1927. Les animaux consacres a'la divinite I'ancienne Lycopolis. Egypt. Serv. 
Antiques, 27, pp. 33-42. 
Gaisler. J., G. Madkour. and J. PelikAn 

1972. On the bats (Chiroptera) of Egypt. Acta Sc. Nat. Brno, 6, pp. 1-40. 

1973. On the bats of Egypt. Prirodoved. Pr. Ustavu Cesk. Akad. Ved. Brno, 6, 
pp. 1-40. 

Gauthier-Pilters, H. 

1967. The fennec. Afr. Wildl. 21, pp. 117-125. 
Gautier, E. F. 

1935. Sahara the great desert. Columbia Univ., New York, xviii+264 pp. 
Gentry, A. W. 
1964. Skull characters of African gazelles. Ann. Mag. Nat. Hist., (ser. 13), 7, pp. 
353-382. 
Geoffroy Saint-Hilaire. E. 
1818. Description de I'Egypte. Histoire naturelle, description des mammiferes qui 
se trouvent en Egypte, Vol. II. Paris, pp. 94-144. 
Geoffroy Saint-Hilaire. E. and V. Audouin 
1829. Mammiferes. In Vol. II of 'Description de I'Egypte. De L'Imprimerie 
Imp^riale, Paris, pp. 733-743. 
Geoffroy Saint-Hilaire, I. 
1827. Memoire sur quelques espdces nouvelles ou peu connues du genre 

Musaraigne. Mem. Mus. Hist. Nat. Paris, 15, pp. 117-144. 
1847. Vie, travaux et doctrine scientifique d' fitienne Geoffroy Saint-Hilaire, 
Librarie Society Geologique de France. 479 pp. 
Ghobrial. L. I. 
1976. Observations on the intake of sea water by the dorcas gazelle. Mammalia, 
40, pp. 489-494. 
Ghobrial, L. I. and J. L. CloudsleyThompson 
1966. Effect of deprivation of water on the dorcas gazelle. Nature, 212, p. 306. 



562 FIELDIANA: ZOOLOGY 

GiDLEY. J. W. 

1914. Preliminary report on a recently discovered Pleistocene cave deposit near 
Cumberland. Maryland. Proc. U.S. Nat. Mus.. 46, pp. 93-102. 
GlEGENtJACK. R. F., Jr. 

1968. Late- Pleistocene history of the Nile Valley in Nubia (unpublished Doctoral 
Dissertation. Yale Univ.). 

Gray. J. E. 

1868. Revision of the species of Hyrax, founded on specimens in the British 
Museum. Ann. Mag. Nat. Hist., (ser. 4). 1, pp. 35-41. 
Groves, C. P. and D. L. Harrison 
1967. The taxonomy of the gazelles (genus Gazella) of Arabia. J. Zool., London, 152, 

pp. 381-387. 
Hackel. D. B., H. E. Lebovitz. L. A. Frohman. E. Mikat. and K. Schmidt Nielsen 
1967. Effect of caloric restriction on the glucose tolerance and plasma insulin of 
the sand rat. Metabolism, 16, pp. 1133-1139. 
Hackel. D. B., E. Mikat, H. E. Lebovitz, K. Schmidt Nielsen. E. S. Horton, and 
T. D. Kinney 
1967. The sand rat {Psammomys obesus) as an ex{>erimental animal in studies of 
diabetes mellitus. Diabetologia, 3, pp. 130-134. 
Hahn. H. 

1934. Die Familie der Procaviidae. Zeitschr. Saugetierk., 9, pp. 207-358. 
Halm. A. and E. Tchernov 

1974. The distribution of myomorph rodents in the Sinai Peninsula. Mammalia, 
38, pp. 201-223. 
Haltenorth, T. 
1953a. Die Wildkatzen der alten Welt: eine ubersicht die Untergattung Felis. 

Geest and Portig, Leipzig. 166 pp. 
1953b. Lebende arabische Sandkatze, FeUs margarita (Loche, 1858). Saugetierk. 
Mitt., 1, pp. 71-73. 
Halton. L. 

1935. An Egyptian game reserve. Sheltering migratory ibex. Field, 166, p. 1520. 
Happold. D. C. D. 

1966a. The mammals of Jebel Marra, Sudan. J. Zool., London, 149, pp. 126-136. 
1966b. Breeding periods of rodents in the Northern Sudan. Rev. Zool. Bot. Africa, 

74 (3-4). pp. 357-363. 
1967a. Biology of the jerboa, Jaculus jaculus butleri (Rodentia. Dipodidae). in the 

Sudan. J. Zool.. London. 151, pp. 257-275. 
1967b. Additional information on the mammalian fauna of the Sudan. Mammalia, 

31, pp. 605-609. 
1967c. Gerbillus {Dipodillus) campestris (Gerbillinae. Rodentia) from the Sudan. J. 

Nat. Hist.. 1, pp. 315-317. 
1967d. Guide to the natural history of Khartoum Province. Part III. Mammals. 

Sudan Notes Rec.. 48, pp. 1-22. 
1968a. The Sudanese jerboa. Animals. 10 (9), pp. 432-433. 
1968b. Observations of Gerbillus pyramidum (Gerbillinae. Rodentia) at Khartoum, 

Sudan. Mammalia. 32, pp. 44-53. 

1969. The mammalian fauna of some jebels in the northern Sudan. J. Zool., 
London. 157. pp. 133-145. 

1970. Reproduction and development of the Sudanese jerboa. Jaculus jaculus 
butleri (Rodentia. Dipodidae). J. Zool.. London. 162, pp. 505-515. 



OSBORN&HELMY: MAMMALS OF EGYPT 563 

Harding-King. W. J. H. 

