USNMB 2751968 Unit

SMITHSONIAN
INSTITUTIO N
MUSEUM
O F
NATURAL
HISTORY
UNITED STATES NATIONAL MUSEUM BULLETIN 275
The Rodents of Libya
Taxonomy, Ecology
and
Zoogeographical Relationships
GARY L. RANCK
Curator, Mammal Identification Service
Division of Mammals, U.S. National Museum
SMITHSONIAN INSTITUTION PRESS

WASHINGTON, D.C.
1968
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Contents
Page
Acknowledgments 1
Introduction 3
Physical description of Libya 6
General features 6
The coastal plain 7
The coastal escarpment 7
Pre-Saharan or steppe 7
Hamadas 8
Sand seas 9
Serirs 9
Fesh-fesh 10
Depressions 10
Mountainous areas of the interior 10
The Cyrenaican Plateau 11
The Tripolitanian Gebel 11
Geology 12
Tripolitania and Cyrenaica 12
Fezzan 13
Volcanic activity 14
Climate 14
Libyan interior 14
Arid or desert type 14
Semiarid or steppe type 15
Mediterranean littoral 16
Coastal uplands 17
Additional meteorological data 17
Phytogeography 18
General features of the Saharan flora 18
Phytogeographic characteristics 19
Systematic composition 19
Endemism 20
Adaptation to the desert environment 20
The vegetative cover of Libya 21
Coastal plain 21
Cyrenaican Plateau 22
Tripolitanian Gebel 23
Saharan steppe 23
Sand seas, hamadas, and serirs 23
Depressions or sebchets 24
Oases 24
The grass cover of Libya 25
Mammalian Faunal Areas 26
Mediterranean Faunal Area 27
Saharan Steppe Faunal Area 30
Saharan Desert Faunal Area 30
VI CONTENTS
Page
Speciation and geographical variation 34
Geographic variation in response to selective factors of the habitat . . 35
Modes of speciation 36
Sympatric speciation 37
Geographic speciation 39
Clinal variation 39
Geographic isolates 40
Zoogeographical considerations 44
Effects of the Pleistocene on the dispersal of Saharan and Libyan
rodents 44
Zoogeographical relationships of the rodents of the Sahara and South-
west Asia 53
Conclusions 61
Systematic treatment 62
Plan of treatment 62
Gazetteer of principal collecting localities 67
Family Cricetidae 69
Subfamily Microtinae 69
Genus Microtus Schrank 69
Microtias mustersi Hinton 69
Subfamily Gerbillinae 73
Genus Gerbillus Desmarest 73
Gerbillus aureus aureus Setzer 87
Gerbillus aureus favillus Setzer 89
Gerbillus aureus nalulensis, new subspecies 90
Gerbillus eatoni eatoni Thomas 95
Gerbillus eatoni inflatus, new subspecies 97
Gerbillus eatoni versicolor, new subspecies 98
Gerbillus gerbillus aeruginosus, new subspecies 103
Gerbillus gerbillus discolor, new subspecies 106
Gerbillus gerbillus gerbillus (Olivier) 109
Gerbillus gerbillus latastei Thomas and Trouessart 110
Gerbillus gerbillus psammophilous, .new subspecies 112
Gerbillus pyramidum hamadensis, new subspecies IIS
Gerbillus pyramidum tarabuli Thomas 121
Gerbillus amoenus vivax (Thomas) 125
Gerbillus campestris brunnescens, new subspecies 133
Gerbillus campestris dodsoni (Thomas) 136
Gerbillus campestris haymani Setzer 139
Gerbillus campestris patrizii (de Beaux) 141
Gerbillus campestris wassifi Setzer 143
Gerbillus grobbeni Klaptocz 146
Gerbillus henleyi henleyi (de Winton) 147
Gerbillus kaiseri Setzer 149
Genus Pachyuromys Lataste 153
Pachyuromys duprasi natronensis de Winton 153
Genus Merioncs Illiger 157
Meriones caudatus amplus, new subspecies 165
Meriones caudatus caudatus Thomas 169
Mi r tones caudatus confalonierii de Beaux 172
CONTENTS VII
Page
—
Family Cricetidae Continued
—
Subfamily Gerbillinae Continued
—
Genus Meriones Illiger Continued
Meriones caudatus luridus, new subspecies 173
Meriones crassus tripolius Thomas 176
Meriones libycus auratus, new subspecies 183
Meriones libycus azizi Setzer 185
Genus Psammomys Cretzschmar 187
Psammomys obesus obesus Cretzschmar 190
Psammomys obesus tripolitanus Thomas 192
Psammomys vexillaris vexillaris Thomas 192
Family Spalacidae 194
Genus Spalax Giildenstaedt 194
Spalax ehrenbergi aegyptiacus Nehring 194
Family Muridae 199
Genus Rattus Fischer 199
Rattus norvegicus (Berkenhout) 199
Rattus rattus (Linnaeus) 200
Genus Mas Linnaeus 203
Mus musculus Linnaeus 203
Genus Acomys Geoff roy 208
Acomys cahirinus viator Thomas 208
Family Gliridae 212
Genus Eliomys Wagner 212
Eliomys quercinus cyrenaicus Festa 214
Eliomys quercinus denticulatus, new subspecies 216
Eliomys quercinus tunetae Thomas 219
Family Dipodidae 220
Genus Jaculus Erxleben 220
Jaculus deserti favilhis Setzer 224
Jaculus deserti fuscipes, new subspecies 226
Jaculus deserti rarus, new subspecies 228
Jaculus deserti vastus, new subspecies 230
Jaculus jaculus arenaceous, new subspecies 234
Jaculus jaculus collinsi, new subspecies 236
Jaculus jaculus cufrensis, new subspecies 238
Jaculus jaculus tripolilanicus, new subspecies 240
Jaculus jaculus whitchurchi, new subspecies 242
Jaculus oricntalis orientalis Erxleben 244
Genus Allactaga Cuvier 248
Allactaga letradactyla (Lichtenstein) 248
Family Hystricidae 251
Genus Hystrix Linnaeus 251
Hystrix cristata Linnaeus 251
Family Ctenodactylidae 252
Genus Clenodactylus Gray 252
Ctenodactylus gundi gundi (Rothman) 254
Ctenodactylus gundi vali Thomas 259
Literature cited 260
Plates 265
Acknowledgments
For the collecting of specimens and for aid in some aspects of the
field work, indebted to James H. Shaw and Henry W. Setzer.
I am
My sincere appreciation is extended to the following staff members
and technical personnel of the United States National Museum,
Smithsonian Institution, and of the Bird and Mammal Laboratories,
United States Fish and Wildlife Service, for their assistance in various
ways: Terry K. Cederstrom, Helen S. Gaylord, Arthur M. Greenhall,
Theodore A. Heist, David H. Johnson, Burrowes T. Lovinggood,
Mary W. Mann, Richard H. Manville, John H. Miles, Jr., John L.
Paradiso, and Lyman B. Smith.
I would like to acknowledge the assistance of Donald J. Ortner,
Museum Technician, Division of Physical Anthropology, for informa-
tion regarding statistical computations; and Jack Scott and other
members of the Photographic Laboratory, Museum of Natural
History, for the reproduction of the maps and charts which appear
in this publication.
Many members and technical personnel from other
additional staff
divisions of the United States National Museum have also given
freely of their advice and services.
For the acquisition of various items of field equipment and assist-
ance in all of the field work in Libya, I thank the following officers
of the 64th Engineering Battalion, Base Topographic Survey, Wheelus
Air Force Base, Tripoli, Libya: Lt. Col. Thomas Whitchurch, Major
Guy Brunnacci, Captain William McMullen, and Chief Warrant
Officer Lynn Walters. In addition, I would like to acknowledge the
assistance of the many other officers and enlisted men of the 64th
Engineering Battalion, who aided in the field work in Libya.
I desire to express my appreciation to the following individuals for
their contributions enabling me to operate more efficiently in Libya:
Dr. Mohammed Ayub, Director of Antiquities for the Fezzan, Libya;
Randy Bowmar, Director of United States Overseas Mission, Fezzan
District, Libya; Aiad Ali Elwinsi, guide and interpreter; James R.
Jones and Gordon Chase Tibbits, hydrologists of the United States
Geological Survey and assigned to the United States Overseas Mis-
sion, Libya; Howard Meadows, Agricultural Advisor for the Fezzan,
United States Overseas Mission, Libya.
For permission to study specimens in their collections and for the
loan of comparative materials, I thank Dr. G. C. Corbett and Mr.
G. W. Hayman of the Mammal Room, British Museum (Natural
1
2 U.S. NATIONAL MUSEUM BULLETIN 275
History), London; and Dr. Francis Petter, Curator of Mammals,

Museum National d'Histoire Naturelle, Paris.
I am particularly indebted to Dr. Henry W. Setzer, Associate
Curator, Division of Mammals, United States National Museum,
Smithsonian Institution, for bis efforts in arranging for my participa-
tion in the Libyan project, his untiring criticism of the manuscript,
and for his continual encouragement and support of this research.
To Professor Stephen D. Durrant, Department of Zoology and
Entomology, University of Utah, and to Dr. Charles O. Handley, Jr.,
Curator in Charge, Division of Mammals, United States National
Museum, extend my deepest gratitude for their helpful suggestions
I
aid reading of the manuscript.

critical
I would like also to acknowledge the time and efforts of the follow-
ing staff members of the Department of Zoology and Entomology,

University of Utah: Drs. George F. Edmunds, Seville Flowers,
Kimball Harper, and Harold F. Hirth.
To my wife Sherri, I am especially grateful for her understanding
patience and constant encouragement during the preparation of this
work, for typing the preliminary and final manuscript, and for the
preparation of the illustrations which appear in the text.
Field work in Libya in 1961 and 1962 was supported by Grant
DA-MD-49-193-62-G35 awarded to the Smithsonian Institution by
the U.S. Army Medical Research and Development Command, Office
of the Surgeon General.
I am most
grateful to the Division of Education and Training,
Smithsonian Institution, for a Predoctoral Internship in 1965-1966
which enabled me to conduct the research necessary for the prepara-
tion of this publication.
The computations were performed on a Mathatron-Model
statistical
8-48S in Natural History Building, United States National
the
Museum, Smithsonian Institution.
An essential portion of this research was submitted as a thesis in
partial fulfillment of the requirements for the degree of Doctor of
Philosophy at the University of Utah.
Introduction
Libya is a vast expanse of semiarid and arid desert of variable

topography located in North Africa bordering on the Mediterranean
Sea. A large portion of the Libyan coastline follows the broad in-
dentation of the Gulf of Sirte. Libya is bordered on the east by Egypt
(United Arab Republic), on the southeast by the Sudan, and on the
west by Tunisia and Algeria. The republics of the Chad and Niger
form various portions of the southern marginal limits. The latitudinal
limits range from 19° north latitude at the extreme southeastern
portion to 33° north latitude at northern extremity in Cyrenaica.
its
Longitudinally, Libya lies between 9° and 25° east longitude. This

large country comprises approximately 680,000 square miles of pri-
marily desert terrain and is divisible into three political provinces.
The province of Cyrenaica (approximately 330,000 sq. mi.) encom-
passes roughly the entire eastern half of Libya, and the provinces of
Tripolitania (approximately 136,000 sq. mi.) and Fezzan (approxi-
mately 213,000 sq. mi.) include the northwestern and southwestern
quadrants respectively.
The surface configuration of Libya is quite varied with elevations
ranging from below sea level in the Sebchet el Gheneien near Gialo
and in the depressions surrounding Giarabub and Bahr el Tubat to
over 10,000 feet above sea level in the outliers of the Tibesti Mountains
of extreme southwestern Cyrenaica. In Libya, elevation tends to
increase gradually from north to south, the deserts of the interior
generally much higher than the coastal areas; a similar gradient in
elevation seen in an east-west transect, the eastern portion ap-
is
preciably lower than the west. Abrupt local and regional changes in
relief occur throughout Libya owing to the juxtaposition of low-lying
plains, high plateaus,and mountain complexes.
The
pattern of vegetative cover in Libya follows rather closely the
physiographic features. The flora of the low-lying coastal plain is
particularly distinct from that of the pre-Saharan (steppe) and Saharan
zones of the interior. Likewise the Cyrenaican Plateau, because of its
higher elevation and more abundant rainfall, has a flora unlike that of
the coastal plain and interior deserts.
Permanent vegetative cover of the Saharan interior of Libya is
sparse, and in large portions of the hamadas, serirs, and sand seas, it
is entirely lacking. Where

vegetation occurs in these interior deserts,
it is usually either sporadic and tends to be localized in sandy de-
pressions of the hamadas, or occurs as narrow fringes bordering the

3
4 U.S. NATIONAL MUSEUM BULLETIN 2 75
larger wadis (dry watercourses). Vegetation also occurs sparingly

along the margins of the sand seas. Some of the mountainous com-
plexes support scattered growths of grasses, and others are almost
devoid of vegetation. The oases of the Libyan Sahara support ex-
tensive groves of date palms and other agricultural crops and appear
as verdant areas in the midst of their barren surroundings.
The climate of most of Libya
is typically Saharan and extremely
arid but more humid in the coastal areas. As a result of their higher
is
elevations and greater amounts of rainfall, the Tripolitanian Gebel

and the Cyrenaican Plateau are local exceptions and have cooler
climates.
Explorations in Libya date from the end of the 18th century, but
zoological information did not begin to accumulate until the end of
the 19th century. Among the earliest publications relating to the
rodent fauna of Libya and adjacent countries are those of Yarrel
(1831), Loche (1867), Lataste (1881 and 1887), Nehring (1897), and
Sordelli (1899). The 20th century marked the beginning of any real
definitive studies of Libyan rodents. Most prominent among these
workers during the early part of this century are: Thomas (1902-
1925), Klaptocz (1909), Andreucci (1914), Ghigi (1920), Festa (1921
and 1925), Hartert (1923), Hinton (1926), and de Beaux (1928, 1932,
and 1938). More recently Zavattari (1934 and 1937), Heim de Balsac
(1935), Rode (1948), and Toschi (1951 and 1954) have published
papers dealing with various aspects of the Libyan rodent fauna.
Setzer (1956 and 1957) is the most recent worker on Libyan mammals.
Prior to the present study, relatively few specimens of Libyan
mammals were available, and the systematic composition and distri-
bution of the rodent fauna were accordingly poorly known. Specimens
collected by the Whitaker Expedition to Tripolitania and the Fezzan
in 1901 (reported by Thomas in 1902), those collected by de Beaux's
expeditions to Giarabub and Cufra in 1928 and 1932, and those ob-
tained by Toschi in 1951 and 1954, constituted the most significant
Libyan rodents prior to 1955.
collections of
This study of Libyan rodents is based primarily on specimens ob-
tained by the author during the period from October 1961 to July
1962.
Field work consisted of approximately 10 months of intensive
collecting in various parts of the three provinces of Libya. During
this period efforts were made to obtain adequate series of representa-
tive rodents from all types of habitats.
The duration of each collecting trip ranged from week to 3 months.
1
One to 14 days were spent at eacli collecting site. The majority of

specimens were obtained by using approximately 200 museum special
RODENTS OF LIBYA O
traps per night, baited with moistened oatmeal and set in a conven-
tional fashion. Occasionally, rajttrapswere used to take large rodents,
such as jirds (Meriones) and sand rats (Psammomys) In a few instances
.
jirds (Meriones), gerbils (GerbUlus) and house mice (Mus) were pur-
chased from local residents of the oases. Mole rats (Sjmlax) were
obtained using Macabee gopher traps. Diurnal rodents, such as sand
rats (Psammomys) and gundis (Ctenodactylus) were shot with a 16-
,
gauge shotgun, using number 9 shot or a .32-caliber auxiliary barrel

in the 16-gauge shotgun, using .32-caliber shells loaded with dust shot.
The most effective method for collecting sand rats (Psammomys) in-
volved shooting them along the roadside while riding on the front of
the Land Rover.
Actual field work consisted of 16,175 trap nights which yielded in
excess of 3,100 specimens, of which some 1,758 were retained as
specimens. One hundred sixty-five days and nights were spent in the
field, of which 124 were spent collecting.
Significant collections were also obtained by Henry W. Setzer in the
fall of 1955, while engaged in a preliminary survey of Libyan mammals,
and again in the spring of 1961, while on a trans-Saharan trip to the

Tibesti Mountains. Additional specimens were collected by James H.
Shaw, who accompanied the author on a collecting trip to northern
Cyrenaica in the spring of 1962. Additional specimens were borrowed
from the British and Paris museums for comparative purposes. The
present study is thus based on a total of 2,026 specimens representing
28 species and 48 subspecies.
Many areas in Libya are still not represented by specimens, however,
and many of the conclusions regarding the systematics and distribution
of certain taxa are thus conjectural. For example, the occurrence in
the Coastal Plain Province of several subspecies of GerbUlus eatoni
Thomas is not consistent with recognized concepts of speciation.
When specimens become available from more localities on the coastal

plain and permit a more comprehensive study, the Coastal Plain
Province will probably prove to be divisible into a number of sub-
provinces, each having a typical rodent fauna.
The present study is the first attempt to provide a comprehensive
treatment of the taxonomy and distribution of Libyan rodents based
upon actual specimens and field experience. This account is not to be
regarded as a final work on the systematics and distribution of Libyan
rodents but is, instead, a progress report limited to information
available at the present time. Much remains to be learned about specia-
tion and distribution of the Libyan rodent fauna, and it is hoped that
this paper will serve as a point of departure for future research.
Physical Description of Libya

General Features
Libya is divisible into several rather broad physiographic elements.

These include the coastal plain adjacent to the Mediterranean Sea, the
coastal escarpment delineating the inland margin of the coastal plain,
the broad zone of sand and gravel plains of the pre-Saharan region
south of the coastal escarpment, and the hamadas, serirs, and sand
seas of the Saharan interior. In many areas, isolated mountain com-
plexes, escarpments, and local rocky outcroppings are widespread and
form a large part of the desert landscape. In other areas, near the
coast and in the interior, large, dry watercourses (wadis) dissect the
otherwise featureless plains. These wadis are a typical feature of the
Libyan Sahara and, in many areas, provide the only visible contrast
in an otherwise monotonous landscape. In northern Cyrenaica, the
deep canyons and high tablelands of the Cyrenaican Plateau, and in
Tripolitania, the high valleys and rocky headlands of the Gebel
Nefusa form physiographic features differing markedly from the low-
lying surrounding terrain.
Figure 1. — Physical features of Libya.

RODENTS OF LIBYA
The Coastal Plain
The Libyan coastal plain varies in width from a few hundred

meters in some places in northern Cyrenaica to perhaps 50 or 60
kilometers at its widest portion in northwestern Tripolitania. This
plain stretches uninterruptedly across northern Tripolitania and along
the southern margins of the Gulf of Sirte, but in northern Cyrenaica,
it is markedly reduced, and in some places completely obliterated,
owing to the encroachment of the escarpments of the Cyrenaican

Plateau and Gebel Achdar. In most areas the coastal plain is slightly
elevated above the sea, and extensive sandy beaches occur near the
coastline. Occasionally, larger wadis emerging from the coastal escarp-
ment cause local irregularities in the surface of the coastal plain, but
normally it is relatively flat and rather featureless. In many areas
along the Gulf of Sirte, huge coastal dunes extend for great distances
along the seaward margins. Smaller, more localized dunes occur near
the coastal escarpment in the broad coastal plain of northwestern
Tripolitania.
In northern Tripolitania, where the coastal plain is frequently 40
or 50 kilometers in width, the higher, more inland portions are gently
undulating and support extensive olive groves and vineyards. Near
Misurata and Tauorga, along the western margin of the Gulf of Sirte,
the coastal plain is also quite wide and is partially submerged along
the seaward portions, forming an extensive salt marsh known as the
Sebchet el Tauorga.
The Coastal Escarpment

A coastal escarpment varying in height from a few meters to a
thousand meters is usually present, separating the coastal plain from
the higher deserts of the interior. These escarpments reach their most
impressive size in northern Cyrenaica and northwestern Tripolitania
where they mark the northernmost limits of the Cyrenaican Plateau
and the Tripolitanian Gebel (Gebel Nefusa). In many areas near the
Gulf of Sirte, the coastal escarpments are reduced to a line of indis-
tinct cliffs on the inner margin of the coastal plain. In other areas
the escarpment is lacking entirely, and the coastal plain grades
almost imperceptibly into transitional desert farther inland.
In some areas of Cyrenaica, the lower terraces of the coastal escarp-
ment are sometimes a hundred meters above the level of the sea
and form rocky headlands along their seaward margins.
Pre-Saharan or Steppe
A broad belt of pre-Saharan desert is typical of vast areas of Libya

located between the humid Mediterranean littoral and the truly
Saharan desert of the interior. These transitional deserts are generally

irregular in profile and are frequently dissected by numerous dry
watercourses (wadis) which drain northward into the Mediterranean.
Some of the larger wadis, such as the Wadi Bey, Wadi Soffegin, and
Wadi Zemzem of northern Tripoli a nia, form characteristic topo-
t
graphic features with their deep canyons, boulder-strewn slopes and

broad, eroded bottoms. Gravel plains (hamadas) of small extent are
not uncommon in these marginal deserts and are frequently inter-
spersed with mud pans reminiscent of playas. Sandy plains and dune
areas are present also but are usually sporadic and localized.
Farther southward, in the Sahara proper, these gravel plains and
sandy areas are of much greater extent and form discrete physio-
graphic entities which are known regionally by specific names, such
as the Hamada el Hamra of southwestern Tripolitania, the Idehan
Murzuch of the Fezzan, and the vast Sand Sea of Calanscio of eastern
Cyrenaica.
Hamadas
The terrain of most of the interior of Libya is typically Saharan,
being composed primarily of vast areas of pebble deserts or "hamadas"
interspersed among desolate sand seas or "ramleh." Mountainous
areas or "gebels" and rocky scarps frequently interrupt the otherwise
featureless landscape. Contrary to popular belief, these hamadas are
the dominant physical feature of the Sahara, the sandy plains (serirs)
and sand seas forming probably less than 20 percent of the total
land area.
The hamada desert is a distinct physiographic type characterized
by extensive gravel plains formed of pebbles of various sizes lying
on top of a rather firm substrate. The size of the individual particles
varies considerably in the different types of hamada, ranging from
small pebbles to rather coarse rock-strewn surfaces. Regional differ-
ences in the character of the are sometimes striking. For
hamadas
example, the Hamada cl Hamra
southwestern Tripolitania is of
of
the fine-grained type and provides probably the smoothest, most
featureless surface of any of the Libyan hamadas. The hamadas
farther south are much coarser in composition, especially those sur-
rounding the volcanic extrusions of the Gebel el Ilarug el Asued of
centra] Libya and the numerous localized hamadas which occur
between the oases of Traghen, Umm el Araneb, Meseguin, and
Zuila in the Fezzan. Some of these Fezzanese hamadas could be
likened to vast boulder fields.
The overall coloration of thesehamadas is generally gray, but

local variations of rod, yellow, and brown occur. The coarse hamadas
associated with volcanic extrusions are normally dark gray or even
RODENTS OF LIBYA 9
black. The majority hamadas are usually flat and lack any
of these
contrasting physical features. The coarser hamadas tend to have
more undulating; surfaces, and occasionally local mounds or rocky
outcroppings are present. In general, the hamadas show little change
in relief, being of comparable elevation throughout their full expanse.
Some of the larger hamadas stretch uniterruptedly for several hun-
dred kilometers and in this respect are remote and desolate.
Sand Seas
Although occupying a smaller total land area than the hamadas,
the sand seas or "ramleh" are one of the most distinctive physical
features of the Sahara. They occur sporadically in the interior of
Libya where they are contiguous with the hamadas and frequently
interdigitate with them. Small and localized sandy plains and dune
areas are present in the Libyan interior, but the principal sand seas
are represented by the Sand Sea of Calanscio of east-central Cy-
renaica; the Sand Sea of Rebianna, which is continuous with the
latter and extends to the northern outliers of the Tibesti Mountains
of extreme southern Cyrenaica; the great Idehan Murzuch of the south-
ern Fezzan and the Idehan Ubari of the southwestern Fezzan, which
includes virtually all south of the Hamad a de Tinrhert
of the region
to the Wadi Irauen and Wadi el Agial and includes the area from
Sebha west to the Algerian border. The "Ramleh Zellaf," immediately
north of Sebha, is the easternmost portion of this immense sand sea.
Normally these sand seas have gently undulating surfaces, but
frequently extensive dune areas occur. The individual dunes vary in
size, but some reach almost mountainous proportions, sometimes
exceeding 250 meters in height. Some of the dunes of the Sand Sea
of Calanscio are reputed to be even higher. In many areas the dunes
are relatively stable, but in others they are constantly changing
shape and shifting their positions. Firmness of the sand varies mark-
edly from place to place and according to the time of day, being
firmer during the cool, early morning hours than in the hotter periods
of midday.
Serirs
Occasionally, bedrock is covered with a veneer of sand and gravel

of varying depth. When these areas are quite extensive, they are
known as "serirs" and differ from typical sand seas by their more
uniform surfaces and the coarser, more heterogeneous nature of the
individual particles. The Serir of Calanscio, of central Cyrenaica,
typifies this physiographic type and includes the vast plain extending
from Gialo Oasis south to Tazerbo Oasis and extending from the
285-134 — 68^—2
western margins of the Sand Sea of Calanscio to the eastern limits

of the Gebel el Harug el Asued.
Fesh-fesh
In some areas of Libya, the surface layer is composed of extremely

softand unstable materials similar in texture to silt or loess deposits.
These "fesh-fesh" deserts are usually restricted in size but occasionally
cover large areas. This type of desert is most abundant in areas of low
elevation and along the margins of depressions such as the Sebchet el
Gheneien, north of Gialo, and Giarabub Oasis. The hamadas and
serirs sometimes grade imperceptibly into local "pockets" of fesh-fesh.
This intercalation of serir and fesh-fesh occurs frequently in the
western part of the Serir of Calanscio near the eastern limits of the
Gebel el Harug el Asued.
Depressions
The depressions associated with Giarabub Oasis and Bahr el Tubat

and the Sebchet el Gheneien north of Gialo are physiographically
distinct from the surrounding terrain. These depressed areas are
located great distances from the sea but are below sea level. These
low-lying areas of Cyrenaica represent the westernmost limits of the
great east-west depression which links together Siwa Oasis, theQattara
Depression, and the Wadi Natroun of northern Egypt. Most of these
low-lying areas have extensive salt marshes in their lowest portions,
and the larger depressions frequently contain saline lakes of con-
siderable size. Bahr
Tubat, east of Giarabub, is typical of this type
el
of shallow salty lake, which has thick encrustations of salts along its
margins, which in some places extend several hundred meters from
the waters edge. Smaller saline lakes occur in the oases of Tazerbo,
Cufra and Bzema of southern Cyrenaica, and isolated pockets, con-
taining some open water with marginal encrustations of salt, are of
sporadic occurrence in the central Fezzan.
Mountainous Areas of the Interior

Scattered throughout the interior of Libya are numerous rocky
escarpments, large wadis with steep rocky margins, and extensive
desert mountain ranges (gebels). Most of the mountainous areas of
the Libyan hinterland are characteristically rugged with sharp con-
tours and have broad canyons with angular rock formations and
precipitous walls. Isolated rock formations, which reach huge
some of
proportions, occur irregularly throughout the Fezzan and southern
Cyrenaica.
The mountain complexes of Libya are volcanic extrusions
larger
and are surrounded by great expanses of coarse, boulder-sized talus.
RODENTS OF LIBYA 11
The Gebel es Soda and the Gebel el Harug

Asued of central Libya
el
are mountains of this type. Both have been regarded

of these gebels
as the "Black Gebel" because of the color of the lava of which they
are composed. Other mountain complexes of the Libyan interior which
reach sizable proportions include the Acacus Mountains near Ghat in
the extreme southwestern Fezzan and the Gebel Archenu, Gebel
Uweinat, and the outliers of the Tibesti Mountains in southern
Cyrenaica. In many areas of the eastern Fezzan and southern Cyre-
naica, rocky formations are present in the form of buttes and mesas in
the midst of hamadas or sand seas. Cufra Oasis is surrounded by a
complex of steep-sided gebels, and Bzema Oasis, which is located deep
within the Sand Sea of Rebianna, is situated at the base of an imposing
monolith.
The Cyrenaican Plateau

The massif Cyrenaican Plateau of extreme northern Cy-
of the
and deep canyons, differs strikingly
renaica, with its high rolling slopes
from all other physical features of Libya. The Cyrenaican Plateau
proper encompasses the region of Cyrenaica from Benghazi in the
west to Martuba in the east. Its southern terminus is near El Mechili
where it grades imperceptibly into the inland hamada. The Gebel el
Achdar, which comprises the northernmost and highest portions of
the plateau, extends almost to the Mediterranean Sea where it
terminates in a rocky terraced escarpment. These escarpments pro-
vide a sharp contrast in relief to the low-lying hamadas and coastal
plain. In many places the coastal "gebel" encroaches onto the coastal
plain causing its virtual elimination. Deep canyons with rugged
precipitous slopes have their origin in the interior of the plateau and
dissect the coastal escarpment.
The highest point in the plateau is 2,844 feet above sea level lo-
cated on the Gebel el Achdar between Slonta and El Faidia. The
higher portions of the Cyrenaican Plateau have characteristic broad,
rolling slopes,and the canyons are less rocky and have less precipitous
margins than those nearer the coast. Some areas of the plateau are
almost flat and allow sizable agricultural endeavors. The Barce Valley,
in the western part of the plateau, is a basin of internal drainage and
is unique in Libya in this respect. During periods of unusually high
rainfall, fresh water accumulates in the bottom of the valley to a

depth of several feet and thus forms a rather sizable lake.
The Tripolitanian Gebel
The Gebel Nefusa of northwestern Tripolitania is less clearly de-

fined by discrete physical elements. It includes the high tablelands
extending roughly from the Tunisian border in the west to the vicinity
of Cussabat near the western edge of the Gulf ofSirte. The southern
portions of the gebel grade imperceptibly into the hamadas and
transitional desert north of the Hamada el Hamra. A decreasing
gradient in elevation runs from north to south, thus the highest
portions of the gebel are on its northern margins where it terminates
abruptly in the coastal escarpment.
The average elevation of the Gebel Nefusa exceeds 2,000 feet and
in many areas approaches 3,000 feet. The highest point in the gebel
(3,176 feet above sea level)is located between Gharian and Bern* Ulid.
South of the rugged coastal escarpment the interior of the Gebel

Nefusa transforms into gently, undulating tablelands with broad
valleys and smooth slopes. Several major and numerous lesser wadis
have their origins in the interior of the Tripolitanian Gebel. Chief
among these are the Wadi Soffegin, which drains eastward into the
Gulf of Sirte; the Wadi Tmasia, which has its origin in the gebel
north of the Wadi Soffegin; and the Wadi Caam, which arises in the
northeastern portions of the gebel and empties into the Mediterranean
Sea east of Tripoli.
Geology
Tripolitania and Cyrenaica
This discussion of the major geological features of Libya is based
on information given by Raymond Furon in "Geology of Africa"
(1960).
A succession of Mesozoic and Paleozoic sandstones, ranging from
the Permian to the Jurassic, outcrops at El Azizia on the Gefara Plain
south of Tripoli. These systems are continuous with those of Tunisia.
The "Continental Intercalaire," of Triassic, Jurassic, and Cretaceous
age, outcrops at the base of the coastal escarpment near Horns,
Tripolitania, and extends westward nearly to the Tunisian border.
This formation is composed of sandstones and clays with numerous
plant remains, including silicified wood. The "Continental Intercalaire"
extends south to the Tibesti Mountains, but in southern Tripolitania
and the northern Fezzan, it is overlain by the Cretaceous marine
limestones of the Hamada el Hamra and the Hamada de Tinrhert.
Marine transgressions invaded Tripolitania and Cyrenaica during
the Cretaceous and at successive periods of the Tertiary. The deposits
of the Cretaceous form the Hamada el Hamra and the Hamada de
Tinrhert in Libya and the Tademait Plateau of the Algerian Sahara.
These marine limestones are highly fossilil'erous, containing a variety
of marine organisms (corals, nerineas, and radiolarians), and extend
south to the cliffs on the northern margin of the Fezzan Cuvette.
To the east they are overlain by Tertiary deposits.
RODENTS OF LIBYA 13
The Eocene transgression outcrops widely north and east of the

Gebel es Soda and also appears near the coast on the Gebel el Akhad
east of Benghazi. At El Fogaha, south of the Gebel es Soda, and on
the Gebel el Achdar near Derna, the succession is complete, ranging
from sediments of lower Eocene to upper Eocene.
Another transgression advanced to the Tibesti Mountains during
the middle Eocene. These sediments, comprising 200 meters of lime-
stone overlying the Paleozoic sandstones or resting directly upon
the Pre-Cambrian, represent the Sirtica Trough, whose shorelines are
found today at a height of 600 meters above sea level on the northern
mountains.
outliers of the Tibesti
Oligocene sediments indicate a deep gulf south of the Gulf of Sirte,
which did not reach as far south as that of the Eocene.
The Miocene Sea included parts of Cyrenaica, and some lower
Pliocene beds are found at Agedabia near the southeastern margins
of the Gulf of Sirte.
Fezzan
The geological structure of the Fezzan is relatively simple, being
primarily a large cuvette (basin) of Nubian Sandstone (Cretaceous)
overlying the Paleozoic. The Fezzanese cuvette is divisible into two
large structural components: a southern synclinal zone which is an
extension of the covering of the Ahaggar Mountains of southeastern
Algeria and the Tibesti Mountains of the northern Chad, and a
northern zone consisting of the Cretaceous Hamada de Tinrhert in
the west and Eocene sediments in the east. Extensive volcanic ex-
upon portions of the northern
trusions rest zone.
The morphology of the western Fezzan determined by a large
is
east-northeast west-southwest anticline, which is connected in the
southwest to another anticlinal zone near Ghat. This anticline extends

from north of Serdeles to north of Edri. Farther west, it grades into
the Fezzan synclinal cuvette and forms an artesian basin. The northern
and southern limits of the anticline outcrop, respectively, as south-
facing escarpments near El Hasi north of Edri, and as north-facing
escarpments in the southern parts of the Wadi es Sciati.
In the western Fezzan, geological succession generally runs from
Devonian to Lower Cretaceous and includes fossiliferous limestone,
sandstones, and marls containing brachiopods, crinoids, and silicified
wood.
The northeastern Fezzan is composed of Cretaceous rocks which
disappear beneath Tertiary deposits farther east. Farther south, strata
range from Pre-Cambrian to Cretaceous, but Paleozoic sediments
are dominant.
Near Murzuch the Cretaceous Nubian Sandstone outcrops, forming

the Murzuch Cuvette. The "Murzuch Limestones" (Cretaceous) form
a series of limestone plateaus of lacustrine origin and contain gastro-
pods. The plateaus are deeply eroded and are overlain by deep
Quaternary deposits.
East of the basaltic Gebel el Harug el Asued, marine deposits are
covered by the "Continental Terminal," which contains the remains
of saurians, tortoises, fishes, and silicified woods.
The Gebel Uweinat is a mountain massif located near the Libyan,
Egyptian, and Sudanese borders. Basally, it is formed of middle
Pre-Cambrian crystalline schists, which have been intruded by
granite, which forms the greater part of the Gebel. Following mag-
matic activity, a basal layer of Carboniferous sandstones was de-
posited, followed by several layers of continental sandstones containing
silicified wood.
Volcanic Activity
Volcanic activity occurred in three regions of Libya: the Gebel

Gharian (Gebel Nefusa) south of Tripoli, the Gebel es Soda south of
Socna, and the Gebel el Harug el Asued of southeastern Tripolitania
and the northeastern Fezzan. These eruptions began in the Oligocene
and continued until the Recent. Volcanic extrusions of the Gebel el
Harug el Asued cover an area of 25,000 square kilometers and are
represented by two major basaltic divisions. The first eruptions
inundated the Eocene Plateau and today form an extensive lava field.
A plateau of scoriaceous basalt of more recent origin occupies the
center of the massif. The most recent basaltic flows have been de-
posited by volcanic cones, which still persist but are inactive. At
present, there is no volcanic activity in any of these areas in Libya.
Climate
Libyan Interior
Two climatic subdivisions of the Libyan interior can be recognized:
an arid or desert type, and a semiarid or steppe type.
Arid or Desert Type

The Saharan portion of Libya is typical of the low latitude desert
as regarded by Trewartha (1937), and unless stated otherwise, the
following meteorological data relating to Libya was obtained from
this source.
In the true Saharan portions of Libya, rainfall is always meager
and extremely variable from year to year. In these areas, the variabil-
ity of rainfall shows a 40+ percent departure from the normal, so
it is meaningless to speak of a "typical" rainfall curve for the Libyan
Sahara. Many areas in the interior of Libya receive less than one inch
RODENTS OF LIBYA 15
of rain per year, and some parts of the Fezzan receive no rainfall at
all, sometimes extending over many years.
Most precipitation results from violent convectional showers,
which usually are limited to small areas. During the winter, there
are occasional widespread rains of a cyclonic rather than convectional
origin in the northern portion of the Libyan Sahara.
Skies are most frequently clear in the Libyan interior, particularly
during the winter months (over much of the Sahara, December and
January have a cloudiness of only 1/10).
Relative humidity is almost always low in the Saharan interior
of Libya (12-30% for the midday hours) and evaporation is extremely
,
high. Relative humidity as low as 2 percent, with temperatures of

100° F., has been recorded from the Egyptian Sahara, and doubtless
similar conditions prevail in the Libyan Sahara. Even though the
desert air is physiologically dry and does not approach saturation,
absolute humidity is quite high because the hot desert air contains
a considerable quantity of water vapor. Potential or actual evapora-
tion, owing to high temperatures and low relative humidity, is exces-
sive, often being 20 or more times the amount of precipitation.
In the Libyan interior it is the excessively high temperature in
summer, rather than the cold winter temperature, which is responsible
for the marked differences between the seasons. Dry air, cloudless
skies, and bare, dry earth which produce relatively large temperature
differences between the extreme months are also responsible for pro-
ducing marked temperature variations within a 24-hour period.
During the high-sun period (summer) intense diurnal heat prevails.
In the northern Sahara average daily maxima of 99° are followed by
minima approximating 71° F. The highest air temperature (136.4° F)
ever recorded anywhere in the shade under standard conditions was
from El Azizia on the Gefara Plain south of Tripoli.
During the period of low-sun (winter), the days are still warm;
the daily maxima usually average 60° to 70° F, but occasionally reach
80° F.
As is true with the rest of the Sahara, the sun largely controls
local weather, resulting in a great deal of similarity between successive
days.
The Saharan portion ofLibya tends to be windy owing to the
lack of obstruction of the moving air by the sparse vegetative cover.
Also the rapid daytime heating of the lower air tends to convectional
overturning. Nights are generallymuch less windy owing to the rapid,
nocturnal convectional cooling of the surface air.
Semiarid or Steppe Type
A region of low latitude steppe climate characteristically bounds

the true Saharan portion of Libya to the north and is a transitional
beltbetween it and the humid climate of the Mediterranean littoral.

These steppe areas in Libya are located on the margins of the trade
winds and subtropical highs and are hence nearer to the humid climates.
These areas are subjected to brief periods of winter, rain-bearing winds
and their associated storms (the westerlies and their cyclonic storms),
which cause them to be semiarid rather than arid.
At Benghazi, near the northern limits of the steppe region of
Cyrenaica, no rainfall occurs during the months of June, July, and
August, while December and January receive moderate amounts of
rain. The steppe thus has almost all of its rain in the winter; the
yearly total is 11.9 inches. The amount of precipitation varies from
year to year from a recorded high of 24.3 inches to a low of 6.8 inches.
Variability of rainfall is as great in the steppe region as in the true
Saharan region. Because rain falls in showers of comparatively short
duration, the weather is prevailingly sunny.
Mediterranean Littoral
Most portions of coastal Libya have a Mediterranean or dry-summer
subtropical climate. This climate, according to Trewartha, is character-
ized by most precipitation in the winter
three principal features:
season; warm to extremely hot summers and mild winters; and a
high percentage of sunshine for the year, especially in the summer.
This Mediterranean climate in coastal Libya results from the
dought-producing dry subtropical highs and tradelike winds on the
south and the humid westerlies with their cyclonic storms on the north.
As a result of the north-south shifting of these wind belts, these
Mediterranean latitudes are joined at one season to the dry tropics
and at the opposite season to the humid middle latitudes.
A uniform summer and a variable winter climate characterize these
coastal areas. The climate of coastal Libya is transitional between
the low-latitude steppe and desert and the cool, humid climates
farther poleward.
It is the relative warmth of the Mediterranean Sea in winter, and
the resulting low-pressure trough coincident with it, that attracts
cyclonic storms in the winter season. The Mediterranean climate of
coastal Libya is thus assured of a temperature regimen in which
cold weather is largely absent. Winter months have average tempera-
tures of between 40° F and 50° F, and the summer months between
70° F and 80° F, and mean annual ranges of 20° to 30° F are common.
At Tobruch, on the northeastern Cyrenaican coast, the average
January temperature is 56° F, while the average for July is 79° F.
The average annual temperature at Benghazi on the coastal plain on
on the Gulf of Sirte is 69° F with averages of 55° in January and 79°
in August.
RODENTS OF LIBYA 17
On the coastal plain, rainfall generally approaches 15 to 25 inches.

