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BIOLOGY
JUL 1 9 1982
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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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32
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, 0 and 44
mm. for the same period. South of 29° N lat., the average annual
rainfall is 0 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 0 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
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88
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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OSBORN & HELMY: MAMMALS OF EGYPT
105
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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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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,
^^Vi
Jlw^^^E
^^m^B^Br
ft ^
•f3
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^^^^^^^^^K
^ 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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^^^^^Ha *^^^^
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4 —
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.
144
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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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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
I
I
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• • 3 OO
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o
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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
I
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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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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 .
0
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
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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.
s
I
SI
e
^
o
O
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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W w t m
M.MUSCULUS
S EHRENBC0GI
U
\y
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
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"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
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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.
ita^^^
Fk; 105. Live specimen oiJaculus jaculus jaculus. Note that longest vibrissae are
contacting ground.
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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
30
25
20
15
«0
5 10
Q
J5
10
WES TERN DESERT :
FLAVILLUS H
JACULUS Q
i:
pur
±L
18 20 22 24 26 28 56 58 60 62 64 66 68 mm
SINAI AND EASTERN DESERT
SCHLUETERI Q
BUTLERI [— 1
ij^
Jl
a
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.
Age determination.— Specimens are considered adult when all per-
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).
498
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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* 33* 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.
>^
(°)
0
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 0 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 0 Shim (Kom Aushim)
EL FAIYUM
29
34
30
55
OSBORN & HELMY: MAMMALS OF EGYPT
545
Locality
Governorate
NLat.
E Long.
Kom 0 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 0 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
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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
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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.,
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1^1
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