1925. Mysteries of the Libyan Desert. Seeley, Service and Co. Ltd., London. 
384 pp. 
Hardy. E. 

1949. The Palestine leopard. J. Soc. Preserv. Fauna Emp., 55, pp. 16-20. 
Harper. F. 

1940. The nomenclature and type localities of certain Old World animals. J. 
Mammal., 21. pp. 191-203, 322-332. 
Harrison. D. L. 

1964. The mammals of Arabia, Vol. I. Ernest Benn Ltd., London, xx -1-192 pp. 

1967. Observations on some rodents from Tunisia, with the description of a new 
gerbil (GerbiUinae: Rodentia). Mammalia, 31, pp. 381-389. 

1968. The Mammals of Arabia, Vol. II. Ernest Benn Ltd., London, xiv-l-pp. 
193-381. 

1972. The mammals of Arabia, Vol. III. Ernest Benn Ltd., London, xvii-l-pp. 
382-670. 

Hart. H. C. 
1891. Some account of the fauna and flora of Sinai, Petra, and Wady Arabah. 
Watt, London, x-f 255 pp. 
Hartert. E. 
1913. Expedition to the central western Sahara. Ill: Notes on ruminants and 
other large mammals. Nov. Zool., 20, pp. 33-37. 
Hassan. M. I. 
1953. Physical elements of agricultural land use in the Nile Delta. Bull. Soc. 
G6ogr. D'figypte, 26, pp. 227-241. 
Hassan. M. S. and A. Hegazy 
1968. Rats injurious to agriculture and their control (In Arabic). Tech. Bull. 
U. A. R. Min. Agric, Cairo, No. 28, pp. 1-31. 
Hassib. M. 
1951. Distribution of plant communities in Egypt. Fac. Sci. Fouad I Univ., Cairo, 
BuU. No. 29, pp. 59-261. 
Hassinger. J. D. 

1973. A survey of the mammals of Afghanistan. Fieldiana, Zoo., 60, xi-l-195 pp. 
Hatt. R. T. 

1959. The mammals of Iraq. Misc. Publ. Univ. Mich., No. 106, 113 pp. 

Hauser. W. 
1951. Beobachtungen an einigen Tierformen Angolas in der Natur und nach dem 
Fang, 3: Klippschliefer, Procavia capensis Pall. Zool. Garten Leipzig, 18, pp. 
208-210. 
Hayman. R. W. 
1948. The Armstrong College zoological expedition to Siwa Oasis (Libyan Desert) 
1935. Mammalia. Proc. Egypt. Acad. Sci., 4, pp. 38-43. 
Hefny. M. B. 
1953. Two climatic maps of the Nile Basin and vicinity. Bull. Soc. R. Geogr. 
figypte, 26, pp. 182-192. 

Heim de Balsac. H. and J. J. Barloy 
1966. Revision des Crocidures du groupe flavescens-occidentalis-manni. 
Mammalia, 30, pp. 601-633. 



564 FIELDIANA: ZOOLOGY 

Heim de Balsac. H. and M. Lamotte 
1957. Evolution et phylog^nie des Soricid^s Africain^. (suite et fin). Mammalia. 
21. pp. 15-49. 
Heim dk Bai^ac. H. and P. Mkin 
1971. Les musarai^es momifiees des hypogees de Thebes. Existence d'un 
metalophe chez Crocidurinae (sensu repenning). Mammalia. 35. pp. 220-244. 
Hemmek. H. 
1974. Studien zur systematik und biologie der Sandkatze. Z. Kolner Tjoo, 17. 
pp. 11-20. 
Hemmer. H.. p. Grubb. and C. P. Groves 
1976. Notes on the sand cat. Felis margarita Loche. 1858. Z. Sauegetierk.. 41. 
pp. 286-303. 
Hemprich. F. W. and C. G. Ehrenberg 
1828-1833. Symbolae physicae seu icones et descriptiones mammalium. Decas 1. 
1828 {plates published in 1828; part of text not until 1833). Decas 2. 1830 
(published in 1833; fide C. D. Sherborn). 
Henderson. K. D. D. 

1965. Sudan Republic. 256 pp. Praeger. New York. 
Heuglin. Th. von 

1877. Reise im Nordost Afrika, Vol. 2, iv+304 pp. Westermann. Brannschweig. 
Hills. E. S. (ed.) 

1966. Arid lands. Methuen and Co.. Ltd.. London, xviii + 461 pp. 
Hinton. H. E. and A. M. S. Dunn 

1967. Mongooses. Oliver and Boyd, London, vii+144 pp. 
Holland. F. W. 

1868. On the Peninsula of Sinai. J. Roy. Geogr. Soc., London, 38. pp. 237-257. 
Hollister. N. 
1918. East African mammals in the United States National Museum. Part I. 
Insectivora. Chiroptera, and Carnivora. U.S. Nat. Mus. Bull., 99. 194 pp. 
Hoogstraal. H. 
1960. Faunal exploration as a basic approach for studying infections common to 
man and animals. East Afr. Med. J., 33 (1). pp. 275-282. 

1962. A brief review of the contemporary land mammals of Egypt (including 
Sinai). 1: Insectivora and Chiroptera. J. Egypt. Publ. Hlth. Assn., 37 (4). pp. 
143-162. 

1963. A brief review of the contemporary land mammals of Egypt (including 
Sinai), 2: Lagomorpha and Rodentia. J. Egypt. Publ. Hlth. Assn.. 38(1). pp. 1-35. 

1964. A brief review of the contemporary land mammals of Egypt (including 
Sinai), 3: Carnivora. Hyracoidea, Perissodactyla and Artiodactyla. J. Egypt. 
Publ. Hlth. Assn.. 39 (4). pp. 205-239. 