The most characteristic feature of the climate of coastal Libya is the
pronounced summer drought. If this amount of rain fell during the
hot summers when evaporation is high, semiarid conditions would
result. Coming as it does, however, in the cooler season, much less is
evaporated, and Mediterranean climate of Libya can be described as

subhumid rather than as semiarid.
A peculiarity of the climate of northern Tripolitania is the "Ghibli,"
a hot, desert wind which can cause temperatures to rise 30° to 40° F
in both summer or winter. The "Ghiblis" raise the temperatures of
the coastal plain well above 110° F, and sometimes the strong winds
persist for several days carrying large quantities of dust into the
coastal regions.
Coastal Uplands
On the Cyrenaican Plateau and Tripolitanian Gebel, the summers

are more moderate than those of the coastal plain. These plateau
areas of Tripolitania and Cyrenaica, by virtue of their higher eleva-
tions, receive more rainfall than the surrounding areas. The Gebel
Nefusa of northern Tripolitania receives a yearly average of 15 to
20 inches, most of this falling during a comparatively short winter
period. The Cyrenaican Plateau has approximately 30 inches of rain
distributed rather uniformly throughout the year.
Additional Meteorological Data
Meteorological data obtained during field work in 1961 and 1962

provides supplemental information regarding the nature of the Libyan
climate. In the Fezzan, during the period from Dec. 8, 1961, to
Feb. 1962, the daytime temperatures remained quite high with
10,
average maximum temperatures of 94° F (60°-117° F), but at night
the temperature sometimes dropped below freezing, and the average
minimum temperature was 36.1° F (27°-52° F). The temperature
range for a 24-hour period sometimes exceeded 80° F. On the Gebel
Nefusa of northwestern Tripolitania, for a 4-day period in early
March 1962, the averages and extremes of the maximum and mini-
mum temperatures were 91° F (83°-98° F) and 52.5° (45°-59° F).
Averages and extremes of maximum and minimum temperatures at
Cufra and Tazerbo oases for the period Apr. 1, 1962, to Apr. 17, 1962,
were 98.4° F (83°-110° F) and 53.3° F (44°-68° F). During this same
period the average relative humidity (taken in the evening) was 21
percent. For the periods of May 21 to May 25, June 1 to June 6, and
from June 12 to June 21, 1962, the averages and extremes of the
maximum and minimum temperatures and the average relative
humidity, at various localities along the Cyrenaican and Tripolitanian
coastal plains, were, respectively: 113.7° F (108°-120° F), 62° F (53°

F-74° F), and 74.2 percent. The above temperatures were taken during
the "summer drought" on the coastal plain and probably represent
the highest figures that would be obtainable during the year. Those
from the interior were taken during the winter or low-sun period.
Direct comparison of the data from the coastal plain with that
obtained in the Fezzan and southern Cyrenaica would lack validity,
as the two sets of data represent, respectively, the two extremes in
the annual cycle of the Libyan climate.
The above data indicate that the Saharan interior is markedly
more arid than the coastal region and undergoes greater temperature
variation for any given 24-hour period.
Phytogeography
General Features of the Saharan Flora
The following information regarding the phytogeographic charac-

teristics of the Sahara, including Libya, has been compiled from:
Ozenda, "Flora du Sahara" (1958), which includes a comprehensive
treatment of the flora of the northern and central Sahara; Zavattari,
"Prodromo della Fauna della Libia" (1934), which treats briefly the
regional Libyan flora; and Rattray, "The Grass Cover of Libya"
(1960), in which Libya is divided into several regions according to the
dominant type of grass cover.
According to Ozenda, the flora of North Africa, including the
Sahara, comprises portions of two major floral empires: the Holarctic
Floral Empire which includes all of Europe, North Africa (including
the northern Sahara), and the greater part of Asia; and the Paleo-
tropical Floral Empire which encompasses the central and southern
Sahara and the remainder of Africa.
Coastal Libya falls within the Mediterranean Floral Region, and
the Saharan interior of Libya belongs to the Saharo-Sindien Floral
Region, which also includes northern Arabia and southern Iran, and
extends as far east as the Sind Desert of West Pakistan. The flora of
the Mediterranean Floral Region and the Saharo-Sindien Floral
Region, and hence that of Libya, has most of its affinities with the
Holarctic Floral Empire.
The Saharo-Sindien floral element is always dominant and contains
three-fourths of all the plant species that occur in the Sahara. The
Mediterranean affinities are naturally well developed along the north-
ern margins of the Sahara in Libya. A Mediterranean element exists
even in the mountains of the Central Sahara, but these plants are
Mediterranean only in origin, not in their biology.
.
RODENTS OF LIBYA 19
Phytogeographic Characteristics
Ozenda summarizes the phytogeographic characteristics of the

Saharo-Sindien Floral Region, which includes most of Libya, as fol-
lows :
(1) Great poverty of species (of the nearly 1,500 species of plants
present, 1,000 occur in the Sahara). In the Sahara, there are 150 species
of vascular plants per 10,000 square kilometers, as opposed to 1,000 to
2,000 in an area of comparable size in Europe. In the tropics the num-
ber of species increases to 3,000 or 4,000 per 10,000 square kilometers.
(2) Extreme sparseness of individual plants, the vegetation being
thinly scattered.
(3) Monotony of the country and of the plant groups (one type or
species of plant may occupy a vast area).
(4) Absence of characteristic systematic groups above the genus
(there are no families or tribes that are most typical of the Sahara, but
several species and genera are typical of the region)
(5) The presence of a large number of endemic species.

(6) Human economy based on the culture of the date palm.
Cryptogamic vegetation is relatively sparse in the Sahara and is
represented by approximately 100 species of mushrooms, lichens,
algae, liverworts, and mosses; they comprise approximately 12 to 18
percent of the floral composition. Bacteria and filamentous mushrooms
have developed even in barren soils which contain no phanerogamic
vegetation.
Systematic Composition
The Gramineae, Leguminosae, and the Compositae are considered by

Ozenda to represent the dominant families of the Sahara. Even though
these three families represent about 35 to 40 percent of the floral com-
position of the Sahara, they are not the most typical Saharan families.
The Chenopodiaceae with two endemic genera, Fredolia Coss. and Dur.
and Nucularia Batt. the Cruciferae represented by numerous genera
;
and species; and the Zygophyllaceae including the genus Fagonia L.,
are the most typical families of plants in the Libyan flora. Representa-
almost nonexistent in the Mediterranean
tives of the latter family are
littoral. Cruciferae and Chenopodiaceae, however, have numerous
The
representatives in the Mediterranean region and the entire Holarctic
Floral Empire.
Generic dominance, with the number of species in each genus of the
plants of the northern Sahara, including Libya (Fezzan), is as follows:
Astragalus L., 8; Fagonia L., 8; Aristida L., 8; Launaea Cassini, 7;
Reseda L., 6; Salsola L., 6; Plantago L., 6; Tamarix L., 5; Euphorbia L.,
5. These genera belong to the families Gramineae, Leguminosae, Com-
positae, Zygophyllaceae, Resedaceae, Chenopodiaceae, Plantagin-

aceae, Tamaricaceae, and Euphorbiaceae.
Owing few species (one or two) representing each genus of the
to the
Cruciferae in the Sahara, this family is not included with the above
account of generic dominance, which is based on only those genera con-

taining five or more species. Because of the large number of Saharan
genera comprising the Cruciferae, it is still regarded as one of the
dominant families.
Endemism
In the Libyan Sahara, the presence of vast spaces almost unfit for
life constitute barriers to the dissemination of species, and endemism is
particularly well developed. Some plants are endemic to particular
geographic formations (mountains, sand seas, and hamadas). Other
species have a wider range of endemism and are endemic to the entire
Sahara; therefore, all degrees of endemism are possible.
Specific endemism attained in the Sahara is in the neighborhood of
25 percent of the total component plant species (162 endemic species
of vascular plants out of a total of 650 species). Ozenda lists 35 species
of endemic vascular plants in the northern Sahara, many of which
occur in Libya.
Adaptation to the Desert Environment

The problem of adaptation of plants to a desert climate involves
primarily an adjustment to long dry periods. A large proportion of
desert plants shorten their fife cycle and thereby suppress their aerial
parts during periods of drought. Others maintain their aerial parts
but have an arrangement of anatomical devices which tend to diminish
water loss by evaporation.
In the Sahara, two types of vegetation can be distinguished accord-
ing to the length of their life cycles: temporary vegetation and
permanent vegetation.
The temporary vegetation is divisible into annual plants and the
geophytes. Following the rains, annual plants appear suddenly and
develop with amazing rapidity, completing their life cycle, including
flowering and fruiting, before the soil becomes dry. The length of the
vegetative cycle is variable among the different species but generally
is about four months. Some species of the genera Boerhavia L. and
Tribvlus L. complete their development from germination to the
production of seeds in 8 to 15 days. Additional examples of Saharan
ephemerophytes include Convolvulus fatmensis Kunze, Launaea glome-
rate (Cass.) Hook, Schismus barbatus (L.) Thell., and Plantago albicans
L. These ephemeral plants are important as sources of food for
domestic animals.
RODENTS OF LIBYA 21
Geophytes are plants which are able to survive periods of drought

by the development of enlarged subterranean organs such as bulbs
and rhizomes. These are represented chiefly by members of the
Umbelliferae.
Permanent vegetation has adapted to the aridity of the desert by
the development of anatomical and morphological structures which
increase the amount of water absorption and decrease evaporating
surfaces. In general, a high level of absorption is insured by the
maintenance of greater hypertonicity in the roots than in the aerial
parts. The roots of these desert plants are frequently elongated and
specialized to absorb water more efficiently in any given type of soil.
The problem of water retention in desert plants has been solved by
a reduction in the evaporating or transpirating surfaces, a reduction
in the speed of evaporation, and by the accumulation of water in the
tissues. Species belonging to the family Chenopodiaceae and to the
genera Ephedra L. and Calligonum L. have minute flowers and are
even apetalous. In the case of Anabasis articulata Moq., Tamarix
gallica L., and Calligonum, the leaves drop off during the dry season.
In other plants (sclerophy tes) the epidermis is covered with a thick
,
cuticle formed of sclerified cells, which retard significantly the speed

of evaporation or transpiration. Succulent plants which store water in
the tissues are represented in the Sahara by relatively few species
belonging to the genera Mesembryanthemum L., Euphorbia, and
Caralluma R. Br.
In the northern Sahara, including the steppe and deserts of Libya,
there are approximately 20 species of trees or arborescent plants.
These include such well-known genera as Ephedra, Cupressus L.,
Phoenix L., Ficus L., Acacia Mill., Pistacia L., Tamarix, Calotropis
R. Br., Olea L., Calligonum, Rhus L., and certain arborescent genera of
the Chenopodiaceae.
The Vegetative Cover of Libya
Coastal Plain. and density of plants of the

Floral composition
Libyan coastal plain varies considerably from region to region de-
pending upon the character of the soil and upon the season. Generally,
the vegetative cover of the coastal plain is denuded by overgrazing
but occasionally it is quite dense and uniform. The most typical
pattern consists of extensive areas of small hillocks supporting bush-
like plants. Emergent grasses and annuals are sometimes interspersed
among these hillocks. Zavattari
(1934) states that the following
genera are the most typical of the coastal plain: Hyparrhenia Anderss.,
Imperata Cyrill., Agropyron L., Cyperus L., Pancratium L., Silene L.,
Euphorbia, and Ononis L. In sandy areas of the coastal plain, the
vegetative cover is sparser and consists of larger, bushlike species
such as Calligonum. The large coastal dunes also support a meager

vegetative cover.
In those portions of the coastal plain where salt concentration is
relatively high, halophytic species are dominant. Certain species
such as Crithmum maritimum L., Lotus argenteus Webb, Beta maritima
L., Dactylis glomerate, L., and Agropyron junceiforme (A. and D.
Love), indicate transitional areas in the gradient of salt concentra-
tion of the soil. Considerable areas of salt marshes exist in Tripoli-
tania near Zuara, Misurata, Tauorga, Agedabia, and in various
other localities along the coastal plain of northern Cyrenaica ad-
jacent to the Mediterranean Sea. These areas are characterized by
typical halophytic genera, which include Salicornia L., Suaeda Forsk.
Arthrocnemum Moq., Halocnemum Marsch.-Bieb., and Salsola L.
The coastal plain near Tocra in Cyrenaica consists of a broad
bush-covered plain which differs markedly from the typical vege-
tative cover elsewhere on the coastal plain. In coastal Tripolitania,
large wadis dissect the coastal plain and frequently are bordered
by thorny, bushlike perennials, some of which occur on large
hummocks.
Many areas of the coastal plain, particularly those of northwestern
Tripolitania, have been modified by man and support extensive
groves of olives and date palms and vineyards. Eucalyptus trees
and bull-thorn acacia have also been planted along many of the
roadsides. The vegetative cover of large portions of the Gefara plain
of northwestern Tripolitania is not typical of that of the coastal
plain and covered instead with a shrub-steppe type of vegetation.

is
Cyrenaican Plateau. The lush and varied flora of the Cyrenaican

Plateau is predominantly of Mediterranean affinities and consists
of a variety of arborescent species, grasses, and other herbaceous
plants. Most of the canyons and broad slopes of the plateau are
covered with dense chaparral consisting of several genera of treelike
forms. These genera include primarily Quercus L., Viburnum L., and
Juniperus L. Some of the larger wadis, such as the Wadi el Kuf,
support occasional stands of Pinus L. and Cupressus L. In the higher
portions of the plateau, especially along the northern portions, a
savanna type of vegetative cover is present, consisting of grassy
plains with occasional juniper or cypress trees. In those areas where
agriculture is widespread, the grass cover is interrupted by open
fields and cultivated areas. Numerous other genera of plants occur
on the plateau either as understory vegetation or as perennials as-
sociated with agricultural areas. Narrow strips of mesophytic vege-
tation border the few permanent streams which drain from the
plateau, and small pockets of mesophytic plants occur near springs
RODENTS OF LIBYA 23
and other sources of water. The genera most frequently found in

these mesic habitats are: Typha L., Potamogeton L., Phragrnites Trim,
and Cyperus L.
Tripolitanian Gebel. The composition of the Tripolitanian
floral
Gebel (Gebel Nefusa) differs markedly from that

of the Cyrenaican
Plateau and has greater affinities with the flora of the Saharan steppe.
The vegetative cover of the Tripolitanian Gebel is sparser and con-
sists of a smaller assemblage of species. Grasses, chiefly Stipa L.,
constitute the dominant element in the plant cover. Other plants
present on the Gebel, which are considered more typical of the
Saharan steppe, include Teucrium L., Lavandula L., and Rhamnus L.
Olive trees (Oka) occur in the less rugged areas of the Gebel, and date
palms (Phoenix dactylifera L.) are of localized occurrence on the
terraces of the coastal escarpment.
Saharan steppe. The Saharan steppe, or transitional desert of Libya,
extends as a broad between the denser
belt of scrub desert vegetation
plant cover of the coastal plain and the extremely sparse vegetative
cover of the Saharan interior. This semiarid region contains plant
species typical of both the Mediterranean and Saharo-Sindien Floral
Regions. In any given area of transitional desert, the vegetation tends
to be rather uniform and frequently large areas are dominated by
specific types. Several different types of steppes can thus be dis-
tinguished in Libya based upon the dominant plant occurring in each.
Chief among these types of steppe are the Asphodelus L. steppe, the
Artemisia L. steppe, and the Aristida-Stipa L. steppe. In the steppe
region south of the Cyrenaican Plateau and near the Gulf of Sirte,
the genera Asphodelus and Lygewm L. are common. The most widely
spread plant genus of the Libyan steppe is Aristida. The vegetative
cover of this semiarid region sometimes extends uninterruptedly for
200 kilometers or more but becomes more sporadic and localized near
the northern margins of the Sahara proper. In this steppe region,
acacia trees (usually Acacia radianna Savi) are not uncommon and
occur either singly or as small groups along the margins of the larger
wadis.
Sand seas, hamadas, and In the sand seas the grass, Aristida
serirs.
pungens Desf., is the dominant herbaceous plant. In areas where the
sand has become stabilized, plant cover is characterized by the follow-
ing arborescent genera: Ephedra, Retama Boiss., Genista DC, and
Cailigonum. Cyperus and Mottkia Lehm. are the most abundant
herbaceous plants.
The sparse vegetative cover of the hamadas consists of several
characteristic generawhich include Haloxylon deBunge, Fagonia, and
Fredolia Coss. and Dur. Following periods of rain, a variety of annual
plants develop and sometimes extend over large areas of the hamadas.
These annuals are represented by Erodium L'Herit, Lifago Schweinf.
and Muschl., Convolvulus L., and Urginea Steinheil.
In the serirs, where the surface layer is sandy, grasses belonging to
the genus Aristida constitute the major vegetative cover. The floral
cover of these sandy plains is generally denser and more varied than
that of the hamadas. The following genera are most common Andro- :
cymbium Willd. and Asphodelus, representing forms with underground

bulbs or tubers; Daucus L. and Ammodaucus Coss. and Dur., which are
annuals; Cornulaca Del., a thorny chenopod; and Randonia Coss., a
bushlike perennial. Wild gourds, Colocynthis vulgaris (L.) Schrad.,
occur in both hamadas and serirs.
Depressions or sebchets. The large saline depressions or sebchets in
various parts of Libya have a distinctive flora consisting of salt-tolerant
representatives of the Chenopodiaceae, such as Salsola, Traganum
Del., Atriplex L., Cornulaca, and Suaeda. Other families are repre-
sented by Zygophytlum L. and Frankenia L.
Sixty-five species of herbaceous and woody plants are recorded from
the vicinity of Giarabub Oasis, many of which belong to well-known
salt-tolerant genera and are capable of existing under conditions of
marked The most salt- tolerant of these include Chenopodium
alkalinity.
L., Arthrocnemum, and Salsola. In some of the larger sebchets, several
successional stages are apparent in the flora extending from areas of
high-salt concentrations to those of lower concentrations. Along the
margins of the depressions where the soil is less saline, tamarix and
date palms are sometimes abundant. Near the saline lake of Bahr el
Tubat in eastern Cyrenaica, dense growths of Phragmites occur on
the moist, sandy-clay soils near the margins of the zone of true
halophytic plants.
Oases. Owing to their high water tables, the oases in the interior of
Libya support a lush flora entirely unlike that of other areas. Large
groves of date palms (Phoenix dactylifera) are the dominant vegeta-
tive feature of the oases, but numerous other plants of agricultural
significance, such as figs, wheat, and alfalfa, are also present. On the
periphery of the oases, and to a lesser extent in the interior, tamarix
is relatively common. This outer zone of tamarix is typical of almost
all larger oases in Libya. In these outlying areas tamarix forms
clones of considerable size on sandy soils and occurs sparingly on top
of large sandy-clay hummocks. Frequently, Calligonum occupies these
same hummocks. The hummocks supporting Tamarix and Calligonum
frequently occur for 20 kilometers or more beyond the palm groves
of the oases.
In many large oases, such as Cufra, Tazerbo, and Bzema, small
lakes (mostly saline) occupy the central portions and are bordered
RODENTS OF LIBYA 25
by dense growths of sedges and other mesophytic plants. In those

oases where the water table is near the surface, sedges (Cyperus and
Scirpus L.) occur over an appreciable portion of the oasis. Besides
sedges, the most abundant species of emergent vegetation are Poly-
pogon monspeliensis (L.) Desf., Erianthus ravennae (L.) Beauv.
Imperata cylindrica (L.) Beauv. Phragmites communis Trim, Desmo-
stachya bipinnata (L.) Stapf., Juncus bujonius L., Juncus maritimus
Lam., and Typha angustifolia L. At the oasis of Brach in the Fezzan,
fresh water is abundant and supports dense growths of Phragmites.
In the broad, shallow wadis of the Fezzan, date palms and tamarix
occur almost uninterruptedly for stretches of 50 kilometers or more
and, in effect, have all of the floral characteristics of the isolated
oases. The oases in the Wadi
es Sciati, north of Sebha, and those of
Murzuch, Traghen, Umm el Araneb, Meseguin, and Zuila are linked
together by scattered palms and tamarix.
In the Wadi el Agial, southwest of Sebha, groves of date palms
extend uninterruptedly for over 150 kilometers.
The grass cover of Libya. The grass cover of Libya has been divided
by Rattray (1960) into six regions based on the dominant type of
grasses present. The Phalaris L. type is associated with a woodland
habitat and is limited to the uplands of the Cyrenaican Plateau.
Several other species of grasses are also commonly associated in this
Phalaris type of grass cover. These include Hordeum bulbosum L.,
Poa bulbosa L., Lolium rigid um Gaud., Oryzopsis miliacea (L.) Aschers.
and Schweinf., and Dactylis glomerata.
Rattray includes the coastal fringes and littoral deserts of Libya
within the Hyparrhenia type of grass cover. Because of intensive
utilization in this region, the vegetation has been reduced to a shrub
steppe, but in protected areas, Hyparrhenia hirta (L.) Stapf. is the
dominant grass. The grasses Oryzopsis miliacea and Cynodon dactylon
(L.) Pers. are also common in the Hyparrhenia zone which occurs at
altitudes ranging from sea level to over 600 feet and receives from
8 to 16 inches of rainfall during the winter season.
According to Rattray, a small portion of the Gefara Plain of
northwestern Tripolitania belongs to the Stipa lagascae Roem. and
Schult.-tSîpa parviflora Desf. type of grass cover. This is a sparsely
growing perennial grass cover interspersed among a shrub-steppe
type of vegetation consisting of Ziziphus lotus (L.) Desf., Artemisia
campestris L., and Rhanterium suaveolens Desf. This type of grass
cover occurs at elevations ranging from sea level to 650 feet and with
a winter rainfall of 6 to 10 inches.
The Stipa tenacissima L.-Lygeum spartum Loefl. type of grass cover
represents a perennial grassland on the Gebel Nefusa and adjoining
portions of the Tripolitanian Gebel. This association usually occurs
285-134 O —68 3
;
with a discontinuous cover of small shrubs, primarily Artemisia.

The large grass Stipa tenacissima, known locally as "esparto" grass,
is harvested and exported on a fairly large scale in some parts of coastal
Libya.
Grass cover of the Arislida type is dominant in all desert areas of
Libya. The Aristida plumosa L.-A obtusa Del. -A acvtifolia Trin. and
Rupr.-A ciliata Desf.-A pungens Desf. "type" is typical of the pre-
Saharan or transitional desert area of northern Tripolitania and
Cyrenaica. These grasses are also associated with a steppe type of
vegetation consisting of small, scattered groups of acacia and desert
shrubs. Stipa lagascae, S. capensis Thunb., and S. barbata Desf. are
present but are not common in this zone.
In the Libyan Sahara, where the annual rainfall is less than two
inches, the Aristida spp.-Panicum turgidum Forsk.-Lasiurus hirsutus
Boiss.-Pennisetum divisum (Gmel.) Henrard "type" of grass cover is
most characteristic. In the Sahara, these grasses are associated with
scattered acacias and shrubs including such genera as Anabasis L.,
Haloxylon, and Cornulaca. Frequently vast areas are devoid of vegeta-
tion, and only an ephemeral cover of grasses and herbs appears follow-
ing a period of rainfall.
Mammalian Faunal Areas

Libya has three primary faunal areas based on the distribution of
the rodent fauna. These areas coincide roughly with the major physi-
ographic, vegetative, and climatic features of the country. These
faunal areas are divisible into regional provinces according to the
kinds of rodents most typical of each. These areas (with the provinces
in parentheses)
are: Mediterranean (Coastal Plain, Cyrenaican
Plateau) Saharan steppe (Transitional Desert, Tripolitanian Gebel)
;
and Saharan Desert (Cyrenaican Desert, Fezzanese Desert).

Each faunal area is characterized by certain species (or subspecies)
restricted to it or which occur only marginally in an adjacent faunal
area. Members of a given subspecies are usually confined to the same
province, but occasionally representatives of a widely ranging sub-
species can occur in both provinces of a faunal area. Representatives
of genetically plastic subspecies frequently occur in more than one
faunal area, but these are exceptional cases.
Owing to the diverse topography of Libya, a greater number of
mammalian faunal zones could be expected. The aridity of the Saharan
interior apparently renders most areas unfit for life, and as a result,
only the most adaptable species of rodents are able to survive. Be-
cause the Saharan portion of Libya comprises most of the land area
and because widespread distribution of most of the desert
of the
species, the faunal areas are correspondingly large.
RODENTS OF LIBYA 27
Figure 2. — I, Mediterranean Faunal Area; IA, Coastal Plain Province; IB, Cyrenaican
Plateau Province; II, Saharan Steppe Faunal Area; IIA, Transitional Desert Province;
IIB, Tripolitanian Gebel Province; III, Saharan Desert Faunal Area; IIIA, Cyrenaican
Desert Province; III B, Fezzanese Desert Province.
Although the Mediterranean littoral and steppe regions of Libya

cover a much smaller area than the Saharan portion, they are more
varied, both climatically and physiographically. As a consequence,
these northern portions of Libya support a more varied rodent fauna
and are thus divisible into a greater number of faunal provinces.
Portions of these faunal provinces, and in some cases, the entire
province, possess rodents which demonstrate a high degree of en-
demism; these areas act as centers of differentiation for several kinds
of rodents. Genetic fixation among populations of Libyan rodents
lias arisen either through long periods of spatial (geographic) isolation
or as a more local response to variations in the character of the climate,
soil, and plant cover.
Mediterranean Faunal Area

The Mediterranean Faunal Area comprises all the coastal areas of
Libya, including the Cyrenaican Plateau. This faunal area is composed
of the rather discrete Coastal Plain Province and the Cyrenaican
Plateau Province. The Coastal Plain Province includes the Mediter-

ranean littoral from the seaward margins of the coastal plain to the
coastal escarpment and northernmost edge of the hamadas of the
Saharan steppe. The Cyrenaican Plateau comprises the massif of the
Cyrenaican Plateau including the Gebel el Achdar.
Owing to the mild climate of these coastal areas and the greater
amounts of rainfall and higher humidity, vegetative cover is denser
and more varied than in other parts of Libya. This is particularly
true of the Cyrenaican Plateau, which receives more rainfall than any
other region of Libya and has a flora differing strikingly from that of
the surrounding lowlands.
The ecologic and climatic distinctness of the Mediterranean Faunal
Area is reflected in the greater number of kinds of rodents that occur
there. Of the 56 kinds of rodents that occur in Libya, 31 are found
in the Mediterranean Faunal Area —
55 percent of the total Libyan
rodent fauna.
The following species of rodents are restricted to the Mediterranean
Faunal Area: Microtus mustersi; Gerbillus eatoni; Gerbillus henleyi;
Meriones libycus; Psammomys obesus; Spalax ehrenbergi; and Allactaga
tetrad actyla.
The and Rattus norvegicus are also restricted
species Rattus rattus
to the coastal areas of Libya, but because of their commensal nature
and tendency to be disseminated by man, they are not included in the
above list. Because of its doubtful validity, Gerbillus grobbeni, which
was described from specimens collected near Derna, has not been
included.
Local differentiation of rodent populations in the Mediterranean
Faunal Area has resulted in the formation of 16 subspecies representing
11 species. The majority of these subspecies (12) are restricted to the
Coastal Plain Province and include Gerbillus aureus favillus ; Gerbillus
eatoni eatoni; Gerbillus eatoni inflatus; Gerbillus eatoni versicolor;
Gerbillus campestris wassifi; Gerbillus henleyi henleyi; Meriones caudatus
conjalonierii; Meriones libycus auratus; Meriones libycus azizi; Psam-
momys obesus obesus; Psammomys obesus tripolitanus ; and Jaculus
deserti favillus.
The vole, Microtus mustersi, and the following subspecies occur in
both the Cyrenaican Plateau Province and the Coastal Plain Prov-
ince: Gerbillus campestris brunnescens; Sjmlax ehrenbergi aegyptiacus:
Eliomys quercinus cyrenaicus; and Jaculus orientalis orientalis. With
the exception of Jaculus orientalis orientalis, all are more abundant in
the Cyrenaican Plateau Province. The subspecies Gerbillus campestris
brunnescens and Spalax ehrenbergi aegyptiacus are the most typical
rodents of the Cyrenaican Plateau and are common in almost all
types of habitat and at all elevations.
RODENTS OF LIBYA 29
The Cyrenaican Plateau, in view of its unique ecologic and climatic

regimen, has a surprisingly- depauperate rodent fauna. The absence of
endemic subspecies in an area so distinct ecologically is most surprising.
The most logical explanation for this lack of differentiation in the
populations of rodents occurring here may be the close proximity of the
coastal plain and, in many places, the continuity of similar habitats
between and the uplands of the plateau, which have allowed for
it
genetic interchange between populations of rodents occurring in both

areas.
The Coastal Plain Province has a richer rodent fauna than the
Cyrenaican Plateau probably because of its larger area and its greater
diversity of habitats. Although rainfall is more seasonal on the coastal
plain, falling mostly during a comparatively short winter period,
there are much greater local fluctuations in daily and yearly temper-
atures. Most inland areas of the coastal plain are contiguous with
the Saharan steppe and tend to be somewhat arid. Some portions of
the coastal plain, in floral composition and climate, do not differ
appreciably from the Saharan steppe. This ecologic and climatic
diversity has increased significantly the number of habitats available
to mammals, and the rodent fauna is accordingly more diversified.
The most characteristic species of rodents of the Coastal Plain
Province are: Gerbilbis eatoni; Gerbillus henleyi; Meriones libycus;
Psammomys obesus; Jaculus orientalis; and Allactaga tetradactyla. All
of these species except Jaculus orientalis, which occurs on the Cyrenai-
can Plateau and Gebel Nefusa, are restricted to the coastal plain.
Most of these coastal species have developed subspecies in response
to local differences in the character of the coastal plain. Thus Gerbillus
eatoni is divisible into three subspecies, G. e. eatoni, with a broad
range covering the southern margins of the Gulf of Sirte; G. e. inflatus,
which is confined to the coastal plain of extreme northeastern Cyrena-
ica; and G. e. versicolor, whose range includes the coastal plain of the
eastern margins of the Gulf of Sirte.
The sand rat, Psammomys is represented by two subspecies,
obesus,
P. o. obesus and P. o. which are confined, respectively, to
tripolitanus,
the coastal plains of northeastern Cyrenaica and the Gulf of Sirte.
Meriones libycus is also divisible into two coastal subspecies.
The range of Meriones libycus auratus includes all coastal areas of
Tripolitania and those areas adjoining the southern margins of the
Gulf of Sirte, while Meriones libycus azizi is confined to the coastal
plain of northern Cyrenaica.
The species Gerbillus henleyi and Jaculus orientalis inhabit large
portions of the Libyan coastal plain but have not differentiated
sufficiently in Libya to warrant division into subspecies. The raono-
typic species, Allactaga tetradactyla, likewise has not formed subspecies.
In Libya, this species is known only from a few specimens from El

Agheila and Gheminez on the Cyrenaican coastal plain; therefore,
its taxonomic status is not well established.
Subspecies confined to the Libyan coastal areas representing
populations on the periphery of the ranges of their respective species
are: Gerbillns campestris wassifi; Meriones caudatus confalonierii;
Eliomys qiiercinus cyrenaicus; and Jaculus desertifavillus.
Because of the encroachment of the Saharan steppe onto portions
of the coastal plain,some subspecies, which typically inhabit the
Saharan Steppe Faunal Area, occur marginally in the Coastal Plain
Province. Chief among these intrusions from the steppe regions are
representatives of the following subspecies : Gerbillns campestris
dodsoni; Gerbillns gerbillns psammophilons; Gerbillns pyramidnm
hamadensis; Pachyuromys dnprasi natronensis; Meriones caudatus
luridus; and Jaculus jaculus whitchurchi.
Conversely, members of typical coastal subspecies penetrate short
distances into the Saharan Steppe Faunal Area. Examples of coastal
subspecies which are found occasionally along the northern margins of
the steppe region are: Gerbillns ealoni eatoni; Gerbillns henleyi henleyi;
Meriones libycus auratus; Spalax ehrenbergi aegyptiacns; and Jaculus
deserti javillus.
From the above examples, apparent that elements of two
it is
different faunal aieas overlap in some parts

of coastal Libya.
Distributions of the porcupine (Hystrix cristata), the large Egyptian
jerboa (Jaculus orientalis), and the small gerbil (Gerbillns kaiseri)
do not conform to other distributional patterns of rodents in Libya.
known from both the Cyrenaican Plateau and the Gebel
All three are
Nefusa, which belong, respectively, to provinces of the Mediterranean
and Saharan Steppe Faunal Areas. Such a discontinuous distribution
for members of the same species strongly opposes modern concepts
of speciation. It is most likely, therefore, that the populations of
J. orientalis and on the Gebel Nefusa are relicts of formerly
G. kaiseri
continuous populations. The porcupine, owing to its reputed adapta-
bility and tenacity, represents a genetically plastic species which
has been affected less by ecologic and climatic barriers and thus has
not followed the distributional patterns of other Libyan rodents.
Saharan Steppe Faunal Area

The Saharan Steppe Faunal Area comprises the steppe region of
Libya located between the inner margins of the coastal plain and the
Saharan interior; it is situated between the Mediterranean Faunal
Area and the Saharan Faunal Area. The Saharan Steppe Faunal Area
is divisible into two distributional provinces, the Transitional Desert
Province and the Tripolitanian Gebel Province. The Transitional

RODENTS OF LIBYA 31
Desert Province includes the Saharan steppe and is the larger province
of the two. The Tripoli tanian Gebel Province consists of the Gebel
Nefusa, Gebel Tigrinna, and the mountainous coastal escarpment of
northwestern Tripolitania. Because the rodents of the Gefara Plain
resemble those of the nearby gebel areas, this portion of the Tri-
politanian coastal plain is included in the Tripoli tanian Gebel Province.
Twenty-four kinds of rodents occur in the Saharan Steppe Faunal
Area, representing 17 species and 19 subspecies. In contrast to the
rather large number of species restricted to the Mediterranean Faunal
Area, the Saharan Steppe Faunal Area has only Psammomys vexillaris
and Ctenodactylus gundi restricted to it. Besides these two, other species
of rodents occurring in this faunal area are: Gerbillus amoenus;
Gerbillus aureus; Gerbillus campestris; Gerbillus gerbillus; Gerbillus
kaiseri, Gerbilluspyramidum, Pachyuromys duprasi; Meriones caudatus;
Meriones crassus; Mus musculus; Acomys cahirinus; Eliomys guercin us;
Jaculus deserti; Jaculus jaculus; and Hystrix cristata. Of the above
species, Gerbillus aureus, Pachyuromys duprasi, Meriones caudatus,
Jaculus deserii, and Ctenodactylus gundi are most characteristic of the
Saharan Steppe Faunal Area. Gerbillus campestris, Gerbillus gerbillus,
and Jaculus jaculus are relatively common but are more abundant
farther south in the Saharan Faunal Area.
The Tripolitanian Gebel Province is distinctive in having restricted
to it: Gerbillus aureus aureus; Gerbillus aureus nalutensis; Eliomys
quercinus tunetae; Jaculus jaculus tripolitanicus; and Ctenodactylus
gundi gundi.
The following subspecies are either confined to the Transitional
Desert Province in Libya or occur only marginally in neighboring
faunal areas: Gerbillus campestris haymani; Gerbillus gerbillus gerbillus;
Gerbillus gerbillus latastei; Gerbillus pyramidum hamadensis; Meriones
caudatus caudatus; Meriones caudatus lurid us; Psammomys vexillaris
vexillaris; Jaculus jaculus whitchurchi; and Ctenodactylus gundi vali.
Giarabub is the northernmost of the Saharan oases and has floral
and edaphic elements resembling those of the more southern oases of
Cyrenaica. For this reason and because of its geographic position on
the northern portion of the Libyan Desert, Giarabub Oasis would be
expected to have a rodent fauna related to that of the Saharan Faunal
Area to the south. Instead, rodents from this oasis are related more
closely to the rodent fauna of the Saharan Steppe Faunal Area to the
north. Apparently the Sand Sea of Calanscio and the Serir of Calanscio
have served to isolate rodent populations of Giarabub Oasis from those
to the south. To the north, suitable habitat, although localized, is
sufficiently widespread to allow for the dispersal of rodents.

The subspecies Gerbillus gerbillus gerbillus and Gerbillus campestris
haymani apparently reach their westernmost distributional limits at
Giarabub. Representatives of both also occur farther east at Siwa

Oasis in the Western Desert of Egypt. In northern Egypt, the great
chain of depressions composed of the Wadi Natroun, Qattara Depres-
and Siwa Oasis probably has acted as a dispersal corridor through
sion,
which members of these two subspecies have extended their range into
Libya.
Saharan Desert Faunal Area

The Saharan Desert Faunal Area comprises all of Libya south of
the Saharan Steppe Faunal Area. This faunal area is limited entirely
to the truly Saharan portions of Libya and is divisible into an eastern
Cyrenaican Desert Province and a western Fezzanese Desert Province.
Owing to the insular distribution and relative isolation of popula-
tions of rodents in the Saharan Desert Faunal Area, a high degree of
endemism is found. Although no full species are restricted to this
faunal area, 11 of the 16 subspecies which occur here are endemic.
This high proportion of endemic subspecies is a most distinctive feature
of this area.
The most widespread species of rodents in this faunal area include
Gerbillus gerbillus, Gerbillus campestris, and Jaculus jaculus. These are
all typical desert species and are common also in the steppe and littoral
desert regions of Libya.
The Fezzanese Desert Province is confined in Libya to the Fezzan
but includes adjacent portions of southeastern Algeria and parts of
northern Niger and the Chad. This province is bordered on the north
and east by the mountainous barriers of the Hamada de Tinrhert and
the volcanic massifs of the Gebel es Soda and the Gebel el Harug el
Asued. It is limited on the west by the mountain complex of Tassil-n-
Ajjer and the Ahaggar Mountains of southeastern Algeria and on the
south by the Plateau de Mangueni of northern Niger and the Tibesti
Mountains of northern Chad. All these surrounding, rocky mountain-
ous areas provide unsuitable habitat for most species of rodents oc-
curring in the Fezzan. Genetic exchange with populations outside of
the Fezzan, therefore, has been of rare occurrence and a large number
of endemic subspecies have arisen. Endemic rodents in the Fezzanese
Desert Province are: Gerbillus gerbillus discolor; Gerbillus pyramidwm
tarabuli; Meriones caudatus amplus; Eliomys quercinus denticulatus;
and Jaculus jaculus arenaceous.
The Cyrenaican Desert Province is limited to the central and south-
ern portions of Cyrenaica and includes the desolate areas of the Sand
Sea and Serir of Calanscio and the isolated oases of southern Cyrenaica.
This province is characterized also by a largely endemic fauna and has
restricted to it: Gerbillus gerbillus aeruginosus; Gerbillus campestris
patrizii; Jaculus deserti rarus; Jaculus deserti vastus; Jaculus jaculus
collinsi; and Jaculus jaculus cufrensis.
RODENTS OF LIBYA 33
Several subspecies of genetically plastic species are distributed over

wide geographic areas in Libya and do not conform to faunal areas
and provinces. These Libyan subspecies, which occur in more than
one faunal area, are: Gerbillus amoenus vivax; Gerbillus campestris
dodsoni; Gerbillus gerbillus psammophilous; Meriones crass us tripolius,
and Acomys cahirinus viator.
Gerbillus campestris dodsoni occupies the largest range of any sub-
species of Libyan rodent and is found in all four provinces of the
Saharan and Saharan Steppe Faunal Areas. In the Fezzan and southern
Cyrenaica, this subspecies is largely confined to the oases where it is
usually concentrated in the more mesic portions. In the Tripolitanian
steppe and the Gebel Nefusa, it is almost always confined to rocky
habitats. Less commonly, members of this subspecies occur in sandy
habitats. By virtue of this apparent wide ecological tolerance, G. c.
dodsoni is able to occupy this large geographic area.
The species Gerbillus amoenus and Meriones crassus require similar
habitats and have almost identical ranges in Libya, including the
Libyan Fezzan and portions of the Tripolitanian steppe. Each is
represented in Libya by a single subspecies.
The range of Gerbillus gerbillus psammophilous includes a vast area
in Cyrenaica extending from the interior oases of Tazerbo and Bir el
Harasc to the margins of the coastal plain. Representatives of this
subspecies occur near Agedabia on the inner margins of the Cyrenaican
coastal plain, at Gasr es Sahabi in the Cyrenaican steppe, and at Gialo,
which is one of the northernmost of the Libyan Saharan oases. The
range of this subspecies includes portions of all three major Libyan
faunal areas.
In Libya, the spiny mouse, Acomys cahirinus viator, is known from
three widely separated localities which include El Giof of Cufra Oasis
in southern Cyrenaica, the oases of Socna and Bir Fergian of south-
eastern Tripolitania, and the oasis of El Barcat in the Fezzan. It is
doubtful that populations of these mice having such a strikingly dis-
continuous distribution (polytopic) actually belong to the same sub-
become available from more localities in
species. Until larger series
Libya and permit a more accurate analysis of the morphological
characters of these spiny mice, their taxonomic status should be
considered provisional.
The house mouse, Mus musculus, is the most widespread of Libyan
rodents and is the only species which occurs in all faunal areas and
provinces. These mice occur as commensals in many coastal cities and,
by the agency of man, have colonized almost all of the isolated oases
of Cyrenaica and the Fezzan. They occur also as wild or feral popula-
tions in almost every type of habitat. On the coastal plain and the
Cyrenaican Plateau, where the vegetative cover is relatively uniform
and dense, house mice are practically ubiquitous. In the more arid
Saharan steppe and Sahara Desert, they are more localized and usually
confined to the more mesic areas of the larger oases. Occasionally,
house mice are obtained, along with jerboas, gerbils, and jirds, from
the sandy periphery of the oases, but these habitats are probably
suboptimal.
Speciation and Geographical Variation
Most of the species of Libyan rodents satisfy the requirements of

the multidimensional, polytypic species being composed of groups of
local populations that "actually or potentially" interbreed with each
other. In some genetically plastic species, such as Jaculus jaculus and
Gerbillus gerbillus, there are varying degrees of gene flow the among
various local populations resulting in pronounced phenotypic vari-
ability.Other species, which are more restricted ecologically, such as
Gerbillus pyramidum, Eliomys quercinus, and Meriones caudatus, are
composed of discontinuous allopatric populations, many of which are
geographic isolates and thus only potentially capable of interbreeding.
Members of each species of Libyan rodents share in a common gene
pool and do not outbreed with members of other contiguous or
sympatric species. By virtue of these two biological characteristics
they conform to modern concepts of "biological species."
The species Gerbillus gerbillus and Jaculus jaculus serve to illustrate
the concept of polytypic species. Each has a wide distribution in
Libya and has developed several subspecies in response to selective
pressures of the local environment. The pattern of distribution of the
subspecies of Jaculus jaculus and Gerbillus gerbillus is essentially
continuous, the range of one subspecies overlapping with that of the
contiguous subspecies. In some cases, zones of intergradation are
demonstrable.
In Libya, several species of rodents show varying degrees of morph-
ological differentiation, and in some cases, certain populations have
progressed to the level of subspecific distinctness. Libyan species which
best demonstrate this geographic variation and the number of sub-
species comprising each are: Ctenodactylus gundi, 2; Eliomys quercinus,
3 ; Gerbillus aureus 3 ; Gerbillus campestris 5 ; Gerbillus eatoni 3 ; Gerbillus
gerbillus 5; Gerbillus pyramidum 2; Jaculus deserti 4; Jaculus jaculus 5;
Meriones caudatus 4; Meriones libycus 2; and Psammomys obesus 2.
All of the above species are polytypic and are represented by at
least two subspecies. Some species of Libyan rodents, however, are
remarkably uniform, morphologically, over extensive areas of Libya
and, as a consequence, are not divisible into subspecies. Species of
rodents which are either monotypic or represented in Libya by a
single subspecies are: Gerbillus amoenus; Gerbillus henleyi; Gerbillus
kaiseri; Jaculus orientalis; Allactaga tetradactyla; Microtus mnstersi;
RODENTS OF LIBYA 35
Pachuromys duprasi; Acomys cahirinus; Hystrix cristata; and Psam-

momys vexillaris.
Most of the above species are more highly specialized than the
polytypic species discussed earlier; consequently, they are less variable
genetically and show a more limited range of phenotypic expression.
In such genetically impoverished species, natural selection has fewer
opportunities to alter genotypic and phenotypic composition of the
population, and morphological characteristics of these species tend
to remain relatively uniform, sometimes over large areas. Usually
the geographic ranges of these genetically static species tend to be
small because of the narrow ecological tolerance of the individuals
comprising the population. This tendency for more specialized forms
to have smaller geographic ranges is exemplified by Spalax ehrenbergi
Gerbillus kaiseri, Gerbillus eatoni, and Gerbillus henleyi, all of which
have ranges confined to northern Cyrenaica or areas near the Libyan
coast. On the other hand, species such as Gerbillus gerbillus, Gerbillus
campestris, and Jaculus jaculus, which have broad genetic constitu-
tions, occur virtually throughout Libya.
Geographic Variation in Response to Selective Factors of the Habitat