Hoogstraal. H., M. N. Kaiser. R. A. Ormsbee. D. J. Osborn. I. Helmy. and 
S. Gaber 

1967. Hyalomma (Hyalommina) rhipicephaloides Neumann (Ixodoidae: Ixodidae): 
Its identity, hosts, and ecology, and Rickettsia conori, R. prowazekia, and 
Coxiella bumeti infections in rodent hosts in Egypt. J. Med. Entomol.. 4, pp. 
391-400. 

Hoogstraal. H., K. Wassif. I. Helmy. and M. Kaiser 

1968. The Cheetah, Acinonyx jubatus Schreber. in Egypt. Bull. Zool. Soc. 
Egypt. 21. pp. 63-68. 



OSBORN&HELMY: MAMMALS OF EGYPT 565 

HooGSTRAAL. H., K. Wassif. and M. N. Kaiser 

1955. New mammal records from the Western Desert of Egypt. Bull. Zool. Soc. 
Egypt. 12, pp. 7-12. 

1957a. Results of the NAMRU-3 Southeastern Egypt expedition, 1954, 1: Intro- 
duction, itinerary, and environmental conditions. Bull. Zool. Soc. Egypt, 12, 
pp. 7-12. 

1957b. Results of the NAMRU-3 Southeastern Egypt expedition, 1954, 6: 
Observations on non-domesticated mammals and their ectoparasites. Bull. Zool. 
Soc. Egypt, 13, pp. 52-75. 
HOWELLS. V. 

1956. A naturaUst in Palestine. Andrew Melrose, London. 192 pp. 
Hughes, A. R. 

1954. Hyaenas versus australopithecines as agents of bone accumulation. Amer. 
J. Phys. Anthropol.. 12, pp. 467-486. 

Hume. W. F. 
1906. The topography and geology of the Peninsula of Sinai (south-eastern 

portion). National Printing Department, Cairo. 280 pp. 
1921. The Egyptian wilderness. Geogr. J., 58, pp. 249-276. 

1925. Geology of Egypt, Vol. 1 and 2, pt. 1 and 2. Gov. Press, Ministry of Finance, 
Cairo, xiv + 408, btv + 300 pp., xxxv + pp. 301-688. 
Hurst. H. E. 

1910. A journey from Wadi Haifa to Aswan. Cairo Sci. J., 4, pp. 8-10. 
Innes. W. 

1932. Nos mammiferes rongeurs (Rodentia). Bull. Inst. D6sert figypte, 14, pp. 1-61. 
Jarvis. C. S. 

1937. Three deserts. Dutton, New York, ix -1-313 pp. 
Jayakar. a. S. G. (transl.) 
1906. Ad-Damiri's hayat al-hayawan (A zoological lexicon). Vol. I. Luzac & Co., 

London, xxx + 875 pp. 
1908. Ad-Damiri's hayat al-hayawan (A zoological lexicon), Vol. II, pt. 1. Luzac & 
Co., London. 604 pp. 
Jennison. G. 
1937. Animals for show and pleasure in ancient Rome. The Univ. Press, 
Manchester, xiv-*- 209 pp. 
Kasim. M. 

1912. A meteorological expedition to Siwa. Cairo Sci. J., 6, pp. 76-84. 
Kassas, M. 

1952. Habitat and plant communities in the Egyptian desert, I: Introduction. 
J. Ecol., 40, pp. 342-351. 

1953. Habitat and plant communities in the Egyptian desert, II: The features of 
a desert community. J. Ecol., 41, pp. 248-256. 

1955. Rainfall and vegetation belts in arid northeast Africa. Plant Ecology, 
Proc. Mont. Symp. (UNESCO), pp. 49-50. 

1956. The mist oasis of Erkwit, Sudan. J. Ecol., 44, pp. 180-194. 

1966. Plant life in deserts. In Hills, E. S., ed.. Arid Lands. Methuen and Co., 
Ltd., London, pp. 145-180. 
Kassas. M. and W. A. Girgis 
1964. Habitat and plant communities in the Egyptian desert, V: The limestone 
plateau. J. Ecol., 52, pp. 107-119. 



566 FIELDIANA: ZOOLOGY 

1965. Habitat and plant communities in the Egyptian desert. VI: The units of a 
desert ecosystem. J. Ecol.. 53, pp. 715-728. 

1970. Habitat and plant communities in the Egyptian desert, VII: Geographical 
facies of plant communities. J. Ecol., 58, pp. 335-350. 

1972. Studies on the ecology of the Eastern Desert of Egypt, I: The region 
between latitude 27 ° 30 ' and latitude 25 ° 30 ' N. Bull. Soc. Geogr. Egypte, 41-42, 
pp. 43-72. 
Kassas. M. and M. Imam 

1954. Habitat and plant communities in the Egyptian desert. III: The wadi bed 
ecosystem. J. Ecol., 42, pp. 425-441. 

1959. Habitat and plant communities in the Egyptian desert, IV: The gravel 

desert. J. Ecol.. 47, pp. 289-310. 
Kassas. M. and M. A. Zahran 
1962. Studies on the ecology of the Red Sea coastal land, I: The district of Gebel 

Ataqa and El-Galala El-Bahariya. Bull. Soc. Geogr. Egypte, 35. pp. 129-175. 
1965. Studies on the ecology of the Red Sea coastal land, II: The district from 

El-Galala El-QibUya to Hurghada. Bull. Soc. Geogr. Egypte, 38. pp. 155-193. 
1967. On the ecology of the Red Sea littoral salt marsh, Egypt. Ecol. Monogr., 

37. pp. 297-316. 