Morphological variation in rodent populations in Libya usually does
not conform to climatic or ecogeographical rules, such as Bergmann's
Ride and Allen's Rule. In some species, trends in morphological
gradients are directly opposite those predicted by the rules. These
ecogeographical rules actually have a broad application and are nor-
mally apparent only in faunas spanning entire continents. Conforma-
tion, therefore, cannot really be expected on such a small scale.
In Libya, those species that do conform probably do so by coincidence.
Geographic variation in response to selective factors of the habitat
is most apparent in the cryptic coloration of rodents living on differ-
ent types of substrate. Coloration of many subspecies and popula-

tions of Libyan rodents closely corresponds to the color of the sub-
strate on and in which they live.
The subspecies of Gerbillus campestris and Meriones caudatus
demonstrate this substrate adaptation very well. On the Cyrenaican
Plateau, representatives of Gerbillus campestris brunnescens are all
of a dark chestnut dorsal color, reflecting the dark color of the sub-
strate. Farther south and west in the desert areas of Libya, gerbils
representing Gerbillus campestris dodsoni are markedly paler in dorsal
color in response to the pale color of the sand and desert soils. The
dorsal color of G. c. dodsoni is quite varied and several distinct color
patterns are sometimes present within the same local population.
This polymorphism in dorsal color is discussed in greater detail below.
A similar adaptation to color of the substrate is shown by subspecies

of Meriones caudatus. In the Fezzan, the pale and uniform dorsal color
of jirds referrable to Meriones caudatus amplus may reflect the pre-
vailing pale coloration of the desert substrate. To the north, popula-
tions of Meriones caudatus caudatus and M. c. confalonierii which
inhabit the darker, more grayish soils of the steppe and coastal plain
are correspondingly darker in color with a stronger suffusion of black
on the dorsum.
Cryptically colored individuals in a population of rodents where
predation is widespread probably have a selective advantage over
those less cryptically colored. In most cases a predator, whether it be

an owl, fox, or jackal, would most likely select individuals that were
most discordant with the color of their surroundings.
To determine whether or not these "substrate races" are the result
of predator selection would require much more detailed research,
but it is the most plausible explanation at present.
The adaptive significance of this cryptic coloration in Libyan
rodents may be related to climatic factors, such as humidity, aridity,
and solar radiation or may reflect relative density or sparseness of
vegetative cover.
The subspecies Gerbillus campestris dodsoni shows a high degree of
polymorphism in dorsal color throughout its wide range in Libya. In
the same local population, individuals range in color from uniformly
tan or buff to almost dark brown and demonstrate varying degrees of
streaked or variegated patches in the pelage. Members of this sub-
species are highly polymorphic in dorsal color in some areas (Tazerbo
and Brach Oases) and almost uniform in others (Gebel es Soda).
This type of color polymorphism in Gerbillus campestris dodsoni
may be the result of "balanced polymorphism" resulting from a
balance among the genotypes AA, Aa, and aa. This balance is subject
to geographic variation, but polymorphism is insured by the selective
superiority of heterozygotes over homozygotes.
Several subspecies of Gerbillus gerbillus all possess orangish dorsal
color, which doubtless has a broad adaptive significance in the sandy
areas in which these gerbils live. Constant genetic interchange among
widely distributed local populations of these gerbils, however, has
prevented the formation of distinct discontinuous "substrate races" as
in Gerbillus campestris and Meriones caudatus.
Modes of Speciation
Gradual speciation, as opposed to instantaneous speciation or

macroevolution, has doubtless been the prevailing mode of speciation
among Libyan rodents. Of the two types of gradual speciation,
geographic speciation has played the primary role in the evolutionary
RODENTS OF LIBYA 37
history of the Libyan rodent fauna. Sympatric speciation, or speciation

without geographic separation, may be operative in Libyan rodents,
but evidence to support geographic speciation is far greater.
Sympatric speciation. The presence in Libya of closely related sym-
patric species, differing more in ecological or biological characteristics
than in morphological characters, is suggestive of a type of sympatric
speciation. The following pairs of closely related species of Libyan
rodents have greater ecological than morphological differences and, in
this respect, resemble sibling species:
Gerbillus eatoni and Gerbillus gerbillus

Jaculus deserti and Jaculus jaculus
The above pairs are sympatric over portions of their range in Libya
yet do not interbreed. Isolating mechanisms, which segregate each
member of the species pair, involve habitat selection and behavioral
characteristics which usually have a genetic basis and are not to be
construed with geographic isolation, a topic that will be discussed later.
Habitat isolation contributes significantly to spatial segregation of
these species of Libyan rodents; in the present study, examples will be
limited to this type of isolating mechanism.
In many areas of coastal Tripolitania and Cyrenaica, Gerbillus
eatoni and Gerbillus gerbillus occur together without loss of genetic
identity. In these coastal habitats, G. gerbillus is almost exclusively
confined to localized sandy areas of the coastal plain or to the barren
margins of the nearby hamada, and G. eatoni seems to prefer the hard
clay substrate of the coastal plain. Vegetative cover of the sandy areas
and hamadas is generally extremely sparse, whereas that of the
majority of the coastal plain is relatively dense. This habitat segrega-
tion is largely instrumental in the prevention of introgression of the
gene pools of the respective species. In the absence of this habitat
exclusion, G. eatoni, owing to its more static gene pool, would doubtless
be "swamped out" by the more widely ranging and more adaptable
G. gerbillus.
The dipodils, Gerbillus amoenus and Gerbillus campestris, also illus-
trate the principle of habitat isolation very well. These two species
are sympatric throughout most of Libya and show no evidences of
hybridization. Gerbillus campestris is highly polymorphic, both crani-
ally and in dorsal color, whereas G. amoenus is remarkably uniform
morphologically. The latter species is normally confined to sandy
margins of the oases or to gravel plains of the steppes. The larger
G. campestris shows a wide ecological tolerance and occupies a wide
variety of habitats, including the sparsely vegetated gravel plains of
the steppes, the margins of sand seas, the sandy environs of the
oases, the dense chapparal vegetation of the Cyrenaican Plateau,
and adjacent densely vegetated coastal plain. The preferred habitat of
G. campestris, however, seems to be the fissures and recesses of rocky

outcroppings. Because of this broad range of ecological tolerance, G.
campestris is the most widespread rodent in Libya. In this respect it
parallels rather closely the ecological and genetic plasticity of the
deer mouse, Peromyscvs maniculatus (Wagner) of North America.
,
When G. amoenus and G. campestris inhabit the same oasis, the

latter is almost invariably found to occupy the interior, more lush
portions of the oasis, and G. amoenus is usually confined to the drier,
peripheral zones. Thus, habitat exclusion is clearly operative.
Habitat exclusion is further demonstrated by these two species
along the coastal escarpment where G. campestris is confined to rocky
outcroppings or rock-strewn uplands, and G. amoenus invariably
occupies the intervening pockets of sand and gravel.
Two sibling species of jerboas, Jaculus deserti and Jaculus jaculus,
are sympatric over large parts of the Libyan interior but show subtle
habitat preferences. Jaculus deserti occurs in the outlying hamadas
and sparsely vegetated margins of wadis, whereas J. jaculus is con-
fined more to margins of sand seas and dune areas associated with
oases. Only rarely were both species found to occur in the same local
area, and then the two species were never collected from the same
portion of the trapline.
Sympatry of species of Libyan rodents is not linlited to these few
examples. In many areas of Libya, several species of genetically and
morphologically distinct rodents occur in the same geographic area
and demonstrate marked habitat exclusion.
In some oases, Gerbillus gerbillus, Gerbillus pyramidum, Gerbillus
campestris, Gerbillus amoenus, Meriones caudatus, Jaculus jaculus,
and Mus musculus are all present. In these oases, each species or
species pair tends to inhabit a particular niche with consequent ex-
clusion of the other species. The dense growth of sedges and other
mesophytic plants of the interior of these oases are occupied exclu-
sively by Gerbillus campestris and Mus musculus. In open sandy areas
and around bases of palm trees, Gerbillus gerbillus and Gerbillus
pyramidum are more abundant. Meriones caudatus is confined to
sandy hummocks supporting shrubby young palms, and Jaculus
jaculus and Gerbillus amoenus are confined largely to peripheral areas
of oases in the zone of tamarix and outlying dunes.
Many additional factors probably prevent genetic exchange between
closely related sympatric species of Libyan rodents. Some of these
additional isolating mechanisms include behavioral barriers to mating,
differences in the time of the breeding season, and mechanical incom-
patabilities, such as differences in the size and shape of the external
genitalia.
RODENTS OF LIBYA 39
Geographic speciation. In Libyan rodents, far more evidence is

available to support speciation caused by geographic isolation. Be-
cause evolution is operative at the level of the local population, it is
the degree of isolation of these local populations and the resulting

cessation of gene flow between them that is of greatest significance
in interpreting evolutionary progress. In some of the examples of
geographic speciation discussed below, the subspecies themselves act
as incipient species, but it is the geographic isolate which is the main
focus of geographic speciation. In many instances geographic variation
is clinal.
Clinal variation.Among Libyan rodents, the species Gerbillus ger-
billus, Meriones caudatus, and Gerbillus amoenus demonstrate clinal
variation. All of these species show progressive increase or decrease
in the size of either external or cranial features. Other morphological
and physiological characters of these three species doubtless undergo
clinal changes, but detection of these characters requires extremely
refined techniques, and, in some cases, living specimens would be
required.
In eastern Cyrenaica, populations comprising the subspecies Ger-
psammophilous and Gerbillus
billus gerbillus gerbillus, Gerbillus gerbillus ,
gerbillus aeruginosas demonstrate a regular north to south progression

in diminution of cranial and body size. Individuals of the largest body
and cranial size are found in representatives of G. g. gerbillus in
Giarabub Oasis near the northernmost limits of the range of this
species in Libya. The trend of decreasing body and cranial size con-
tinues in a progressive fashion from north to south in populations
of these gerbils from Gialo, Gasr es Sahabi, and Tazerbo. The extreme
in small body and cranial size is shown by members of G. g. aeruginosus
from Cufra and Bzema Oases of southeastern Cyrenaica.
A progressive increase in body and cranial size is clearly notice-
able among populations of jirds extending southward from the coastal
areas of Tripolitania to the interior of the Fezzan. Because of the
relatively few localities from which specimens were examined, this
character gradient lacks the uniformity of the cline in populations
The differences in external and cranial dimen-
of Gerbillus gerbillus.
sions between representatives of Meriones caudatus confalonierii from
coastal Tripolitania and members of M. c. ampins from the Fezzan
are of the degree of specific distinctness. Populations of the two
extremes of this character gradient have thus diverged much more
than those of the cline in Gerbillus gerbillus. Apparently the more
diverse physiography and more varied edaphic conditions of western
Libya have proven to be a more severe deterrent to gene flow than
has the more uniform terrain of eastern Cyrenaica.
The degree of smoothness of a cline gives some indication of the

amount and regularity of gene flow between successively contiguous
populations. Because environmental factors vary from region to
region, phenotypic responses to the selective pressures of these environ-
ments also vary. Genetic exchange between neighboring populations,
however, tends to smooth out phenotypic characters that result from
this selection, and clinal patterns tend to be gradual and uniform.
In Libya, clinal patterns of geographic variation generally run from
north to south or vice versa. The dipodil, Gerbillus amoenus, is an ex-
ception in having a pattern of clinal variation extending generally
from east-northeast to west-southwest. This east-northeast west-
southwest cline, showing progressively increasing cranial size, is de-
monstrable among populations of G. amoenus extending from the Nile
Delta to eastern Algeria. Members of this species from northern Egypt
and northern Cyrenaica have small, gracile skulls. In the Fezzan,
representatives of G. amoenus have markedly larger and more robust
skulls, while specimens examined from eastern Algeria are even larger
cranially than any of those from Libya.
In all examples of clinal variation in Libyan rodents, only the broad
features of clinal progression are observable, and these clinal gradients
may be far more irregular than indicated. Regardless of local differ-
ences in clinal pattern, clines in Libyan rodents serve to demonstrate
general trends in morphological variation. In many cases, a clinal
population represents a stage in speciation, the morphological differ-
ences among populations being attributable to differences in their
genetic constitution.
Geographic isolates. Subspecies of Libyan rodents representing geo-
graphic isolates have developed rather striking phenotypic characteris-
tics as a result of their altered genetic constitutions. These subspecies
are: Gerbillus gerbillus aeruginosus; Gerbillus pyramidum hamadensis;
Gerbillus campestris patrizii; Meriones caudatus amplus; Psammomys
obesus tripolitanus; Psammomys Eliomys quercinus
vexillaris vexillaris;
cyrenaicus; Eliomys quercinus denticulatus; Jaculus deserti vastus;
Jaculus jaculus cujrensis; Ctenodactylus gundi gundi; and Ctenodac-
tylus gundi vali.
Ranges of most of the above subspecies are located on the periphery
of the range of their respective species. Many of these peripheral
isolates have undergone marked morphological differentiation and are
well on the way toward becoming full species. It is probable that in
Libya most new species have arisen as a result of the genetic diver-
gence of peripheral isolates.
The range of Gerbillus pyramidum hamadensis is limited to the
coastal plain of the Gulf of Sirte and littoral deserts and steppes of north-
ern Tripolitania. The nearest populations of gerbils belonging to this
species are those of G. p. tarabuli, 200 kilometers farther south in the
RODENTS OF LIBYA 41
Fezzan. This population of G. p. hamadensis represents the northern-

most occurrence of the species Gerbillus pyramidum in Libya and as is
sometimes typical of populations on the margins of a species range, it
is sporadic and localized in occurrence. This subspecies has diverged
strikingly morphologically from the nearest related populations in the

Fezzan and approaching full species status. There is no evidence of
is
intergradation between G. p. hamadensis and G. p. tarabuli to the south.

Apparently the mountainous areas of the Gebel es Soda and Gebel el
Harug el Asued and the vast plateau of the Hamada el Hamra, which
are located between the ranges of these two subspecies, act as absolute
barriers to dispersal.
The distinct Meriones caudatus amplus of the Fezzan is another
typical peripheral isolate which near to achieving species distinct-
is
ness. In the case of M. c. amplus, parental populations are to the north

and east, rather than to the south as in Gerbillus pyramidum
hamadensis.
The species Psammomys obesus is distributed along the coastal
areas of the eastern Mediterranean, northern Egypt, and portions of
coastal Libya. The subspecies Psammomys obesus tripolitanus of the
coastal plain and littoral deserts along the Gulf of Sirte in Cyrenaica,
Libya, represents the westernmost populations of the species Psammomys
obesus in North Africa. The Cyrenaican Plateau is interposed between
the range of P. o. tripolitanus and Egyptian populations representing
the nominate subspecies. The amount
morphological divergence of
in populations of P. o. demonstrated
tripolitanus is far less than that
by Gerbillus pyramidum hamadensis or Meriones caudatus amplus.
The Cyrenaican Plateau has acted more as a filter in retarding gene
flow between populations of Psammomys obesus than as an absolute
barrier.
It is possible that Psammomys vexillaris vexillaris represents a
species evolved through the agency of geographic isolation. Although
Psammomys vexillaris somewhat superficially resembles Psammomys
obesus, it is clearly a separate species and is readily separable from
P. obesus by its markedly smaller cranial size and external dimensions.
Some time in the past, peripheral populations of the coastal species,
P. obesus, may have become
separated from the parental population.
In time, this isolated population developed different morphological
and ecological characteristics as a reflection of its altered genetic
constitution. The continuous action of selection has resulted in the
evolution of additional isolating mechanisms, both ecological and
behavioral, and this inland population of sand rats is now both
spatially and reproductively isolated from the parental population.
The two subspecies of the gundi, Ctenodactylus gundi gundi and
Ctenodactylus gundi vali, have probably had an evolutionary history
similar to that of Psammomys obesus and Psammomys vexillaris, except

285-134 O —68 4
that in Ctenodactylus, the isolated population has not yet attained

full species status.
Ctenodactylus gundi vali represents a population that originally
became isolated from a population of C. g. gundi and has since con-
tinued to evolve independently in the more inland areas of Libya.
As far as is known, the range of C. g. gundi in Libya is limited to
rocky escarpments of the Gebel Nefusa and that of C. g. vali to
larger wadis and occasional rocky outcroppings of the steppe region
of northeastern Tripolitania. Both subspecies are clearly allopatric,
and it is doubtful that the two forms would interbreed if they were
to make contact. Morphological differences between the two popula-
tions are more on the level of subspecific distinctness. For the present,
it seems best to regard Ctenodactylus gundi vali as an incipient species
closely approaching full species status.
The dormice of Libya, by virtue of their apparent rarity and
disjunct distribution, present a distributional pattern differing
markedly from that of other Libyan rodents and demonstrate clearly
the effects of geographic isolation on evolutionary processes.
The three subspecies of the Libyan dormouse, Eliomys quercinus,
are geographically, morphologically, and ecologically distinct and
illustrate the modern concept of the biologically defined polytypic
species. In the past, the subspecies E.q. tunetae and E. q. cyrenaicus
have at one time or another been regarded as full species by taxonom-

ists. Until very recently the genus Eliomys was considered to repre-
sent a polytypic genus composed of several distinct species distributed

in Europe {Eliomys quercinus), North Africa {Eliomys munbyanus
and Eliomys lerotinus), and the Middle East {Eliomys melanurus).
These various isolated, morphologically distinct "species," in addition
to a new subspecies, Eliomys quercinus denticulatus from the Fezzan,
are now regarded as subspecies of a single polytypic species, Eliomys
quercinus.
The discontinuous distribution of these subspecies of E. quercinus
is suggestive of a pattern expected for full allopatric species. In most
cases, these allopatric subspecies show an amount of morphological
distinctness typical of sympatric species. Because species normally
consist of a series of continuous or discontinuous local populations
and each population is composed of individuals with a different
genetic constitution, a polytypic species would be expected to contain
individual populations differing in phenotypic expressions. The
morphological dissimilarity among the various "allopatric subspecies"
of E. quercinus is not suggestive of specific differences but expresses,
instead, a highly variable species gene pool. The ranges of Eliomys
quercinus tunetae and Eliomys quercinus cyrenaicus include the Gebel
Nefusa and the Cyrenaican Plateau respectively, while Eliomys
RODENTS OF LIBYA 43
quercinus denticulatus is confined to the Fezzan. Such a discontinuous

distribution for members of the same species is not in agreement with
modern evolutionary concepts. Mayr (1931) has proposed the term
semispecies for such widely separated allopatric members of the
same species which normally differ so strikingly in morphological
characters that reproductive isolation can almost be assumed. If
one subscribes to Mayr's classification, all of the dormice of Libya,
which are presently recognized as subspecies of a single species,
Eliomys quercinus, would be regarded as morphologically distinct
semispecies of a single, monophyletic superspecies.
The subspecies Gerbillus gerbillus aeruginosus, Gerbillus campestris
patrizii, Jaculus deserti vastus, and Jaculus jaculus cufrensis also
represent geographic isolates, but their ranges are located well within
the limits of the ranges of their respective species, rather than on the
periphery of the species range as in the above examples. Three of these
subspecies occur only in the isolated oases of Cufra and Bzema in
southern Oyrenaica, and Jaculus deserti vastus is known only from the
Gebel el Harug el Asued of central Libya. All of these subspecies
represent geographically isolated populations which are physically
and genetically isolated from neighboring populations. As a result of
this isolation, exchange with neighboring populations is
genetic
impossible. Thus, these populations have been subjected to a rapid
repatterning of their gene pools with consequent reduction of vari-
ability and an increase in the rates of morphological differentiation
and genetic fixation. Those subspecies restricted to Cufra Oasis are
generally smaller in body and cranial size than other representatives
of their respective species and, in this respect, represent a stage in
clinal variation. Jaculus deserti vastus is significantly larger in body
size than other representatives of this species in Libya. These rodent
popidations in Cufra and the Gebel el Harug el Asued illustrate
clearly the significance of geographic isolation in the evolutionary
process.
In summary, habitat segregation could have led to the splitting
of the gene pool of parental species with subsequent development of
other biological and behavioral differences. Spatial isolation of these
populations has tended to further increase these differences, resulting
originally from ecological (or habitat) isolation.
The populational structure of Libyan rodents is characterized by
two primary structural components: geographically isolated popula-
tions and, less frequently, a series of contiguous populations that
show progressive changes in morphological characters (clines). In
some cases, where the ranges of two subspecies meet, there are zones
of intergradation containing individuals intermediate in morphological
characters.
Zoogeographical Considerations
Effects of the Pleistocene on the Dispersal of Saharan and Libyan Rodents

No account of Libyan (or Saharan) mammals should minimize the
effects of the Pleistocene upon them and its obvious role in having
influenced their present and past distribution. Henry Fairfield Osborn
(1915, p. 225) summarized the importance of this period in North
Africa by stating "Thus in no region of the world have more profound
changes occurred during and since Pleistocene times than in Africa
north of the Sahara Desert." In fact, it is probably true that most
of the present distributional patterns of Libyan rodents are directly
related to Pleistocene climates.
The Pleistocene was characterized by widespread glaciation in
the northern parts of the world. As a response to broad climatic
fluctuations during this period, four successive glaciations (in the
northern hemisphere) alternated with milder interglacial periods
following the withdrawal of the ice sheets. Estimates vary regarding
the duration of the Pleistocene, but it probably endured for perhaps
1,000,000 years. The time withdrawal of the last continental

of the
glaciation is also not firmly fixed, and estimates range from 10,000
to 25,000 years.
Widespread glaciation in northern and southern latitudes was not
the only manifestation of prevailing cold during the Pleistocene.
Temperatures in the North Temperate Zone are thought to have been
significantly lower than those of the present. This lowering of the
temperature was probably synchronous over most of the Northern
Hemisphere (Flint, 1947). During the intervening interglacial periods,
climates were apparently warmer and sometimes drier than those of
the pluvial periods.
As a result of lower prevailing temperatures, the amount of rainfall
increased significantly over much and there was an
of the world,
accordant shifting of the temperate rain zones into the low latitude,
subtropical deserts.
These changes in the Pleistocene climate produced manifold changes
in the physical configuration in many regions of the Northern Hemi-
sphere. Lakes in the arid portions of all continents increased in size
and became reduced inrhythm with the advance and recession of
the successive glacial stages. The Dead Sea, the great saline lakes
of Asia (the Caspian, Aral, and Lake Balkash), and the large lakes of
East Africa have lacustrine deposits high above their present shore-
lines,indicating marked expansion during the Ice Age, followed by
shrinking during dry interglacial periods.
As glacial'ice accumulated or receded on the continents, there was
a synchronous lowering or raising of sea level throughout the world.
RODENTS OF LIBYA 45
Thus, continents having broad continental shelves were increased

The magnitude of this fluctuation
or decreased considerably in area.
of sea level has been estimated to have reached 100-120 meters
during the height of glaciation (Darlington, 1957; after Kuenen,
1950).
Even today, there are abundant indications of Pleistocene glacia-
tion. The higher peaks of the Caucasus and Elburz Mountains near
the Caspian Sea in Southwest Asia contain glacial remnants. Portions
of the Alps of central Europe are enveloped in ice, and the Pyrenees
still
of northern Spain have small East Africa has residual glaciers

glaciers.
on Mount Kenya, Mount Kilimanjaro, and on the Ruwenzori Range,
all of which are almost astride the equator.
In response to these marked changes in temperatures during the

Pleistocene, marked changes took place in the underlying patterns of
floral and faunal distributions. In the Northern Hemisphere, the
unglaciated areas south of the "ice front" were profoundly affected
by periodic changes in Pleistocene climate. Widespread migrations of
plants and annuals occurred as a result of the gradual shifting of
climatic zones along the glacial front. During the moist pluvial
periods forest dwelling animals dispersed southward, while during the
dry interpluvial periods animals typical of the steppe and desert
regions migrated northward. Throughout the Pleistocene, there was a
north-south shifting of the flora and fauna in concert with the four
successive advances and withdrawals of the ice sheets.
During the pluvial periods, with increased rainfall throughout
the world, forests and prairies expanded, and during interglacial
periods steppesand deserts increased in area. Some of this repatterning
of floraland faunal elements resulted in development of geographic
discontinuity owing to complex advance, retreat, readvance, and
interspersion of migrating populations.
Pleistocene climatic changes affected especially the Mediterranean
region of North Africa. The Saharan and central portions were also
profoundly affected. Evidence is widespread in North Africa and the
Sahara to support climatic fluctuations during the Pleistocene.
According to Flint (1947; after Brooks, 1932), in the Saharan region
of North Africa, two moist stages, interpreted on the basis of fossil
mammals, are separated by a period of aridity indicated by an accumu-

gypsum
lation of in the soil. The numerous dry watercourses (wadis)
in the Sahara suggest that permanent streams were present during
a pluvial stage when the amount of rainfall was greater than it is
today.
The occurrence in northern Africa of fossil mammals characteristic
of markedly different climatic zones supports other evidences that
climatic patterns shifted in relation to the colder, moister pluvial

and warmer, drier interpluvial stages of the Pleistocene.
The most recent study of the effects of the Pleistocene in Libya
is that of McBurney and Hey (1955), entitled "Prehistory and Pleisto-
cene Geology in Cyrenaican Libya." They consider the terraces on the

northern side of the Gebel Achdar of extreme northern Cyrenaica to
represent "ancient wave-cut platforms," and the cliffs to be also of
marine origin. They therefore recognize six (or possibly seven) differ-
ent shorelines corresponding to the number of terraces and the nature
of their deposits. They propose two deductions (p. 68) regarding the
manner of formation of these ancient shorelines. The first of these
postulates involves a series of "successively lower altitudes at which
sea-level halted, for a considerable length of time in each case, before
reaching its present These altitudes are estimated by
position."
McBurney and Hey 140-200
(p. 68) to consist of successive levels at:
meters (possibly representing two separate stages), 79-90, 44-55,
35-40, 15-25, and 6 meters. The second postulate considers the period
in which these terraces were formed to have been one of "almost com-
plete tectonic stability."
This evidence from northern Cyrenaica indicates that sea-level
fluctuations from the highest level in Pliocene times to the lowest
6-meter level were punctuated by a series of intermediate shorelines
probably corresponding to the pluvial and interpluvial climatic changes
of the Pleistocene. McBurney and Hey conclude (p. 137) that during
the Pleistocene the climate of the Gebel Achdar "was influenced
profoundly," especially during the last continental glaciation (Wiirm
glaciation).
In addition to establishing the pattern and sequence of ancient
shorelines in Cyrenaica, McBurney and Hey (1955) conducted ex-
tensive diggings in the Wadi Derna and in two caves in northern
Cyrenaica. The fossil mammals unearthed as a result of these investi-
gations were reported by Bate as an appendix to the archeological
text by McBurney and Hey (1955). These fossil mammals are largely
represented by fragments, but 15 species have been identified, most
of which shed interesting light on the Pleistocene mammalian fauna
of northern Cyrenaica and North Africa.
Of particular interest to zoogeographical considerations in Libj'a
was the discovery in northern Cyrenaica of fossils of the field mouse,
Apodemus Kaup; the dormouse, Eliomys; a gerbil Gerbillus, and a vole.
Microtus. All except Apodemus occur in Cyrenaica today, and these
fossils probably represent earlier Pleistocene forms of the species
occurring today in Cyrenaica. The field mouse, Apodemus, is now
and northern Morocco.
restricted in Africa to Algeria
The remains Cyrenaican vole were described as a new
of the fossil
species, Microtus cyrenae, by Bate (in McBurney and Hey, 1955,
RODENTS OF LIBYA 47
p. 280). These fossil voles from Cyrenaica and a fossil Ellobius Fischer
from the Pleistocene deposits in Algeria and Tunisia constitute the
only known fossil records of the Microtinae in Africa. Members of the
genus Ellobius are now restricted to Eurasia, but in Cyrenaica, a
modern vole (Microtus mustersi) has persisted into modern times.
According to Bate, this Pleistocene vole, which is known from 63
specimens consisting of teeth and fragments of jaws, differs from the
modern Microtus mustersi primarily in having shorter and narrower
cheek teeth with smaller reentrant angles. Bate concluded that M.
cyrenae belongs to the Microtus guentheri group, which also includes
Microtus mustersi and M. philistinus of Palestine.
According to Bate, the remains of voles were distributed through
four successive levels of the Hagfet ed Dabba Cave, which would
indicate an occupation of considerable duration in Cyrenaica and
suggests an early Pleistocene arrival of voles into Cyrenaica.
This discovery of fossil voles in Cyrenaica, coupled with the present
distribution of members of the M. guentheri group in the Middle
Eastern countries, supports my hypothesis (and also that of Bate)
that much of the Eurasiatic fauna of North Africa entered by way of
the eastern Mediterranean countries and the Isthmus of Sinai some-
time during the Pleistocene.
Probably the most unequivocal example of a Pleistocene relict is the
vole, Microtus mustersi, of the Cyrenaican Plateau and adjacent coastal
plain. These Libyan voles are the only living representatives of the
subfamily Microtinae on the African continent and clearly represent a
marginal intrusion of the Eurasian rodent fauna into North Africa.
Today the nearest populations of Microtus occur in the Middle East
and in Turkey. During the pluvials, suitable moist habitats probably
were continuous along the coastal areas of northern Egypt, thus facili-
tating the dispersal of voles. Apparently, owing to the return of arid
conditions during the following interpluvial period, the coastal areas
of Egypt became dry, as they are today, and no longer afforded suit-
able habitats.
The Cyrenaican Plateau, by virtue of its higher elevation and more
mesic habitats, must have served as a refugium for voles throughout
the arid interpluvial periods. With the gradual return of less arid
conditions to North Africa and Cyrenaica, voles reentered the Libyan
coastal plain, while the Egyptian coastal plain still remains dry and
unfit for them.
In addition to fossil rodents, several other genera of fossil mammals
were discovered in these Pleistocene deposits of northern Cyrenaica.
These include Crocidura Wagler, Vulpes Oken, Panthera Oken, Bos
Linnaeus, Ammotragus Blyth, Gazella Blainville, Hippotigris Smith,
and Rhinoceros Linnaeus. Of these genera, only the shrew (Crocidura)
and the fox (Vulpes) are found in the region today, and they are con-
fined to the more arid regions. Today, members of the genus Hip-
potigris are restricted to Africa south of the Sahara, and those of the
genus Panthera have a wide distribution, occurring in parts of central
and southern Africa and in central, Southwest, and Southeast Asia.
The genus Rhinoceros is today restricted to Southeast Asia.
Most fossil mammals from other parts of North Africa are related to
species or genera now found only in the warmer tropical or subtropical
regions of Africa or Asia. Pleistocene fossils from North Africa are
listed by Osborn (1915) and include six species of elephants, including
the mastodon; two species of rhinoceroses; horses, representing ances-
Ethiopian ass; camels, showing no
tral zebras; wild asses, similar to the
evidences of domestication; and cattle and buffalo. According to
Osborn, the disappearance of the buffalo from North Africa at the
beginning of the Recent Period was attributable to the increasingly
dry conditions and the result of destruction by early man. Other large
mammals known as fossils from North Africa consist of gnus (Conno-
chaetes Lichtenstein) ; several species of the water buffalo (Bubalus H.
Smith) nine species of gazelles (Gazella) oryx (Oryx Blainville) reed-
;
; ;
buck (Redunca H. Smith); several kinds of eland (Taurotragus Wag-

ner) dwarf antelopes or duikers (Cephalophus H. Smith) wild sheep
; ;
similar to the extant North African Barbary sheep (Ammotragus lerviu

Pallas) and wild goats (Capra Linnaeus).
;
Fossil hippopotami have been found in ancient Pleistocene stream-

beds, one form being annectent to the existing hippopotamus of the
Nile River. Other fossils include wild boar (Sus Linnaeus) and wart
hogs (Phacochoerus G. Cuvier).
Fossil carnivores from the Pleistocene of North Africa include
lions, leopards, hyaenas, jackals, and wolves. A fossil bear is known
from Pleistocene deposits of Algeria, and machairodonts (sabre-
toothed tigers) from the Pleistocene of Egypt. Fossil giraffe have been
found associated with Paleolithic stone implements, and giraffe and
a species of deer are depicted in rock drawings of Neolithic age in
Algeria.
Judging from this impressive list of fossil mammals, much of the
mammalian fauna of North Africa and probably also that of the
Sahara, during the Pleistocene phi vials, resembled in composition
the fauna which today is typical of the high African plains and of the
tropical lowlands of central and southern Africa.
The buffalo (Bubalus), the rhinoceros (Rhinoceros), and the wild
goat (Capra) clearly represent contingents of the Asian fauna into
the Pleistocene of North Africa.
It is well established that animal (mammalian) populations do not
preserve their distributional patterns indefinitely but are continually
expanding and/or contracting in accordance with the prevailing
RODENTS OF LIBYA 49
climatic regimen. At present, the Sahara Desert provides a decided

deterrent to nondesert rodents, and local barriers even prevent the
movements of desert rodents. Certainly, this was not the case through-
outmuch of the Pleistocene.
From the above discussion it becomes evident that the climate of
the Sahara during the Pleistocene, although somewhat arid, fluctuated
in relation to climatic changes resulting from successive pluvial and

interpluvial periods. During the moist pluvial periods, rodents now
characteristic of the African veldt and savanna were able to disperse
northward. Conversely, during the dry interpluvials, rodents of the
steppes and deserts extended their ranges into the subtropical and
tropical regions of Africa. The present distribution and composition
of the African rodent fauna are probably a direct result of this inter-
mingling and interdigitation of populations during Pleistocene time.
The present widespread distribution in Africa of the gerbils (Ger-
billus) and spiny mice (Acomys), which are both restricted to arid
habitats, probably is the result of widespread shifting of mammalian
faunas during the dry interglacial periods. During interpluvials,
various arid regions in Africa, now disjunct and separated by inter-
vening areas of veldt and savanna or even tropical rain forest, probably
were continuous and would have permitted genetic interchange
among populations of gerbils that had previously been separated.
Unfortunately the fossil record of African rodents is meager, and
many the conclusions regarding distributional patterns of the
of
Libyan and Saharan Pleistocene rodent faunas are conjectural. It
seems reasonable to assume that at some period during the Pleisto-
cene, the central and southern African rodent fauna, exclusive of
that of the Congo Basin, dispersed far northward, probably to the
northern limits of the continent.
Several extant species provide almost incontestable evidence of
this widespread, northward displacement of this rodent fauna. The
Barbary ground squirrel, AUantoxerus getulus Linnaeus, and the
Barbary striped mouse, Lemniscomys barbarus Linnaeus, of northern
Morocco and Algeria clearly represent residual populations of the
central African rodent fauna that advanced to the northern limits of
the continent during a Pleistocene pluvial. Both of these species have
all of their affinities with rodents occurring south of the Sahara.
Lemniscomys barbarus is considered by some to be conspecific with
populations of Lemniscomys Trouessart as far away as East Africa,
and AUantoxerus getulus is the only sciurid in North Africa; the
nearest other African squirrels occur south of the Sahara.
In early Pleistocene times there were supposed broad land connec-
tions between Europe and North Africa and during the later Pleisto-
cene, there were periodic fluctuations of sea levels occasioned by the
successive melting and accumulations of glacial ice during pluvial

and interpluvial periods. During these times, rodents of European
affinities could have easily entered North Africa by following the
moist habitats appearing along the then exposed continental shelves
or land bridges.
The mole rat, Spalax ehrenbergi, the field mouse, Apodemus, and
the dormouse, Eliomys quercinus, are all clearly of European affinities
and probably reached North Africa (and Libya) in this manner. The
mole rat, Spalax, apparently migrated to North Africa following a
dispersal corridor along the coastal regions of the Middle Eastern
countries. An eastern entry of the mole rat into North Africa is sup-
ported by the fact that today they do not occur west of the Cyre-
naican coastal plain of Libya. The direction from which the dormouse,
Eliomys, entered North Africa is less distinct. Dormice are presently
irregularly distributed along the coastal and steppe regions of North
and Northwest Africa from Spanish Sahara to Sinai; they could have
entered Africa by Avay of the Iberian Peninsula (Gibraltar) or could
have followed the dispersal route of Spalax and entered Africa by
way of the eastern Mediterranean countries. Dormice, field mice
(Apodemus), and mole rats must have extended their ranges south-
ward into Africa during one of the Pleistocene pluvials, because
these species are not typically desert forms.
The fat-tailed sand rat, Pachyuromys duprasi, is typically an in-
habitant of the well vegetated portion of the Saharan steppe. Yet,
in Libya, a specimen was collected from the Wadi er Rueis near the
Gebel el Harug el Asued, located several hundred kilometers within
the Saharan interior. This disjunct population of fat-tailed sand rats
likewisemay represent a relict of the Pleistocene that moved south-
ward when cooler, pluvial climates prevailed in the Sahara and with
the return of aridity became stranded in this localized pocket of
suitable habitat.
The genera Meriones, Psammomys, Jacidus, and Allactaga probably
entered North Africa from Asia sometime late in the Pleistocene.
All are typical of arid steppes or deserts and presumably evolved in
the great steppes of central and southwest Asia during the Tertiary.
The jirds (Meriones) and the jerboas (Jacubis and Allactaga) must
have extended their ranges southwestward from Southwest Asia
during a dry interpluvial period and entered North Africa at a time
when the Sahara was even more widespread than now. The sand rat
(Psammomys) being less restricted to arid habitats, probably entered
,
North Africa along the coastal regions of the Middle East, Sinai, and
Egypt as they are presently defined. The genus Psammomys is today
largely restricted to the coastal plain and the adjoining steppe of
RODENTS OF LIBYA 51
Libya and is similarly restricted throughout its range in North Africa

and the Middle East.
Recent entry of the genera Meriones, Jaculus, and Allactaga into
Africa is indicated by their present distributional patterns. None is
known from Africa south of the Sahara, and of the three genera, only
Jaculus occurs virtually throughout the Sahara. The four-toed jerboa,
Allactaga tetradactyla, is the only representative of the genus Allactaga
in North Africa where it is limited to the coastal fringes of northern
Egypt, and in Libya it occurs only as far west as the coastal plain of
the Gulf Thus,
of Sirte. this species apparently represents a recent
arrival to the African rodent fauna. The more widespread distribution
of jerboas of the genus Jaculus in the Sahara may reflect their greater
ecological tolerance and may not indicate an earlier arrival into
North Africa.
The genus Meriones, represented in Libya and North Africa by
three species, M. caudatus, M. crassus, and M. libycus, and possibly
by M. shawi of doubtful validity, has a peripheral distribution in
relation to the Sahara. Members of this genus occur in all African
countries bordering on the Mediterranean Sea and in the northern
Sudan, Mauritania, and Spanish Sahara. With the exception of
occasional encroachments into the northern Sahara, such as in the
Libyan Fezzan, members of this genus inhabit a comparatively small
portion of the Sahara.
Although suitable habitats are widespread in the isolated oases
of southern Cyrenaica, such as Tazerbo and Cufra, jirds (Meriones)
are unknown from this portion of Libya. Apparently, the great
tracts of sand and sandy plains comprising the Sand Sea and Serir
of Calanscio, and farther south, the Sand Sea of Rebianna, prevent
the dispersal of members of this genus.
The distribution in North Africa, and particularly Libya, of the
large gerbil, Gerbillus pyramidum, agrees closely with that of the
genus Meriones.
The fact that these two kinds of rodents have been unable to
surmount these present-day physical barriers, may indicate a recent
entry for them into North Africa.
In considering faunal movements between Asia and Africa during
the Pliocene-Quaternary period, the Nile River, because of its great
length, large volume,and perennial nature, cannot be overlooked as
a possible barrier. In modern times, the Nile River, in combination
with the manmade Suez Canal, acts as a serious deterrent to the
movements of rodents, particularly desert rodents. The movements
ofsome species are retarded and, in some cases, curtailed by the Nile.
The genera of North African rodents that appear to be most restricted
in their movements by the Nile include Pachuromys and Sekeetamys

Ellerman. The effect of the Nile as a barrier to dispersal during modern
times is further indicated by the following species of North African
rodents which are represented by different subspecies occurring on
opposite sides of the river: Gerbillus henleyi; Meriones crassus;
Psammomys obesus; Jaculus jaculus; and Gerbillus gerbillus.
Judging from what is known of the past distribution of North African
rodents, however, the barrier effect of the Nile has not been pro-
nounced. The manner in which rodent populations in the past managed
to surmount this apparent barrier is largely unknown, but it may be
related to the developmental history of the river.
According to Said (1962, p. 24), an ancient pre-Miocene precursor
of the Nile is evidenced by fluviatile sands and gravels that overlie
the Eocene in Egypt. The modern Nile is therefore thought to represent
a "much diminutive representative" of this ancient river. If this
larger river was not the actual precursor of the Nile, it apparently
followed a similar course and must have had a similar effect on the
dispersal of rodents.
Opinions differ regarding the developmental history of the Nile.
Said (1962) stated that the valley of the Nile in its present form was
established following a regional uplift near the middle Miocene. The
mechanics of the origin of the Nile Valley are not thoroughly under-
stood, but available evidence indicates that a crustal disturbance
probably established the course of the Nile. Some students feel that
the Nile developed primarily as an erosional feature following a north-
south syneline, while others favor a trough-fault origin. Recent
drilling in the Nile Valley supports the theory that the Nile was eroded
along a line of faulting and rifting (Said, 1962).
Most students of African geology agree that during the period
between the end of the Cretaceous and middle Pleistocene important
tectonic events resulted in the warping and uplifting of large portions
of the African continent. This period was characterized by much fault-
ing and deformation of the land surface. Toward the end of this long
period of violent crustal disturbances, a Pliocene-Quaternary deforma-
tion produced additional movements along the large faults of the
Red Sea and the great Rift Valleys of East Africa (Furon, 1960).
These crustal disturbances presumably had pronounced effects in
North Africa, including the Nile Valley.
In eastern Egypt, it is possible that these tectonic disturbances
caused regional and local deformation of the surface configuration
sufficient to alter the course of the Nile. Several large wadis, such as
the Wadi Araba and the Wadi Ghuweibba, originate east of the Nile
Valley and drain easterly into the Gulf of Suez. During a period of
crustal disturbance one of these wadis could have usurped the function
RODENTS OF LIBYA 53
of the Nile Valley, and the river would thus have entered the Gulf of
Suez, rather than emptying into the Mediterranean Sea as it does
today.
Other less tenable hypotheses include the diversion of the Nile
along the great depressions of northern Egypt and entering the Medi-
terranean near the present city of Benghazi, Libya, on the Gulf of
Sirte. It has also been suggested that the freshwater source for the
chain of oases stretching irregularly across the central Sahara from
Egypt to Morocco represents an "underground Nile" and thus a
remnant of a previous course followed by the Nile River. The similarity
of the fish fauna of these oases, many of which are today separated by
vast tracts of desert, tends to support this hypothesis.
These latter changes in the course of the Nile are strictly conjectural,
but some similar phenomenon is required to explain the present dis-
tribution of Saharan and southwest Asian rodents. In the absence
of the Nile River as a barrier, the Arabian Peninsula, the Middle
Eastern countries, and the Isthmus of Sinai would have provided a
dispersal corridor allowing free interchange of Afro-Eurasian popula-
tions of rodents.
To establish breeding populations of desert rodents on the west side
of the modern Nile in northwestern Egypt and northern Libya, a
faunal immigration of relatively short duration would have been
required. Following the reestablishment of the Nile along its present
course, these "founder populations" of desert rodents in Egypt and
Libya would have been isolated from the ancestral Eurasian popula-
tions and could have provided the "genetic pool" for the ensuing
dispersal of these rodents throughout large portions of North Africa
and the Sahara.
Zoogeographical Relationships of the Rodents of the Sahara and

Southwest Asia
During this study of the rodents of Libya it became necessary to
examine specimens from adjacent parts of the Sahara and from the
desert areas of Southwest Asia. As the study progressed, rather
striking similarities were noted in the composition of the fauna of
such widely scattered areas as the Saharan portions of North Africa,
the deserts of Saudi Arabia, and the dry portions of the Middle
East and Southwest Asia. Several genera of rodents occur exclusively
in these arid portions of North Africa and Southwest Asia, and many
others are widespread and occur only marginally in adjacent parts of
Eurasia and Africa. Most of these genera have little taxonomic
relationship with the rodents of marginal areas.
In view of the high degree of endemism shown by the rodent fauna
of these desert regions of North Africa, Saudi Arabia, and Southwest
;
Asia, it seems appropriate to depart from conventional zoogeo-

graphic concepts and designate this arid belt of steppe and desert
as a distinct zoogeographic subregion of the Palaearctic Realm.
The name Saharo-Sindien Faunal Region is proposed for this new
subregion of the Palaearctie, which coincides rather closely with the
Saharo-Sindien Floral Region comprising the broad belt of steppe
and desert vegetation occurring over this same geographic area.
More specifically, the Saharo-Sindien Faunal Region, referred to
hereafter as the Saharo-Sindien Region, is delimited as: The Saharan
portions of North Africa comprising the region from the Mediterranean
coast southward to the broad belt of Saharan steppe of north-central
Africa; the northern and central Sahara including Egypt, Libya,
Tunisia, Algeria, Morocco, Spanish Morocco, and Spanish Sahara;
the central and southern portions of the Sahara comprising portions
of the Sudan, Chad, Niger, Mali, and Mauritania, and transforming
farther south into the broad steppe region which extends into the
central Sudan, southern Chad, northern Nigeria and Senegal; all of
Saudi Arabia except possibly the extreme southwestern tip; the
steppes and deserts of Sinai, Jordan, Syria, southeastern Turkey,
and Iraq; the dry portions of Iran which comprise many desert and
steppe areas, and the central plateau which includes the desolate
Dasht-i-lut and Dasht-i-Kavir of eastern Iran; Baluchistan in both
Iran and West Pakistan the dry areas of northwestern West Pakistan
;
Afghanistan, excluding the central mountain complex; the Turkmen

Plains, including the Kara and Kyzyl Kum Deserts; and the
Kum
great steppe and desert region east of the Caspian Sea, comprising
the dry regions of the Ust Urt and the Russian provinces of Kazakh-
stan, Uzbekistan, and Turkmenia,
The most striking feature of the Saharo-Sindien Region is the
prevailing aridity and the sparseness of the rodent fauna. This de-
pauperate fauna consists of only 1 1 genera, surprisingly few considering
the great size of the region. All of these genera are especially adapted
to survive under dry conditions and even during periods of pro-
longed drought.
Of the 1 1 genera of rodents occurring in the Saharo-Sindien Region,
almost all are endemic or, at best, occur only marginally in adjacent
portions of Eurasia, India, or Africa. Genera most typical of the
Saharo-Sindien Region include Ctenodactylus, Massoutiera Lataste,
Allactaga, Jacidits, Thomas, Gerbilhos, Pachyuromys,
Calomyscus
Sekeetamys, Meriones, Psammomys, And Rhombomys Wagner. The
genera Gerbillvs, Meriones, and Jaculns occur virtually throughout the
entire Saharo-Sindien Region and are the most typical groups.
The genera Tatera Lataste and Acomys, although widespread in the
RODENTS OF LIBYA 55
Saharo-Sindien Region, are probably commoner in Africa south of

the Sahara.
Some of the 11 genera are restricted to either the African or Asian
portions, but none is confined exclusively to Saudi Arabia. Genera
endemic to North Africa include Ctenodactylus, Massoutiera, and
Pachyuromys. The Asian portion of the Saharo-Sindien Region is
characterized by having the genera Calomyscus and Rhombomys
restricted to it.
Some genera of rodents discussed above occur in adjacent parts of

the Eurasian (Palaearctic), Oriental, and Ethiopian (Africa south of
the Sahara) regions but usually are not widespread and probably
represent marginal invasions. On the other hand, many typical
Eurasian, Indian, and central African genera encroach upon various
parts of the Saharo-Sindien Region. In most cases, these faunal
encroachments have taken place in the more lush habitats provided
by major river systems, mountain complexes, and coastal areas or
have been linked with changes in the landscape wrought by man.
The nonrodent mammalian fauna of this geographic area provides
strong supportive evidence for the hypothesis of a distinct Saharo-
Sindien Faunal Region. Of great importance is the presence in the
Saharo-Sindien Region of 23 endemic species of large (nonrodent)
mammals. Mammals that are restricted to the Saharo-Sindien Region
are:
Sand fox Vulpes ruppelli Schinz

Blanford's fox Vulpes cana Blanford
Fennec Fennecus zerda Zimmermann
Libyan striped weasel Poecilictis libyca Ilemprich and Ehren-
berg
Sand cat Felis margarita Loche
Persian fallow deer Dama mesopot arnica Brooke
Addax Addax nasomaculatus Blainville
Arabian oryx Oryx leucoryx Pallas
Scimitar-horned oryx Oryx tao H. Smith
Dorcas gazelle Gazella dorcas Linnaeus
Slender-horned gazelle Gazella leptoceros F. Cuvier
Korin or red-fronted gazelle Gazella rufifrons Gray
Dama gazelle Gazella dama Pallas
Wild goat Capra hircus Linnaeus
Barbary sheep Ammotragus lervia Pallas
Lesser Moroccan hare Lepus atlanticus de Winton
Arabian hare Lepus arabicus Ehrenberg
Afghan pik;i Ochotona rufesccns Cray
Afghan hedgehog Hemiechinus megalotis Blyth
Ethiopian hedgehog Paraechinus aethiopicus Ehrenberg
Flower's shrew Crocidura floweri Dollman
Egyptian pygmy shrew Crocidura religiosa Geoff roy
Egyptian giant shrew Crocidura olivieri Lesson
;
Many additional species of mammals occur in parts of the Saharo-

Sindien Region but are not restricted to it. These species include the
jackal, Canis aureus Linnaeus; Indian grey mongoose, Herpestes ed-

wardsi Geoffroy; striped hyaena, Hyaena hyaena Linnaeus; Pallas's
cat, Felis manul Pallas; caracal lynx, Felis caracal Schreber; leopard,
Panthera pardns cheetah, Acinonyx jubatus Schreber;
Linnaeus;
Asiatic wild ass (Onager), Equus hemionus Pallas; Persian gazelle,
Gazella subgutturosa Guldenstaedt; mountain gazelle, Gazella gazella
Pallas; ibex, Capra ibex Linnaeus; markhor, Capra falconeri Wagner;
Algerian hedgehog, Erinaceus algirus Duvernoy and Lereboullet;
long-eared hedgehog, Hemiechinus auritus Gmelin Brant's hedgehog,
;
Paraechinus hypomelas Brandt, and the pale grey shrew, Crocidtira

pergrisea Miller.
Being volant, bats are naturally less restricted by physical barriers
and accordingly do not usually conform to standard mammalian
distributional patterns. several genera of bats are
Nevertheless,
rather characteristic ofSaharo-Sindien Region, and several
the
species occur exclusively in these parts of Asia and Africa. Bats most
typical of this region belong to several families which are widely
distributed throughout the temperate and tropical parts of the Old
World. The genera of bats which are most common in the Saharo-
Sindien Region are Rousettus Gray, Rhinopoma Geoffroy, Taphozous
Geoffroy, Nycteris Cuvier and Geoffroy, Rhinolophus Lacepede,
Asellia Gray, Triaenops Dobson, Tadarida Rafinesque, Myotis Kaup,
Eptesicus Rafinesque, Pipistrellus Kaup, Otonycteris Peters, and
Plecotus Geoffroy.
Although none of the above genera are endemic to the Saharo-Sin-
dien Region, the following species are restricted to the area: Rousettus
Win ton; Rhinopoma microphyllum Briinnich;
arabicus Anderson and de
Taphozous nudiventris Cretzschmar; Rhinolophus acrotis Heuglin
Triaenops persicus Dobson; Eptesicus walli Thomas; Eptesicus isabel-
linusTemminek; Pipistrellus deserti Thomas, and Pipistrellus ariel
Thomas.
The above lists show an amount of endemism in the large mammals
and other nonrodent groups comparable to, and in some cases exceed-
ing, that shown by the rodents. This peculiarity of the Saharo-Sindien
mammalian fauna is probably largely attributable to the prevailing
aridity of the region and the presence of vast tracts of desolate steppe
and desert.
The genera of rodents which define the Saharo-Sindien Faunal
Region differ as to origin and dispersal. Some are clearly of Asiatic
origin, others show a closer relationship to the African fauna, while
some appear tohave developed in situ and bear little resemblance to
the fauna of either Asia or Africa.
RODENTS OF LIBYA 57
Today the genus GerbiUus is widely distributed throughout the

steppes and deserts of Southwest Asia and is practically ubiquitous
in the Sahara. Gerbils also occur sporadically throughout most of the
African continent wherever desert or steppe conditions prevail and
are represented by a broad spectrum of species. The greater number
of species occurring south of the Sahara may reflect the more disjunct
nature of suitable habitat in these less arid portions.
Owing to the greater number of species in Africa, probably the
African continent has been the center of differentiation for members
of the genus GerbiUus. The occurrence of gerbils in Southwest Asia,
Saudi Arabia, and the Middle East therefore must be the result of
the northward and eastward dispersal of this group through the
Isthmus of Sinai sometime early in the Tertiary. This hypothesis of an
early northward and eastward dispersal for members of this genus is
supported by the discovery of fossil GerbiUus from the Pliocene of

Asia (Darlington, 1957).
The genus Acomys is widely distributed in Africa but is limited in
Asia to the steppe and desert regions south of the Caucasus Moun-
tains and as far east as the Sind Desert of West Pakistan. Spiny mice
occur also in the intervening deserts of Saudi Arabia and the Middle
East. Because these mice are more restricted ecologically, their dis-
persal must have required long periods of time. As with GerbiUus, the
African continent probably has served as the center of origin for spiny
mice, and the populations of the Middle East and Southwest Asia
represent invasions of the African fauna into Asia.
Although the genus Jaculus is ubiquitous in the Sahara, it clearly
represents an intrusion of the jerboan fauna of central and Southwest
Asia into North Africa. Jerboas are unknown from Africa south of the
Sahara.
The four-toed jerboa, Allactaga tetradactyla, is restricted to the
coastal areas of northern Egypt and northeastern Libya and is the
only representative of the genus in North Africa. This genus occurs
widely in temperate Asia ranging throughout Southwest Asia and
extending north and east into the cold steppes of Russian and Chinese
Turkestan and is represented by several species.
The genera Jacuius and Allactaga are the only African representa-
tives of the Dipodidae. Temperate Eurasia, however, has 10 additional
dipodid genera and a broad assemblage of species. Furthermore, the
fossil record of Asia contains four genera of dipodid rodents extending
as far back as the Miocene (Darlington, 1957). No fossil dipodids are
known from North Africa.
The present distribution of the jerboan fauna of temperate Eurasia,
strengthened by the rather extensive fossil record, provides convincing
285 134 () us — r.
evidence of an Asian origin for the family Dipodidae and the two
genera occurring in Africa.
The genus Meriones is clearly of Asiatic origin and probably repre-
sents a recent arrival of the Eurasian fauna into North Africa. Mem-
bers of the genus Meriones (jirds) are widely distributed in North
Africa in Egypt, Libya, Algeria, Tunisia, and Morocco but are con-
fined more to the northern portions of these countries. In Asia, several
species belonging to this genus are widely distributed in parts of
Transcaucasia, Russian Turkestan, Chinese Turkestan, Iran, Afghani-
stan, Baluchistan, and occur as far east as Manchuria and Mongolia.
The genus Tatera, although unknown from the entire North African
coast and the Sahara proper, occurs extensively in Southwest Asia as
the monotypic Tatera indica Hardwicke. In Africa south of the Sahara
the genus is known polytypically. This hiatus in distribution of the
genus is a zoogeographical enigma which is not readily soluble.
Africa may have served as the center of dispersal for the genus
Tatera, or the genus may represent a comparatively recent invasion
into Africa. The static nature of the genus in Asia, however, may be
the result of prolonged isolation of a peripheral isolate which shows
signs of genetic impoverishment and evolutionary decline. A static
gene pool is indicated by the maintenance of a single monotypic
species over such a large geographic area. The fact that in many parts
of the range of Tatera indica it is closely associated with human
agriculture suggests that this Asian species has indeed lost some of its
original genetic variability.
In view of the extreme discontinuity in the range of the genus
Tatera, the Asian and African complexes must have become isolated
early in the Tertiary and have since evolved independently.
The greater number of African species of Tatera, the more variable
nature of the populational gene pool, and the general similarity of its
range to those of other genera thought to have evolved in Africa, seem
to favor an African center of origin for the genus Tatera, but until the
fossil record provides something more conclusive, I prefer to regard
the geographic origin of Tatera with reservations.
In the preceding pages, discussion has dealt with those genera
common to both the Saharan and Asian portions of the Saharo-
Sindien Region. Other genera characteristic of this newly proposed
zoogeographic region are confined either to the Saharan or Asian
portions.
The family which comprises the genera Cteno-
Ctenodactylidae ,
Blyth and Felovia Lataste, is proba-

dactylus, Massovtiera, Pectinator
bly the most unusual Saharan family. The genera Ctenodactyhts and
Massoutiera are exclusively Saharan, whereas Pectinator and Felovia
RODENTS OF LIBYA 59
approach, respectively, only the southeastern and southwestern mar-

gins of the Sahara.The members of this distinctive family are regarded
by most systematists as being related to the hystricomorph rodents
and therefore have few relatives in Africa. Other hystricomorph genera
in Africa, such as Hystrix, Atherurus Cuvier, Thyronomys Fitzinger,
Pedetes Illiger, and Petromys A. Smith, are, except Hystrix, confined to
Africa south of the Sahara. That the genus Pectinator is "doubtfully
represented in the Pliocene of India" (Darlington, 1957, p. 392)
strengthens the hypothesis of a Saharo-Sindien Faunal Region.
The fat-tailed sand rat, Pachyuromys duprasi, is apparently con-
fined to the Saharan steppe of Egypt, Libya, Tunisia, Mauritania,
and Algeria. This unique, monotypic genus is probably most closely
related to Meriones but unlike the latter genus has undergone an in
situ development in North Africa.
The range of the sand rat, Psammomys obesus, is usually more coastal
than other jirdlike rodents and includes coastal northern Sudan in
addition to North Africa, the Isthmus of Sinai, and adjacent Israel
and Arabia. The genus Psammomys, although a significant element of
the rodent fauna of North Africa, is related to the genera Meriones
and Rhombomys and is probably of Asiatic origin.
Sekeetamys calurus Thomas, the bushy-tailed jird, known only from
Sinai, Israel, and eastern Egypt, is probably the most localized in
distribution of any of the species of Saharo-Sindien rodents. This
species has apparently undergone a rapid in situ development in this
restricted areaowing to its narrow ecological tolerances.
Three monotypic genera, Rhombomys, Calomyscus, and Nesokia, are
confined almost exclusively to the Asian portion of the Saharo-Sindien
Region. Rhombomys ranges from Turkmenia and eastern Iran into
Chinese Turkestan and Mongolia; Calomyscus is found in the dry
foothill areas of Turkmenia, Iran, Afghanistan, and Baluchistan;
the range of Nesokia extends from India and Baluchistan to Arabia
and Egypt.
In addition to the endemic and characteristic genera and species of
the Saharo-Sindien Region, several kinds of rodents have intruded
into the peripheral desert areas of Southwest Asia from Europe, the
high latitude steppes and deserts of Russia, and the subtropical parts
of the Oriental region, while some typical European and African
rodents have penetrated into peripheral Saharan portions of North
Africa.
The squirrel Spermophilopsis Blasius; the dipodid genera Salpingotus
Vinogradov, Pygeretmns Gloger, Paradipus Vinogradov, Dipus Zim-
mermanu, and Stylodipus G. Allen; and the mole-vole Ellobius are
representatives of the central Asian fauna with Eurasian affinities and

occur marginally in the Saharo-Sindien Region.
The palm squirrel Funambulus occurs in Baluchistan and represents
an encroachment of the Indian fauna into the extreme southeastern
portion of the Saharo-Sindien Region.
Examples European rodent fauna which have entered North
of the
Africa include the dormouse, Eliomys quercinus; the Palestine mole
rat, Spalax ehrenbergi; the vole, Microtus mustersi; and the field
mouse, Apodemus sylvaticvs Linnaeus. As discussed earlier, these
species probably entered North Africa during one of the pluvials of
the Pleistocene.
The Barbary ground squirrel, Atlantoxerus getulus, the only North
African squirrel, and the Barbary striped mouse, Lemniscomys barbarus,
are notable exceptions in the North African fauna and are related to
the fauna of central and southern Africa. These two species are limited
respectively to the Algerian Atlas and to the less arid parts of Algeria
and Morocco. The genus Atlantoxerus monotypic, but Lemniscomys
is
is composed Ellerman and Morrison-

of several central African species.
Scott (1951) consider this North African species of Lemniscomys to be
conspecific with one of the central African species.
Some African rodents which encroach into portions of the southern
Saharan steppe include: the sciurids Euxerus Thomas and Heliosciurus
Trouessart; the dormouse Graphiurus Smuts; the cricetid genera
Taterillus Thomas and Desmodilliscus Wettstein, and numerous murid
genera including Arvicanfhis Lesson, Mastomys Thomas, Praomys
Thomas, and Grammomys Thomas.
The various penetrations of the European and African rodent faunas
into marginal portions of the Sahara and the intrusions of the Eurasian
and Indian faunas into portions of Southwest Asia do not refute the
hypothesis of a Saharo-Sindien Faunal Region. The indigenous rodent
fauna (and the nonrodent fauna) of these low latitude deserts of the
Old World differs markedly from that of adjoining Eurasia, India, and
Africa and clearly justifies the formation of this new zoogeographic
region.
The rodent fauna of the Russian steppes and the high latitude
deserts of Manchuria and Mongolia contains some genera which occur
in the northern portions of Southwest Asia, particularly in the vast
steppe region east of the Caspian Sea. Probably it was this arid region
east of the Caspian Sea which served as the center of differentiation
for most of the rodents now comprising the Saharo-Sindien Faunal
Region. The rodent faunas of the Gobi Desert, the Takla Makan of
Sinkiang, and other cold deserts of central Asia probably represent an
outgrowth of the Saharo-Sindien fauna which has since become
adapted to cold, arid climates.
RODENTS OF LIBYA 61
Conclusions
This study of Libyan rodents is based primarily on specimens

obtained by the author during the period from October 1961 to July
1962 and is the first attempt to provide a comprehensive treatment
of the taxonomy and distribution of the Libyan rodents based upon
actual specimens and field experience.
Fifty-six kinds of rodents comprising 7 families, 14 genera, 28
species,and 4S subspecies occur in Libya. In this report, 20 subspecies
are described as new, and 3 forms, Gerbillus aureus, Meriones caudatus,
and Jaculus deserti, which had previously been regarded as subspecies,
are elevated to full species.
Species belonging to the genera Gerbillus, Meriones, and Jaculus
show the widest range of geographic and individual variation found
among the Libyan rodent fauna. The polytypic species Gerbillus
campestris, Gerbillus gerbillus, and Jaculus jaculus, each represented
in Libya by 5 subspecies, are the most widely distributed and the
most genetically variable of any of the Libyan rodents.
Libyan rodents usually do not conform to climatic or ecogeographical
rules, such as Bergmann's or Allen's rule, and geographic variation in
response to selective factors of the environment is most apparent in
the cryptic coloration of the "substrate races" of Gerbillus campestris
and Meriones caudatus.
The populational structure of Libyan rodents is characterized by
two primary components. These are geographical isolates and, less
frequently, a series of contiguous populations showing progressive
change in morphological characters (clines). It is probable that
in Libya most species have arisen as a result of the genetic divergence
of peripheral isolates. Gradual speciation, as opposed to instantaneous
speciation or macroevolution, has been the prevailing mode of specia-
tion for Libyan rodents. Of the two types of gradual speciation, geo-
graphic, rather than sympatric, has played the primary evolutionary
role.
Three faunal areas, which coincide roughly with the major physi-
ographic, vegetative, and climatic features, are recognizable in Libya
based on the distribution of the rodent fauna. Each of these faunal
areas is two regional provinces according to the kinds
divisible into
of rodents most typical of each. The faunal areas of Libya and their
provinces are: The Mediterranean Faunal Area which is comprised
of the Coastal Plain Province and the Cyrenaican Plateau Province;
the Saharan Steppe Faunal Area consisting of the Transitional
Desert Province and the Tripolitanian Gebel Province; and the
Saharan Desert Faunal Area which includes the Cyrenaican Desert
Province and the Fezzanese Desert Province.
Local differentiation of Libyan rodents indicates the presence of

four areas of potential endemism. These areas are the Cyrenaican
Plateau and coastal plain of northern Cyrenaica, the Gebel Nefusa,
the Fezzan, and Cufra Oasis.
The distribution and composition of the Saharan (and Libyan)
rodent fauna has been profoundly influenced by the widespread
shifting of mammal populations during the Pleistocene.
Owing to the similarities of the rodents of North Africa and South-
west Asia, and because of the high degree of endemism shown by
them, the Saharo-Sindien Faunal Region is here proposed as a new
zoogeographic region for this arid belt of steppe and desert. The
genera of rodents defining the Saharo-Sindien Faunal Region differ
as to origin and dispersal. The genera Allactaga, Jaculus, Calomyscus,
Meriones, Psammomys, and Rhombomys are clearly of Asian origin.
The genera Gerbillus and Acomys are apparently of African origin.
The genera Pachyuromys, Massoutiera, and Ctenodactylus probably
have developed in situ in North Africa.
The arid region east of the Caspian Sea possibly has served as the
center of differentiation for most of the rodents comprising the
Saharo-Sindien Faunal Region.
Interchange of Afro-Eurasian rodent faunas probably took place
at a time when the Nile River was less effective as a disperal barrier
than it is today.
Systematic Treatment
Plan of Treatment
The phylogenetic arrangement of the taxonomic categories from

suborder to genus follows Simpson (1945). The species of each genus
and their component subspecies are arranged alphabetically. The major
genera are introduced by a detailed summary of their taxonomic
history, and in the case of Gerbillus, Meriones, and Jaculus, keys are
provided for the identification of the species. Owing to the lack of
standardization of vernacular names of North African rodents and the
inconsistency of their use by workers in the past, they have been
omitted as formal headings and are employed only in a general sense.
The following procedure is employed in the accounts of species
and subspecies
1. The currently accepted binomial name combination in agree-
ment with the "International Code of Zoological Nomenclature"
(1961) is followed on the same line by the name of the author.
2. Original description: The original name
succeeded by the
is
name on
of the author, a reference to the original account, the date
which the foregoing account was published, and the type locality in
parentheses.
RODENTS OF LIBYA 63
3. General distribution of species: The known distribution of the

species in North Africa, Europe, and Southwest Asia is given under
this heading.
4. Distribution in Libya: The entire range of the species in Libya,
including political provinces or portions thereof, based upon specimens
examined and published records.
5. Distribution of the subspecies in Libya: Each subspecies is
arranged alphabetically and its range in Libya briefly outlined

according to physiographic features and political boundaries.
6. Published records: In this section all known records from the
literature are summarized for each political province. For each province
the records are arranged chronologically, and when two or more
localities arerepresented they are listed in alphabetical order. The
author's name and the date of publication appear in parentheses
following each locality record or series of localities.
Comparisons: Comparisons are limited to those species which
7.
are easily confused with each other and separable only by subtle
characters. Because most of the Libyan species of rodents are quite
distinct morphologically, this section is frequently omitted.
8. Remarks: The taxonomic history of the species is covered in
this section, and problems and events relating to systematics and
dispersal are discussed.
9. Ecological observations: This section deals with the habitat
requirements of the species and discusses other species of rodents

which share the same habitat.
After the general discussion at the species level, the following pro-
cedure is employed in the treatment of the subspecies:
1. The currently accepted trinomial name combination is followed
on the same line by the name of the author.
2. Original description: The original name is succeeded by the
author, a reference to the original account, the date on which the
foregoing account was published, and the type locality in parentheses.
Specimens examined: Specimens examined, unless otherwise
3.
qualified, are skins and skulls. The total number examined is given
first; this is followed by the name of the province, the exact locality
of capture, and the exact number from this locality. Collecting sites
from Cyrenaica are listed first, followed by those from Tripolitania
and the Fezzan. Within the provinces, collecting sites are arranged in
sequence from north to south. Those having the same latitudinal
coordinate are arranged from east to west.
Sight and verbal records and records of occurrence from the literature
are included, respectively, in the sections on "remarks" and "published
records." Tn a few cases, specimens were examined from the mammal
collections of the British Museum (Natural History), London, and
Museum National d'Histoire Naturelle, Paris; these are indicated,

respectively, by the abbreviations BM
and MNHN. Unless otherwise
stated, all other catalog numbers represent specimens in the collections
of the Division of Mammals, United States National Museum, Smith-
sonian Institution, Washington, D.C.
The distribution maps, in most cases, indicate localities from which
I examined specimens. These are indicated by solid black circles. The
few localities showing published records are indicated by a solid black
triangle. The shaded portions of the maps (vertical and horizontal
lines) indicate the range of a given species or subspecies in Libya, as
judged from known records of occurrence. No species or subspecies
occurs in all parts of the shaded area but is limited to areas of suitable
habitat.
4. Measurements: All measurements are given in millimeters and
are of adults unless otherwise indicated. Given first are the four
standard body measurements: Total length (from the tip of the
rostrum to the end of the last caudal vertebra) length of tail (from
;
the proximal end of the first caudal vertebra to the distal end of the
last caudal vertebra) length of hind foot (from the proximal end of
;
the calcaneus to the end of longest claw) length of ear (from the base
;
of the notch to the distal end of the pinna). Cranial measurements,

taken by dial calipers, follow the external measurements. When the
measurements of more than three specimens are given, they are listed
as average and extremes. If there are three specimens or less, their
measurements are listed individually, and the museum catalog number
is given for each.
5. Diagnosis: Capitalized color terms are those of Ridgway
"Color Standards and Color Nomenclature" (1912). Noncapitalized

color terms are employed in those instances where a detailed color
description âs deemed unnecessary. Whether the description repre-
sents winter or summer pelage is specified in those subspecies where
seasonal pelage colors differ significantly. In most cases detailed
descriptions involving the gross features of the skull and, in some
cases, characters useful in specific and subspecific diagnosis are given.
6. Comparisons: Specimens from Libya are compared with those
representing marginal subspecies to determine their true taxonomic
position. Whenever possible, paratypical and topotypical specimens
are used in these comparisons to better validate the assignments. In
some instances, comparisons were made with all subspecies comprising
a given species, which frequently necessitated examination of speci-
mens from Egypt, Sudan, Tunisia, and Algeria.
7. Remarks: All systematic problems and enigmas are considered
under this heading. Evidences of intergradation with other subspecies
and their interpretation are within the province of this section. Also
RODENTS OF LIBYA 65
considered are the implications of physiography and geography and

their roles in generating evolutionary developments and morphological
differences. Plausible avenues of dispersal are considered whenever
affinities and relationships with other populations are held unac-
countable. Range extensions are also included here.
8. Ecological observations: This section provides information on
behavior and some aspects of natural history with emphasis on habitat
preferences. Considerable detail is given to the types of plants and
the nature of the substrate forming the various habitats.
9. Description of new subspecies: The format for describing new
subspecies is identical to that used for the accounts of recognized

subspecies except the newly proposed subspecific name is followed by
the designation of the holotype. Data relating to the holotype are
recorded in the following sequence: Sex; an indication as to whether
the holotype represents skin, skull or both; museum of repository
(USNM = United States National Museum); catalog number; exact
collecting locality; date of collection; name and original field number
of the collector.
10. Analysis of variation and the diagnostic characters of the
species and subspecies: In an effort to determine the extent of indi-
vidual and geographic variation within the various species and sub-
species of Libyan rodents and to ascertain the measurable characters
which distinguish them, specimens were selected from localities
which represented the most typical representatives of the particular
taxon and were then treated statistically. In some instances, on the
specific level, it was necessary to pool specimens from several localities
in order to increase sufficiently the size of the sample and thus in-
crease statistical reliability. Generally, however, the analyses are
based on specimens from a single locality. In determining the amount
of morphological divergence and variation within and between sub-
species, paratypical or topotypical specimens were used whenever
possible. In all of the analyses and comparisons, the sexes have been
treated separately to prevent erroneous conclusions arising from
sexual dimorphism.
Before taxonomic variation could be determined for a population
of rodents, the amount of nongeographic variation had to be taken
into account. In this latter category are included random genetic
variation inherent in the gene pool of the species or subspecies and
the differences resulting from growth, age, sex, seasonal changes,
pathological conditions, and other kinds of variation related to known
stimuli. To prevent misleading interpretation of the variation associ-
ated with geographical distribution, extreme care was taken to insure
that all specimens in the sample were of comparable age, and those
suspected of showing seasonal characters, pathological conditions, or
other abnormalities were not used. Series of specimens were analyzed

by standard statistical methods which involved computation of the
arithmetic mean, standard deviation, and standard error of the mean.
In some instances, in which the absolute size of a character is the
same in two taxa, scatter diagrams are employed to demonstrate
differences in the relative size of this character.
Graphical methods employed here are patterned after those given
by Hubbs and Hubbs (1953). The range of variation is shown by a
heavy horizontal line, and the mean is indicated by a sharply pointed
triangle.
The black portion of each bar comprises two standard errors of the
mean on either side of the mean. The entire bar (black and white
portions) constitutes one standard deviation on either side of the
mean. In samples of four specimens, two standard errors and one
standard deviation have the same value, and the entire bar is there-
fore black. Samples of less than four specimens were not treated
statistically. In these cases, the range of variation is shown by a
heavy horizontal line, the mean is indicated by a sharply pointed
triangle, and the bar is omitted. A single triangle indicates that only
one specimen is represented. The standard deviation (S.D.) indicates
8-
RODENTS OF LIBYA 67
the dispersion about the mean, and the standard error (S.E.) indicates
the degree of reliability of this dispersion.
According to Hubbs and Hubbs (1953), a large amount of overlap
of the dark bars (S.E.) indicates a low reliability of the observed
difference between two samples, and any appreciable separation of
these bars indicates a high reliability. These same authors recommend
plotting one standard deviation on either side of the mean to indicate
probable subspecific difference by a nonoverlap of the bars. This
method indicates an 84 percent separation of the specimens by the
character being analyzed.
Gazetteer of Principal Collecting Localities
Cyrenaica
Agedabia, 10 km S
Agedabia, 20 km SW
El Agheila, 65 km WNW
El Agheila, 5 km W
Augila
Bardia, 5 km W
Benghazi, 8 km N
Bzema Oasis
Fort Capuzzo, 10 km SW
Coefia, 2 km N
Derna, 5 km SE
Ain el Gazala, 11 km E
Gerdes, 10 km N
Gheminez
Gialo
Gialo, 150 km S
Giarabub
Giarabub, 24 km SSE
El Giof, Cufra Oasis
Bir el Gobi, 60 km S
Gubba, 12 km NW
Bir el Harasc
Gebel el Harug el Asued, 200 km SE Zella
El Hauuari, Cufra Oasis
Wadi el Kuf, 13 km WSW Messa
Maraua, 7 km E
Merg (=Barce)
Messa, 35 km W
Wadi er Rueis, 340 km WNW Tazerbo
Oasis
Gasr es Sahabi
Susa (= Apollonia), 11 km SW
Tazerbo Oasis
Tocra, 2 km W
Tocra, 20 km SW
Gazetteer of Principal Collecting Localities — Continued

Cvrenaica — Continued
Locality
number
Tobruch, 20 km E (31° 58 N, 24° 08 E) 14
Bahr el Tubat, 21 km ESE Giarabub (29° 36 N, 24° 53 E) 25
Ain Zueia, Gebel Uweinat (21° 53 N, 24° 50 E) 37
Tripolitania
Bir Allagh, 55 km SW
Wadi Bey, 45 km W Bu Ngem
Bu Ngem, 30 km S
El Cusher
Cussabat, 5 km W
Derg, 5 km E
Gharian, 25 km N
El Gheddahia, 7 km S
Ain Hammam
Hun, 2 km SW
Gebel Limhersuk
Marble Arch, 15 km WNW
Mizda, 5 km N
Nalut, 40 km ENE
Rumia, 20 km E
Rumia, 3 km W
Sinauen, 40 km N
Sirte, 20 km E
Sirte, 5 km E
Sirte, 12 km W
Socna, 5 km S
Gebel es Soda, 60 km S Socna
Zliten, 12 km W
El Abiad, 60 km SW Sebha
Umm el Araneb
Brach
Edri
El Gatrun
Ghat, 20 km N
Goddua, 26 km N
Goddua
Meseguin
Murzuch, 28 km E
Murzuch
Sebha
Serdeles, 55 kin SSW
Temenhint, 30 km NK Sebha
Traghen
Ubari, 75 km W
RODENTS OF LIBYA 69
Family Cricetidae
Subfamily Microtinae
Genus Microtus Schrank

Microtus tniistersi Hinton
Microtus mustersi Hinton, Ann. Mag. Nat. Hist., ser. 9, vol. 1<S, p. 305, September
1926 (Merj [=Barce], Cyrenaica).
General distribution of species. Libya, on the Cyrenaican

Plateau and adjacent coastal plain.
Specimens examined. Six, from Cyrenaica: 10 km SW El Faidia,
1 and skeleton); Merg ( = Barce), 2 (BM;
(skin, skull, 1 skin only); 5
km W Tocra, 3.
Published records in Libya. Cyrenaica: Barce (Hinton, 1926).

;
Measurements. The measurements of two immature males,

325019 and 325021, and of an adult male, 325020, from 5 kilometers
west of Tocra, Cyrenaica, are respectively: Total length 131, 130, 130;
length of tail 30, 25, 31 length of hind foot 18, ?, 18; length of ear 1 1,
;
10, 12; condyloincisive length of skull 24.6, 25.8, 26.5; greatest breadth
across zygomatic arches 14.4, 14.5, 15.9; length of nasals 6.7, 6.9, 6.7; least
interorbital breadth 3.7, 3.9, 3.8; crown length of upper molariform
toothrow 5.9, 6.4, 6.4; greatest breadth across braincase 11.4, 11.7, 12;
length of incisive foramina 4.4, 4.5, 4.5.
Diagnosis. Entire dorsum uniformly colored Buckthorn Brown,
becoming slightly paler on flanks and sides and forming a distinct con-
trast with the white-tipped hairs of the venter all hairs of dorsum and
;
venter Plumbeous basally; pinnae of ears short and inconspicuous,

Prout's Brown, sparsely haired, and with several long brownish hairs
partially covering ventrolateral surfaces; eye ring absent; vibrissae
relatively short and fine and composed of both light and dark individ-
ual hairs; dorsal surfaces of forelegs, hindlegs, and feet light tan;
ventral surfaces of fore and hind feet naked with prominent carpal and
palmar tubercles; front feet with four functional digits with claws, the
first digit reduced to a rudimentary tubercle; hind feet with five func-
tional digits with claws; tail noticeably short and distinctly bicolored
dark brown above and buff below. Skull: Medium in size; compact
and angular; zygomata heavy with a uniform lateral curvature; inter-
orbital breadth markedly constricted; suprameatal portion of auditory
bulla enclosed by processes of the supraoccipital and temporal bones;
parietal ridges markedly reduced and inconspicuous; interparietal
transversely elongated nasals short and moderately flaring anteriorly
;
upper incisors markedly prognathous; dorsal margin of foramen

magnum with prominent notch; audital portions of auditory bullae
conspicuously inflated ventrally; pterygoid processes and external
pterygoid fossae heavily fenestrated; hamulae large and applied to
anteromedial portions of auditory bullae; molariform toothrows long
and individual teeth with prominent inner and outer salient angles;
anterior palatine foramina relatively short and narrow; posterior
palatine canals small and inconspicuous.
Comparisons. Compared to representatives of Microtus guentheri
Danford and Alston from the Valley of Ali Bey Dere, near Piringcikov,
16 kilometers west of Istanbul, Turkey, the specimens from Libya
differ strikingly in having markedly smaller, less angular skulls, nar-
rower rostra, more ventrally inflated auditory bullae, more fragile
zygomata, narrower basioccipitals, markedly smaller incisors, and the
third molar nearly as large as the second, rather than being signifi-
cantly smaller as in guentheri. In external characters, the specimens
RODENTS OF LIBYA 71
from Libya are significantly larger, have less hair on the ears, are more
brightly colored dorsally (Buckthorn Brown as opposed to Snuff
Brown), have a greater suffusion of white ventrally, more orangish-
buff on the dorsal surfaces of the fore and hind feet, and have more
prominently bicolored tails.
In M. m,ustersi the tail is markedly longer than the hind foot and
almost one fourth the length of the head and body, rather than only
slightly larger than the hind foot and one-fifth the length of the head
and body as in M. guentheri, and the sole of the hind foot is naked for a
greater distance posterior to the base of the toes.
In the number of palmar and plantar pads and in the pattern and
number of the molar prisms and reentrant angles, the Libyan speci-
mens resemble M. guentheri and indicate relationship with the latter
species.
Remarks. Hinton (1926) described M. mustersi as a species distinct
from Microtus philistinus Thomas of Palestine. According to Hinton,
M. mustersi is closely allied to M. philistinus, but differs in slightly
darker (less grayish) dorsal color, slightly narrower choanae, slightly
smaller auditory bullae, less reduction of the third molar posteriorly,
and less salient temporal ridges with a wider interval between them.
Hinton regarded these differences as somewhat tenuous but stated
that "craniologically" the specimens representing both Af. in ustersi and
M. philistinus, and upon which his comparisons are based, were all
subadult and that fully adult specimens probably would show more
marked differences.
Ellerman and Morrison-Scott (1951) regarded M. mustersi and
M. philistinus as conspecific and relegated them both to subspecific
rank under M. guentheri. Toschi (1954) also considered M. mustersi
as a subspecies of M. guentheri. I agree with Ellerman and Toschi in
regarding M. mustersi as a member of the "M. guentheri" group but
feel that other morphological differences between the two are too
great to suggest conspecificity, and here M. mustersi is regarded as a
species distinct from M. guentheri.

Hinton's description of Microtus mustersi from Cyrenaica in 1926
constituted the first record of occurrence of a microtine rodent for the
African continent, even though members of this group were widespread
throughout Europe and Asia. The type series of At. mustersi obtained
by Chaworth-Musters in 1926 from Barce, a specimen collected by
H. W. Setzer in 1955 from 10 kilometers southwest of rDl Faidia, and
the three specimens that I obtained from 5 kilometers west of Tocra
are all from Cyrenaica and constitute the only known representatives
of the subfamily Microtinae in Libya.
All specimens from Cyrenaica are either topotypes or near topotypes

of M. mustersi. Those from the coastal plain near Tocra, however, are
the farthest from the type locality, but two specimens are clearly
immature and are unsuitable for comparative purposes. The third is
of questionable adult status but

near or of comparable age to those
is
specimens with which it is being compared from Merg (= Barce) and

El Faidia. The specimen from Tocra is identical in color to M. mustersi.
In cranial characters it resembles rather closely a topotype of M.
mustersi from Barce but differs in slightly smaller size of skull and
external dimensions, shorter and narrower anterior palatine foramina,
slightly smaller pterygoid foramina, and slightly more inflated
mastoidal portions of the auditory bullae.
From the specimen from near El Faidia, this specimen from near
Tocra differs in the same characters as set forth above, except the two
specimens are comparable in the length of the anterior palatine
foramina.
These differences between the voles of the coastal plain and those
from the higher slopes of the plateau are too slight to suggest sub-
specific differences and indicate that M. mustersi is a monotypic
species whose range includes all of the Cyrenaican Plateau and the
adjacent coastal plain wherever suitable habitat occurs.
Ecological observations. The three localities in Cyrenaica
where voles are known to occur differ markedly in the character of
the habitat. The type locality at Barce, according to Hinton (1926),
is at an altitude of 300 meters and in a basin of interior drainage
about 20 miles from the coast. Microtus inhabited open burrows with
well marked "runs" in the cornfields. The ground was described as
being "hard-baked." Near El Faidia, Setzer (1957) described the
terrain as hilly and rocky, and a single specimen was obtained from
the moist north-facing slope where mosses were growing. This locality
is significantly higher in elevation than the Barce Valley and is near
the highest point of the Cyrenaican Plateau. In contrast to the high

valleys and mountain slopes near Barce and El Faidia, the site near
Tocra is located on the lowest level of the coastal plain about one-half
kilometer from the Mediterranean Sea. This particular portion of the
coastal plain has a dense vegetative cover consisting primarily of
interwoven patches of tamarix, large sedges, coarse grasses, and a
species of bushlike woody perennial with purple flowers. All these
plant-types form rather discrete communities composed almost
exclusively of a single dominant type of plant. All voles were taken
the same night from the dense understory of the bushlike perennials.
Large series of Mus musculus and Gerbillus campestris and several
specimens of Crocidura russula Hermann were also obtained from this
RODENTS OF LIBYA 73
bushlike growth. The areas of tamarix, sedges, and grasses were

trapped intensively, but yields were quite poor.
This portion of the coastal plain, with its lush and profuse vege-
tative cover, is not typical of most of the Cyrenaican coastal plain,
and probably at these lower elevations voles are of very sporadic and
local occurrence. The range of M. mustersi, in Libya, therefore prob-
ably is largely confined to the higher and more typically montane
areas of the Cyrenaican Plateau and the Gebel Achdar, which provide
more abundant and continuous habitat.
The occurrence of voles in Cyrenaica poses some very interesting
zoogeographic problems, particularly since they are not known from
coastal Egypt, the only conceivable dispersal route. It is possible that
the population of voles in Cyrenaica are relicts from the period when
the more boreal climates of the Pleistocene prevailed over North
Africa. The Cyrenaican Plateau by virtue of its higher elevation has
retained at least a vestige of these boreal elements, whereas the
remainder of coastal Africa has become much drier and more sparsely
vegetated and no lons-er contains habitats suitable for voles.
Subfamily Gerbillinae
Genus Gerbillus Desmarest

Lataste (1881, p. 506) divided the genus Gerbillus into two genera,
Gerbillus and Dipodillus Lataste, and separated the latter from
Gerbillus by the number of plantar pads, the pattern of infolding of
enamel on the surfaces of the teeth, and by characteristics of the
auditory bullae. According to Lataste, typical members of Gerbillus
had one carpal pad and no plantar pads, while representatives of
Dipodillus had five carpal and six plantar pads. Lataste used the
character of naked feet to separate two species, Gerbillus garamautis
Lataste and Gerbillus hirtipes Lataste. Heptner (1937) and Ellerman
(1941) suppressed Dipodillus as a full genus. The latter author stated
that members two genera could not be distinguished by cranial
of the
characters and that there was a lack of constancy in the characters
used by Lataste to distinguish between the two genera. Later, Wassif
(1956) and Setzer (1957) regarded Dipodillus as a subgenus of the
older prevailing Gerbillus and used the presence or absence of hair on
the hind feet to assign their specimens to the two subgenera. The
present author uses this same character to separate the subgenera
Dipodillus and Gerbillus, each represented in Libya by four species,
285 134 O —68 6

A
RODENTS OF LIBYA 75
C —
A
RODENTS OF LIBYA 77
./
4 _§ -
Figure 8. — Statistical comparison of length of hind foot of the species of Gerbillus. Notation
remains the same as in figure 5.
14 |
^ I ?
c _
Figure 9. — Statistical comparison of least interorbital breadth of the species of Gerbillus.