1971. Plant life on the coastal mountains of the Red Sea, Egypt. J. Indian Bot. 
Soc., 50A., pp. 571-589. 

Keimer. L. 

1942. Quelques representations rarissimes de Mustelides conserves sur de bas- 
reliefs de I'ancien empire. De L'Institut Francais D"Arch. Oriental, pp. 1-29. 

1944. L'Orycterope dans L'Egypte ancienne. Etudes Egyptologie, VI, pp. 1-20. 

1949. Le pore- 'epic dans L'Egypte ancienne. Ann. Service Antiqitee Egypte, 
49. pp. 393-415. 

1955. Interpretation de plusieurs passages contenus dans les "Histories" 
d'H6redote. BuU. Inst, figypte. 36, pp. 455-476. 

KemalelDin. H. 

1928. L'exploration du Desert de Libye. Geographie, 1928, pp. 171-183, 320-336. 
Khairat. O. 

1954. Hunting in Ancient Egypt. Egyptian Shooting and Fishing Club, Cairo. 
13 pp. 

KiRMIZ. J. P. 

1962. Adaptation to desert environment. Butterworths. London, xi-l-168 pp. 
1965. The jerboa and the desert. Sci. J.. Nos. 64-65, pp. 47-51. 
KlTTO. J. 
1841. Palestine: The physical geography and natural history of the holy land. 

Charles Knight & Co.. London. 438 pp. 
Klunzingkr. C. B. 
1878. Upper Egypt: Its people and its products. Scribner. Armstrong and Co.. 

New York, xv + 408 pp. 
KocK. D. 
1970a. Die Verbreitungsgeschichte des Flusspferdes, Hippopotamus amphibius 

Linn^, 1758. im unteren Nilgebiet. Ein Beitrag zur Zoogeographie Nordafrikas. 

Saugetierk. Mitt., 18. pp. 12-25. 
1970b. Zur Verbreitung der Mendesantilope, Addax nasomaculatus (De Blainville, 

1816), und des Spiessbockes, Oryx gazella (Linn^ 1758) im Nilgebiet. Ein Beitrag 

zur Zoogeographie Nordafrikas. Saugetierk. Mitt.. 18, pp. 25-37. 



OSBORN&HELMY: MAMMALS OF EGYPT 567 

1971. Zur Verbreitung von Mahnenschaf und Steinbock im Nilgebiet. Ein Beitrag 
zur Zoogeographie Nordost-Afrikas. Saugetierk. Mitt., 19, pp. 28-39. 

Kruuk. H. 

1968. Hyenas, the hunters nobody knows. Nat. Geogr., 134, pp. 44-57. 

1976. Feeding and social behavior of the striped hyaena (Hyaena vulgaris 
Desmarest). E. Afr. WUdl. J., 44, pp. 91-111. 

KULLMAN. E. 

1965. Die Saugetier Afghanistans. 1. Carnivora, Artiodactyla, Primates. Sci. 
Quart. J. Fac. Sci. Kabul, pp. 1-17. 
Lawrence. B. 

1967. Early domestic dogs. Z. Saugetierk., 32, pp. 44-59. 
Lawrence. B. and W. H. Bossert 

1967. Multiple character analysis of Canis lupus, latrans and familiaris, with a 
discussion of the relationships of Canis niger. Amer. Zool., 7, pp. 223-232. 
Lay. D. M. 
1967. A study of the mammals of Iran. Fieldiana: Zool., 54, pp. 1-282. 

1972. The anatomy, physiology, functional significance and evolution of 
specialized hearing organs of the Gerbillinae rodents. J. Morphol., 138, pp. 
41-120. 

Lay, D. M., K. Agerson, and C. F. Nadler 
1975. Chromosomes of some species of Gerbillus (Mammalia: Rodentia). Z. 
Saugetierk., 40, pp. 141-150. 
Lay. D. M., and C. F. Nadler 

1969. Hybridization in the rodent genus Meriones. I: Breeding and cytological 
analyses of Meriones shawi (m.) x Meriones libycus (f.) hybrids. Cytogenetics, 
8, pp. 35-50. 

1972. Cytogenetics and origin of North African spalax (Rodentia: Spalacidae). 
Cytogenetics, 11, pp. 279-285. 
Legouix, J. P., F. Petter, and A. Wisner 
1954. Etude de I'audition chez des Mammif6res a bulles tympaniques hyper- 
trophiees. Mammalia, 18, pp. 262-271. 
Lewis. R. E., J. H. Lewis, and D. L. Harrison 
1965. On a collection of mammals from northern Saudi Arabia. Proc. Zool. Soc., 
London, 144 (1), pp. 61-74. 

Lhote, H. 
1946. Observations sur la repartition actuelle et les moeurs de quelques grands 
mammif^res du pays Tuareg. Mammalia, 10, pp. 26-56. 

Like. A. A. and E. MiKi 
1967. Diabetic syndrome in sand rats. IV: Morphologic changes in islet tissue. 
Diabetologia, 3, pp. 143-166. 

Lindsay, J. 
1965. Leisure and pleasure in Roman Egypt. Muller, London, xiv+482 pp. 

LODER. E. G. 
1894. On the "Reem" antelope of Algeria. Proc. Zool. Soc., London, 1894, pp. 
473-476. 
Long. G. A. 
1955. The study of natural vegetation as a basis for pasture improvement in the 
Western Desert of Egypt. Bull. Inst. D6sert 6gypte, 5, pp. 18-48. 



568 FIELDIANA: ZOOLOGY 

LoRET. V. and C. Gaillard 

1908. La faune momifi^ de I'ancienne Egypte. Arch. Mus. Hist. Nat.. Lyon. 8, 
pp. i-vii. 1-200. 
Lydekker. R. 

1916. Catalogue of the Ungulate mammals in the British Museum (Natural 
History), Vol. 5. British Museum, London, xlv + 207 pp. 
MacDougal. D. T. 