Notation remains the same as in figure 5.
B _ o
C _ o"
_ 20
Figure 10. — Statistical comparison of upper molariform toothrow of the species of the
subgenus Gerbillus: A, G. aureus; B, G. eatoni; C, G. gerbillus; D, G. pyramidum.
RODENTS OF LIBYA 79
36
* Gerbillus aureus
O Gerbillus pyramidum
35
34
Q
ooo
o
^ o
-. 33
32
oo
o
30
* *
* *
*
#*
# *
29
**
28
12 CROWN LENGTH
3
OF UPPER
4
MOLARIFORM
5
T00THR0W
6
of upper molariform toothrow relative to oc-

Figure 11— Comparison of crown length
cipitonasal length of Gebillus aureus and Gerbillus pyramidum.
RODENTS OF LIBYA 81
95
85
* #
*
#
- 80
*
3 75 #
70
65
• Gerbillus henleyi
-jfe- Gerbillus kaiseri
60
70 75 80 85 90
65
LENGTH OF HEAD AND BODY
Figure 13. — Comparison of length of tail relative to length of head and body of Gerbillus
henleyi and Gerbillus kaiseri.
S2 U.S. NATIONAL MUSEUM BULLETIN 2 75
O Gerbillus henleyi
# Gerbillus kaiseri
27
26
~ 25
i 24
23
22 o
o o
21
7 8 9 10
LENGTH OF AUDITORY BULLA
Figure 14. — Comparison of length of auditory bulla relative to occipitonasal length of

Ccrbdlus henleyi and Gerbillus kaiseri.
RODENTS OF LIBYA S3
32 O Gerbillus amoenus
* Gerbillus campestris
*
31 *
*
**
***
**
**
30
*
#*
29
28
27
26
O
oo
o
25
ooo
ooopo
o
24
13
11 12
10
LENGTH OF AUDITORY BULLA
to occipitonasal length of
Figure IS.— Comparison of length of auditory bulla relative
Gerbillus amoenus and Gerbillus campestris.

the former consisting of G. amoenus, G. campestris, G. henleyi, and

G. kaiseri and the latter composed of G. aureus, G. eatoni, G. gerbillus,
and G. pyramidum.
Without doubt, gerbils are the most variable morphologically and
the most ecologically tolerant of all the Libyan rodents. They have
adapted to practically every available environmental situation. The
ability of these gerbils to exploit such a wide array of ecological
opportunities indicates a broad genetic constitution. This marked
adaptability has enabled them to occupy a vast geographic range in
Libya and has contributed to the maintenance of a common mor-
phological pattern among the widely scattered populations.
Owing broad genetic constitution of the species of the genus
to the
Gerbillus, there is wide variation in color and in cranial and external
characters of the component subspecies. Individual subspecies vary
markedly in any given character and generally are distinguished
from each other by an aggregate of characters rather than by specific,
measurable cranial differences. In gerbils, the usual criteria for recog-
nizing subspecies, such as differences in color and cranial and ex-
ternal dimensions, are employed, but these differences are usually
much more subtle than in other genera of Libyan rodents.
Owing to this genetic plasticity in gerbils, the usual 84 to 93 percent
rule (1 to 1.5 standard deviations on either side of the mean) for
separating the members of two subspecies by a given character is
seldom realized, and the percentage of overlap in a character is
usually far greater.

The ranges of many of the subspecies of gerbils in Libya are con-
tiguous, and gene exchange is of common occurrence. These inter-
grading populations contain genes from both subspecies and contribute
significantly to the maintenance of genetic variability among the
populations of gerbils in Libya.
In most populations of a given species, and particularly in those
subspecies which are geographic isolates, genetic fixation occurs
rapidly and trends develop toward morphological distinctness.
Key to the Subgenera of Gerbillus
Plantar surfaces of hind feet hairy Gerbillus

Plantar surfaces of hind feet naked Dipodillus
.
RODENTS OF LIBYA 85
Key to the Specie? of the Subgenus Gerbillus
1. Skull prominently domed; anterior palatine foramina markedly enlarged.

G. eatoni
Skull not prominently domed; anterior palatine foramina not markedly en-
larged 2
2. Occipitonasal length of skull less than 30 mm; length of upper molariform
toothrow less than 4 mm G. gerbillus
Occipitonasal length of skull usually more than 30 mm; length of molari-
form toothrow usually more than 4 mm 3
3. Tail with distinct brush; the latter never black G. pyramidum
Tail without distinct brush; the latter frequently black G. aureus
Gerbillus aureus Setzer
Gerbillus pyramidum aureus Setzer, Proc. Biol. Soc. Wash., vol. 69, pp. 170-180,
Dec. 31, 1956 (12 km W
Zliten, Tripolitania Province, Libya).
General distribution of species. Libya; range probably also

includes coastal areas of Tunisia.
Distribution in Libya. Coastal plain and littoral deserts of
northern Tripolitania.
Distribution of the subspecies in Libya.
Gerbillus aureus aureus. Tripolitania: Gebel Nefusa and the
coastal plain between Azizia and the Gulf of Sirte.
Gerbillus aureus favillus. Tripolitania: Vicinity of Sirte; range
probably includes much of the coastal plain of the Gulf of Sirte.
Gerbillus aureus nalutensis. Tripolitania: Coastal plain of extreme
northwestern Tripolitania
Comparisons. This species can be distinguished from Gerbillus
pyramidum, which it most closely resembles, by its markedly smaller
skull, smallerbody, shorter, less tufted tail, generally darker color,

shorter and wider posterior palatine canals, wider anterior palatine
foramina, and longer molariform toothrows relative to occipitonasal
length.
From Gerbillus gerbillus, G. aureus differs in larger body and skull,
larger molariform teeth, markedly less tufted tail, larger anterior
palatine foramina, more robust skull, heavier zygomata, and generally
darker, less orangish dorsal color.
Gerbillus aureus can be readily separated from Gerbillus eatoni by
its larger body, larger and heavier skull, larger molariform teeth,
smaller anterior palatine foramina, longer posterior palatine canals,
smaller and less inflated auditory bullae, and markedly more flattened
braincase.
Remarks. Setzer described Gerbillus pyramidum aureus ( = Gerbillus
aureus aureus) (1956, p. 179) and Gerbillus pyramidum favillus
(= Gerbillus aureus favillus) (1956, p. 180) as new subspecies of G.
pyramidum. In naming these new subspecies, he had only topotypical
RODENTS OF LIBYA 87
In color and overall body size, members of G. aureus from Libya

closely resemble representatives of Gerbillus pyramidum hirtipes
Lataste from Ain Sefra, southwestern Algeria, but differ from the
latter in having; noticeably longer tails. Cranially, the two groups
are of comparable size, but can be distinguished by the more massive,
compact skulls, heavier zygomata, and wider rostra of the Algerian
specimens. Ellerman and Morrison-Scott (1951), in keeping with their
objective of simplifying nomenclature, relegated Lataste's Gerbillus
hirtipes (1882) to subspecific status under the more widely distributed
Gerbillus pyramidum. At this time Tripolitania and the coastal areas
of western Libya were not represented by specimens, and Gerbillus
aureus was unknown. In the majority of external and cranial characters
these specimens from Algeria are much closer to G. aureus than to G.
pyramidum; in fact, they bear little resemblance to the latter. When
more localities in Algeria are represented by specimens of G. aureus and
G. pyramidum and when topotypes of G. p. hirtipes are available for
comparison, the latter probably be reinstated as a full species,
will
distinct from either pyramidum but more closely re-
G. aureus or G.
lated to the former, or aureus and hirtipes may be regarded as con-
specific, in which case the older hirtipes will take precedence.
Ecological observations. Vegetated dunes are apparently the
preferred habitat of this gerbil, but they occur also in the littoral
deserts where dunes are usually lacking. In the Gebel Nefusa, the
habitat consists of rolling uplands with rather dense vegetative cover,
and sand, if present, is usually localized.
Gerbillus aureus aureus Setzcr
Gerbillus pyramidum. aureus Setzer, Proc. Biol. Soe. Wash., vol. 69, pp. 179-180,
Dec. 31, 1956 (12 km W Zliten, Tripolitania Province, Libya).
Specimens examined. Forty, from Tripolitania: 12 km W

Zliten, 12; 25 km N Gharian, 8; 20 km E Rumia, 8 (1 skin only) ; 3 km
E Rumia, 10; 12 km S Chicla, 2.
Measurements. Averages and extremes of six adult males and
measurements of two adult females, 302073 and 302075, from the
type locality, are: Total length 236.8 (223-244), 222, 230; length of
tail 131.8 (127-137), 125, 129; length of hind foot 30.7 (30-31), 30,
30; length of ear 15.2 (13-18), 13, 13; occipitonasal length of skull
30.7 (29.9-31.6), 29.2, 30; length of auditory bulla 11.1 (10.9-11.2),
10.7, 10.9; crown length of upper molariform toothrow 4.1 (4-4.3), 4.1,
4.1; greatest breadth across zygomatic arches 16 (15.6-16.5), 15.6,
15.9; least interorbital breadth 5.7 (5.5-6), 5.4, 5.4; breadth of ros-
trum at level of antorbital foramina 3 (2.9-3.1), 2.8, 2.9; greatest
length of nasals 11.7 (11.4-11.9), 11, 11.5.
Diagnosis. Members of this subspecies show pronounced variation

in dorsal color,which ranges from uniform, brilliant Ochraceous-Buff
in some specimens to more subdued Tawny-Olive in others. This
darker color results from a strong admixture of dark-tipped hairs.
These darker specimens are characterized as follows: Mystacial,
rostral, and circumoral areas and supraorbital patch Light Buff; post-
auricular region Clay Color heavily suffused with black; eye ring
black; pinna of ear dark distally (Hair Brown) with fringe of buff-
colored hairs on anterior margin; anteromedial surface of pinna Cin-
namon-Buff; vibrissae short, with occasional dark hairs; lateral
surface of forearm with buffy patch; fore and hind feet and legs
densely haired dorsally and ventrally and each bearing five digits with
claws; relatively long and indistinctly bicolored Cinnamon-Buff
tail
dorsally and Light Ochraceous-Buff ventrally, becoming more sub-

dued distally and appearing Light Ochraceous-Buff; terminal pencil
relatively reduced and Hair Brown; entire underparts white. The
lighter, more brilliantly colored specimens have white rostral, mysta-
cial, and circumoral areas, much lighter vibrissae, more distinct
postauricular patches, a smaller supraorbital patch, more conspicu-

ously bicolored and lack the buffy patch on the forearm. Skull:
tails,
Medium in size, somewhat gracile and narrow in dorsal aspect;

upper molariform toothrow relatively short and individual teeth
small; anterior palatine foramina large; auditory bullae small and
ventrally inflated; zygomatic arches almost parallel to orbital surface.
Comparisons. From the type series of Gerbillus aureus favillus,
Gerbillus aureus aureus differs in larger size and larger cranial measure-
ments, being comparable in crown length of upper molariform tooth-
row, least interorbital breadth and breadth of rostrum at level of
antorbital foramina. In color, G. a. aureus is darker and more varied
in dorsal color and has a markedly more distinct and darker penicillate
tail. Members of the nominate subspecies have more parallel tooth-
rows, more ventrally inflated auditory bullae, relatively longer poste-

and the zygomatic arches are parallel rather than
rior palatine canals,
bowed medially. Setzer (1956) used most of the foregoing characters
to separate these two subspecies but stated that the skulls of animals
of these two subspecies Mere of comparable size. The present study,
however, shows that representatives of G. a. aureus are significantly
larger in almost all cranial measurements.
For comparison with Gerbillus aureus nalutensis, see account of
that subspecies.
Remarks. This subspecies apparently represents a genetically
heterogeneous group, and the component populations represent a
rather broad assemblage of morphological patterns. Specimens from

the vicinity of Rumia are slightly darker than those from the type
RODENTS OF LIBYA 89
locality and have prominent tufts on the tail.

lighter colored, less
The common "golden" color phase not present in
is also apparently
these gerbils from Rumia. "In contrast, the series from 25 kilometers
north of Gharian consists almost exclusively of gerbils having this
"golden" color. Animals from near Gharian also are wider across the
zygomatic arches and have larger molariform teeth. In the aggregate,
however, many morphological characters are common to all these
different populations and serve to unite them into a single subspecies.
Setzer (1957) assigned a single specimen (no. 302102) from 5
kilometers west of El Agheila, Cyrenaica Province, to Gerbilhis
pyramidum aureus ( = Gerbilhis aureus aureus). This specimen resem-
bles Gerbilhis aureus in color and general size but in the prominent
tufted tail and all other cranial and external characters, it is typical
of Gerbilhis pyramidum to which it is here referred.
Ecological observations. The specimens from near Rumia and
Chicla were taken near the brink of the coastal escarpment. The
habitat here consists of gently rolling valleys and depressions with
rather dense vegetative cover consisting of a wide assemblage of
close-growing perennial and annual species. The ground in this area
is quite rocky, and the soil has a clay or sandy base. Shallow deposits
of smooth sand and small dunes are present, but these are of rare
occurrence. The collecting site north of Gharian is located on the
coastal plain where large, permanently vegetated dunes are present.
I have not visited the type locality near Zliten, but presumably the
habitat resembles that near Gharian.
Gerbillus aureus favillus Setzer
Gerbillus -pyramidum favillus Setzer, Proc. Biol. Soc. Wash., vol. 69, pp. 179-180,
Dec. 31, 1956 (2 km E Sirte, Tripolitania Province, Libya).
Specimens examined. Twenty, from Tripolitania: 5 km E Sirte,

8; 2 km E Sirte, 12.
Measurements. Averages and extremes males and 13 females
of 7
from the above localities are, respectively: Total length 226.1 (216-
240), 220.1 (210-236); length of tail 125.7 (119-135); 119.3 (110-132);
length of hind foot 30.4 (30-31), 30.3 (30-31); length of ear 14.4
(13-15), 13.6 (13-15); occipitonasal length of skull 29.7 (29.3-30.5),
29.4 (28.4-30.1); length of auditory bulla 10.5 (10.3-10.8), 10.5
(10.3-10.9); crown lengthupper molariform toothrow 4.1 (4.1-4.2),
of
4.2 (4-4.3); greatest breadth across zygomatic arches 15.5 (15.1-16.6),
15.7 (15.2-16.3); least interorbital breadth 5.8 (5.5-6), 6 (5.7-6.2);
breadth of rostrum at level of antorbital foramina 3.2 (3-3.3), 3.2
(3.1-3.4); greatest length of nasals 11.4 (11-12), 11.3 (10.4-13.1).
Diagnosis. Upperparts uniformly Clay Color becoming paler on
sides and with admixture of brownish hairs on back; post auricular
285 134 O — <",s 7
patches conspicuous and white; supraorbital patch white; mystacial,

circumoral and pectoral areas, forelegs, hindlegs, feet, and entire
underparts pure white; pinna of ear sparsely haired, darker colored
distally, Cinnamon-Buff on anteromedial surface, and bearing a row
of fine buffy hairson anterior margin; vibrissae short and composed
almost entirely of white hairs; fore and hind feet bearing five digits
with claws; tail relatively short, distinctly bicolored Cinnamon-Buff
dorsally, Pale Pinkish Buff ventrally, and with faint terminal Avella-
neous pencil. Skull: Relatively small, gracile, and flattened in profile;
anterior palatine foramina relatively wide; upper molariform tooth-
rows bowed slightly laterally, relatively short, and individual teeth
small;auditory bullae relatively small; zygomatic arches bowed
slightly medially.
For comparisons with G. a. aureus and G. a. nalutensis, see accounts
of those subspecies.
Remarks. This subspecies is known only from the coastal plain in
the vicinity of Sirte on the western shores of the Gulf of Sirte, but
its range probably includes a much greater area to the east. I was
unable to obtain additional specimens of this subspecies and, conse-
quently, obtained no firsthand data regarding their habitat preferences.
Setzer (1956, p. 181) describes the habitat at the type locality as
consisting of "loose coastal dunes where the vegetation had been
heavily eaten by domestic animals." The pale dorsal color of these
gerbils may result from a genetic response to the color of these sandy
areas.
One specimen, no. 302096, from 5 kilometers east of Sirte has large
molariform teeth and anterior palatine foramina, a flattened brain-
case, and is These characters are typical of
pallid in dorsal color.
G. a. faviUus, andspecimen is referred to that subspecies. This
this
same specimen, however, in size of body, external measurements,
and the small size of the skull closely resembles animals belonging to
GerbiUus (jerbiUus. This parallelism probably does not indicate a direct
relationship with G. gerbillus but more likely is the result of a response
to similar environmental conditions or represents an extreme in the
range of variation for the subspecies G. a.faviUus.
(lerbillus aureus nalutensis, new subspecies
Holotype. Adult male, skin and skull, USNM

321816, from 40 km
ENE Nalut, Tripolitania Province, Libya: obtained Nov. 12, 1961,
by H. W. Setzer, original no. 3113.
Specimens examined. Forty, all from the type locality.
Diagnosis. Upperparts Tawny-Olive with strong suffusion of
blackish hairs in interauricular and subanricular regions and on rum]);
sides and flanks paler than dorsum; eye ring black; postauricular and
RODENTS OF LIBYA 91
supraorbital patches white; mystacial, rostral, circumoral, and

pectoral areas white, the latter tinged with buff; vibrissae short and
with numerous dark hairs; pinna of ear drab-colored distally, basally
approaching color of dorsum and with row of buffy hairs on anterior
margin; fore and hind feet densely haired, white, and with five digits
bearing claws; entire underparts pure white; tail Cinnamon-Buff with
admixture of brownish hairs dorsally, grading to Pinkish Buff ventrally
and terminating in a distinct blackish-brown pencil. Skull: Relatively
large and robust, auditory bullae large and bulbous; molariform
toothrows long, individual teeth large; rostrum wide; braincase wide
and flat.
Measurements. Averages and extremes of 17 adult males and

13 adult females from the type with the measurements of
locality,
the type in brackets, are, respectively: Total length 231.2 (21S-249),
224 (213-238), [234]; length of tail 130.4 (117-142), 125.6 (106-132),
[130]; length of hind foot 29.2 (27-31), 28. S (27-31), [30]; length of
ear 14.9 (14-16), 14.3 (13-15), [15]; occipitonasal length of skull 30.3
(29.1-31.5), 29.5 (28.7-30.3), [30.7]; length of auditory bulla 11.1
(10.5-11.5), 10.9 (10.2-11.4), [11.3]; crown length of upper molariform
toothrow 4.3 (3.9-4.5), 4.3 (3.9-4.4), [4.5]; greatest breadth across
zygomatic arches 16.2 (15.7-17.1), 15.6 (15.1-16.1), [16.3]; least
interorbital breadth 5.9 (5.3-6.3), 5.8 (5.5-6.1), [6]; breadth of
rostrum at level of antorbital foramina 3.3 (2.9-3.4), 3.1 (3-3.4),
[3.4]; greatest length of nasals 11.6 (11.1-12.2), 11.3 (10.6-11.6),
[11.8].
Comparisons. From topotypes of Gerbillus aureus aureus, the
type and paratypes of G. a. nalutensis differ in having more robust
skulls, markedly larger and wider auditory bullae, markedly longer
molariform toothrows, larger individual molariform teeth, wider
braincases, and wider and shorter rostra. In color, G. a. nalutensis is
somewhat darker dorsally with more suffusion of black hairs on the
rump, and the tail has a darker colored pencil.
This new subspecies can be distinguished from Gerbillus aureus
favillus by darker dorsal color, darker (almost black) pencil, markedly
longer molariform toothrows, larger size of individual teeth, much
larger, more bulbous auditory bullae, more domed braincase, wider
rostrum, and markedly larger size of body and cranium.
Remarks. Although a fundamental morphological pattern is
always present in members of this subspecies, they vary considerably
in color and cranial characters. As in the other two subspecies of
G. aureus, dorsal color ranges from the typical dark form, with a
heavy suffusion of black, to the more uniformly colored "golden"
form. Specimens showing all gradations in intermediate color are also
known. Cranially, this subspecies is remarkably polymorphic, partic-
ularly as concerns the size and shape of the auditory bullae, the
relative length and width of the anterior palatine foramina, the
degree of arching of the braincase, the length of the molariform
too throw, and the size of the individual molariform teeth. This poly-
morphism suggests the presence of sibling species within this popula-
tion from Nalut, but overlap of characters occurs too frequently,
and no specific combinations of characters are demonstrable for
individual specimens. Apparently this population from Nalut repre-
sents one of pronounced genetic fluidity which has engendered this
wide array of morphological patterns.
Although this new subspecies is known from only the type locality,
its range doubtless includes the coastal plain and littoral deserts of
southern Tunisia and, in Libya, probably extends much farther

along the coastal plain to the east. In Libya, there are no apparent
barriers to dispersal along the coastal plain, and intergradation
between animals of G. a. nakitensis and those of the nominate sub-
species takes place probably somewhere between Tripoli and Nalut.
Ecological observations. The type locality consists of a series
of vegetated dunes. These dunes are not extensive and occur sporadi-
cally as irregular bands along the coastal plain near the coastal
escarpment. The role of the coastal escarpment in retarding the
velocity of the winds probably has resulted in the formation of these
coastal dunes.
Forty-seven gerbils and five jirds were obtained from a single night
of trapping at 40 kilometers east-northeast of Nalut. The presence
of numerous active burrows plus this large catch suggests a high
population density of rodents. A severe wind storm or "ghibli"
10 2 10 4 10 6 10 8 11 11.2 114 116
Figure 17. — Statistical comparison of length of auditory bulla of the subspecies of Gerbillus
aureus: A, G. a. aureus; B, G. a. favillus; C, G. a. nalulensis.
RODENTS OF LIBYA 93
_ 2 , ?
2 8 29 30 31 32 33 34
Figure 18. — Statistical comparison of breadth of rostrum of the subspecies of Gerbillus

aureus. Notation remains the same as in figure 17.
developed during the night but had little apparent effect on trapping
success.
The name nalutensis is used in reference to the village of Nalut on
the westernmost limits of the Gebel Nefusa and located a short distance
from the type locality.
Gerbillus eatoni Thomas

Gerbillus eatoni Thomas, Proc. Zool. Soc. London, vol. 2, pt. 1, p. 6, October 1902
(El Cusher, Tripolitania Province, Libya).
General distribution of species. Libya; range probably also

includes coastal areas of northern Algeria, Tunisia, and Egypt.
Distribution in Libya. Coastal areas of Tripolitania and
Cyrenaica.
Gerbillus eatoni eatoni. Cyrenaica: Coastal plain and littoral deserts
in vicinity of Agedabia and north to Gheminez; Tripolitania:
Coastal plain of the Gulf of Sirte, probably including the coastal
areas to the west.
Gerbillus eatoni inflatus. Cyrenaica: Coastal plain and littoral
deserts north and east of the Cyrenaican Plateau.
Gerbillus eatoni versicolor. Cyrenaica: Coastal plain of Cyrenaica
in vicinity of Benghazi.
Published records in Libya. Cyrenaica: Vicinity of Benghazi
(Klaptocz, 1909); Tripolitania: Wadi Agarib, Wadi Aggar, El
i
RODENTS OF LIBYA 95
three localities in northern Tripolitania. Setzer (1957) recognized G.

on the characters established by Thomas.
eatoni as a full species largely
He collected both speciesfrom 5 kilometers west of El Agheila, and
in the present study, G. gerbillus and G. eatoni were found to occur
together at several localities in coastal Tripolitania and Cyrenaica
without any evidence of interbreeding. The present study indicates
that G. eatoni is a fidl species and can be distinguished from other
species of Gerbillus by the same diagnostic characters as set forth by
Thomas and by Setzer.
Although this species is not abundant in Libya, and large series
are lacking, local differentiation has occurred sufficiently to warrant
the recognition of three distinct subspecies, each restricted to a
particular portion of the Libyan coastal plain. In Libya, the range of
this species probably includes all of the coastal regions and in many
areas, the transitory desert associated with the coastal escarpment.
It is unknown from the Saharan interior where G. gerbillus is the
dominant species.
Ecological observations. These gerbils are usually found in
areas of loose sand on the coastal plain but frequently occur in habi-
tats lacking sand. In many
areas the coastal plain is reduced to a
narrow fringe beach or is entirely eliminated. Thus, these
of desolate
coastal populations of gerbils sometimes occur in a variety of eco-
logical conditions.
Throughout their range in Libya these gerbils occur with several
species of jirds (Meriones) and jerboas (Jaculus), fat-tailed sand rats
(Pachyuromys) and gerbils of the subgenus Dipodillus.
,
Gerbillus eatoni eatoni Thomas

Gerbillus eatoni Thomas, Proc. Zool. Soc. London, vol. 2, pt. 1, p. 6, October
1902 (El Cusher, Tripolitania Province, Libya).
Specimens examined. Twenty-five, from Cyrenaica: 4 km W

Gheminez, 6; 10 km S Agedabia, 8; 5 km El Agheila, W 4 (2 skin
only); from Tripolitania: 5 km W
Sirte, 4; El Cusher, 1 (subadult
BM) 15 km
; WNW
Marble Arch, 2.
Measurements. Averages and extremes of three males and four
females from 10 kilometers south of Agedabia, Cyrenaica Province,
are, respectively:Total length 214 (208-218), 214.8 (210-222); length
of tail (117-131), 124.5 (122-126); length of hind foot 27.7
124.3
(27-28), 27.3 (26-29); length of ear 14.3 (13-15), 14.5 (13-15);
occipitonasal length of skull 28.5 (28.1-28.9), 28.4 (28.1-28.7); length
of auditory bulla 10.6 (10.5-10.7), 10.5 (10.3-10.6); crown length of
upper molariform toothrow 4 (4-4.1), 4 (3.9-4.2); greatest breadth
across zygomatic arches 15.2 (15-15.3), 15.4 (15.1-15.6) least inter- ;
orbital breadth 5.7 (5.6-5.7), 5.6 (5.4-5.7); breadth of rostrum at

level of antorbital foramina 2.9 (2.8-3), 3 (2.9-3.2); greatest length

of nasals 10.6 (10.6-10.7), 10.7 (10.5-10.8).
Diagnosis. Upper parts Sayal Brown suffused with Bister; tail
Cinnamon-Buff and bicolored owing to strong suffusion of brownish
hairs dorsally; postauricular patches prominent and white; pinna of
ear sparsely haired, Clay Color basally and becoming darker distally;
vibrissae short and formed from about equal numbers of brown and
white hairs; eye ring dark brown; mystacial, rostral and pectoral
areas, flanks and sides Cinnamon-Buff; forelegs, hindlegs, and feet
white dorsally, sparsely haired ventrally and each bearing five digits
with claws; entire underparts white. Skull: Relatively small and
gracile; upper molariform teeth relatively short; anterior palatine
foramina short and wide; posterior palatine canals relatively short;
auditory bullae large and markedly inflated; braincase prominently
arched.
Comparisons. Members of the nominate subspecies can be dis-
tinguished from those representing other subspecies in Libya by
their smaller size and the marked doming of the braincase. For more
detailed comparisons with Gerbillus eatoni inflatus and Gerbillus eatoni
versicolor, see accounts of those subspecies.
Remarks. Only a single, subadult topotype of G. e. eatoni is available
for this study. Whenspecimens of comparable age from various locali-
ties along the Gulf of Sirte are compared to this topotype, they differ
in having lighter dorsal color, less doming of the skull, and relatively
smaller and less inflated auditory bullae. Even though specimens from
near Agedabia, Cyrenaica, do not conform precisely to the topotype
of G. e. eatoni, they constitute a series of sufficient size for comparisons
with other subspecies of G. eatoni in Libya and in the following accounts
will be used to represent the diagnostic characters of typical G. e.
eatoni.
Specimens from near Gheminez represent the northeasternmost
occurrence of gerbils expressing characters typical of G. e. eatoni and,
in the majority of characters, are referable to the nominate subspecies,
but in dorsal color and length of tail, they show intergradation with
G. e. versicolor to the north.
Oneold and aberrant male specimen, no. 325311, from Agedabia,
shows characters of both G. e. eatoni and G. e. versicolor. In its marked
divergence of the upper molariform toothrows, less domed braincase,
and particolored dorsal pelage, it is nearer to Gerbillus eatoni versicolor.
This specimen is here referred to the nominate subspecies because it
resembles the latter in size and degree of inflation of the auditory
bullae and in the majority of other cranial characters.
Ecological observations. The habitat at Agedabia is character-
ized by extensive sandy plains with occasional small dunes. Vegetative
cover is relatively dense and composed of bushy succulent shrubs and
;
RODENTS OF LIBYA 97
other more ephemeral plants. The specimens from near Sirte were
collected from the abundant vegetation of the coastal plain. At this
locality, the soil is composed of firmly packed clay and sand, and
dunes are lacking.
Throughout much of its range in northern Tripolitania, this
subspecies occurs with members of Gerbillus aureus and Gerbillus
gerbillus.
Gerbillus eatoni inflatus, new subspecies
325527, from 10 km
SW Fort Capuzzo, Cyrenaica Province, Libya; obtained June 1, 1962,
by G. L. Ranck, original no. 2203.
Specimens examined. Seven, all from the type locality.
Measurements. Averages and extremes of five adult males and the
measurements of an adult female, 325528, from the type locality,
with the measurements of the type in brackets, are, respectively:
Total length 226 (218-236), 219, [225]; length of tail 125 (121-131),
120, [123]; length of hind foot 27.2 (26-28), 27, [26]; length of ear 15.4
(15-16), 16, [15]; occipitonasal length of skull 30.6 (29.9-31.7), 30.2,
[30.5]; length of auditory bulla 11.1 (10.7-11.6), 11.2, [11.2]; crown
length of upper molariform toothrow 3.9 (3.7-4), 3.9, [3.7]; greatest
breadth across zygomatic arches 16.1 (15.6-16.4), 16.2, [16]; least
interorbital breadth 5.9 (5.5-6.5), 6.1, [5.6]; breadth of rostrum at
level of antorbital foramina 3 (2.8-3.1), 2.9, [3.1]; greatest length of
nasals 11.7 (11.4-12.3), 12, [11.5].
Diagnosis. Middorsal region uniformly colored Sayal Brown grad-
ing to Clay Color on sides (one specimen, 325559, represents a
very old male and is noticeably lighter and more brilliant in dorsal
color with decidedly yellowish hues rather than brownish tones)
subauricular region suffused with blackish hairs; postauricular patch
distinct and white; rostral, mystacial, supraorbital, and pectoral areas,
dorsal surfaces of forelegs, hindlegs, feet, and entire underparts white;
some specimens with a faint buffy patch on lateral surface of forearm;
pinna of ear Antimony Yellow basally and becoming darker distally;
anteromedial surface of pinna covered with ochraceous hairs which
extend laterally to anterior margin of pinna as a distinct tuft; eye ring
black; vibrissae short and formed from equal numbers of light and
dark hairs; fore and hind feet sparsely haired ventrally and with five
digits bearing claws; ground color of tail uniformly Cinnamon with
slight admixture of brownish hairs dorsally and buffy hairs ventrally;
pencil indistinctand Avellaneous. Skull: Relatively large and massive;
auditory bullae conspicuously large and inflated braincase moderately
;
vaulted upper molariform teeth relatively small and toothrow bowing

;
slightly laterally; anterior palatine foramina large and expanded

laterally; posterior palatine canals wide and distinct; pterygoid
fossae shallow.
Comparisons. Only a subadult, topotypical specimen of Gerbillus

eatoni eatoni is available for study, and because no specimens of
Gerbillus eatoni injlatus of comparable age are available, valid com-

parison is not possible. Specimens of G. e. injlatus, however, can be
readily distinguished from those representing the nominate subspecies
from 15 kilometers west-northwest of Marble Arch, Tripolitania
Province, and from 10 kilometers south of Agedabia, Cyrenaica
Province, by their darker, more uniform dorsal color, generally
larger overall size, and conspicuously larger and more inflated auditory
bullae. Members of this subspecies also differ from those representing
G. e. eatoni in having much wider and less domed braincases, wider
and longer posterior palatine canals, relatively shorter upper molari-
form toothrows, more open pterygoid fossae, and relatively shorter
tails.
For comparison with Gerbillus eatoni versicolor, see account of that

subspecies.
Remarks. Representatives of this subspecies can be separated
easily from all by then- markedly larger
other subspecies of G. eatoni
and more inflated auditory bullae and larger size of body and cranium.
The type series was taken from a sandy depression in the hamada
near the brink of the coastal escarpment. In these localized depressions,
the substrate is composed of poorly sorted sand and gravel and
supports a rather uniform plant cover of juniper-like species, large
grasses, and smaller, woody shrubs. Burrows are usually widespread
throughout these areas and suggest relatively high population densi-
ties. Jerboas also occur in this habitat.
Although this subspecies is known only from the type locality, its
range probably includes the coastal plain and littoral deserts of
northern Cyrenaica and those of northern Egypt as far east as the
Nile River Delta.
The name injlatus refers to the pronounced inflation of the auditory
bullae.
Gerbillus eatoni versicolor, new subspecies
Holotype. Adult male, skin and skull, USNM 325298, from 2 km N

Coefia, Cyrenaica Province, Libya; obtained June 15, 1962, by
G. L. Ranck, original no. 2341.
Specimens examined. Twenty-five, from Cyrenaica: 2 km N
Coefia, 16 (2 skin only); 8 km N Benghazi, 9 (1 skin only).
Measurements. Averages and extremes of five adult males and
four adult females from the type locality, with the measurements of
the type in brackets, are, respectively: Total length 222.4 (216-236),
221 (205-236), [236]; length of tail 124.2 (119-134), 121.3 (114-130),
[134]; length of hind foot 27 (26-28), 27.3 (27-28), [28]; length of ear
16 (15-17), 15.8 (15-16), [15]; occipitonasal length of skull 29.5
(28.7-30.3), 29.2 (28.8-30), [30.3]; length of auditory bulla 10.4
RODENTS OF LIBYA 99
(10-10.8), 10.3 (10-10.6), [10.8]; crown length of upper molariform

toothrow 3.9 (3.8-3.9), 3.9 (3.8-4), [3.8]; greatest breadth across
zygomatic arches 15.9 (15.4-16.8), 15.8 (15.4-16.5), [16.8]; least
interorbital breadth 5.8 (5.5-6.3), 5.8 (5.4-6), [6.3]; breadth of
rostrum at level of antorbital foramina 3.2 (3.1-3.4), 3.3 (3-3.4), [3.4];
greatest length of nasals 11.5 (11.4-11.8), 11.4 (11.3-11.6), [11.8].
Diagnosis. Sides and dorsum variegated or particolored ranging
from Pinkish-Cinnamon to Cinnamon; all areas, particularly the
sides, with admixture of lighter colored hairs; postauricular patch
prominent and Light Buff; mystacial, cir cum orbital, and pectoral
areas Pinkish Buff; eye ring black; underparts generally white, but,
in some specimens, with a ventromedian line suffused with Light
Pinkish Cinnamon; dorsal surface of forelegs, hindlegs, and feet Pale
Pinkish Buff; fore and hind feet sparsely haired ventrally and bearing
five digits with claws; pinna of ear relatively long and almost naked,
Cinnamon-Buff basally and Deep Grayish Olive distally, and with
row of delicate buff-colored hairs on anterior margin; vibrissae
relatively short and composed of about equal numbers of white and
brown hairs; tail faintly bicolored Cinnamon-Buff dorsally and
Pinkish Buff ventrally and bearing an indistinct Drab terminal tuft
or pencil. Skull: Relatively large and compact; braincase moderately
inflated; interorbital region with distinct longitudinal, median fossa;
zygomata relatively heavy; upper molariform toothrow relatively
short, straight, yet divergent anteriorly; anterior palatine foramina
largeand widely open; posterior palatine canals markedly wide and
bowed slightly laterad; auditory bullae relatively short and moder-
ately inflated ; nasals relatively long.
Comparisons. This new subspecies differs markedly from specimens
representing Gerbillus eatoni eatoni from 15 kilometers west-northwest
of Marble Arch, Tripolitania Province, and those from 10 kilometers
south of Agedabia, Cyrenaica Province, in darker, more variegated or
particolored dorsal pelage (Cinnamon as opposed to Clay Color), more
anteriorly divergent and shorter upper molariform toothrows, shorter,
narrower, and less inflated auditory bullae, greater length of skull,
shorter and wider posterior palatine canals, longer nasals, and less
doming of the braincase.
From a subadult, topotypical specimen of G. e. eatoni from El
Cusher, Tripolitania Province, specimens of comparable age from
Coefia are paler in dorsal color, have less distinctly bicolored tails,
lack the strong suffusions of brownish hairs on the middorsum, have
relatively and absolutely markedly smaller arid less inflated auditory
bullae, much less inflated braincases, and longer posterior palatine
canals.
From specimens representing Gerbillus eatoni inHatus from 10 kilo-
meters southwest of Fort Capuzzo, Cyrenaica Province, G. e. versicolor
is slightly smaller in overall size, has noticeably shorter and much less
ventrally inflated auditory bullae, smaller and more gracile skull,
larger upper molariform teeth which form straight toothrows rather
than being bowed laterally, narrower and smaller braincase, slightly
shorter nasals, paler, more uniform dorsal coloration (Cinnamon as
opposed to Sayal Brown), and less prominent subauricular patches.
Remarks. Members of this subspecies can be distinguished from
other subspecies of G. eatoni in Libya by their particolored dorsal
pelage, more parallel upper molariform toothrows, and conspicuously
smaller and less inflated auditory bullae.
Specimens from 8 kilometers north of Benghazi are similar to G. e.
inflatus in more uniform color of dorsum and general body size, but
resemble the nominate subspecies in having larger and more domed
braincases and more lateral bowing of the upper molariform teeth. In
the majority of characters, however, they are closer to G. e. versicolor
to which they are here referred. A small series from Gheminez, al-
though similar in color to G. e. versicolor, is clearly referable to the
nominate subspecies in all morphological characters.
The type localities of G. e. versicolor and G. e. inflatus are, geo-
graphically, not too distant, but the Gebel Achdar and the massif of
the Cyrenaican Plateau are interposed between them and provide
unsuitable habitat for members of this species. Little suitable habitat
owing to the encroachment of the coastal
exists along the coastal plain
escarpment. In the past, gene exchange between these two populations
probably has been of rare occurrence, and the two populations have
undergone significant morphological divergence.
The type series was collected from an area of large, sparsely vege-
tated coastal dunes lying between the sea and the coastal plain. It is
doubtful if these gerbils are entirely limited to this type of habitat.
The from near Benghazi occupied the rather densely vegetated
gerbils
coastal plain where the substrate was claylike, and sandy areas or
dunes were entirely lacking. These gerbils probably are not limited to
a particular type of substrate and occur throughout the coastal plain
where a wide variety of soil conditions exist.
The name "versicolor," from the Latin meaning variegated or of
different colors, refers to the particolored dorsal pelage.
Gerbillus gerbillus (Olivier)
Dipus gerbillus Olivier, Bull. Sci. Phil. Paris, vol. 2, p. 121, 1801 (Giza Province,
Egypt).
General distribution ofspecies. Israel, Sinai, Egypt, Sudan,

Uganda, and North Africa south through the Sahara including parts
of Niger, Mauritania, Chad, and Mali.
Distribution in Libya. Almost ubiquitous throughout the coastal
and interior areas of Cyrenaica, Tripolitania, and the Fezzan.
RODENTS OF LIBYA 101

Gerbillus gerbillus aeruginosus. Cyrenaica: Cufra and Bzema
oases and the serirs and oases of extreme southeastern Libya.
Gerbillus gerbillus discolor. Fezzan: Oases, serirs, hamadas, and
wadis linking together the various Fezzanese oases.
Gerbillus gerbillus gerbillus. Cyrenaica: Environs of Giarabub
Oasis.
Gerbillus gerbillus latastei. Tripolitania: Desert areas of central
Tripolitania and the Hamada Hamra.
el
Gerbillus gerbillus psammophilous. Cyrenaica: Coastal deserts

near Gulf of Sirte, Oases of Gialo and Tazerbo, eastern margins of
the Gebel el Harug el Asued, and the intervening Serir of Calanscio.
Published records in Libya. Cyrenaica: Augila, El Giof, Es
Sahabi, Gialo, Giarabub (de Beaux, 1932); Benghazi (Toschi, 1951);
Tripolitania: Ain Hammam, Wadi Aggar, "Attich Loulileh,"
"Loumoulieh," "Shup" (Thomas, 1902); Bu Ngem (Toschi, 1951);
Fezzan: El Baharia, Gara el Hamra, Hatiet el Fachri, Hatiet er
Zeroi, Sebha, Bir el Wasti (de Beaux, 1928) Wadi Tenezoft, Murzuch
;
(Toschi, 1951).
Comparisons. Based on measurements of Gerbillus dallonii Heim
de Balsac as given in the original description (Heim de Balsac; 1936,
Gerbillus gerbillus
I oeruginosus
2 discolor
3 gerbillus
4- lotostei
5. psammophilous
Figure 20. — Distribution of the subspecies of Gerbillus gerbillus.

p. 45), G. gerbillus can be distinguished by its shorter hind feet, longer

rostrum, longer ears, longer molariform toothrow, and markedly
larger size of body and cranium.
Remarks. Previous workers in Libyan mammals recognized little
variation in G. gerbillus, and all of Libya was included within the range
of the nominate subspecies. Much larger series are now available
from Libya, and this species is now known to be represented by five
subspecies and not one as previously supposed.
In many areas of Libya, isolated populations of these gerbils occur
throughout seemingly barren wastelands. These areas support only a
meager vegetative cover, and frequently plant life is entirely wanting.
These widely scattered populations are characterized by common
morphological characters which indicate at least some degree of
genetic relationship. Apparently, seasonal or occasional rains in these
desert areas enable plants, primarily grasses, to undergo their brief
life cycle and produce abundant quantities of seed which allow small
populations of these gerbils to subsist until the next period of mois-

ture. By successive seasonal or yearly storms, populations of repro-
ductively effective size are able to progressively extend their range.
These periods of rainfall may occur within the same year or be spaced
over a period of several years. In this manner, small populations of
gerbils would be able to "bridge" vast areas in the desert which
otherwise would prove insurmountable.
These shifting populations illustrate another significant aspect of
distribution. Failure of seasonal or periodic rains can cause the ex-
termination of "bridging" populations just as their occurrence can
perpetuate such populations. Failure of rains thus sets up isolating
barriers. Occurrence or lack of rain insures a fluctuating pattern of
genetic "input" and "output" over the range of the species and also
makes it probable that the pattern observed now will be altered in
subsequent sampling.
Ecological observations. Members of this species are the most
ubiquitous of the subgenus Gerbillus in Libya. They are not known
to occur in habitats lacking sand and are widely distributed through-
out the oases, serirs, and hamadas of the interior. They are commonest
along the sandy margins of the oases but occur also in the coastal
areas wherever sandy areas are present. They are unknown from the
coastal areas of Tripolitania and are not found in Cyrenaica north of
the Cyrenaican Plateau. In these areas, they are supplanted by
Gerbillus eatoni and members pyramidum group.
of the Gerbillus
Throughout their range inLibya they are sympatric with many
other species of rodents but most frequently occur with Gerbillus
pyramidum and Jaculus jaculus.
Gerbillus gerbillus aeruginosus, new subspecies

Holotype. Adult male, skin and skull, USNM 325146, from El
Giof, Cufra Oasis, Cyrenaica Province, Libya; obtained Apr. 3, 1962,
Specimens examined. Eighty-four, from Cyrenaica: Bzema Oasis,
8 (1 skin only); El Hauuari, Cufra Oasis, 36 (1 skin only); El Giof,
Cufra Oasis, 40 (2 skin only).
Measurements. Averages and extremes of 16 adult males and 8
adult females from the type locality, with the measurements of the
type in brackets, are, respectively: Total length 205 (200-215),
205.4 (200-213), [206]; length of tail 117.7 (109-129), 120 (115-128),
[119]; length of hind foot 29.4 (28-32), 28.4 (27-31), [29]; length of
ear 13.2 (13-14), 12.8 (12-13), [13]; occipitonasal length of skull 28
(27.3-28.8), 27.4 (26.8-27.8), [28.4]; length of auditory bulla 10.3
(9.8-10.8), 10.3 (10.1-10.7), [10.4]; crown length of upper molariform
zygomatic arches 15 (14.6-15.5), 14.7 (14.3-15.1), [14.7]; least inter-
orbital breadth 5.7 (5.4-6.1), 5.7 (5.4-5.9), [5.9]; breadth of rostrum
at level of antorbital foramina 3 (2.7-3.3), 3 (2.9-3.1), [3]; greatest
length of nasals 10.3 (9.8-10.6), 10.1 (9.7-10.6), [10.4].
Diagnosis. Interauricular, subauricular, interorbital, and rostral
areas same color as dorsum and ranging from Ochraceous-Buff, to
Clay Color, to tawny-rust; circumorbital, postauricular spots,
mystacial and pectoral areas, fore and hind limbs, and entire under-
pays white; vibrissae short and white; fore and hind feet sparsely
haired dorsally and ventrally, and each bearing five digits with
claws; pinna of ear almost naked and Pale Orange- Yellow with a row
of buffy hairs on anterior margin; tail long and distinctly bicolored,
ranging from Capucine Buff to Cinnamon dorsally and from Pinkish
Buff to Light Buff ventrally and with a distinct pencil, Light Buff
ventrally, and varying in dorsal color from Avellaneous in darker
colored individuals to Pale Pinkish Cinnamon in paler animals. Skull:
Small and gracile; anterior palatine foramina short and elliptical in
shape; posterior palatine canals short and bowed slightly laterally;
teeth small; zygomata fragile and convergent anteriorly; braincase
slightly inflated and skull rather flattened in profile; auditory bullae
moderately inflated ventrally and partially transparent; basioccipitals
narrow anteriorly; basisphenoid reduced to a narrow rib between
anteromedial edges of auditory bullae; pterygoid fossae large and
forming a depression dorsal to pterygoid hamulae; supraorbital bead
projecting over dorsal rim of orbit.
Comparisons. Compared with topotypical Gerbillus gerbillus
gerbillus from Alexandria Road, 5 kilometers northwest of the Pyramids
of Western Desert Governorate, Egypt, Gerbillus gerbillus

Giza,
aeruginosus differs in smallersize, shorter tail and hind feet, smaller
cranial measurements, heavier zygomata, slightly larger and more

divergent upper molariform toothrows, more laterally expanded
pterygoid fossae, slightly larger lachrymals, and narrower basioc-
cipitals. In color, G. g. aeruginosus is darker dorsally, the tail is less
distinctly bicolored, and the buffy dorsal color extends farther ventrally
and laterally. In general, the range of color in G. g. aeruginosus is
much greater than in the nominate subspecies.