1913. The deserts of western Egypt. Plant World.. 16, pp. 291-303. 
Madkour. G. a. 

1961. The structure of the facial area in the mouse-tailed bat, Rhinopoma 
hardwickei cystops Thomas. Bull. Zool. Soc. Egypt, 16. pp. 50-54. 
Manson Bahr, p. 

1936. Wolves in Palestine and Egypt. Field, 167 (4355). p. 1448. 
Maser. C. O. 

1966. Commensal relationship betwen Acomys and Rousettus. J. Mammal., 47, 
p. 153. 
Mason. M. 

1936. The paradise of fools. Hodder and Stoughton, London. 282 pp. 
McGiNNiES. W. G., B. J. Goldman, and P. Paylore 

1968. Deserts of the world. Univ. Arizona Press, Tuscon. xxviii-*-788 pp. 
Meester. J. and H. W. Setzer (ed.) 

1971. The mammals of Africa— An identification manual. 
Smithsonian Inst. Press, Washington, D.C. vii -I- pagination by parts. 
Meigs. P. 

1966. Desert Uttoral of Egypt. UNESCO, Paris. 144 pp. 
Meinertzhagen. R. 

1954. Birds of Arabia. Oliver and Boyd, London. 624 pp. 
Mendelssohn. H. 

1965. Breeding the Syrian hyrax Procavia capensis syriaca Schreber, 1784. Int. 

Zoo Yearbook, 5. pp. 116-125. 
MiCHENER. G. R. 
1976. Tail autotomy as an escape mechanism in Rattus rattus. J. Mammal., 57. 

pp. 600-603. 
MiGAHiD. A. M. and A. A. Abd el Rahman 
1953a. Studies in the water economy of Egyptian desert plants, 1: Desert climate 

and its relation to vegetation. Bull. Inst. D6sert figypte, 3. pp. 5-24. 
1953b. Studies in the water economy of Egyptian desert plants, II: Soil and water 

conditions and their relation to vegetation. Bull. Inst. Desert Egypte, 3, pp. 

25-57. 
1953c. Studies in the water economy of Egyptian desert plants. III: Observations 

on the drought resistance of desert plants. Bull. Inst. Desert Egypte, 3. pp. 

59-83. 
Migahid. a. M.. a. a. Abd El Rahman. M. El Shafei An. and M. A. Hammouda 

1955. Types of habitat and vegetation at Ras el Hikma. Bull. Inst. Desert Egypte, 
5, pp. 107-190. 

Migahid. A. M. and M. A. Ayyad 
1959a. An ecological study of Ras el Hikma district, I: The climatic environment 

of the vegetation. Bull. Inst. Desert Egypte, 9. pp. 1-45. 
1959b. An ecological study of Ras el Hikma district. III: Plant habitats and 

communities. Bull. Inst. Desert Egypte, 9, pp. 74-98. 



OSBORN&HELMY: MAMMALS OF EGYPT 569 

1959c. An ecological study of Ras el Hikma district, IV: Structure of vegetation 
in the main habitats. Bull. Inst. Desert fegypte, 9, pp. 99-120. 
MiKi. E., A. A. Like. J. S. Soeldner. J. Steinke. and G. F. Cahill. Jr. 

1966. Acute ketotic-type diabetic syndrome in sand rats (Psammomys obesus) 
with special reference to the pancreas. Metabolism, 15, pp. 749-760. 

MiKi. E., A. A. Like. J. Steinke. and J. S. Soeldner 

1967. Diabetic syndrome in sand rats. II: Variability and association with diet. 
Diabetologia, 3, pp. 135-139. 

Miller. G. S. 
1912. Catalogue of the mammals of western Europe. British Museum (Natural 
History), London, xv+1019 pp. 
Missone. X. 

1969. Exp^tion scientifique Beige dans le Desert de Libye Jebel Uweinat 1968- 
1969, V: La Faune. Africa-Tervuren, 15, pp. 117-119. 

1970. Animals of the Libyan Desert. Int. Union Conserv. Nat. Bull., 2, pp. 131-132. 

MiTWALLY. M. 

1953. Physiographic features of the oases of the Libyan Desert. Bull. Inst. Desert 
figypte, 3, pp. 143-163. 

1954. The call of the desert. Bull. Inst. Desert figypte, 4, pp. 125-129. 
MoNAiERY. A. M. El 

1955. Situation of wild mammals in Egypt. Bull. Zool. Soc. Egypt, 12, p. 1 (1954). 

1965. Some notes on the Egyptian cat and its origin. Bull. Zool. Soc. Egypte, 20, 
pp. 71-72. 
MONOD. Th. 
1958. Majabat Al-Koubra. Contribution a I'etude L' "Empty Quarter" Quest 
Saharien. Mem. L'Inst. Francais D'Afr. Noire. No. 52, 406 pp. 
Montagu. I. 
1924. On the burrowing of the rodent, Spalax (Mesospalax). Proc. Zool. Soc, 
London, 1924, pt. IV, pp. 1,153-1,160. 

MONTASIR. A. H. 

1954. Habitat factors and plant distribution in Egypt. Bull. Inst. Desert Egypte, 
4, pp. 36-64. 
Morrison-Scott. T. C. S. 
1939. Some Arabian mammals collected by Mr. H. St. J. B. Philby, C. I. E. Nov. 
Zool., 41, pp. 181-211. 
Morrison-Scott, T. C. S. 
1952. The mummified cats of ancient Egypt. Proc. Zool. Soc., London, 121, pp. 
861-867. 

Mountfort. G. 

1965. Portrait of a desert. Collins, London. 192 pp. 
Murray. G. W. 

1912. The hamada country. Cairo Sci. J., 6, pp. 264-273. 

1930. Egyptian mountains. Alpine J., 42, pp. 226-235. 

1935. Sons of Ishmael. Routledge & Sons Ltd., London, xv+344 pp. 

1950. The Egyptian desert and its antiquity. Paper No. 40, Survey Dept. of 
Egypt, Cairo. 22 pp. 

1951. The Egyptian climate: an historical outline. Geogr. J., 117, pp. 422-434. 
1967. Dare me to the desert. Allen and Unwin Ltd., London. 214 pp. 



570 FIELD1ANA:ZCX)L0GY 

Murray. J. 
1891. Handbook for travelers in lower and upper Egypt 8th ed. rev. John Murray, 
London. 568 pp. 
MUSCHLER. R. 

1912. A manual flora of Egypt, Vol. 2. Friedlander. Berlin, pp. 673-1312. 
Nadlkr. C. F. and D. M. Lay 
1967. Chromosomes of some species of Meriones (Mammalia: Rodentia). Z. 
Saugetierk., 32. pp. 285-291. 
Neoumi. a. 
1949. List of desert animals seen or collected during the periods shown. Bull. 

Zool. Soc. Egypt, 8, pp. 20-21. 
1952. Egyptian mammals (In Arabic). Khalil Ibrahim, Alexandria. n-(-134 pp. 
Nehring. a. 
1901. iJher Dipus schliiteri n. sp. und einige andere Nager aus Palestina. S. B. Ges. 
Nat. Fr.. 1901 (8). pp. 163-176. 
Neumann. O. 

1935. liber afrikanische wildesel. Z. Saugetierk.. 10, pp. 152-153. 
Nevo. E. 
1961. Observations on Israeli populations of the mole rat Spalax e. ehrenbergi 

Nehring. Mammalia, 25, pp. 127-144. 
1969. Mole rat Spalax ehrenbergi: Mating behavior and evolutionary significance. 
Science. 163, pp. 484-486. 
Newbold. D. 
1928. Rock-pictures and archaeology in the Libyan Desert. Antiquity, 2, pp. 
261-288. 

NiETHAMMER. J. 

1959. Die nordafrikanischen Unterarten des Gartenschlafers {Eliomys quercinus). 
Z. Saugetierk., 24, pp. 35-45. 
Omer-Cooper. J. 
1947. The Armstrong College zoological expedition to Siwa Oasis (Libyan Desert) 
1935. General Report. Proc. Egypt. Acad. Sci.. 3, pp. 1-52. 
OSBORN. D. J. 

1968a. Hyena hunt. Bull. Field Mus. Nat. Hist., 39, pp. 2-6, 14. 
1968b. Notes on medicinal and other uses of plants in Egjrpt. Econ. Botany. 22, 
pp. 165-177. 
OsBORN. D. J. and K. V. Krombein 

1969. Habitats, flora, mammals, and wasps of gebel 'Uweinat, Libyan Desert. 
Smithson. Contr. Zool., No. 11, pp. 1-18. 
Palmer. E. H. 

1872. The desert of the exodus. Harper. New York. xvi-f470 pp. 
Paton. D. 

1925. Animals of ancient Egypt. Oxford Univ. Press, London. 37 pp. 
Pearse. C. K. 

1955. Improving Egypt's range forage. Bull. Inst. Desert fegypte, 5, pp. 77-86. 
Pease. A. E. 
1896. On the antelopes of the Aures and Eastern Algerian Sahara. Proc. Zool. 
Soc., London, 1896, pp. 809-814. 
Peeu R. F. 
1939. The Gilf Kebir. In Bagnold, R. A., An expedition to the Gilf Kebir and 
Uweinat. 1936. pt. 4. Geogr. J., 93, pp. 295-307. 



OSBORN&HELMY: MAMMALS OF EGYPT 571 

Pelikan. J., G. Madkour. and J. Gaisler 
1971. On some mammals from Egypt (Insectivora, Lagomorpha, Rodentia). Zool. 
Listy. 20. pp. 307-318. 
Ferret. R. 

1935. Le climat du Sahara. Ann. Geogr.. 44, pp. 162-186. 
Fetermann. a. 
1861. Mittheilungen aus Justus Perthes' Geographischer Anstalt etc. Perthes, 
Gotha. viii+482 pp. 
Fetter. F. 
1953a. Note pr^liminaire sur I'Ethologie et I'Ethologie de Meriones libycus 

(Rongeurs. Gerbillid^s). Mammalia, 17, pp. 281-294. 
1953b. Remarques sur la signification des bulles tympaniques chez les 
Mammiferes. C. R. Acad. Sci., Paris. 237, pp. 848-849. 

1956. Evolution du dessin de la surface d'usure des molaires de Gerbillus, 
Meriones, Pachyuromus et Sekeetamys. Mammalia. 20, pp. 419-426. 

1957. Liste commentee des espies de gerbillides de Palestine. Mammalia, 21, 
pp. 241-257. 

1959. Evolution du dessin de la surface d'usure des molaires des Gerbillides 

Mammalia. 23. pp. 304-315. 
1961. Repartition geographique et 6cologie des rongeurs desertiques (du Sahara 
occidental a I'lran oriental). Mammalia. 25, (special no.), pp. 1-222. 
PococK. R. I. 
1932. The leopards of Africa. Proc. Zool. Soc, London, 1932, pp. 543-591. 
1934. The races of the striped and brown hyenas. Proc. Zool. Soc, London, 1934, 

pp. 799-835. 
1941. The fauna of British India including Ceylon and Burma. Mammalia, Vol. IL 

Taylor and Francis Ltd., London, xii-f 503 pp. 
1951. Catalogue of the Genus Felis. British Museum (Nat. Hist.), London. 
vii+190pp. 
FOMEL. M. A. 
1856. Notes sur la mammalogie de I'Alg^rie. Compt. Rend. Acad. Sci. Paris, 42, 
pp. 652-655. 
Fraakash, I. and P. K. Ghosh (ed.) 