Compared with topotypes of Gerbillus gerbillus asyutensis Setzer
from the beginning of the Wadi el Asyuti, 13 miles southeast of
Asyuti, Eastern Desert Governorate, Egypt, G. g. aeruginosus differs
in darker, more uniform and less variegated dorsal pelage, larger
posterior palatine canals, larger molariform teeth, narrower basioc-
cipitals, less ventral inflation of auditory bullae, larger anterior
palatine foramina, and larger size of all external and cranial measure-
ments.
Specimens of Gerbillus gerbillus sudanensis Setzer from the Anglo-
Egyptian Sudan can be readily distinguished from G. g. aeruginosus
by their paler dorsal color and markedly smaller external and cranial
measurements.
From measurements of Gerbillus gerbillus agag Thomas from El
Fasher, Anglo-Egyptian Sudan, G. g. aeruginosus differs in shorter
tail, slightly larger body size, shorter skull, shorter molariform
toothrow, shorter nasals, and greater breadth across the zygomatic
arches. Only three subadult specimens of G. g. agag are available for
study and, in color, these closely approximate G. g. aeruginosus.
Gerbillus gerbillus aeruginosus differs markedly from gerbils repre-
senting Gerbillus gerbillus andersoni de Winton from the vicinity of
Alexandria, Egypt. The latter are noticeably larger in all cranial
characters, particularly in the size of the anterior palatine foramina,
posterior palatine canals, molariform teeth, and the auditory bullae.
The skull is more domed and narrower in G. g. andersoni,and the
dorsum and ears are conspicuously darker.
For comparisons with Gerbillus gerbillus discolor and Gerbillus ger-
billus psammophilous see accounts of those subspecies.
,
Remarks. In body and cranial size, animals referable to Gerbillus

gerbillus aeruginosus are among the smallest of the species of Gerbillus
gerbillus in Libya. This trend toward small body and cranial size
is apparent in all populations of this species to the east and south.
G. g. asyutensis from central Egypt, G. g. sudanensis, and G. g. agag

from the Sudan, and gerbils representing G. g. aeruginosus from the
Tibesti Area of the Chad, all represent populations of gerbils of small
body and cranial size. These common characters may indicate taxo-
nomic relationships among these scattered populations, and when the

intervening areas are represented by specimens, they will probably
contain intergrades. A clinal gradient of progressive increase in size
probably will be demonstrable from east to west.
In most characters, gerbils from Bzema Oasis are referable to
G. g. aeruginosus, but in greatest length of skull and dorsal color,
they show intergradation with Gerbillus gerbillus psammophilous to
the north. The nearest populations of G. g. psammophilous which
could provide the gene pool causing these intermediate characters
occur in the vicinity of Bir Bu Zarregh and Bir el Harasc north of the
Sand Sea of Rebianna. Apparently, then, this sand sea is not an
absolute barrier but a filter for genetic exchange between animals
psammophilous and G. g. aeruginosus.
strictly referable to G. g.
Animals belonging to G. g. aeruginosus are exceptional in their
wide range of dorsal coloration, and three distinct groups are recog-
nizable, ranging from Ochraceous-Buff, to Clay Color, to tawny-rust.
The majority of individuals are tawny-rust. A genetic response to the
local character and color of the sand in Cufra Oasis may account for
this wide range of dorsal coloration.
One specimen, 319683, from El Giof, Cufra Oasis, differs markedly
from all other specimens of G. g. aeruginosus in having a more promi-
nently vaulted skull (especially the parietals), smaller and less inflated
auditory bullae, and heavier zygomata. In all other morphological
characters this specimen clearly represents Gerbillus gerbillus. In the
marked inflation of the braincase, however, it resembles Gerbillus
eatoni whose range includes the Egyptian and Libyan coasts. This
similarity to G. eatoni is entirely fortuitous, and this specimen is
probably aberrant or represents an extreme in the range of variation
for the subspecies.
Ecological observations. These gerbils prefer the sandy areas
along the periphery of the oasis but occasionally were taken from
areas of open sand within the palm groves. At El Giof, a large series
was obtained from a sandy clearing in the interior of the oasis. Usually
however, Gerbillus pyramidum and Gerbillus campestris are more
abundant in the oasis proper.
At Bzema, El Giof, El Hauuari, and most of the other large oases
of southern Cyrenaica, large saline lakes occupy the interior of the
oasis and form hard precipitates of salt along their margins. These
barren areas are surrounded by a ring of sedges and halophytic plants,
and G. gerbillus occurs here only sparingly; however, Gerbillus cam-
pestris occurs abundantly in these mesic habitats.
The name aeruginosus, from the Latin meaning rust or rusty,
alludes to the rusty dorsal color of most specimens.
285-134 O —68 8
Gerbillus gerbillus discolor, new subspecies
Holotype. Adult male, skin and skull, USNM 322461, from Ghat,
Fezzan Province, Libya; obtained Dec. 18, 1961, by G. L. Ranck,
original no. 1116.
Specimens examined. One hundred sixty-two, from Tripolitania:
2 km SW Hun, 18; 5 km S Socna, 6; Bir Fergian, 10 km S Socna, 2;
from Fezzan: Edri, 11; Temenhint Oasis, 30 km NE Sebha, 12;
3 km NW Sebha, 5; 5 km NW
Sebha, 3 (1 skin only); 10 km SE
Sebha, 4 (skin only); El Abiad, 60 km SW
Sebha, 18 (2 skin only;
2 skull only); 75 km W
Ubari, 11; Meseguin 1; Umm
el Araneb, 2;
6 km NMurzuch, 1; 55 km SSW Serdeles, 8; 20 km Ghat, 1; N

12 km N Ghat, 13 (4 skull only) Ghat, 25 (1 skeleton) El Gatrun, 17;
; ;
El Barcat, 4.
6 adult females from the type locality, with the measurements of
the type in brackets, are, respectively: Total length 211 (200-220),
210.7 (200-218), [213]; length of tail 121.6 (112-130), 124.3 (116-128),
ear 13.4 (13-14), 13.2 (12-14), [13]; occipitonasal length of skull
28.3 (27-28.9), 27.9 (27.1-29), [28.4]; length of auditory bulla 10.5
(10.l-10.7j, 10.3 (10-10.7), [10.6]; crown length of molariform tooth-
row 3.6 (3.5-3.8), 3.6 (3.5-3.6), [3.6] greatest breadth across
;
zygomatic
arches 15.4 (15.2-15.6), 14.6 (14.5-14.8), [15.6]; least interorbital
breadth 5.7 (5.2-6), 5.7 (5.5-5.8), [5.5]; breadth of rostrum at level of
antorbital foramina 3.1 (2.9-3.3), 3 (2.9-3), [3]; greatest length of
nasals 10.6 (10.2-11.1), 10.3 (9.8-10.6), [10.5].
Diagnosis. Upperparts Clay Color, suffused throughout with
brownish hairs imparting a streaked or marbled appearance; sub-
auricular patch prominent and Cinnamon-Buff; pinna of ear sparsely
haired and Clay Color; preauricular patch Pinkish Buff and extending
as a row of fine hairs onto anterior margin of pinna; postauricular
patch, circumorbital, rostral and mystacial areas, dorsal surfaces
of fore and hind feet and entire underparts white (in some specimens,
the venter is Pale Pinkish Buff owing to artificial staining) fore and ;
hind feet with five digits, each bearing a claw; tail distinctly bicolored
Cinnamon-Buff dosally, Light Buff ventraUy and terminating in a
prominent pencil, Avellaneous dorsally and Light Buff ventrally;
vibrissae short and composed of both brownish and whitish hairs.
Skull: Medium in size; braincase moderately arched; auditory
bullae relatively large and inflated: anterior palatine foramina and
posterior palatine canals narrow; zygomata moderately heavy and
convergent anteriorly; posterior lacerated foramina between basioc-
cipital and anteromedial margins of auditory bullae reduced to
slitlike aperatures.
Comparisons. From topotypes of Gerbillus gerbillus gerbillus, the

type and paratypes of G. g. discolor differ in having narrower anterior
palatine foramina and posterior palatine canals, slightly more
inflated auditory bullae, shallower pterygoid fossae, shorter skulls,
shorter nasals, greater zygomatic breadth, and more robust skulls.
Gerbils from the Fezzan are smaller in overall length and length
of tail and hind foot; in color, they are noticeably darker and appear
more streaked owing to a greater admixture of brownish hairs on
dorsum.
Two specimens, 321821 and 321822, from 40 kilometers north
of Sinauen and a single specimen, 321825, from 5 kilometers east
of Derg, Tripolitania Province, are referable to Gerbillus gerbillus
latastei and differ from those of G. g. discolor in having wider anterior
palatine foramina, more gracile zygomata, more vaulted braincases,
and larger external and cranial measurements, being of comparable
size only in least interorbital breadth and length of nasals. In color,
these specimens from Tripolitania are more uniform in dorsal color,
with less suffusion of brownish hairs.
When the type series of G. g. discolor and Gerbillus gerbillus aeru-
ginosa are compared, having slightly paler,
G. g. discolor differs in
more variegated dorsal wider and more inflated auditory bullae,
color,
narrower posterior palatine canals, larger cheek teeth, proportion-
ately smaller and deeper pterygoid fossae, wider basioccipitals, and
markedly larger size of all external and cranial measurements, except
length of hind foot and breadth of rostrum at the level of the antorbital
foramina.
For comparison with Gerbillus gerbillus psammophilous, see account
of that subspecies.
Remarks. Gerbils from Edri, near the northern limits of the range
of G. g. discolor, are similar cranially to topotypes of G. g. discolor
from Ghat, Fezzan. They are intermediate in color, however, between
that of a single representative, 321825, of Gerbillus gerbillus latastei
from Derg, Tripolitania, and topotypes of G. g. discolor. The color of
these specimens from Edri suggests a relationship to G. g. latastei
whose range is to the north. Unfortunately, specimens are not avail-
able from the Hamada el Hamra and the Hamada de Tinrhert which
are located north of Edri and between the ranges of G. g. latastei
and G. g. discolor. Although sandy habitats are not widespread in
these hamadas, localized sandy areas are present and doubtless
support small populations of these rodents. These hamadas, there-
fore, provide an avenue through which gene exchange takes place
between populations of G. g. latastei to the north and those of G. g.
discolor to the south. Gerbils inhabiting these regions may be inter-
grades between these two subspecies.
Representatives of this subspecies from El Gatrun on the eastern

margin of the Idehan Murzuch are paler in dorsal color than those
from elsewhere in the Fezzan. When specimens of Gerbillus gerbillus
become available from the Serir Tibesti and other regions to the
south, they may represent an undescribed subspecies to which the
specimens from El Gatrun will be referable.
Gerbils from El Abiad, 55 kilometers west of Ubari, 75 kilometers
south -southwest of Serdeles, and 12 kilometers north of Ghat are
remarkably uniform in color and cranial characters. Apparently the
sandy beds of the Wadi el Agial, Wadi Irauen, and the Wadi Tenezoft
act as dispersal corridors which insure genetic uniformity among these
widely scattered populations.
Members of this subspecies from the vicinity of Hun, Socna, and
Bir Fergian, Tripolitania Province, show many characters typical of
Gerbillus gerbillus psammophilous, whose range includes most of
Cyrenaica and the coastal areas of the Gulf of Sirte. In the degree of
inflation of the auditory bullae, breadth of the anterior palatine fora-
mina, and the size and shape of the posterior lacerated foramina between
the basioccipital and auditory bullae, they resemble G. g. psammophil-
ous, but in the majority of cranial characters, they are closer to G. g.
discolor to which they are here referred. For additional comments
regarding gene exchange with G. g. psammophilous, see under
"remarks" in account of that subspecies.
Although this subspecies is characterized by generally darker, more
variegated dorsal color, individuals in the population range from uni-
form colors of pale, subdued tones to rather brilliant, ochraceous hues.
Partial albinism is also of occasional occurrence. In gerbils from Sebha
and Temenhint Oases of the central Fezzan, this broad range in dorsal
color is particularly striking. These oases apparently represent areas
in which the genetic character of the population is less rigidly fixed.
These small gerbils are probably the most widely distributed rodent
in the Fezzan. They are most abundant near the periphery of the
oases where the palm groves are less dense and open sandy areas are
more widespread. In the Fezzan, G. g. discolor is sympatric with the
larger, and usually more abundant, Gerbillus pyramidum tarabuli.
Other rodents, such as Gerbillus amoenus, Jaculus jaculus, Meriones
caudatus, and Acomys cahirinus occur with G. gerbillus but usually
are far less abundant.
The range of this subspecies, as far as is known, does not include
coastal Libya. In the coastal areas of northern Tripolitania, members
of the Gerbillus pyramidum group are the dominant gerbils.
The term discolor, from the Latin meaning "of different color or
colors," is used for the subspecies name in reference to the variegated
character of the dorsal pelage.
Gerbillus gerbillus gerbillus (Olivier)
Dipus gerbillus Olivier, Bull. Sci. Soc. Philom., Paris, vol. 2, p. 121, 1801 (Giza
Province, Egypt).
Specimens examined. Twenty-six, from Cyrenaica: Giarabub, 10

(1 skin only); Bahr el Tubat, 21 km ESE Giarabub, 11; 24 km SSE
Giarabub, 5.
Measurements. Averages and extremes of five males and measure-

ments of two females from Giarabub are, respectively: Total length
214 (205-221), 214, 205; length of tail 127 (121-135), 122, 119; length
of hind foot 29 (28-30), 31, 32; length of ear 13 (13-13), 12, 13; occip-
itonasal length of skull 28.7 (28.1-29.4), 28.1, 27; length of auditory
bulla 10.7 (10.2-11.2), 9.9, 10.2; crown length
molariform toothrow
of
3.7 (3.6-3.8), 3.4, 3.6; greatest breadth across zygomatic arches 15.3
(14.5-15.8), 14.7, 14.9; least interorbital breadth 5.9 (5.7-6), 5.6, 5.5;
breadth of rostrum at level of antorbital foramina 3 (3-3.1), 3, 2.8;
greatest length of nasals 10.6 (9.9-11.6), 10.6, 10.4.
Diagnosis. Upperparts Cinnamon-Buff becoming paler on sides
and flanks owing to suffusion of whitish hairs; circumorbital regions,
mystacial and pectoral areas, dorsal surfaces of fore and hind feet, and
entire underparts white; postauricular patch distinct and white;
vibrissae short and composed of brownish and white hairs; pinna of
ear almost naked, Warm Buff, and with row of fine, buffy hairs on
anterior margin; fore and hind feet sparsely haired ventrally and
bearing five digits, each with a claw; tail indistinctly bicolored Light
Ochraceous-Buff above and Light Buff below, and with a prominent
terminal pencil, Tilleul-Buff dorsally and Pale Pinkish Buff ventrally.
Skull: Medium in size and moderately robust; zygomata heavy;
lachrymals large; molariform teeth moderately large; auditory bullae
markedly large and inflated; braincase moderately domed.
Comparisons. From topotypical Gerbillus gerbillus gerbillus from
Alexandria Road, 5 miles northwest of the Pyramids of Giza, Cairo
Area, Egypt, specimens from Giarabub and vicinity differ in having
slightly heavier zygomata, larger lachrymals, larger and more in-
flated auditory bullae, slightly larger molariform teeth, slightly
narrower basioccipitals and shorter overall size of body. In color, the
majority of specimens are closer to Gerbillus gerbillus psammophilous,
whose range is farther west and south. In the majority of cranial
characters and in external measurements, however, they resemble
the nominate subspecies to which they are here referred.
For comparisons with Gerbillus gerbillus aeruginosas and G. g.
psammophilous, see accounts of those subspecies.
Remarks. In cranial characters, these specimens from Giarabub
are indistinguishable from those from Bahr el Tubat and Siwa Oasis,
western Egypt. These populations also do not differ significantly
from those from the type Apparently, the low-lying areas

locality.
associated with the Wadi Natroun, Qattara
Depression, Siwa Oasis,
and Bahr el Tubat provide a continuum of suitable habitat linking
these various populations together.
In some characters, notably heavy zygomata, large and ventrally
inflated auditory bullae, narrow basioccipital and small body size,
these gerbils from Giarabub show evidences of gene exchange with
G. g. psammophilous to the west. Apparently they are able to overcome
the barrier imposed by the Sand Sea of Calanscio in this region of
Libya.
Some from Giarabub are noticeably darker, more orangish
gerbils
in dorsal color,than others. This same feature was observed in jerboas
(Jaculus jaculus) from Giarabub and is presumably an adaptive
response to the different color of the sand in this area.
Ecological observations. Giarabub Oasis is a portion of the
great depression formed from the east-west linkage of the Wadi
Natroun, Qattara Depression, and Siwa Oasis. Although located in
the interior of the desert, all three "sebchets" are below sea level.
Sandy areas of large extent have accumulated here and provide
desirable habitat for these gerbils. Plant cover is more abundant in
these low-lying areas than elsewhere; dense stands of Phragmites are
widespread in the bottoms of the depressions; and Tamarix occupies
the more elevated areas where the soil is firmer and more stable.
Various other species of woody shrubs are also present. Many resemble
the chenopods, woody composites, and thorny perennials of the
western United States. The series from Giarabub was obtained from
some large dunes near the oasis.
A concentration of sandy-clay hummocks within a localized sandy
elevation of the valley floor formed the habitat at the collecting site
24 kilometers south-southeast of Giarabub. The vegetative cover here
was particularly dense and varied.
Bahr el Tubat is a large, quite shallow, saline lake surrounded, in
some places, by dense stands of reeds and sedges. In other areas of the
shoreline, salty deposits of great thickness and extent occur, rendering
the habitat unsuitable for mammals. The specimens from Bahr el
Tubat were taken from a series of vegetated dunes beyond the lake and
slightly higher in elevation.
Gerbillus gerbillus latastei Thomas and Trouessart
Gerbillus gerbillus latastei Thomas and Trouessart, Bull. Soc. Zool. France, ser.
5, vol. 28, pp. 171-174, July 28, 1903 (Kebili, Southern Tunisia).
Specimens examined. Three, from Tripolitania : 40 km N

Sinauen, 2; 5 km E Derg, 1.
Measurements. The measurements of an adult male, 321822,

and female, 321821, from 40 kilometers, north of Sinauen, and of an
adult male, 321825, from 5 kilometers east of Derg are, respectively:
Total length 219, 215, 217; length of tail 128, 130, 130; length of hind
foot 30, 30, 29; length of ear 14, 14, 13; occipitonasal length of skull
crown length
29.3, 28, 28.7; length of tmditory bulla 10.8, 10.5, 10.5;
of upper molariform toothrow 3.8, 3.7, 3.8; greatest breadth across
zygomatic arches ?, 14.6, ?; least interorbital breadth 5.7, 5.6, 6.1;
breadth of rostrum at level of antorbital foramina 3.3, 2.9, 3.1;
greatest length of nasals 10.6, 10.6, 10.6.
Diagnosis. Upperparts lustrous, Clay Color with uniform admixture
of brownish hairs extending over sides of body nearly to the venter
where it has greater suffusion of whitish hairs; subauricular patches
distinct, Cinnamon-Buff and also with admixture of brownish hairs;
mystacial, rostral, preauricular, and circumorbital areas, postauricular
patch, dorsal surfaces of fore and hind feet, and entire underparts
white; pinna of ear almost naked, Ochraceous-Buff, and with row of
buffy hairs on anterior margin; vibrissae short and white with oc-
casional darker hairs; fore and hind feet sparsely haired, with five
digits, each bearing a claw; tail distinctly bicolored Cinnamon-Buff
dorsally, white ventrally, with prominent terminal pencil, Avellaneous
dorsally and white ventrally. Skull: Relatively large, molariform
teeth long; interorbital breadth moderately wide; nasals short,
auditory bullae and posterior palatine canals large zygomata markedly
;
fragile and convergent anteriorly; supraorbital bead prominent;

braincase moderately vaulted.
Comparisons. The two specimens, 321821 and 321822, from near
Sinauen and the single specimen, 321825, from Derg differ from
topotypes of the nominate subspecies in darker color, more streaked
and lustrous dorsal pelage, slightly smaller overall length, slightly
longer ears, slightly longer skulls, longer molariform teeth, narrower
interorbital breadth, slightly shorter nasals, larger auditory bullae,
and longer posterior palatine canals.
Compared topotype (MNHN 57) of Gerbillus gerbillus
to a single
foleyi Heim de Balsac from Beni Abbes, western Algeria, the speci-
mens from Libya are similar in dorsal color, but differ in having
smaller hind feet and auditory bullae, narrower zygomatic breadths,
and slightly shorter nasals.
For comparisons with Gerbillus gerbillus discolor and Gerbillus
gerbillus psammophilous see accounts of those subspecies.
,
Remarks. The specimens from near Sinauen are slightly darker

in dorsal color than the specimen from Derg and also have larger
auditory bullae, shorter posterior palatine canals, and more vaulted
braincases. These differences are too slight to warrant assignment

of these specimens to separate subspecies, and they are here referred
to the same subspecies.
Ellerman and Morrison-Scott (1951) considered G. g. latastei to be a
little-known and dubious form. In the present study comparative
specimens of G. g. latastei are not available, but Thomas and Trouessart
(1903), in the original description, included the following measure-
ments: Length of head and body, 97; length of tail, 110; length of
hind foot, 27.5; length of ear, 12; length of skull from the tip of the
nasals to the extremity of the interparietal, 31; length of nasals, 12.5;
interorbital breadth, 6.3; length of diastema, 8.5; length of theupper
molariform toothrow, 4.4. In the original description latastei was also
characterized as having patches above the eyes and behind the ears,
all underparts pure white, hairs grayish at the base, except the post-
auricular patches which are entirely white, and the dorsal color of the
tail the same color as the back and white below with a small brownish
pencil. According to Thomas and Trouessart, it can be distinguished

from all other gerbils by its brilliant color. The three specimens
from Tripolitania agree closely with the above color description but
are somewhat smaller in most cranial characters and larger in most
external measurements. Owing to inadequate comparative materials,
the assignment of these specimens to G. g. latastei is provisional and
based largely on geographic grounds.
The specimens from near Sinauen were taken from a broad slope in
the hamada where vegetation was sparse and interspersed with small,
transient dunes. The habitat at Derg was similar to that at the above
locality,but the plant cover was more abundant and confined to mar-
gins of a small wadi, and the sand was more sporadic and localized.
This subspecies probably has a much larger distribution in Libya
than the few specimens would indicate, and the range probably
includes a large portion of the Hamada el Hamra, the Hamada de
Tinrhert, and the inner margins of the Tripoli tanian coastal plain.
Gerbillus gerbillus psammophilous, new subspecies
Holotype. Adult male, skin and skull, USNM 325078, from

Gialo Oasis, Cyrenaica Province, Libya; obtained Mar. 16, 1962,
Specimens examined. One hundred eighty, from Cyrenaica: 10
km S Agedabia, 27 (2 skin only) 65 km ; WNW
El Agheila, 1 5 km ;
W El Agheila, 2; Gasr es Sahabi, 12 (6 skull only); Augila, 23; Gialo

Oasis, 58; 150 km S Gialo, 1 Wadi er Rueis, 340 km
; WNW
Tazerbo,
3; El Gezira, Tazerbo Oasis, 16; Tazerbo Oasis, 23; Bir bu Zarregh, 3;
Bir el Harasc, 9; from Tripolitania: 15 km WNW
Marble Arch, 2.

type in brackets, are, respectively: Total length 214.4 (202-220),
208.1 (200-220), [210]; length of tail 122.5 (117-129), 120.3 (113-128),
ear 13.9 (13-14), 13.3 (13-14), [14]; occipitonasal length of skull
28.6 (27-29.9), 28.1 (27.3-29.5), [28.5]; length of auditory bulla 10.4
(9.8-11), 10.3 (9.7-10.9), [10.5]; crown length of upper molariform
zygomatic arches 15.5 (15-16.4), 15.1 (14.7-15.4), [15.7]; least inter-
orbital breadth 5.9 (5.4-6.4), 5.7 (5.3-6.2), [5.7]; breadth of rostrum
at level of antorbital foramina 3 (2.7-3.2), 3.1 (2.9-3.2), [3]; greatest
length of nasals 10.6 (10.4-11.6), 10.5 (10-11.3), [10.6].
Diagnosis. Upperparts uniformly Cinnamon-Buff becoming inter-
spersed with white hairs on sides and pectoral areas; circumorbital
region, rostral area, postauricular areas, and entire underparts white;
fore and hind feet Light Buff dorsally, sparsely haired ventrally, and
each bearing five digits with claws; pinna of ear Warm Buff, finely
haired and with thin row of buffy hairs along anterior margin vibrissae ;
short and usually white, but with occasional darker hairs present; tail
indistinctly bicolored Light Buff ventrally with narrow dorsal line of
Light Ochraceous-Buff terminating in a distinct pencil, Avellaneous
dorsally and Light Buff ventrally. Skull: Medium in size; moderately
robust; auditory bullae moderately inflated; zygomata heavy; basi-
occipital constricted anteriorly and forming distinct foramina near
anteromedial margins of auditory bullae; external pterygoid fossae
forming distinct depressions dorsal to posteriorly flaring pterygoid
hamulae; braincase moderately vaulted.
Comparisons. From topotypes of Gerbillus gerbillus discolor from
Ghat, Fezzan Province, Gerbillus gerbillus psammophilous differs in
having narrower auditory bullae, more anteriorly constricted basi-
occipitals, larger foramina between basioccipital and auditory bulla,
more arched braincase, less divergent pterygoid hamulae, greater
occipitonasal length, greater interorbital breadth, and slightly larger
hind In color, G. g. psammophilous is much paler and less streaked
feet.
dorsally and has a less prominent subauricular patch and a lighter
pencil.
Gerbillus gerbillus psammophilous from topotypical specimens
differs
of Gerbillus gerbillus having heavier zygomata, more
gerbillus in
anteriorly constricted basioccipitals, more distinct foramina between
the basioccipitals and the auditory bullae, smaller, more slitlike
posterior palatine canals, shorter overall length, slightly larger
occipitonasal length, greater breadth across zygomatic arches, longer
nasals,and rostra less decurved distally. These gerbils are similar to

those of the nominate subspecies in dorsal color but appear slightly-
paler owing to a greater suffusion of whitish hairs on sides, more
white around the eyes and rostrum, less prominent subauricular
patches, and a paler pencil.
Gerbils representing the nominate subspecies from the vicinity of
Giarabub, Cyrenaica Province, resemble those referable to G. g.
psammophilous but have larger, more massive skulls, larger, more
inflated auditory bullae, and larger anterior palatine foramina. In
color, they are almost indistinguishable from topotypes of G. g.
psammophilous .
Gerbillus gerbillus psammophilous can be readily distinguished from

topotypical specimens of Gerbillus gerbillus andersoni by much paler,
more orangish dorsal color, shorter anterior palatine foramina, less
vaulted braincase, less ventrally inflated auditory bullae, shorter and
smaller molariform toothrow, more prominent supraorbital bead, less
robust zygomata, and smaller size of all cranial measurements.
From topotypes of Gerbillus gerbillus asyutensis from the Wadi
Asyuti, Egypt, G. g. psammophilous differs in having markedly darker,
less variegated dorsal pelage, shorter posterior palatine canals, wider
basioccipitals, and larger size of all cranial measurements.
Compared with two specimens, 321821 and 321822, representing
Gerbillus gerbillus from 40 kilometers north of Sinauen,
latastei
Tripolitania Province, G. psammophilous has slightly smaller and
g.
more ventrally inflated auditory bullae; smaller molariform teeth;

heavier zygomata shorter occipitonasal length shorter tail markedly
; ; ;
paler, more subdued dorsal color with less suffusion of brownish

hairs; a less distinctly bicolored tail; and vibrissae with fewer dark
hairs.
Paratypes of G. g. psammophilous can be distinguished from the type
and paratypes of Gerbillus gerbillus aeruginosus from El Giof, Cufra
Oasis, by their paler dorsal color and markedly larger size of all
cranial measurements, being similar in length of auditory bulla, crown

length of upper molariform toothrow, and least interorbital breadth.
G. g. psammophilous also has narrower posterior palatine canals,
heavier zygomata, more ventrally inflated auditory bullae, less domed
skull, and less distally decurved rostrum.
Remarks. Members of this subspecies show significant intergrada-
tion with other subspecies whose major ranges are far removed
geographically. Gerbils from Giarabub and Bahr el Tubat, in some
characters, suggest intergradation between G. g. psammophilous and

the nominate subspecies. In color, gerbils from Tazerbo Oasis, Bir el
Harasc, and Bir bu Zarregh show evidences of gene exchange with

G. g. aeruginosus to the south. Specimens from near Hun, Socna, and
Bir Fergian, Tripolitania Province, are intermediate in color and

cranial charactersbetween gerbils from the Fezzan and those from
Cyrenaica.
The Sand Sea and the Sand Sea of Rebianna
of Calanscio to the east
to the south apparently do not act as absolute barriers to the move-
ments of these gerbils. Likewise, the volcanic ramparts of the Gebel el
Harug el Asued of eastern Tripolitania and western Cyrenaica do not
entirely limit the east-west dispersal of this species in Libya. Three
specimens from the Wadi er Rueis, 340 kilometers west-northwest of
Tazerbo are indistinguishable from those from the type locality in
color and cranial characters. A pathway for gene exchange between
populations of gerbils from the Fezzan and Cyrenaica is provided by
the numerous small oases that circumscribe the Gebel el Harug el
Asued. When specimens become available from these areas, they
probably will show characters allying them to populations of gerbils
in both the Fezzan and Cyrenaica.
Cranially, gerbils from the coastal areas near Agedabia, Marble
Arch, and El Agheila are indistinguishable from those from the type
locality in the Saharan interior. In color, however, these animals
from the coast are much paler dorsally with greater suffusion of
whitish hairs. This trend toward darker dorsal color increases pro-
gressively toward the south and reaches its extreme in gerbils from
Tazerbo and Cufra Oases. A single specimen, 325105, from 150 kilo-
meters south of Gialo is intermediate in dorsal color between gerbils
from Tazerbo and Gialo Oases. Similarly, specimens from Gasr es
Sahabi, which is located midway between Gialo Oasis and the coast
are lighter than those from Gialo but darker than those from the
coastal deserts. Thus a north-south, clinal gradient in dorsal color
is demonstrable within populations of gerbils from Cyrenaica. This
gradient in dorsal color reaches its greatest intensity in gerbils in-
habiting the southern oases.
Ecological observations. These gerbils range throughout the
sandy environs of the oases, where they are frequently associated
with debris near the bases of palm trees. They also occur in sandy
margins of wadis and frequently inhabit the margins of the sand
seas or "Ramleh" where unstable, mountainous dunes are wide-
spread. The larger sand seas, however, such as Rebianna and Calanscio,
contain vast areas of constantly shifting dunes which probably are
not inhabited by these mice. Apparently sand seas of small extent
are not barriers to dispersal. The widely scattered distribution of
this subspecies and the collecting sites at 150 kilometers south of
Gialo and from the Wadi er Rueis confirm this.
In the coastal areas of Cyrenaica this species supplanted by
is
Gerbillus eatoni, which prefers the more densely vegetated margins

of the coastal plain.
The subspecies name psammophilous, from the Greek meaning

"sand lover," suggests a predilection of members of this subspecies
for habitats of a sandy character.
Gerbillus pyramidum E. Geoffroy
Gerbillus pyramidum E. Geoffroy, Catalogue des mammiferes du Museum Na-

tional d'Histoire Naturelle, p. 202, 1803 (near Pyramids of Giza, Giza
Province, Egypt).
General distribution of species. Israel, Sinai, Egypt, Libya,

Algeria, Tunisia,and Morocco; southward throughout the Sahara to
central Sudan, Chad (Tibesti Mountain Area) Niger, and Mauritania.
,
Distribution in Libya. Desert and coastal regions of Tripolitania

and throughout the Fezzan. (Only a single specimen is known from
Cyrenaica.)
Gerbillus pyramidum hamadensis. Tripolitania: Coastal areas and
interior deserts of Tripolitania north of the Gebel es Soda and Hamada
de Tinrhert.
Gerbillus pyramidum tarabuli. Fezzan.
...P— ^ L
^J A
^^
Published records in Libya. Cyrenaica: Vicinity of Benghazi

(Ghigi, 1920). The occurrence north in Cy-
of this species this far
renaica is questionable. While I have not examined these specimens,
it seems likely that they would be representatives of Gerbillus eatoni;
Tripolitania: Wadi Agarib, Wadi Aggar, Ain Hammam, Ferdjan,

El Koshby, Linzerat, Wadi Sultan, Tamari-Ferdjan (Thomas, 1902);
Gargaresch, Tripoli (Andreucci, 1914); Ghadames (Perugini, 1929);
Giofra, Tripolitania settentrionale (Zavattari, 1937); Fezzan: God-
dua, Sebha, Umm el Abid, Zeigen (Thomas, 1902) ; Murzuch
(Andreucci, 1914); Sciati, Hofra (Zavattari, 1937).
Comparisons. From Gerbillus aureus, which it somewhat resembles,
this species can be distinguished by its larger skull and body, longer
and more tufted tail, generally lighter color, longer and narrower
posterior palatine canals, and relatively narrower anterior palatine
foramina.
This species can be distinguished from Gerbillus eatoni by its
markedly larger overall size (occipitonasal length usually more than
30 mm), relatively smaller and less inflated auditory bullae, larger
molariform teeth, and less domed, almost flattened braincase.
This species is markedly larger in all respects than Gerbillus gerbillus
and has relatively larger molariform teeth, less inflated auditory
bullae, and less orange color in the dorsal pelage.
Gerbillus pyramidum closely resembles Gerbillus perpallidus Setzer
from Egypt but can be distinguished by its darker color, markedly
more open pterygoid fossae,

larger skull, less inflated auditory bullae,
longer anterior palatine foramina, and much more conspicuous supra-
orbital beads. The foregoing characters were used by Setzer (1958
p. 221) to separate these two species and I concur.
Remarks. These gerbils are widely distributed throughout North
Africa from the Red Sea to the Atlas Mountains and occur virtually
throughout the Sahara; however, there is a conspicuous hiatus in
their distribution in western Egypt and eastern Libya. In Libya,
the oases of Gialo, Giarabub, Tazerbo, and Cufra, and in Egypt, Siwa
Oasis and the low-lying areas of the Qattara Depression contain
abundant suitable habitat for them, but as yet, no specimens are
known from these areas. Apparently, members of this species are
unable to circumvent the barriers imposed by the Cyrenaican Plateau
and the northern part of the Libyan Desert and have been unable to
extend their range into these parts of Libya and Egypt. In Libya, the
Serir of Calanscio and the Sand Sea of Calanscio doubtless serve to
further isolate the oases in southern Cyrenaica.
This species occurs abundantly throughout western Libya in the
provinces of Tripolitania and Fezzan. Presently, these populations
in Libya are linked to those of the Nile Valley in Egypt by a series
of rather discontinuous populations throughout the northern Chad

and west-central Sudan. In the future, when more localities become
represented by specimens, it is probable that a more clearly defined
dispersal route, consisting of several overlapping populations, will
be found to unite the Libyan populations with those of the type
locality in the Nile Valley.
Prior to the present study, Ellerman and Morrison-Scott (1951)
included Algeria within the range of G. p. pyramidum, based, appar-
ently, on specimens from El Golea and In Salah. It is now known
that the range of the nominate subspecies is confined to Egypt (in-
cluding Sinai) and Sudan as far south as Khartoum. The range of
G. p. tarabuli thus is interposed between the areas of occurrence of
these Algerian populations and the type locality. Such a discontinuous
distribution for members of the same subspecies is untenable, and I
suspect that these gerbils from central Algeria represent subspecies
distinct from those in Libya and Egypt.
Gerbillus pyramidum hamadensis, new subspecies

321827, from 5 km
E Derg, Tripolitania Province, Libya, obtained Nov. 14, 1961, by
G. L. Ranck, original no. 886.
Specimens examined. Fifty-five, from Cyrenaica: 5 km W El
Agheila, 1; from Tripolitania: 5 km N Mizda, 3; 12 km W Sirte, 2;
15 km WNW Marble Arch, 3; 30 km S Bu Ngem, 2; 5 km E Derg,
13; 5 km S Socna, 15; Bir Fergian, 10 km S Socna, 17 (16 skin only).
Measurements. Averages and extremes of six adult males and five
type in brackets, are, respectively: Total length 240.2 (235-244), 228
(223-234), [244]; length of tail 141 (137-145), 131.4 (128-135), [145];
length of hind foot 30.2 (29-32), 31 (29-32), [29]; length of ear 14.8
(14-16), 15.2 (14-16), [16]; occipitonasal length of skull 31.6 (31-32),
30.5 (29.7-31.4), [32]; length of auditory bulla 11.7 (11.4-11.9), 11.5
(11.2-11.7), [11.6]; crown length of upper molariform toothrow 4.3
(4.1-4.4), 4.2 (4-4.4), [4.2]; greatest breadth across zygomatic arches
16.7 (15.7-16.9), 16.2 (15.8-16.8), [16.6]; least interorbital breadth
6.5 (6-6.8), 6.3 (6-6.6), [6.8] breadth of rostrum at level of antorbital
:
foramina 3.6 (3.5-3.8), 3.6 (3.5-3.8), [3.6], greatest length of nasals

12.3 (11.8-12.6), 11.7 (11.3-12), [12.6].
Diagnosis. Upperparts Cinnamon-Buff grading to Clay Color and
Tawny-Olive on rump and becoming paler on sides; entire dorsum
lightly washed with brownish hairs which project appreciably beyond
the paler colored underlying hairs; subauricular areas Cinnamon-Buff
with slight suffusion of grayish-brown hairs; supraorbital, rostral,
mystacial, circumoral, and pectoral areas pure white; postauricular
patches distinct and pure white; pinna of ear same color as dorsum,
sparsely haired and with row of buffy hairs along anterior margin;
vibrissae composed of nearly equal numbers of brown and white hairs;
dorsal and ventral surfaces of fore and hind feet white, densely haired
and each bearing five digits with claws; tail indistinctly bicolored,
Pinkish Buff dorsally and Pale Pinkish Buff ventrally, and terminating
in a conspicuous brownish-colored pencil; entire underparts white.
Skull: Relatively small; rostrum wide; molariform teeth relatively
large; anterior palatine foramina markedly narrow and slitlike;
auditory bullae rather bulbous.
Comparisons. From Gerbillus pyramidum pyramidum as known from
several localities in Giza Province, Egypt, Gerbillus pyramidum
hamadensis differs in markedly smaller size of body and skull, relatively
smaller and more slitlike anterior palatine foramina, less ventrally
inflated auditory bullae, less robustzygomata, and smaller lachrymals.
This subspecies is much paler and uniform in dorsal color and has
more prominent postauricular patches and lighter colored pencil.
Compared with topotypes of Gerbillus pyramidum tarabuli, G. p.
hamadensis is noticeably smaller in all cranial and external characters,
being of comparable size only in crown length of molariform toothrows
and breadth of rostrum. In color, members of this new subspecies are
slightly paler, have a grayer cast owing to a greater suffusion of
grayish hairs on dorsum, and have a pencil of a slightly lighter color.
These gerbils can be readily distinguished from representatives of
Gerbillus pyramidum hirtipes Lataste from Ain Sefra, southwestern
Algeria, by paler dorsal color, longer and more thickly tufted tails,
larger auditory bullae and molariform toothrows, and generally larger
size of all other measurable cranial characters.
In general body size and length of hind foot and ear, G. p. hamadensis
resembles Gerbillus aureus nalutensis but differs markedly in having
smaller anterior palatine foramina, longer posterior palatine canals,
more robust zygomata, larger molariform teeth, longer and more
tufted tail, paler and less varied dorsal color with less suffusion of
black on rump, and larger size of all cranial measurements.
Remarks. Members of this subspecies can be distinguished from all
others in Libya by their conspicuously smaller size and paler dorsal
color. Throughout the range of this subspecies, animals vary locally.
Gerbils from 5 kilometers south of Socna, Tripolitania Province, are
more strongly suffused with brown on the back and dorsum of tail,
have slightly darker pencils, slightly longer and wider anterior palatine
foramina, more gracile skulls, less prominent supraorbital ridges, and
are slightly smaller in all cranial measurements. Two specimens,
322097 and 322098, from 30 kilometers south of Bu Ngem, Tripolitania