1975. Rodents in desert environments. The Hague, Junk, xvi+624 pp. 
France. H. D., K. Schmidt-Nielsen, and D. B. Hackel 
1968. Care and breeding of the fat sand rat {Psammomys obesus Cretzschmar). 
Lab. Anim. Care. 18, pp. 170-181. 
Ranck. G. L. 
1968. The rodents of Libya: taxonomy, ecology, and zoogeographical relation- 
ships. U. S. Nat. Mus. Bull., 275, pp. vii+264. 
Reed. C. A. 

1958. Observations on the burrowing rodent Spalax in Iraq. J. Mammal., 39, 
pp. 366-389. 

1964. A natural history study of Kurkur Oasis, Libyan Desert, Western 
Governate, Egypt. Postilla, No. 84, pp. 1-20. 

1966. Organic remains from Yale University Nubian expedition archaeological 
sites D 1-2 IB and W0-2A, with a discussion of the radiocarbon determinations. 
Postilla, No. 102. pp. 35-46. 

Reed. C. A., M. A. Baumhoff. K. W. Butzer. H. Walter, and D. S. Boloyan. 

1967. Preliminary report on the archaeological aspects of the Yale University 



572 FIELDIANA: ZOOLOGY 

prehistoric expedition to Nubia, 1962-1963. Extract from Fouilles en Nubie 
(1961-1963). Antiquities Department of Egypt, Cairo, 12 pp. 
Reed. C. A. and P. F. Turnbull 
1969. Late Pleistocene mammals from Nubia. In van Zinderen Bakker, E. M., ed., 
Palaeoecology of Africa and of the Surrounding Islands and Antarctica, 
Vol. IV. Balkerna, Capetown, pp. 55-56. 
Roberts. A. 
1 951. The mammals of South Africa. Trustees of "The Mammals of South Africa" 
Book Fund, xlviii+700 pp. 
Robinson. P. 
1966. Fossil occurrence of murine rodent {Nesokia indica) in the Sudan. Science, 
154, p. 264. 
Roche. J. 
1960. Nouvelles donnees sur la reproduction des Hyracoides. Mammalia, 26, 
pp. 517-529. 
Rothschild. W. 

1913. On Ovis lervia Pallas and its subspecies. Nov. Zool., 20, pp. 459-460. 
Ruppell. E. 
1826. Atlas zu der Reise im nordlichen Afrika. Erste Abteilung Zoologie der 
Senkenbergischen naturforschenden Gesellschaft. Saugethiere.. Frankfurt 
am Mein. pp. 1-78. 
1829. Reisen in Nubien, Kordofan und dem Petraischen Arabien. Frankfurt am 

Mein. 40 pp. 
1842. Saugetheire aus der Ordnung der Nager, beobachtet im nordostlichen 
Afrika. Mus. Senckenb., Abh.. 3, pp. 91-116. 
Russell. M. 
1831. View of ancient and modern Egypt with an outline of its natural history. 
Oliver and Boyd, Edinburgh. 480 pp. 
Russell. Th. 
1949a. Egyptian service. John Murray, London, xi+294 pp. 
1949b. Desert fauna. Bull. Zool. Soc. Egypt, No. 8, pp. 5-8. 
1951. The fauna of the Egyptian deserts. Bull. Zool. Soc. Egypt, No. 9. pp. 19-21. 
Said. R. 

1%2. The geology of Egypt. Elsevier. Amsterdam-New York. iv-»-357 pp. 
Saint-Girons. M. 
1962. Notes sur les dates de reproduction en captivity du Fennec, Fennecus zerda 
(Zimmermann. 1780). Z. Saugetierk., 27, pp. 181-184. 
Sale. J. B. 
1965. The feeding behaviour of rock hyraxes (genera Procavia andHeterohyrax) in 
Kenya. E. Afr. Wildl. J.. 3, pp. 1-18. 
Sanborn. C. C. and H. Hoogstraal 
1953. Some mammals of Yemen and their ectoparasites. Fieldiana: Zool.. 34, 

pp. 229-252. 
1955. The identification of Egyptian bats. J. Egypt. Publ. Hlth. Assn.. 30 (4), 
pp. 103-119. 
Sandford. K. S. 
1936. Observations on the distribution of land and fresh-water moUusca in the 
southern Libyan Desert. Quart. J. Geol. Soc. London, 92, pp. 201-220. 



OSBORN&HELMY: MAMMALS OF EGYPT 573 

Sandford, K. a. and W. J. Arkell 
1933. Paleolithic man in the Nile Valley in Nubia and upper Egypt. Univ. Chicago 

Oriental Inst. Publ.. Vol. 17. xvii+92 pp. 
1939. Paleolithic man and the Nile Valley in Lower Egypt. Univ. Chicago Oriental 
Inst. Pub.. 46, xix+105 pp. 
Schmidt-Nielsen. B. and K. Schmidt-Nielsen 

1949. The water economy of desert mammals. Sci. Mon., 69, pp. 180-185. 

1950. Evaporative water loss in desert rodents in their natural habitat. Ecology, 
31. pp. 75-85. 

1952. Water metabolism in desert animals. Physiol. Rev., 32, pp. 135-166. 
Schmidt-Nielsen. K. 

1964. Desert animals. Oxford University Press, London. 277 pp. 
Schmidt-Nielsen. K., H. B. Haines, and D. B. Hackel 
1964. Diabetes mellitus in the sand rat induced by standard laboratory diets. 
Science, 143, pp. 689-690. 
Schomber. H. W. and D. KocK 

1960. The wild life of Tunisia, pt. 2. Afr. WUdl, 14 (4), pp. 277-282. 
Schwann, H. 
1905. A list of mammals collected by the Hon. N. C. Rothschild, the Hon. F. R. 
Henley, and Mr. A. F. R. WoUaston in Egypt and the Soudan in January, 
February, and March 1904. Nov. Zool., 12, pp. 1-5. 
SCHWARZ. E. and H. K. Schwarz 
1943. The wild and commensal stocks of the house mouse, Mus musculus 
Linnaeus. J. Mammal., 24, pp. 59-72. 

SCHWEINFURTH. G. 

1874. Notice sur la grande oases du Desert Libyque. Bull. Soc. Geogr., (ser. vii), 
6. pp. 627-634. 
SCLATER. p. L. 

1884. On some mammals from Somali-land. Proc. Zool. Soc, London, 1884, pp. 

538-542. 
1895. On the occurrence of the barbary sheep in Egypt. Proc. Zool. Soc, London, 

1895, pp. 85-86. 
SCLATER, P. L. and O. Thomas 
1894-1900. The book of antelopes, vols. I-IV. Porter, London, xxxv+220 and 

242 pp. 
Setzer. H. W. 
1952. Notes on mammals from the Nile Delta region of Egypt. Proc. U.S. Nat. 

Mus., 102 (3305), pp. 343-369. 

1955. Two new jerboas from Egypt. Proc Biol. Soc. Wash. 68, pp. 183-184. 

1956. Mammals of the Anglo-Egyptian Sudan. Proc. U.S. Nat. Mus., 106 (3377), 
pp. 447-587. 

1957a. An extra tooth in Crocidura. 3. Mammal., 38, pp. 258-259. 

1957b. The hedgehogs and shrews (Insectivora) of Egypt. J. Egypt. Publ. Hlth. 

Assn., 32(1), pp. 1-17. 
1957c. A review of Libyan mammals. J. Egypt. Publ. Health Assn., 32 (2), 

pp. 41-82. 
1958a. The jerboas of Egypt. J. Egypt. Publ. Health Assn., 33 (3). pp. 87-94. 
1958b. The hares of Egypt. J. Egypt. Publ. Health Assn., 33 (5), pp. 145-151. 



574 FIELDIANA: ZOOLOGY 

1958c. The mustelids of Egypt. J. Egypt. Publ. Health Assn.. 33 (6). pp. 199-204. 
1958d. The gerbils of Egypt. J. Egypt. Publ. Health^Assn.. 33 (6). pp. 205-227. 
1959. The spiny mice iAcomys) of Egypt. J. Egypt. Publ. Health Assn.. 34 (3). 

pp. 93101. 
1960a. The baculum of Poecilicitis libyca. J. Mammal.. 41. pp. 138-139. 
1960b. Two new mammals from Egypt. J. Egypt. Publ. Health Assn.. 35 (1). pp. 

1-5. 
1961a. The jirds (Mammalia: Rodentia) of Egypt. J. Egypt. Publ. Health Assn.. 

36(3). pp. 81-92. 
1961b. The canids (Mammalia) of Egypt. J. Egypt. Publ. Health Assn., 36 (3). 

pp. 113-118. 
1963. Notes on some Egyptian rodents. J. Egypt. Publ. Health Assn., 38 (2). pp. 

51-60. 

Shalaby. a. a. 
1962. The effect of injected water on the water economy of the desert mammal 
Jaculus jaculus, 1: The total body water content and water of different tissues. 
Bull. Zool. Soc. Egypt. 17, pp. 1-14. 
Shata. a. 
1955. An introductory note on the geology of the northern jwrtion of the Western 
Desert of Egypt. Bull. Inst. Ddsert figypte. 5, pp. 96-102. 
Shaw. W. B. K. 
1931. Botanical notes: Appendix IV. In Bagnold, R. A.. Journeys in the Libyan 
Desert 1929 and 1930. Geogr. J.. 78, pp. 534-535. 

1933. Libyan Desert: Note on wild life. J. Soc. Pres. Fauna Empire, 20, p. 15. 

1934. The flora of the Libyan Desert: Botanical notes. J. Soc. Pres. Fauna Empire. 
7(1934). pp. 281-289. 

Shaw. W. B. K. and J. Hutchinson 
1931. The flora of the Libyan Desert. Bull. Misc. Inform. Roy. Bot. Gard.. Kew, 4 
(1931). pp. 161-166. 
She H ATA. I. and A. Kawashti 
1966. Comparative drought resistance in summer of camels, donkeys, sheep and 
goats. Bull. Inst. Desert 6gypte. 16, pp. 181-203. 
Sherborn. C. D. 
1897. On the dates of the natural history portion of Savigny's "Description de 
rfigypte." Proc. Zool. Soc. London. 1897, pp. 285-288. 
Shkolnik. a. and A. Borut 

1969. Temperature and water relations in two species of spiny mice (Acomys). 
J. Mammal.. 50, pp. 245-255. 

SONNINI. C. S. 
1807. Travels in Upper and Lower Egypt, Vol. 1-3. John Stockdale, London, 
xix -1-376. vii+368. and vii+314 pp. 
Spitzenberger. F. 

1970. Erstnachweise der Wimperspitzmaus {Suncus etniscus) fiir Kreta und