Province, have slightly smaller anterior palatine foramina, but, in
from gerbils from Socna and Bir Fergian.

color, are indistinguishable
Members from various localities in the coastal plain
of this subspecies
tend to have slightly larger, more bulbous auditory bullae. Those
from 12 kilometers west of Sirte are also darker in dorsal color. This
variation is well within the expected limits of a subspecies whose
range includes such a large geographic area.
The populations from Socna and Bir Fergian show only slight
intergradation with G. p. tarabuli whose range is to the south. Gerbils
from the type locality at Derg show no evidences of interbreeding
with populations to the south. The high escarpments of the Hamada
de Tinrhert in the west and the mountain ramparts of the Gebel es
Soda and the Gebel el Harug el Asued in the east obviously have
proven to be serious deterrents to gene exchange between populations
of these gerbils, and thus the range of this subspecies is limited to
the hamadas, coastal deserts, and coastal plain lying north of these
barriers. In northwestern Tripolitania the range of this subspecies ap-
parently does not attain the Mediterranean Coast. Three specimens
from 5 kilometers north of Mizda constitute the northernmost record of
occurrence of G. pyramidum in Libya. Farther north, Gerbillus aureus
is the dominant gerbil and probably competes with and thus prevents
G. pyramidum from occupying the coastal areas. Farther eastward,

however, along the Gulf of Sirte, G. aureus is known only from Sirte,
while G. p. hamadensis is known from several localities.
Ecological observations. The type series was obtained from the
rather denuded margins of a small wadi in the western portion of
the Hamada el Hamra. This dry watercourse afforded scanty vege-
tative cover, while rocky outcroppings interspersed with bare areas
of sand and gravel were widespread. In the immediate vicinity of the
wadi, Tamarix, Calligonum, and other types of shrubs were the dom-
inant vegetation, but farther out on the hamada, smaller plants
growing close together were commoner. The soil in this area consisted
mostly of sand.
Setzer (1957) describes the habitat at Socna and Bir Fergian as
consisting of a sandy-clay substrate similar to hardpan, which re-
sembles the desert playas of the Western United States. The collecting
sites in the coastal plain near Sirte, Marble Arch, and El Agheila
were characterized by dense plant cover and the soil in these areas
also was sandy-clay. Large, permanent dunes supporting Calligonum
characterized the habitat near Bu Ngem. Judging from the sandy
character of these habitats, this subspecies always requires a sandy
substrate.
The name hamadensis refers to the hamada-like character of the
habitat at the type locality and several other collecting sites of this
subspecies.
Gerbillus pyramidum tarabuli Thomas

Gerbillus pyramidum tarabuli Thomas, Proc. Zool. Soc. London, vol. 2, pt. 1, p. 5,
October 1902 (Sebha, Fezzan Province, Libya).
Specimens examined. Three hundred thirty-one, from Fezzan:

Brach, 22, Edri, 12; Temenhint, 34; 4 km N Sebha, 10; 5 km NW
Sebha, 8; 3 km NW
Sebha, 10; Sebha, 19 (2 BM); 7 km Sebha, SW
5; 10 km SE Sebha, 1 El Abiad, 35 (3 skin only; 4 skeletons) Goddua,
; ;
15 (9 skull only); Meseguin, 33; Umm

el Araneb, 14; Traghen, 12;
6 km N Murzuch, 26; 28 km E Murzuch, 3; Murzuch, 4; 12 km N

Ghat, Ghat, 11; El Gatrun, 46; El Barcat, 10.
1;
adult females from the vicinity of Sebha are, respectively: Total
length 271.4 (261-289), 256.9 (246-277); length of tail 153.8 (143-
165), 145.3 (132-155); length of hind foot 33.1 (31-35), 31.9 (30-35);
length of ear 16.2 (14-17), 15.3 (14-17); occipitonasal length of skull
34.1 (32.5-35.6), 32.9 (31.8-34.2); length of auditory bulla 11.9
(11.4-12.3), 11.6 (11.2-12.3); crown length of upper molariform
toothrow 4.3 (4-4.5), 4.2 (3.9-4.5); greatest breadth across zygomatic
arches 17.8 (17.2-18.6), 17.3 (16.7-17.8); least interorbital breadth
6.8 (6.5-7.4), 6.7 (6.3-7.1); breadth of rostrum at level of antorbital
foramina 3.5 (3.3-3.8), 3.5 (3.3-3.8); length of nasals 13.4 (12.7-14.3),
12.9 (12.4-13.5).
Diagnosis. Upperparts ranging from Light Ochraceous-Buff to
Ochraceous-Buff, becoming paler on sides and flanks; entire dorsum
with grayish cast owing to alternate banding on individual hairs;
this grayish hue more concentrated in middorsal region postauricular ;
and supraorbital patches conspicuous and white; rostral, mystacial,

circumoral, and scapular areas, and entire underparts pure white;
this white extending dorsolaterally beyond the venter and forming
an abrupt line where it merges with the buffy color of the dorsum;
inner and outer surfaces of pinna of ear sparsely haired and Warm
Buff; eye ring black; vibrissae formed from about equal numbers of
light and dark hairs; dorsal and ventral surfaces of fore and hind feet
densely haired, each with five digits bearing claws; tail relatively long,
indistinctly bicolored, Light Ochraceous-Buff above and Pale Ochrace-
ous-Buff below, with prominent pencil (Wood Brown) extending along
dorsum of tail with light admixture of brownish
distal one-third of tail;
hairs. Skull: Large and robust; zygomata heavy; anterior palatine
foramina small; post palatine canals narrow and slitlike; braincase
flattened; supraorbital bead prominent; auditory bullae narrow
transversely and ventrally inflated.
Comparisons. In the shape and proportions of the skull, topotypes
of Gerbillus pyramidum tarabuli from the vicinity of Sebha, Fezzan
Province, Libya, are indistinguishable from specimens of Gerbillus
285-134 O —68 9
.
pyramidum pyramidum Geoffroy, as known from Giza Province,

Egypt. The nominate subspecies is significantly larger, however, in
all external and cranial measurements, particularly in the breadth of
the zygomatic arches and the length of the nasals. The two subspecies
are separable primarily by differences in color, the nominate subspecies
being noticeably darker in dorsal color with greater suffusion of dark-
colored hairs. Typical G. p. pyramidum also have darker and more
conspicuously bicolored tails, darker pencils, darker ears, darker and
less conspicuous postauricular and supraorbital patches, and darker
color around the entire circumorbital area (in many specimens of G. p.
tarabuli, the suborbital and postauricular patches are almost pure
white)
From two specimens (BM, nos. 13.8.6.53 and 13.8.6.60) representing
Gerbillus pyramidum hirtipes from Ain Sefra, southwestern Algeria,
G. p. tarabuli is paler and less brilliant in dorsal color, has a longer tail
with a more pronounced pencil, and is markedly larger in all other
external and cranial measurements.
For comparison with Gerbillus pyramidum hamadensis, see account
of that subspecies.
Remarks. In the original description, Thomas (1902) used the
darker color of G. p. pyramidum to separate these two subspecies, but
stated that, in the size and proportion of the skull, the two forms were
somewhat comparable. Setzer (1957) stated that gerbils representing
G. p. tarabuli were somewhat smaller but darker in color than those
representing G. p. pyramidum from the Nile Valley of Egypt. These
findings are in partial disagreement with mine, probably because the
above comparisons were based on smaller series, as these workers had
fewer specimens available at the time. Also Setzer included specimens
from Socna and Bir Fergian, Tripolitania Province, within G. p.
tarabuli. Gerbils from these localities are now known to represent a
subspecies distinct from G. p. tarabuli and significantly darker in
color.
Ellerman and Morrison-Scott (1951) placed Gerbillus jloweri Thomas
in synonymy under G. p. tarabuli and thus extended the range of
G. p. tarabuli to include portions of eastern Egypt and Sinai. Subse-
quently, Setzer (1958) questioned such a wide distribution for a
subspecies and designated Gerbillus pyramidum jloweri as a separate
subspecies based on its lighter color, more inflated auditory bullae,
smaller molariform teeth, more laterally curved anterior palatine
foramina, and less open pterygoid fossae. It is now known that G. p.
tarabuli is not part of the mammalian fauna of Egypt but is limited
in its distribution to the Libyan Fezzan; in all probability it occurs in
the Saharan Oases and in the vicinity of the Ahaggar Mountains of

southeastern Algeria.
Owing to the barriers provided by the Gebel es Soda, Gebel el Harug
el Asued, and Hamada de Tinrhert, little, if any, outbreeding takes
place with populations of G. p. hamadensis to the north and east. The

vast Idehan Murzuch and Gebel Ben Ghnema prevent interbreeding
with populations of gerbils to the south, so this subspecies has main-
tained its genetic identity.
Members of this subspecies from El Abiad and Temenhint are
darker dorsally than those from nearby Sebha. Specimens from the
vicinity of Murzuch, Goddua, Traghen, Umm
el Araneb, and Meseguin
show an orange-colored dorsal pelage. Animals from Ghat in the

extreme southwestern portion of the Fezzan are darker and more
variegated in dorsal color, with greater admixture of brownish hairs
on the shoulders and rump, and have slightly larger skulls with more
bulbous auditory bullae. Examples from El Gatrun, which is located
a great distance from the type locality at Sebha, possess almost all the
characters typical of tarabuli. These specimens, however, are slightly
paler in dorsal color and have slightly larger auditory bullae and
molariform teeth. Specimens from Brach and Edri in the Wadi es
Sciati are more brilliantly colored (Salmon or Orange-Salmon) than
any other representatives of this subspecies.
This local variation in color is demonstrable in all populations of
this gerbil throughout the Fezzan. In fact, it appears that these gerbils
develop different colored pelage depending upon the type of substrate
on and in which they live. This widespread variation is suggestive of a
broad genetic constitution, which enables these gerbils to respond
readily to the variety of conditions imposed by the prevailing en-
vironment.
An adult female, no. 322196, from El Gatrun is partially albinistic.
This specimen is entirely white except for some brownish hairs on the
distal portion of the tail and a faint suffusion of brown on the dorsum.
Ecological observations. These gerbils are the commonest rodents
of the Saharan Oases and prefer the sandy areas of the periphery and
interior of the oases but also inhabit the vegetated wadis which link
the various oases together. They occur sparingly in the margins of the
hamadas wherever localized sandy areas are present but apparently
do not inhabit the inner portions of these hamadas or the interior of
the desolate sand seas.
Their preferred habitat consists of areas of open sand and the
numerous small vegetated dunes near the oases proper. Date palms
occur in all Saharan oases, and these gerbils are frequently taken from
among the fallen palm fronds and other debris at the bases of the
trees. In the Fezzan, they were rarely taken from habitats lacking
palm trees.
On several occasions in the larger oases, these gerbils were purchased
from local Arabs and presumably had been living in the mud-brick
homes of these people. This is the only gerbil in Libya, known to me,
which shows commensal tendencies. In their requirements, they are
similar to Gerbillus gerbillus, but members of the latter species are less
abundant and more widely distributed owing to their wider range of
ecological tolerances.
Other rodents which occur with G. p. tarabuli include Jaculus jaculus
arenaceous, Gerbillus amoenus vivax, and Acomys cahirinus viator.
Figure 22. — Statistical comparison of length of head and body of the subspecies of Gerbillus
pyramidum: A, G. p. tarabuli; B, G. p. hamadensis.
] d
30
Figure 23.— Statistical comparison of occipitonasal length of the subspecies of Gerbillus
pyramidum. Notation remains the same as in figure 22.

Figure 24. — Statistical comparison of length of nasals of the subspecies of Gerbillus pyra-
midum. Notation remains the same as in figure 22.
Key to the Species of the Subgenus Dipodillus
1. Occipitonasal length less than 23 mm G. henleyi

Occipitonasal length more than 23 mm 2
2. Tail less than 90mm; without terminal brush and unicolorous G. kaiseri . .
Tail more than 90 mm; with prominent terminal brush and bicolored ... 3
3. Occipitonasal length less than 27 mm; auditory bullae large and markedly
inflated ventrally, anterior palatine foramina relatively short . G. amoenus
Occipitonasal length more than 27 mm; auditory bullae small and not markedly
inflated ventrally; anterior palatine foramina relatively long.
G. campestris
Gerbillus amoenus vivax (Thomas)
Dipodillus vivax Thomas, Proc. Zool. Soc. London, vol. 2, pt. 1, p. 8, October 1902
(Sebha, Fezzan Province, Libya).
General distribution of species. Egypt, Libya, and probably

Tunisia, Algeria, and Mauritania.
Distribution in Libya. Desert areas of Tripolitania and the
Fezzan. In Cyrenaica known from the littoral deserts near the Gulf of
Sirte, from Gialo Oasis, and the Gebel el Harug el Asued.
Specimens examined. Eighty-eight, from Cyrenaica: 5 km El W
Agheila, 2; Gialo Oasis, 4; Gebel el Harug el Asued, 200 km SE Zella,
3; from Tripolitania: 12 km Zliten, 2; 12 km WSirte, 1 2 km SW W ;
Hun, 1 5 km S Socna, 3; from Fezzan: Edri, 11 El Abiad, 8 (2 skull

; ;
only); Goddua, 15 (1 skin only, 12 skull only); Meseguin, 2; 6 km N

Murzuch, 6; 28 km E Murzuch, 3; 20 km N Ghat, 1; Ghat, 16; El
Gatrun, 4; El Barcat, 6.
Published records in Libya. Cyrenaica: Gheminez (Festa,
1921); Augila, Cufra, Gialo (de Beaux, 1932); Tripolitania:Ain
Hammam (Thomas, 1902); El Gheddahia (Toschi, 1951); Fezzan:
Sebha (Thomas, 1902); Ghat (Toschi, 1951).
of the ear and eye; rostral and circumoral areas Light Buff; pinna
sparsely haired and with small tuft of buffy hairs on anteroventral
margin; inner surface of pinna Ochraceous-Buff basally, becoming
darker distally and on outer surface approaching Light Grayish Olive;
vibrissae relatively short and formed of both light and dark hairs;
dorsal surfaces of forelegs, hindlegs, feet, and entire underparts white,
the latter, in some specimens, lightly suffused with Buff; fore and hind
feet each with five digits bearing claws; palmar and plantar surfaces
naked, the latter with six metatarsal tubercles; tail relatively long for
the species, sparsely haired with short, hispid hairs, moderately
tufted terminally, and in most specimens, distinctly bicolored Hair
Brown to Wood Brown dorsally and ranging from Light Buff to almost
white ventrally; the dorsum of the tail appears particolored owing to
the interspersion of light and dark hairs and to exposure of the pale
ground color from beneath; this variegated character of the tail is
typical. Skull: Relatively large and robust; auditory bullae markedly
large, bulbous and inflated ventrally; basioccipital narrow and rod-
shaped anteriorly, thus forming distinct elliptical posterior lacerated
foramina between it and the auditory bullae; anterior palatine
foramina short and rectangular; zygomata relatively heavy with
tendency to bow outward posteriorly; braincase moderately domed.
Comparisons. Near topotypes of Gerbillus amoenus vivax from
Goddua, Fezzan Province, Libya, differ from two topotypes of
Gerbillus amoenus amoenus (de Winton) from El Aiyat, Mit Riheina,
Giza Province, Egypt, in having larger and more robust skulls,
noticeably larger, more bulbous and more ventrally inflated auditory
bullae, heavier zygomata, wider zygomatic breadth, slightly larger
molariform teeth, and longer nasals. Specimens from Libya are also
paler in dorsal color, larger in total length, and have longer and slightly
more tufted tails.
Cranially, representatives of G. a. amoenus from Wadi Natroun,

Western Desert Governorate, Egypt, are appreciably larger than the
two topotypes from Giza Province and are comparable to specimens
of G. a. vivax but differ in having shorter and less tufted tails and
markedly smaller and less inflated auditory bullae. Another large
series of G. a. amoenus from near Faiyum, Egypt, has the smallest
auditory bullae of any of the specimens examined from Egypt and,
in this character alone, can easily be distinguished from those of
G. a. vivax from Libya.
This subspecies resembles Gerbillus nanus nanus Blanford from
Kerman Province, Iran but can be distinguished from the latter by
much shorter tail, lighter (less grayish) dorsal color, smaller cranial
size, noticeably smaller anterior palatine foramina, and markedly
smaller and less inflated auditory bullae.
From a single specimen of Gerbillus mackilligini (Thomas) from

Wadi Kansisrob, Sudan Government Administrative Area, Egypt,
specimens of G. a. vivax are noticeably smaller in overall size, more
variegated in dorsal color, and have shorter, more particolored and
less tufted tails, smaller hind feet, smaller and less inflated auditory
bullae, shorter anterior foramina and posterior palatine
palatine
canals, less expanded braincases, and are smaller in all measurable
cranial characters.
Compared to Gerbillus campestris dodsoni, G. a. vivax is much
smaller cranially and in overall size, has a noticeably shorter
and considerably larger and more inflated auditory
less tufted tail,
bullae, and much shorter anterior palatine foramina.
Remarks. Originally, Thomas (1902, p. 8) described Dipodillus
vivax from Sebha, Fezzan Province, Libya, and stated that it was
closely related to Dipodillus quadrimaculatus Bodenheimer and
Dipodillus amoenus de Winton of the D. quadrimaculatus group of Egypt
but differed from both by its decidely larger auditory bullae, narrower
basioccipital, and more uniform "ochraceous buffy" color. He thus
considered D. vivax as distinct from D. amoenus and regarded D. vivax
as the Tripolitanian representative of the quadrimaculatus group of
Egypt.
Later, Ellerman and Morrison-Scott (1951), Toschi (1954), and
Setzer (1957) regarded D. amoenus and D. vivax as conspecific but
included them as subspecies of Gerbillus dasyurus (Wagner), whose
range had previously been limited to Sinai and areas to the north
and east. The above workers had only a few specimens, and their
assignments probably were based largely upon geographic grounds.
In the present study, large series of topotypical G. dasyurus from
Sinai and amoenus from Egypt are available; the two groups were
G.
more compared, and the following differences are noted:
critically
Specimens of G. dasyurus are significantly larger in all cranial and
external measurements, have longer, denser, and more lustrous fur,
appear more variegated dorsally, and have longer and noticeably
more tufted tails, more inflated braincases, relatively narrower rostra,
longer and more slitlike anterior palatine foramina and wider basioc-
cipitals. It is evident that G. amoenus is a species distinct from G.
dasyurus and that the latter is confined, as was previously thought,
to areas east of the Nile River.
Toschi (1954) assigned specimens from Gheminez to G. dasyurus
amoenus (=G. amoenus amoenus) and those from other widely scattered
localities, including Cufra Oasis, to G. dasyurus vivax ( = G. a. vivax).
Setzer (1957), with reservations, included the few specimens available
to him within the subspecies G. a. vivax and did not recognize the
nominate subspecies as occurring in Libya.
Wassif (1956), in reviewing the dipodils of Egypt, recognized that

gerbils representing G. amoenus and G. dasyurus were not of the same
species and reinstated the former as a full species with a range con-
fined to Egypt west of the Nile. He predicted that specimens from
Tripolitania and other localities in Libya also probably represented
a variety of G. amoenus. The present author is in agreement with
Wassif in considering G. amoenus as a full species, and G. dasyurus
is no longer considered as a part of the Libyan mammalian fauna.
Specimens from the Fezzan are slightly larger in size of body, and
their skulls have slightly larger and more inflated auditory bullae
than those from Tripolitania and Cyrenaica. Within the Fezzan, the
various populations are remarkably similar in cranial and external
characters. Specimens from El Gatrun are slightly larger than topo-
typical specimens from near Sebha, and a large series from Ghat
and El Barcat are correspondingly smaller. To the north in Tripoli-
tania, specimens from Zliten and Socna are comparable in size to
typical G. a. vivax but have somewhat smaller auditory bullae.
Specimens from farther east in Cyrenaica and nearer the type locality
of G. a. amoenus are appreciably smaller in size and possess the
smallest auditory bullae of any representatives of this subspecies in
Libya. This diminution in bullar size in these easternmost populations
of gerbils suggests intergradation with G. a. amoenus. A gradient,
extending from west to east and showing progressive decrease in size
and degree of inflation of the auditory bullae, is apparent in popula-
tions of this gerbil in Libya and Egypt. Gerbils with small auditory
bullae from the Nile Delta thus represent the easternmost limits of
expression of this character, and those with larger auditory bullae
from the Fezzan are near the westernmost limits of this character
gradient. It appears that the subspecies G. a. amoenus and G. a.
vivax actually represent the two extremes in this clinal gradient.
It is not uncommon for populations at each end of a cline to develop
dissimilarities of sufficient magnitude to warrant designation as
distinct subspecies, especially when the clinal character extends over
such a wide geographic area.
While recognizing that the foregoing variation and clinal pattern
exists within and between populations of this gerbil in Libya, and
even though the easternmost populations show some characters
typical of G. a. amoenus, I prefer to assign all of the specimens from
Libya to G. a. vivax.
Four specimens from Biskra and a single specimen from Beni
Abbes, Algeria, are within the geographic range ascribed to Gerbillus
garamantis (Lataste). In cranial characters, these specimens are
strikingly similar to those of G. a. vivax from western Tripolitania
and the Fezzan. In cranial measurements, however, gerbils from Libya,
including near topotypes of 67. a. vivaxfrom Goddua, Fezzan, are

slightly smaller in length of skull and have slightly shorter nasals
and anterior palatine foramina. The length of the skull of the type
specimen of G. garamantis, as given by Lataste (1881), is also larger
than typical G. a. vir.ax. Gerbils from both countries agree closely
in external measurements, except for the slightly longer tail in those
from Libya. In color, the populations are almost identical, but speci-
mens from Algeria have longer, more lustrous, silkier fur. This
character is particularly striking in the specimen from Beni Abbes.
The specimens from Biskra differ in having appreciably longer and
more inflated auditory bullae. The foregoing differences are not of
sufficient magnitude to distinguish species but are more typical of the
subtle characters used to distinguish populations at the infraspecific
level. Furthermore, the enlarged auditory bullae of the specimens
from Biskra probably represent an extension of the east-west cline
apparent in populations of 67. a. amoenus and 67. a. vivax to the east.
When the Algerian populations have been studied more thoroughly
and topotypical material from Ouargla becomes available for com-
parison, these gerbils in Algeria and Libya, which are currently known
as 67. garamantis and 67. amoenus, respectively, probably will be
regarded as conspecific and differ only as subspecies. Because of
priority, the current 67. amoenus will become a synonym of 67. gara-
mantis or be relegated to subspecific rank under the latter.
When Lataste named 67. garamantis in 1881, Dipodillus campestris
(= Gerbillus campestris) and Gerbillus simoni Lataste were the only
North African gerbils known within the dipodil group. Comparative
specimens were largely unavailable from Libya and Egypt, and at that
time, these Algerian specimens were clearly distinct from all others
available to him. Ellerman and Morrison-Scott (1951) included
67.garamantis as a subspecies of Gerbillus nanus and thus extended
the range of the species far westward along North Africa. These
Algerian populations were several thousand miles distant from the
nearest representatives of 67. nanus east of the Nile River. The above
authors offered no explanation for this great hiatus in distribution of
members of the same species. Setzer (1952) assigned three specimens
from Kom Aushim, Giza Province, Egypt, to Gerbillus nanus gara-
mantis but thought it unlikely that gerbils from the Nile Valley and
Algeria belonged to the same subspecies. Wassif (1956) referred these
specimens from Kom Aushim to 67. a. amoenus and suspected that
67. nanus did not occur in Egypt. The present study indicates that
67. nanus is also not present in the mammalian fauna of Libya.
Ecological observations. These dipodils occur in virtually all

types of habitats in Libya. Specimens obtained at Ghat, Edri, El
Gatrun, Meseguin, and near Murzuch came from areas of loose sand
covered by fallen fronds at the bases of date palms. At Goddua this

specieswas taken from the loose sand of the palm groves and the
sandy-clay soils of other agricultural areas within the oasis. The
specimens from the Gebel el Harug el Asued were collected from rather
impervious clay soils in a badly eroded large wadi. At other localities
in Libya these gerbils were obtained in habitats ranging from the
dense plant cover of the coastal plain to almost barren hamadas.
They seem to prefer areas of loose sand or substrates of a sandy char-
acter; only rarely are they found in large, permanent dune areas.
The collection of specimens of this species was fortuitous, and
their presence in a given area could never be predicted with certainty.
Adequate series were obtained only after several days of continuous
trapping.
In many areas in Libya, members of this species occur sympatrically
with those of other species of gerbils, but they are almost always less
abundant.
Gerbillus campestris Levaillant
Gerbillus campestris Levaillant, Atlas Expl. Sci. Alg. Mamm. pi. V, fig. 2., 1857
(Philippeville, Province of Constantine, Algeria [Lataste, 1881]).
General distribution of species. Western and coastal Egypt;

Libya, Algeria, Morocco; range probably also includes northern por-
tions of Sudan, Chad, Niger, and Mauritania.
Distribution in Libya. Widespread throughout Cyrenaica, Trip-
olitania, and the Fezzan (currently unknown from Gialo Oasis).
Gerbillus campestris brunnescens. Cyrenaica: Cyrenaican Plateau
and adjacent Mediterranean littoral.
Gerbillus campestris dodsoni. Cyrenaica, Tripolitania, and the
Fezzan: Widespread throughout Tripolitania and the Fezzan, but in
Cyrenaica is limited to the southern oases of Tazerbo and Bzema.
Gerbillus campestris haymani. Cyrenaica: Vicinity of Giarabub
and Bahr el Tubat.
Gerbillus campestris patrizii. Cyrenaica: El Hauuari and El Giof
of Cufra Oasis.
Gerbillus campestris wassifi. Cyrenaica: Coastal plain of extreme
northeastern Cyrenaica.
Published records in Libya. Cyrenaica: Giarabub (de Beaux,
1928); El Giof (de Beaux, 1932); Tripolitania: Wadi Agarib, Ain
Hammam, Wadi Nefed, Sirte, Tamari-Ferdjan (Thomas, 1902);
FEzzAn: Goddua, Umm
el Abid (Thomas, 1902).
Comparisons. Gerbillus campestris can be distinguished readily

from the other dipodils in Libya by its much larger size, longer
anterior palatine foramina, and markedly smaller and less inflated
auditory bullae.
;
of the Fezzan, and a large portion of southern Cyrenaica. As a result

of the present study, five subspecies of 67. c. campestris are now known
to occur in Libya.
Ecological observations. Members of this species are the most
widely distributed of all Libyan rodents and occur in virtually all
habitats. In the larger oases where sandy areas and date palms are
widespread, gerbils belonging to the subgenus Gerbillus are more
abundant, and representatives of G. campestris are confined more to
the mesic habitats of the sedge pockets in the interior of the palm
groves or occur beyond the oasis proper in the zone of tamarix and
acacia. Apparently, G. campestris is not able to compete with these
sand-loving gerbils and is forced to inhabit these marginal habitats.
In a few cases, however, these gerbils occupied all habitats within the
oasis.Rocky areas appear to be preferred above all others; wherever
rocky outcroppings, or talus are present, these rodents occur in
cliffs,
abundance.
Gerbillus campestris brunnescens, new subspecies
Holotype. Adult male, skin and skull, USNM 302180, from 5 km

SE Derna, Cyrenaica Province, Libya; obtained Nov. 9, 1955, by
H. W. Setzer, original no. 2724.
Specimens examined. One hundred twenty-one, from Cyrenaica:
27 km E Apollonia, 3; 11 km SW Susa (= Apollonia), 5; 12 km SW
Apollonia, 2 (1 skin only); 12 km S Apollonia, 2; 5 km NW
Labrag,
6 (1 skin only) km W Labrag, 1 12 km NW Gubba 3 (1 skin only)
; 4 ;
3 km E Derna, 8; 5 km SE Derna, 27 (1 skin only) Wadi el Kuf, 13 ;
km WSW Messa, 7; 10 km SW El Faidia, 6; 7 km NE Slonta, 1; 5 km

W Tocra, 29 (1 skin only); 20 km SW Tocra, 16; 10 km N Gerdes, 3;
2 km N Coefia, 2.
Measurements. Averages and extremes of 15 males and 9 females
from the type locality, with the measurements of the type in brackets,
are, respectively: Total length 244.8 (228-262), 236 (223-252),
[248]; length of tail 143.1 (132-150), 137.8 (127-152), [148]; length of
hind foot 28.2 (27-30), 27.3 (27-28), [28]; length of ear 17.6 (17-19),
17.2 (17-18), [18]; occipitonasal length of skull 30.3 (28.9-31.6), 29.5
(28.9-30.2) [30.2]; length of auditory bulla 9.3 (8.9-9.7), 9.3 (9-9.5),
[9.3]; crown length of upper molariform toothrow 3.9 (3.8-4.1), 4 (3.9-
4.2), [3.9]; greatest breadth across zygomatic arches 15.6 (14.7-16.6),
14.9 (14.4-15.2), [15.5]; least interorbital breadth 5.5 (5.2-5.9), 5.4
(5.2-5.7), [5.7]; breadth of rostrum at level of antorbital foramina 3.2
(3.1-3.4), 3.2 (3.1-3.3), [3.3]; greatest length of nasals 11.6 (11-12.8),
11.3 (10.9-11.7), [11.2].
Diagnosis. Upperparts Sudan Brown becoming paler on sides and
scapular areas; all parts of dorsum with strong suffusion of blackish-
brown hairs, being particularly concentrated on rump; subauricular

region with strong admixture of blackish hairs; preauricular and post-
auricular patches conspicuous and white; eye ring black; mystacial,
circumoral, and pectoral areas Light Buff; pinna of ear sparsely haired,
Ochraceous-Tawny basally and Mummy Brown distally, and with
distinct tuft of grayish on antero ventral margin; vibrissae
hairs
relatively short and formed from equal numbers of light and dark hairs;
forearm with distinct buffy patch and otherwise white throughout;
hindlegs with conspicuous Dark Olive patch on ventral surface,
otherwise white throughout; fore and hind feet white, each bearing
five digits with claws; palmar and plantar surfaces of fore and hind
feet entirely naked, the latter Mars Brown in color; underfur of all
dorsal pelage Deep Plumbeous; entire underparts white with faint
buffy suffusion in some specimens; tail relatively long, distinctly bi-
colored, Verona Brown dorsally, almost white ventrally and slightly
penicillate. Skull: Small and gracile; auditory bullae small and mildly
inflated ventrally; zygomatic breadth narrow and zygomata fragile;
braincase slightly vaulted.
Comparisons. Compared with a single specimen of Gerbillus
campestris campestris from near Oran, Algeria, the type and para types
of Gerbillus campestris brunnescens are paler in dorsal color, have
slightly more tufted tails, much longer tails, more suffusion of dark
color around eyes, larger bodies, and cranially are larger and more
robust, with wider rostra, wider interorbital breadths, and larger
cranial measurements, especially length of the upper molariform
toothrow.
Members of this new subspecies may be distinguished from topo-
types of G. c. dodsoni from Ain Hammam, Tripolitania Province, and
a large series from Brach, Fezzan Province, by their darker (more
brownish) and more uniform dorsal color, longer and markedly less
tufted tails, greater overall length of body, slightly shorter ears, less
robust skulls, markedly smaller and less inflated auditory bullae, and
more vaulted braincases.
Compared with topotypical G. c. haymani from Siwa Oasis, Western
Desert Governorate, Egypt, and a small series representing haymani
from Bahr el Tubat, Cyrenaica Province, Libya, G. c. brunnescens is
markedly darker in dorsal coloration, has a less tufted tail, smaller
hind feet and ears, and a more gracile skull which is markedly smaller in
all cranial characters, being closest in the width of the rostrum and the
length of the upper molariform toothrow.

From topotypes of G. c. wassifi from the Libyan Plateau, near Salum,
Western Desert Governorate, Egypt, G. c. brunnescens differs in its
darker dorsal color, smaller overall size, relatively longer tail, smaller
ears, smaller skull, narrower anterior palatine foramina, narrower
zygomatic breadth, shorter nasals, and shorter and less inflated

auditory bullae.
This new subspecies can be readily distinguished from Gerbillus
campestris patrizii by its darker, more uniform dorsal coloration and
noticeably larger size of body and cranium, being comparable only in
length of the auditory bullae and length of upper molariform toothrow.
Remarks. Members of this subspecies can be distinguished from
all others in Libya by their darker, more uniform dorsal color and
somewhat longer and less tufted tails. Setzer (1957) assigned specimens
from the Cyrenaican Plateau to G. c. dodsoni. His assignment was
based solely upon some notes characterizing the type specimen of
G. c. dodsoni in the British Museum and upon favorable comparison
with the original description of Dipodillus dodsoni as given by Thomas
(1902). In the present study, more specimens are available from the
Cyrenaican Plateau, and topotypes of dodsoni, in addition to large
series from various other localities in Libya, are on hand. Thus, the
taxonomic status of these specimens from the Cyrenaican Plateau can
be determined more accurately.
The range of this subspecies is confined to the uplands of the Cyre-
naican Plateau and the adjacent littoral areas. Intergradation between
G. c. brunnescens and G. c. wassifi of extreme northeastern Cyrenaica
and coastal Egypt is apparent in specimens from the coastal areas
near Derna and Apollonia. In their pale dorsal color, large cranial
size, and wide anterior palatine foramina, they resemble G. c. wassifi,
but in the size and degree of inflation of the auditory bullae and in all
other cranial characters, they are closer to G. c. brunnescens and are
assigned to this subspecies. The vast areas of central Cyrenaica, in-
cluding Gialo Oasis and the Gebel el Harug el Asued, and the littoral
deserts and coastal plain along the Gulf of Sirte are not represented by
specimens and thus create a decided gap in distribution. The two
specimens from near Coefia, representing the southernmost record of
distribution for G. c. brunnescens, show no evidences of gene exchange
with G. c. dodsoni, whose range is farther to the south and west.
Within populations of this subspecies in Cyrenaica, local variation
in dorsal color is apparent. The pattern of this color gradient is too
irregular to suggest clinal variation and probably a genetic response
from the coastal plain,
to the local character of the substrate. Gerbils
where the soils aredarker, tend be darker
to and more uniform in
dorsal color than those from the plateau. This dark dorsal color
reaches its extreme in specimens from the vicinity of Tocra.
Ecological observations. These rodents are the most abundant
gerbils in the Cyrenaican Plateau and were regularly taken in trap-
lines. Other dipodils were collected on the plateau, but they were
fewer in number. Members of this subspecies occupy almost all of the
available habitats on the plateau and coastal plain ranging from the
exposed uplands with a scanty vegetative cover to the dense plant
cover of the larger wadis. They are perhaps most abundant in the
chaparral vegetation of the hillsides and throughout the dense vegeta-
tion of the coastal plain. They also occupy the rocky cliffs and talus
of the wadis and the rocky outcroppings along the coastal escarpments.
They were never collected from coastal dunes or other habitats com-
posed exclusively of sand. In these habitats they are supplanted by the
more sand-loving Gerbillus eatoni.
The name brunnescens, meaning dark, dusky, or brown, alludes to
the brownish color of the dorsal pelage of members of this subspecies.
Gerbillus campestris dodsoni (Thomas)
Dipodillus dodsoni Thomas, Proc. Zool. Soc. London, vol. 2, pt. 1, p. 7, October
1902 (Ain Hammam, Tripolitania Province, Libya).
Specimens examined. One hundred thirty-seven, from Cyr-

enaica: Tazerbo Oasis, 44; El Gezira (Tazerbo Oasis), 15; Bzema
Oasis, 9 (1 skin only); from Tripolitania: 5 km Cussabat, 6; W
20 km N Gharian, 3; 20 km E Rumia, 2; 12 km S Chicla, 2; Ain
Hammam, 3 (BM); Gebel es Soda, 60 km S Socna, 4; from Fezzan:
Brach, 31; Temenhint, 2 (1 skull only); 75 km W
Ubari, 2; 20 km
N Ghat, 1; Ghat, 1.
21 adult females from Tazerbo Oasis, Cyrenaica Province, are,
respectively: Total length 223.1 (205-238), 222.9 (211-254); length
of tail 126.6 (118-140), 127.9 (111-148); length of hind foot 26.9
(25-29), 26.7 (25-28); length of ear 16 (15-17), 15.7 (14-17); oc-
cipitonasal length of skull 30.4 (28.6-31.7), 30.5 (28.7-32.1); length
of auditory bulla 9.7 (9.2-10.1), 9.7 (9.2-10.1); crown length of upper
molariform toothrow 3.9 (3.7-4.2), 4 (3.8-4.3); greatest breadth
across zygomatic arches 15.8 (14.6-16.5), 16 (15.1-16.5); least in-
terorbital breadth 5.5 (5-6), 5.4 (4.7-6); breadth of rostrum at level
of antorbital foramina 3.2 (3-3.4), 3.2 (3-3.4); length of nasals
11.9 (10.6-12.6), 11.4 (11-13).
Diagnosis. Dorsal color ranging from Cinnamon-Brown, Sudan
Brown, and Verona Brown to Ochraceous-Tawny, all frequently
occurring in animals of the same local population; most specimens
with strong suffusions of darker hairs on dorsum, resulting in a varie-
gated or marbled appearance; other specimens uniformly colored;
in most specimens postauricular patches indistinct and suffused
with buffy hairs, and in others almost white; circumorbital areas
generally Light Buff with admixture of brownish hairs, in some
specimens darker and approaching color of dorsum; eye ring black;
scapular regions heavily suffused with brownish or buffy hairs, which
sometimes extend to forearm; vibrissae relatively long and longer

hairs dark brown and lighter ones usually white; pinnae of ears
sparsely haired, becoming darker distally, ranging in color from
Drab to Fuscous; circumoral areas, dorsal surfaces of forelegs, hind-
legs, feet, entireunderparts and, in some specimens, ventral surface
of tail pure white; fore and hind feet with five digits bearing claws;
palmar and plantar surfaces naked, the latter with six metatarsal
tubercles; tail medium in length with distal one-third forming prom-
inent brownish pencil; tail usually conspicuously bicolored varying
from almost pure white to Avellaneous ventrally and from Sayal
Brown to Warm Sepia dorsally this bicoloration less striking in darker
;
specimens. Skull: Medium in size; frontal bone with a distinct fossa

near the nasofrontal suture; zygomata strong and bowing outward
posteriorly; braincase flattened; auditory bullae noticeably large and
bulbous.
Comparisons. Compared to a single adult specimen of Gerbillus
campestris campestris from near Oran, Algeria, specimens from Brach,
Fezzan Province, and Tazerbo Oasis, Cyrenaica Province, are larger
in overall size,have markedly longer and more terminally tufted
tails, darker, more variable, variegated dorsal color, longer vibrissae,
and larger measurable cranial characters.
In dorsal color, these same specimens resemble topotypes of Gerbillus
campestris rozsikae Thomas from Biskra, Algeria but differ from this
subspecies in being larger in size of body and cranium and having
longer, more conspicuously tufted tails, shorter anterior palatine
foramina, and larger, more ventrally inflated auditory bullae.
For comparisons of these gerbils from Brach and Tazerbo Oases with
Gerbillus campestris brunnescens, G. c. wassifi, G. c. haymani, and G. c.
patrizii, see accounts of those subspecies.

Remarks. These specimens from Tazerbo and Brach, although
clearly referable to G. c. dodsoni, differ from topotypes of the latter
from Ain Hammam, Tripolitania Province, in having slightly darker
dorsal coloration, more tufted tails, larger auditory bullae, wider
rostra, and Other representatives
slightly wider interorbital region.
from various localities throughout Cyrenaica, Tripolitania, and the
Fezzan also differ from topotypical G. c. dodsoni in subtle characters.
These tenuous differences, however, do not exceed the variation
expected within populations of the same subspecies.
In 1902, Thomas described Dipodillus dodsoni {= Gerbillus campes-
tris dodsoni) from Ain Hammam, Tripolitania Province, Libya, as
distinct from Dipodillus campestris of the coastal areas of Algeria.

The two species were distinguished by the larger size, more "desert
colour," and more prominently tufted tail of D. dodsoni. He assigned
specimens, now known as Gerbillus campestris rozsikae, from Biskra,
285-134 O — 68 10
Algeria, to D. dodsoni and included all of Tripolitania and portions

of the Fezzan within the range of D. dodsoni. The present work
indicates that the range of G. c. dodsoni is confined to western and
central Libya and perhaps includes portions of eastern Algeria and
Tunisia.
Gerbils from the Gebel es Soda and the Gebel Nefusa are paler
in dorsal color and have larger, more inflated auditory bullae than
those from other localities in Tripolitania. Representatives of this
subspecies from near Cussabat are darker in dorsal color and have
less tuftedtails than those of populations to the south. Specimens
from near Ghat, Temenhint, and Ubari, in the Fezzan, have markedly
more tufted tails (sometimes the pencil occupies one-half the total
length of the tail), less streaked dorsal pelage, slightly larger auditory
bullae, and are slightly larger in general size than those from Brach
farther north. Except for slightly larger auditory bullae in gerbils
from the Fezzan, those from Tazerbo Oasis in Cyrenaica and from
Brach in the Fezzan, although widely separated geographically, are
indistinguishable in color and cranial characteristics. All of the
specimens here assigned to G. c. dodsoni possess prominently tufted
tails and have large, inflated auditory bullae, which are the most
diagnostic characters of this subspecies.

No specimens of G. campestris are available from the oasis of Gialo,
but because this species is widespread to the north and occurs abun-
dantly in the remote oases farther to the south in Cyrenaica, probably
they occur here also. Gialo is located between the ranges of G. c.
brunnescens and G. c. dodsoni, and when specimens of G. campestris
become available from Gialo, they may possess characters of both
subspecies.
With the exception of the large series from Brach, only six additional
specimens of this subspecies were obtained from the Fezzan. These
few specimens do not provide a reliable index to their actual numbers.
Suitable habitat is of widespread occurrence throughout the various
wadis, serirs, and oases of the Fezzan, and any barriers to dispersal
are of local extent. It is probable that these gerbils occur in all the
major oases in the Fezzan and also those of southern and central
Cyrenaica. This species has not been recorded from the Tibesti
Mountains but probably occurs there.
Evidences of gene exchange with other subspecies are lacking,
probably owing to the largely disjunct distribution of this subspecies
and also to the lack of specimens from most of the marginal areas;
hence, comparisons are not possible. Intergradation with G. c. brun-
nescens probably takes place somewhere along the littoral areas of
the Gulf of Sirte, and gene exchange with G. c. rozsikae and the
nominate subspecies may take place somewhere among populations
of these gerbils in eastern Tunisia and Algeria. Members of this

subspecies have no physical contact with those of G. c. wassiji of
the Mediterranean littoral, and it is doubtful if gene flow is a com-
mon occurrence between animals of G. c. dodsoni and G. c. haymani,

whose range includes the isolated depressions and the oases of Giara-
bub and Siwa far to the east.
Ecological observations. In the Gebel Nefusa, these gerbils
inhabit the crevices and fissures of the rocky outcroppings along
canyons that descend onto the coastal plain. Those from the Gebel
es Soda were taken among the coarse extrusions of volcanic material
near the highest point of the gebel. The large series from Brach was
obtained from moist areas supporting dense growths of sedges and
grasses associated with agriculture. At Temenhint Oasis, two speci-
mens were obtained from the sand surrounding date palms. Near
Ubari, two additional specimens were obtained from the practically
denuded margins of a shallow wadi which supported sparse growths
of acacia and occasional dried grasses. Near Ghat the habitat was
typical "hamada" desert without any visible plant cover.
At the oases of Bzema and Tazerbo, these gerbils occur abundantly
in the dense pockets of sedges that occupy the low-lying interior
portions of the palm groves. In fact, at Tazerbo, trap yields exceeded
85 percent and were the highest recorded for Libya.
A specimen from Tazerbo Oasis, an adult male, 325387, is strikingly
different in color from other specimens from this locality. It is uni-
formly gray throughout except for the tail, which is strongly suffused
with brown.
One specimen, an adult female, 325403, from Bzema Oasis is heavily
suffused with white on the dorsum, the tail has uniform admixtures
of white hairs throughout, and the ears and feet are extremely pale.
This specimen is the only example of albinism or partial albinism
observed in this species.
(Jerbillus campestris haymani Setzer

Gerbilius campestris haymani Setzer, Journ. Egypt. Publ. Health Assoc, vol. 33,
no. 6, pp. 208-209, 1958 (Siwa Oasis, Western Desert Governonite, Egypt).
Specimens examined. Four, from Bahr el Tubat, Cyrenaica

Province, Libya.
Measurements. Measurements of two adult males, 325501 and
325502, and two adult females, 325503 and 325504, from the above
locality are, respectively: Total length 268, 240, 239, 241; length of
tail 160, 141, 136, 140; length of hind foot 29, 30, 27, 28; length of
ear 17, 17, 17, 17; occipitonasal length of skull 32, 31.2, 31.5, 31.5;
length of auditory bulla 10.3, 9.8, 10, 10.1; length of upper molari-
form toothrow 3.9, 3.8, 4.2, 4.2; greatest breadth across zygomatic
,
arches 16, 16.4, 16.5, 16; least interorbital breadth 5.6, 5.4, 5.9, 5.5;
breadth of rostrum at level of antorbital foramina 3.1, 3.1, 3.4, 3.3;
length of nasals 12.6, 13.2, 12.5, 12.5.
Diagnosis. Middorsal region Clay Color to Tawny-Olive, becoming
paler on sides and flanks and approaching Cinnamon-Buff; entire
dorsum with uniform suffusion of brown hairs; circumorbital areas
Pale Pinkish Buff, strongly suffused with black hairs; postauricular
patches Pale Pinkish Buff; circumoral area, dorsal surfaces of fore and
hind feet and legs, and entire underparts white; upper arms with
indistinct buffy patch; fore and hind feet each with five digits bearing
claws; palmar and plantar surfaces of feet entirely naked, the latter
with six metatarsal tubercles; vibrissae relatively long with equal
numbers of white and brown hairs; pinnae of ears sparsely haired
Ochraceous-Buff basally and Hair Brown distally; tail relatively
long and indistinctly bicolored, being somewhat darker dorsally with
greater suffusion of brown hairs and with inconspicuous Drab pencil.
Skull: Large and robust; zygomata heavy; braincase flattened;
auditory bullae large; supraorbital ridges poorly defined; nasals long;
and rostrum wide.
Comparisons. From topotypes of Gerbillus campestris dodsoni from
Ain Hammam, Tripolitania Province, Libya, and representatives of
G. c. dodsoni from Tazerbo Oasis, Cyrenaica Province, specimens from
Bahr el Tubat are paler and more uniform in dorsal color; have longer,
less bicolored, and less tufted tails; proportionately smaller and less
inflated auditory bullae; and are larger in all measurements.
Compared to topotypes of Gerbillus campestris haymani from Siwa
Oasis, Egypt, specimens from Bahr el Tubat, Cyrenaica Province,
have slightly longer tails, smaller ears, slightly shorter molariform
teeth, and narrower zygomatic arches. These subtle differences, how-
ever, fall well within the range of variation for members of G. c.
haymani, and because these specimens from Bahr el Tubat are much
larger cranially and in body size from all other contiguous subspecies
they are clearly referable to G. c. haymani.
For comparisons with G. c. brunnescens, G. c. patrizii, and G. c.
wassifi, see accounts of those subspecies.
Remarks. Hayman (1949), somewhat dubiously, referred specimens
from Siwa Oasis in western Egypt to DipodUlus dodsoni but recognized
that these specimens differed from typical representatives of DipodUlus
dodsoni from Tripolitania Province in having slightly paler and less
rufous tails with slaty rather than brownish pencils. Ellerman and
Morrison-Scott (1951) assigned these same specimens to Gerbillus
campestris campestris and suggested that D. dodsoni was probably a
synonym of G. c. campestris and thus extended the range of the
nominate subspecies to include most of coastal and interior Libya and
a portion of western Egypt. Later, Wassif (1956) recognized that these

gerbilsfrom Siwa Oasis were distinct from those of the Mediterranean
littoral and assigned the coastal population to G. c. campestris and
those of the interior to G. c. dodsoni. Setzer (1958) concluded that the
from Siwa Oasis and those from the coastal areas were distinct
gerbils
from each other and from both G. c. dodsoni and G. c. campestris;
accordingly, as new subspecies, he described the interior population as
G. haymani and the coastal population as G. c. wassifi.
c.
It is now known that G. c. haymani represents a subspecies which

can be distinguished from all other subspecies of Gerbillus campestris
by its larger size, and its range is limited to the low-lying, sandy
depressions surrounding Bahr el Tubat and Siwa Oasis. When speci-
mens become available from Giarabub Oasis in Cyrenaica Province,
Libya, and the Qattara Depression of Egypt, they probably also will
be referable to G. c. haymani.
This small series was taken in the midst of an extensive growth of
Phragmites adjacent to the large saline lake of Bahr el Tubat. The
substrate in this habitat was hard, impervious, sandy clay which was
furrowed with cracks and crevices and covered with encrustations of
whitish salts.
Gerbillus campestris patrizii (de Beaux)
Dipodillus dodsoni patrizii de Beaux, Ann. Mus. Civ. Stor. Nat. Genova, vol. 55,
pp. 379-381, 1932 (El Giof, Cufra Oasis, Cyrenaica Province, Libya).
Specimens examined. Thirty-two, from Cyrenaica: El Hauuari,

Cufra Oasis, 15; El Giof, Cufra Oasis, 17.
adult females from the type locality are, respectively: Total length
211.8 (183-233), 213 (192-225); length of tail 118.4 (97-131), 121.7
(110-127); length of hind foot 26 (23-28), 26.1 (24-27) ; length of ear
15.9 (15-18), 15.7 (14-17) ; occipitonasal length of skull 29 (27.6-30.3),
28.6 (26.1-28.8); length of auditory bulla 9.3 (8.5-10), 9.3 (8.4-9.5);
crown length of upper molariform toothrow 3.8 (3.4-4), 3.8 (3.3-4);
greatest breadth across zygomatic arches 15.2 (14.3-15.8), 15.2
(14-16.2) least interorbital breadth 5.2 (4.8-5.4), 5.2 (5-5.6) breadth
; ;
of rostrum at level of antorbital foramina 3.1 (2.8-3.3), 3.1 (2.8-3.3);

length of nasals 11.2 (10.5-11.7), 10.9 (9.7-11.4).
Diagnosis. Representatives of this subspecies are extremely
Ochraceous-Tawny through
variable in dorsal coloration ranging from
Avellaneous, Cinnamon, and Tawny-Olive. Nearly all specimens have
strong suffusions of dark colors on the middorsal region and rump,
which impart a streaked appearance to the entire dorsum. This
variegated dorsal color is perhaps the most diagnostic and unifying
character of this subspecies. Other characteristics of this subspecies
are as follows: Postauricular patches indistinct and suffused with buff;
eye ring black; mystacial, rostral, and scapular areas Light Buff;
vibrissae relatively long with a preponderance of black hairs; ears
relatively long and pinnae sparsely haired, Antimony Yellow basally
and Saccardo's Umber distally, with small tuft of buffy hairs on
anterior margin; dorsal surfaces of forelegs, hindlegs, and feet white;
ventral surfaces of hindlegs with strong suffusions of plumbeous-
colored hairs; palmar and plantar surfaces of feet entirely naked, the
latter with six distinct metatarsal tubercles; fore and hind feet each
with five digits bearing claws; tail relatively short, distinctly bicolored
in most specimens, ranging from tan to dark brown dorsally and from
a medium buff to almost white ventrally (in some darkly colored
specimens, the tail is almost uniformly colored throughout) tail ;
markedly penicillate colored. Avellaneous to dark brown on distal

one-third; basal portions of all dark colored hairs Deep Plumbeous;
underparts white with occasional suffusion of buff. Skull: Noticeably
small and compact for the species; auditory bullae small, but mod-
erately inflated ventrally; zygomata heavy and narrow in breadth;
interorbital breadth narrow; braincase, notably the parietals, flattened.
Comparisons. Members of this subspecies can be readily dis-
tinguished from those representing Gerbillus campestris haymani and
Gerbillus campestris wassiji from farther north in Cyrenaica and
western Egypt by the markedly smaller size of body and cranium,
darker, more variable dorsal color, and more prominently tufted tail.
In general characters, these gerbils from Cufra Oasis most clearly
resemble those of Gerbillus campestris dodsoni, whose range is to the
west and north but differ in their smaller body size, shorter tail, and
smaller size of all cranial measurements.
For comparison with Gerbillus campestris brunnescens, see account
of that subspecies.
Remarks. De Beaux (1932, p. 379) described Dipodillus dodsoni
patrizii (—Gerbillus campestris patrizii) as a new subspecies separable
from Dipodillus dodsoni dodsoni (— Gerbillus campestris dodsoni) pri-
marily by its smaller size. Setzer (1957) placed G. c. patrizii in syn-

onymy under G. c. dodsoni and stated that the characters used by
de Beaux in separating patrizii from dodsoni were all typical of sub-
adult specimens of the latter. Setzer examined no specimens of typical
G. c. patrizii, however, and apparently based this assignment solely
on comparisons with the characters given in the original description.
In cranial and external measurements, the large series of adidt topo-
types collected by the present author agree closely with those given
by de Beaux in the original description. In studying these gerbils
from Cufra, extreme care was taken to select only fully adidt speci-
mens. The present study indicates that gerbils from Cufra, primarily
because of their diminutive size, warrant reinstatement as a distinct

subspecies.
Within series of these gerbils, all gradations of body size are noted,
ranging in length from 230 mm
to well below 200 mm. This wide
range may reflect degenerative changes in the genotype of the popu-
lation engendered by the agency of "drift" resulting from a long
period of absolute physical isolation without any outbreeding with
marginal populations. Specimens from nearby Bzema and Tazerbo
Oases, although similar in color to those from Cufra, are clearly re-
and cranial characters to G. c. dodsoni and
ferable in their larger size
show no evidences gene exchange with G. c. patrizii.
of
An adult male, 325348, from El Hauuari, is uniformly gray and
lacks the typical variegated character of the dorsal pelage. This
specimen probably represents an aberration in color without any
genetic significance.
Ecological observations. The from both El Hauuari and
series
El Giof were collected among sedges and other mesophytic plants that
occur as a fringe of dark green vegetation surrounding the low-lying
saline lakes in the interior of these oases. In most areas, these sedges
form almost impenetrable masses; consequently, trapping is usually
limited to the periphery of the habitat. In these areas the substrate
is extremely hard and impregnated with salts from adjoining lakes.
These gerbils were never taken from the sandy areas of the oases
proper. In these areas, Gerbillus gerbillus and Acomys cahirinus are
the dominant rodents.
Gerbillus campestris wassifi Setzer
Gerbillus campestris wassifi Setzer, Journ. Egypt. Publ. Health Assoc, vol. 33
no. 6, pp. 209-211, 1958 (Libyan Plateau, near Salum, ±200 ft., Western
Desert Governorate, Egypt).
Specimens examined. Five (1 skin only), from 5 km Bardia, W

Cyrenaica Province, Libya.
Measurements. Measurements of an adult male, 325509, from
the above locality are: Total length 245; length of tail 135; length of
hind foot 30; length of ear 17; occipitonasal length of skull 30.4;
length of auditory bulla 9.3; crown length of upper molariform tooth-
row 4.1; greatest breadth across zygomatic arches 16; least inter-
orbital breadth 5.5; breadth of rostrum at level of antorbital foramina
3.2; length of nasals 12.
Diagnosis. Dorsum Ochraceous-Tawny becoming lighter on sides
and rostral areas, all parts with strong suffusions of darker hairs;
pre- and postauricular spots and supraorbital patches conspicuous and
white with faint admixture of bmTy hairs; pectoral areas with indis-
tinct buffy patch; eye ring black; cireumoral areas, dorsal surfaces
of forelegs, hindlegs, feet, and entire underparts white; ventral surface
of hindlegs and thighs with conspicuous plumbeous-colored patches,

fore and hind feet naked ventrally, each with five digits with claws;
vibrissae relatively long and formed of equal numbers of light and dark
hairs; ears long, sparsely haired, Ochraceous-Buff basally and Brown-
ish Olive distally and with distinct tuft of buffy hairs on anteroventral
margin; tail prominently bicolored, dorsally same color as dorsum,
but with stronger suffusions of brown hairs, ventrally almost white;
tail with indistinct pencil; immature specimens of this subspecies
markedly paler and more uniform in dorsal color, approaching

Cinnamon-Buff, and lacking the strong suffusions of dark hairs.
Skull: Large and robust; molariform teeth large; auditory bullae
moderately inflated; anterior palatine foramina wide; zygomatic
breadth wide; nasals long and posterior palatine canals wide and
conspicuous.
Comparisons. From topotypes of Gerbillus campestris haymani
from Siwa Oasis, Western Desert Governorate, Egypt, this specimen
from Bardia differs in darker dorsal color with stronger suffusion of
brown, markedly shorter and more prominently bicolored tail with a
much reduced terminal brush, smaller auditory bullae, and smaller
size of all other cranial details, being of comparable size only in the
crown length of the molariform too throw.
Compared to topotypes of Gerbillus campestris dodsoni from Ain
Hammam, Tripolitania Province, Libya, this specimen from Bardia is
paler and more uniform in dorsal color, has markedly smaller auditory
bullae, wider interorbital breadth and rostrum, and less conspicuously
tufted tail.
For comparisons with Gerbillus campestris brunnescens and Ger-

billus campestris patrizii, see
accounts of those subspecies.
Remarks. This specimen from Bardia differs from typical G. c.
wassifi from Salum, Egypt, in having larger hind feet, a less promi-
nently tufted tail, more grayish hairs on the head, slightly smaller
auditory bullae, and slightly narrower anterior palatine foramina. In
view of the geographic proximity of Bardia to the type locality of
wassifi in Egypt and the continuity of habitat between them, even
these subtle differences would not be expected. Thus, this single speci-
men may not possess characters typical of the whole population. These
differences do not depart markedly from those considered typical for
G. c. wassifi, and specimens from Libya are included with this
subspecies.
In reviewing the dipodils of Egypt, Wassif (1956) assigned speci-
mens from coastal Egypt to Gerbillus campestris campestris; the type
locality of which is far to the west at Philippeville in the Province of
Constantine on the Algerian Coast. He had no specimens of typical
G. c. dodsoni but referred gerbils from Siwa Oasis, Egypt, to this
subspecies.
He stated further that these coastal gerbils differed from those of

the interior in their smaller size, denser fur, less tufted tails, and less
variable dorsal pelage. Later, Setzer (1958) also recognized these
differences and named Gerbillus campestris wassifi, with a range
ascribed to northwestern coastal Egypt, and described Gerbillus
campestris haymani representing the interior populations of Siwa
Oasis. He concluded that by virtue of its smaller size and prominently
biocolored G. c. wassifi was more closely related to G. c. dodsoni
tail,
than to G. haymani and attributed these similarities to the con-

c.
tinuity of suitable habitat between the ranges of G. c. dodsoni and

G. c. wassifi. At that time, however, the range of G. c. dodsoni
was thought to include northern Cyrenaica and thus to be contiguous
with that of G. c. wassifi. Northern Cyrenaica is now included within
the range of a new subspecies, G. c. brunnescens, and it probably was
to representatives of this subspecies, then known as G. c. dodsoni,
that Setzer (1958) alluded in discussing the affinities of G. c. wassifi.
It is now known that G. c. wassifi is more closely related to G. c.
brunnescens than to either G. c. dodsoni or G. c. haymani. The range

of G. c. campestris is thought to be confined to coastal Algeria and
Tunisia and possibly northwestern Libya.
In Libya, the range of this subspecies is confined to the Mediter-
ranean littoral of northeastern Cyrenaica at least as far west as Derna,

where intergradation with G. c. brunnescens occurs.
The specimens from Bardia were collected among some localized
concentrations of thorny, bushy perennials growing from residual
mounds or elevations in the bottom of a recently flooded and
denuded wadi.
30
Figure 28. — Statistical comparison of length of auditory bulla of the subspecies of Gerbillus
campestris. Notation remains the same as in figure 27.
Gerbillus grobbeni Klaptocz
Gerbillus (Dipodillus) grobbeni Klaptocz, Zool. Jahrb. Syst., vol. 27, p. 252, 1909
(Derna, Cyrenaica, Libya).
Remarks. No specimens of this gerbil were obtained in the present

study. Judging from measurements given in the original description,
this species most likely represents Gerbillus amoenus. Zavattari (1934)
reported G. grobbeni from Libya, but mentioned no specimens other
than those from the type locality. Later,Ellerman and Morrison-Scott
(1951) suggested that G. grobbeni probably represented Gerbillus nanus
Blanford. Toschi (1954) referred specimens from Derna and the Wadi
el Kuf to G. grobbeni but considered the latter as a subspecies of
Gerbillus dasyurus (Wagner). Setzer (1957) tried unsuccessfully to
obtain topotypes of this species but made no comments regarding its
probable status.
In the present work, gerbils representing the subgenus Dipodillus
were obtained from both the type locality at Derna and from the
Wadi el Kuf. These gerbils clearly represent Gerbillus campestris and
Gerbillus kaiseri, neither of which approach very closely the measure-
ments and description of G. grobbeni as given by Klaptocz in the
original description.
This species is considered to be of doubtful validity but is being
retained until such time as specimens are available to establish its
true status.
.
Gerbillus henleyi henleyi (de Winton)
Dipodilhts henleyi de Winton, Nov. Zool., vol. 10, p. 284, August 1903 (Zaghig,
Wadi Natroun, Egypt).
General distribution of species. Egypt, Sinai, Libya, and
Algeria; range probably also includes Tunisia.
northern Cyrenaica and the Gulf of Sirte.
Specimens examined. Six, from Cyrenaica: 11 km E Ain el
Gazala, 2; 20 km E Tobruch, 3 (1 skin only); 65 km WNW El

Agheila, 1
;
Diagnosis. Diminutive in size; pelage short and silky; upperparts

Buckthorn Brown, uniformly suffused with darker hairs and becoming
paler on sides and flanks; postauricular and supraorbital patches
distinct and white; subauricular areas with strong admixture of black
hairs; circumoral areas, portions of the cheeks, dorsal surfaces of
forelegs, hindlegs, feet, and entire underparts pure white; vibrissae
delicate and formed from both light and dark colored individual hairs
fore and hind feet extremely small, naked ventrally and each bearing
five digits with claws; tail Pinkish Buff, relatively long and with faint
terminal tuft, and appearing bicolored owing to moderate suffusions
of dark brown hairs dorsally; pinnae of ears extremely short and
sparsely haired outer surfaces of pinnae approaching color of dorsum
;
internal surfaces somewhat bicolored, becoming darker distally and

approaching Light Drab. Skull: Small and gracile; rostrum relatively
short zygomata fragile molarif orm teeth small and diverging slightly
; ;
anteriorly; braincase markedly expanded and appearing almost bul-

bous; auditory bullae noticeably large and markedly inflated ventrally;
basioccipital triangular-shaped posteriorly and projecting anteriorly
between auditory bullae as a narrow rod posterior lacerated foramina
;
large and distinct.

Comparisons. Compared to a topotype of Gerbillus henleyi makrami
Setzer from Bir Kansisrob, Sudan Government Administrative Area,
Egypt, these specimens from Libya are larger in all respects and have
more prominently domed braincases.
From topotypical Gerbillus henleyi mariae (Bonhote) from the vicinity
of Cairo, Egypt, these gerbils from Libya differ in having slightly
shorter skulls, smaller and less inflated auditory bullae, wider inter-
orbital breadths and rostra, shorter nasals, wider basioccipitals ante-
riorly, and smaller size of all external measurements except length of
ears. In color, G. h. mariae is noticeably darker with a greater suffusion
of grayish tones.
These specimens from Libya are almost indistinguishable from
topotypes of G. h. henleyi from coastal Egypt but differ in slightly
larger size of body, interorbital breadth and width of rostrum, and

smaller occipitonasal length of skull. In overall size of body and occipi-
tonasal length they are nearer to G. h. mariae, but because they share
more characters with the nominate subspecies and have a range con-
tiguous with populations of the latter in western Egypt, they are here
referred to G. h. henleyi.
Remarks. The diminutive size of members of this species serves to
distinguish them from all other gerbils in Libya. Until as recently as
1955, this distinctive species was unknown from Libya. Earlier col-
lectors in Libya tended to concentrate their efforts in the interior
oases and consequently neglected the coastal areas to which these
gerbils are largely confined. Even today the is poorly known

species
and is represented by comparatively few specimens from widely
scattered localities.
Wassif (1956) assigned specimens from western coastal Egypt to
Gerbillus henleyi jordani (Thomas), whose range is far to the west
in the Central Plateau of Algeria. He had no specimens of G. h.
jordani and based this assignment on similarities in size, particularly
in the length of the auditory bullae. Later, Setzer (1958) referred
all Egyptian specimens from west of the Nile River to G. h. henleyi
and asserted that G. h. jordani did not occur in Egypt. The inclusion
of the Libyan specimens within G. h. henleyi extends the range of
the nominate subspecies even farther westward and indicates that
G. h. jordani is not a part of the Libyan fauna.
In the original description of Dipodillus jordani (= Gerbillus henleyi
jordani), Thomas (1918) distinguished it from Dipodillus henleyi
( = Gerbillus henleyi henleyi) on the basis of the larger size of the
former. This larger size of G. h. jordani was further indicated by
Setzer, who compared members of the two subspecies at the British

Museum. Measurements made by the author on specimens from
Libya and of topotypical specimens of G. h. henleyi from coastal
Egypt are larger, rather than smaller as would be expected, than

those of the type specimen of G. h. jordani. For the present, until
much larger series of these gerbils become available from Algeria
and western Libya to provide more reliable comparative material,
I prefer to regard G. h. jordani as a subspecies of uncertain status
confined to the Central Plateau of Algeria.
Ecological observations. These small gerbils are apparently
confined to the coastal plain and littoral deserts of northern Libya.
Two specimens were obtained from near Ain el Gaza! a along the
coastal highway where a series of large, exposed hummocks supported
sparse growths of thorny perennials. These hummocks were riddled
with larger burrows, presumably those of jirds (Meriones), as two of
the latter were taken in the same trapline. Near Tobruch, three
specimens were obtained from the broad coastal plain. At this locality,
the plant cover was almost complete and continued uninterruptedly
for several kilometers. According to Setzer (1957), the habitat near
El Agheila is characterized by a "pebbly desert which is slightly
raised above the coastal plain."
Gerbillus kaiseri Setzer
Gerbillus kaiseri Setzer, Journ. Egypt. Public Health Assoc, vol. 33, no. 6, pp.
214-216, 1958 (Mersa Matruh, Western Desert Governorate, Egypt).
General distribution of species. Coastal areas of Libya and

Egypt as far east as the Nile River; range probably also includes
portions of Tunisia and Algeria.

Cyrenaica and the uplands of the Gebel Nefusa and Gebel Tigrinna
in Tripolitania.
Specimens examined. Twenty, from Cyrenaica: 3 km E Derna,
2; 2 km N Coefia, 6; 8 km N Benghazi, 2; 20 km E Tobruch, 5;
from Tripolitania: 20 1 km E Rumia, 12 km S Chicla, 4.
;
Measurements. Measurements of one adult male, 325028, and

two adult females, 325027 and 325550, from 20 kilometers east of
Tobruch, Cyrenaica Province, are, respectively: Total length 164,
169, 159; length of tail 83, 81, 77; length of hind foot 21, 21, 20;
length of ear 12, 12, 12; occipitonasal length of skull 25.8, 26.3, 24.4;
length of auditory bulla 8.4, 8.6, 8.1 crown length of upper molariform
;
toothrow 3.5, 3.8, 3.5; greatest breadth across zygomatic arches

14.2, 14.2, breadth
13.5; least interorbital 5.1, 4.7, 4.6; breadth of
rostrum at level of antorbital foramina 2.7, 2.6, 2.6; length of nasals
9.6, 9.7, 9.4.
Diagnosis. Color of the dorsum varying widely according to
geographic locality and ranging from Light Ochraceous-Buff, Cin-
namon-Buff, or Tawny Olive to Sayal Brown; complete dorsum
Gerbillus kaiseri
Figure 30. — Distribution of Gerbillus kaiseri.

interspersed with numerous brown-tipped hairs which render a

variegated or particolored aspect to the pelage; in most specimens
Plumbeous underfur exposed dorsally; region surrounding eyes
heavily suffused with black and gray, in some specimens extending
into subauricular area as a distinct patch; postauricular patches
white and somewhat limited in size; rostral area Cinnamon-Buff;
vibrissae relatively short, hairs having origin near the eye black
whereas those arising nearer the tip of the rostrum white; pinnae of
ears short, sparsely haired, Ochraceous-Buff basally and Hair Brown
distally and with prominent tufts of silky, buffy hairs arising from
the anterior margins; circumoral areas, pectoral region, dorsal sur-
faces of forelegs, hindlegs, feet, and entire underparts white; fore
and hind feet naked ventrally and each bearing five digits with claws;
tail noticeably short, except for suffusions of brown
unicolorous,
hairs dorsally, and somewhat paler in color than that of the dorsum;
a pencil or terminal tuft is lacking. Skull: Small; anterior palatine
foramina long and wide; posterior palatine canals distinct and rel-
atively wide; auditory bullae small and slightly inflated ventrally;
basioccipital present as a rather broad triangular plate between the
auditory bullae and forming an elongated foramen on its lateral
margins; zygomata relatively fragile; braincase moderately domed;
in very old adults a distinct longitudinal fossa present on the medial
surface of the frontals extending forward to the level of the infra-
orbital foramina.
Comparisons. Specimens from Libya somewhat resemble a spec-
imen (MNHN, no. 1789) of Gerbillus simoni Lataste from Guelt es
Stel, Algeria, but differ in noticeably longer tails, less robust skulls,
much more fragile zygomata, and paler, shorter, less lustrous fur.
Unfortunately, the auditory bullae, which are reputedly of taxonomic
significance in distinguishing G. simoni from G. kaiseri, are badly
broken in this specimen of G. simoni from Algeria, and comparisons
based on this character are not possible.
Members of this species can be readily distinguished from all other
dipodils in Libya by their markedly shorter and almost unicolorous
tails and the absence of a well-defined terminal tuft.
Remarks. Wassif (1956) assigned specimens from coastal Egypt to

Gerbillus simoni. This assignment was based on resemblances to the
measurements and descriptions of G. simoni as given by Lataste
(1881). Prior to this time, no specimens of G. simoni were known from
outside Algeria. Earlier, Ellerman (1941) included G. simoni and G.
henleyi within the G. simoni group but stated that G. simoni could
readily be distinguished from G. henleyi and all other members of the
group by its smaller auditory bullae. Later Ellerman (1949) and
Ellerman and Morrison-Scott (1951) considered G. simoni as a sub-
species of Gerbillus dasyurus (Wagner). The latter assignment has no

morphological or taxonomic justification, as the two forms have
nothing in common to suggest conspecificity. Setzer (1958), although
recognizing that these gerbils from Egypt and those from Algeria were
related, considered the Egyptian specimens to represent a new species,
G. kaiseri, distinguished from G. simoni primarily by its significantly
longer tail. He did not compare the crania of the two species but in-
ferred that those of G. kaiseri would be larger and suspected those of
G. simoni to be about the same size as those of G. henleyi. Judging
from the single specimen of G. simoni from Guelt es Stel, Algeria, the
two species are somewhat comparable in cranial size, but G. simoni
appears to be more robust, particularly in having heavier and more
massive zygomatic arches. Compared to G. henleyi, both G. kaiseri
and G. simoni are markedly larger cranially, have much shorter and
more unicolorous tails, proportionately smaller auditory bullae,
larger ears, and larger size of body.
The present series from Cyrenaica and Tripolitania constitutes the
first record of occurrence of G. kaiseri in Libya. Gerbils from the
Cyrenaican coast differ from topotypes and near topotypes of G.

kaiseri from Egypt in their slightly darker dorsal color and shorter
tails, but in all other characters the two populations are almost
indistinguishable. Specimens from the Gebel Nefusa in Tripolitania,

although geographically closer to the type locality of G. simoni in
Algeria, possess all characters typical of G. kaiseri and show no evi-
dences of interbreeding with G. simoni. It is now clear that G. kaiseri
and G. simoni represent two distinct species with separate ranges
ascribed to each. The range of the former includes the littoral areas
and coastal escarpments of Libya and Egypt, and G. simoni is con-
fined to the high plateaus of the Algerian Atlas.
Within Libya the various discontinuous populations are remarkably
uniform in color and cranial characters. Specimens from Derna, how-
ever, are slightly darker than those from elsewhere in Cyrenaica,
and four specimens from Tripolitania have smoother, more sparsely
haired tails than any of the other populations of this gerbil in Libya.
Adequate series are still lacking and a more thorough analysis of the
differentiation among the various populations will have to be deferred
until a later date. Itdoubtful that populations distributed over such
is
a wide geographic area represent the same subspecies.

Ecological observations. In Cyrenaica these gerbils are confined
narrow coastal plain and adjacent lowlands. Appar-
to the relatively
ently they prefer habitats lacking sand. Traplines set throughout
coastal dunes failed to take any of these gerbils. Farther west in
Tripolitania four specimens were collected from the highest portions
of the Gebel Nefusa in the upper reaches of narrow, grassy valleys
that eventually drain through the escarpment into the coastal plain.
In these high, windy valleys vegetation is rather abundant and
consists largely of grasses and other herbs.
These gerbils were never collected in large series and were usually
taken less frequently in traps than other species. Other species of
rodents occurring sympatrically with this species include Gerbillus
campestris, Gerbillus eatoni, Gerbillus henleyi, Gerbillus aureus, Ger-
billus amoenns, and Jaculus orientalis.
The sporadic distribution of these gerbils in coastal Libya is un-
accountable, as suitable habitats are present throughout virtually
allof coastal Libya.
This species is unknown from the coastal areas of Tripolitania and
the coastal plain of the Gulf of Sirte; however, more extensive col-
lecting in these areas will probably reveal its presence.
Genus Pachyuromys Lataste

Pachyuromys duprasi natronensis de Win ton
Pachyuromys dupresi (sic) natronensis de Winton, Nov. Zool. vol. 10, p. 285,
1903 (Bir Victoria, on way to Wadi Natroun from the Nile, Egypt).
General distribution of species. North Africa, including Egypt,

Libya, Tunisia and Algeria, and southward into Mauritania.
Distribution in Libya. Hamadas of Tripolitania, Cyrenaica, and
the northern Fezzan, and interior portions of the coastal plain of the
Gulf of Sirtein Cyrenaica and Tripolitania.
Specimens examined. Twenty-four, from Cyrenaica: 65 km WNW
El Agheila, 1 Wadi er Rueis, 340 km
; WNW
Tazerbo Oasis, 1 from ;
Tripolitania: 20 km E Rumia, 9; 3 km W
Rumia, 2; 12 km S Chicla,
5; 7 km S El Gheddahia, 1 5 km E Sirte, 2; 55 km SW Bir Allagh,
;
1; 15 km WNWMarble Arch, 1 5 km E Derg, 1. ;
Published records in Libya. Cyrenaica: El Agheila (de Beaux,

1932); Zauia Mechili (de Beaux, 1938); Fezzan: Bir el Fatia (Toschi,
1951).
Measurements. Averages and extremes of six adult males and the
measurements of two adult females from 20 kilometers east of Rumia,
Tripolitania, respectively: Total length 165.5 (158-175), 155,
are,
153; length of 55 (51-58), 51, 52; length of hind foot 23.5 (23-24),
tail
22, 22; length of ear 17.2 (17-18), 15, 15; condylobasal length of
skull 31.5 (30.8-32.1), 29.8, 29.7; greatest length of skull 37.7 (36.6-
39.2), 35.8, 34.8; greatest breadth across auditory bullae 21.7 (21.2-
22.3), 20.5, ?; crown length
of upper molariform toothrow 4.9 (4.6-5),
4.7, 4.7; least interorbital breadth 6.1 (5.9-6.3), 6, 6.1 greatest length ;
of nasals 12.9 (12.3-14), 11.9, 11.7; length of auditory bulla 18 (17.4-

18.7), 16.9, 16.9; depth of auditory bulla 14.2 (13.8-14.5), 13.5, 12.8;
greatest breadth across zygomatic arches 19.3 (18.7-19.9), 17.9, 18.5.
285-1.-M O— 68 11
ffllTTTTTTTrr^ LIBYA
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same color as the pelage of the dorsum, but with faint suffusions of
whitish hairs on ventral surface. Skull: Medium in size; flattened
dorsoventrally and wedge-shaped; parietal ridges poorly defined;
suprameatal triangles large and usually completely enclosed by
enveloping processes of the supraoccipital and temporal bones;
auditory bullae conspicuously enlarged and inflated ventrally; audital
portions of bullae extending posterior to mastoidal processes and
covering almost one-half of ventral surface of skull; meatal processes
markedly enlarged and projecting farther laterally than the squamous
portions of the temporal bones to which they are firmly adpressed;
mastoidal portions of bullae also markedly enlarged and bulbous,
and projecting conspicuously beyond the level of the occiput; pterygoid
processes relatively short; hamulae firmly applied to anterior surface
of the auditory bullae; anterior palatine foramina and posterior
palatine canals markedly enlarged and bowing laterally; zygomata
relatively delicate and slightly convergent anteriorly; lachrymals
relatively small.
Comparisons. This species resembles Meriones libycus but can
easily be distinguished from it and from all other jirds by its markedly
shorter and heavier tail, larger auditory bullae, and more wedge-
shaped body.
From a near topotype of Pachyuromys duprasi duprasi Lataste from
Ghardaia, Algeria, specimens from 20 kilometers east of Rumia,
Tripolitania, and those from various localities around the Gulf of Sirte,
differ in having more gracile and less robust skulls (particularly more
delicate zygomata), markedly larger posterior palatine canals, and
longer and narrower rostra. Most of the Libyan specimens are also
darker in dorsal color and smaller in external and cranial dimensions,
being of comparable or larger size in length of ears, crown length of
upper molariform toothrows, length of nasals, and length of auditory
bullae.
A specimen (skin only, BM, no. 39.2168) of Pachyuromys
single
duprasi faro ultiThomas from Mecheria, western Algeria, is markedly
darker in dorsal color than the Libyan specimens and has silkier, more
lustrous pelage. Cranial comparisons are not possible at this time, but
presumably P. d. jar o ulti differs significantly from Libyan specimens,
as the range of the latter is confined to the High Plateau of
northern Algeria and is. geographically, far removed from the Libyan
populations.
Remarks. Although the specimens from near Rumia, Tripolitania,
are clearly referable to Pachyuromys duprasi natronensis, they differ
from topotypes of the latter from Bir Victoria, Western Desert
Governorate, Egypt, in being larger in condylobasal length of skull,
and in having slightly larger auditory bullae, slightly wider zygomata,
and longer These specimens from Rnmia are also darker in dorsal
ears.
color, including the tail,and have a more marbled appearance dorsally,
owing to a much stronger suffusion of brownish hairs.
Compared to specimens from Rumia, specimen, 321831, from 5
kilometers east of Derg and another specimen, 321832, from 55 kilo-
meters SW of Bir Allagh, Tripolitania, have a more streaked pelage,
and the dorsal lip of the external auditory meatus is more inflated
and more closely applied to the squamous portion of the temporal.
In the above characters, these two specimens resemble P. d. duprasi of
Algeria, but in all other characters, they are closer to P. d. natronensis.
Toschi (1951) referred a specimen from Bir el Fatia, Fezzan Province,
to P. d. duprasi. He apparently based this assignment largely on
geographic grounds, as the measurements of this specimen are much
closer to those of topo types of P. d. natronensis.
Specimens from the coastal plain of the Gulf of Sirte in Tripolitania
are the palest in dorsal color and have the smallest hind feet of all
representatives of this species in Libya.

In view of the rather wide range of this species in Libya, specimens
from the various localities are remarkably uniform morphologically.
These similarities suggest a relatively constant genetic interchange
among by the continuity of
the various populations rendered possible
suitable habitat throughoutmost of coastal Libya. With the exception
of two specimens, 321831 and 321832, from western Tripolitania, a
single specimen from Bir el Fatia in the Fezzan (Toschi, 1951), and
a specimen, 325582, from the Gebel el Harug el Asued of central
Cyrenaica, this species in Libya is almost exclusively restricted to
localities near the coast. The comparatively few specimens available
at the present time cannot serve as a reliable index to their distribution
in Libya, and their range doubtless includes a much greater portion
of the interior than presentlyis supposed.
A single adult male specimen, 325582, from the Wadi er Rueis,

Gebel el Harug el Asued, central Cyrenaica, is the southernmost record
for this species in Libya, and it differs strikingly from all other rep-
resentatives of this species in Libya by much
paler dorsal color and
markedly smaller size of all measurements. Unfortunately, the pelage
of this specimen is badly worn, and the skull is badly broken rendering
certain cranial measurements unavailable. If this specimen is typical
of those comprising the population in this region of Cyrenaica, it
clearly represents a new subspecies. The jerboas (Jaculus deserti

vastus) of this isolated region have also undergone significant changes
in size and cranial dimensions. A moderate amount of morphological
divergence is not uncommon among populations of geographic isolates.
Ecological observations. The range of fat-tailed sand rats, in
Libya, closely corresponds to that of several other species of rodents,
.
particularly Meriones libycus, Gerbillus eatoni, and Psammomys obesus.

These unusual rats are most abundant in the transitional
deserts which run roughly parallel to the more lush coastal plain.
They seem to prefer the flat, rock-strewn surfaces of the hamadas and
the margins of the shallow, dry watercourses which dissect them. In
these areas, the surface is covered with coarse pebbles and occasionally
with larger boulders, and the vegetative cover is usually sparse and
localized. The collecting localities near Rumia and Chicla in Tri-
politania are located on the broad uplands near the brink of the
coastal escarpment where vegetative cover is denser, and the ground
is rockier with occasional outcroppings of larger rocks.
Setzer (1957, p. 60) reported the habitat near El Agheila as con-

sisting of a "rock shingle type of desert," and suspected that local,
terrestrial snails provided a source of food for fat-tailed sand rats.
Genus Meriones Illiger
The first records of members of this genus in Libya are those of

Thomas (1902, p. 8), who, reporting on specimens obtained by Edward
Dodson of the Whitaker Expedition to Tripoli (1901), referred
specimens from various localities in Tripolitania and the Fezzan to
Meriones shawi (Rozet) and Meriones schousboei (Loche). Thomas
noticed "characteristic differences" in the size of the auditory bullae
but mentioned that the two species were almost indistinguishable in
external characters and would probably prove to be "mutually
exclusive." He considered M. schousboei as the Barbary representative
of the Meriones erythrourus (Gray) group, the latter being widely
distributed throughout the Middle East and Southwest Asia.
Later, Thomas (1919), in an confusion of names
effort to clarify the
applied to this group, divided the genus into four groups based on
the relative size and shape of the auditory bullae and the suprameatal
triangle. Members of group "a" with large auditory bullae and large
suprameatal triangles included Meriones pelerinus Thomas, Meriones
crassus Sundevall, and Meriones pallidus Bonhote. Group "b," con-
sisting of animals with large auditory bullae but comparatively small
suprameatal triangles, included Meriones libycus Lichtenstein and M
schousboei. Those possessing both small auditory bullae and triangles
formed group "c," which consisted of Meriones isis Thomas and M.
shawi. The last division, group "d," consisted of those types having the
smallest auditory bullae and included Meriones syrius Thomas, Meri-
ones charon Thomas, Meriones ambrosius Thomas, and Meriones
blackleri Thomas, whose ranges were confined to Asia Minor and
Southwest Asia. The majority of the above species were described as
new at this time.
3.0 32 3.4 3.6 3.8 4.0 4.2
Figure 32. — Statistical comparison of breadth of rostrum of the species of Meriones:

A, M. caudalus; B, M. crassus; C, M. libycus.
Figure 33. — Statistical comparison of length of palate of the species of Meriones. Notation
12 13 14 15
Figure 34. — Statistical comparison of length of audital portion of auditory bulla of the
species of Meriones. Notation remains the same as in figure 32.
100 110 120 130 140 150
Figure 35. — Statistical comparison of length of head and body of the species of Meriones.
Notation remains the same as in figure 32.
13 14 15
Figure 36. — Statistical comparison of length of nasals of the species of Meriones. Notation
In Libya, local representatives of groups "a," M. pallidas tripolius

Thomas; "b," M. libycus caudatus Thomas; and "c," M. shawi were
recognized. Meriones pallidus tripolius from the Gebel Limhersuk,
Tripolitania, and M. libycus caudatus from Bir Ferdjan ( = Fergian)
El Hammam (=Ain Hammam) were described as new subspecies
in the same paper.
In Algeria, Meriones richardi (Loche) represented the "shawi"
group north of the Atlas. Populations to the south of the Atlas were
referred to Meriones gnyoni (Loche) and M. schousboei, which Thomas
considered as being "doubtfully separable" from libycus.
Ellerman (1941) divided the genus Meriones into three subgenera,
Parameriones Heptner, Cheliones Thomas, and Meriones, based on the
degree of hairiness of the sole of the hind feet. The latter subgenus
contained all of the North African species, consisting primarily of
M. M. schousboei, and M. erythrourus. These three species,
libycus,
in addition to M. grandis Cabrera, M. trouessarti Lataste, M. kozlovi
Satunin, M. longifrons Lataste, M. ismahelis Cheesman and Hinton,
M. arimalius Cheesman and Hinton, and M. charon, constituted the
"libycus" group. Ellerman (1941) included all of the Libyan jirds
in the single species, M. libycus. He considered M. shawi as a synonym

of M. I. and relegated M. crassus to subspecific status. Meriones
libycus
erythrourus, whose range included Russia and Southwest Asia, was
given specific rank but thought possibly to represent only a subspecies
of M. libycus.
In a later paper, Ellerman (1947) further divided the genus into
two additional subgenera, Sekeetamys Ellerman and Pallasiomys
Heptner, based upon both the hairiness of the hind feet and the
relative size of the auditory bullae, and placed all the forms with
large bullae, including all of the Libyan forms, in the subgenus
Pallasiomys. In addition, M.was elevated to specific rank and
crassus
M. shawi withdrawn from synonymy and established as a subspecies
of M. libycus. Thus the Libyan representatives of this genus were
treated as two full species, M. libycus and M. crassus.
Later in the year Ellerman and Chaworth-Musters in "A Revision
of the Genus Meriones" (1947), because of the sympatry of M. I.
shawi and M. I. caudatus, separated M. shawi specifically from both
M. crassus aud M. libycus. The above authors separated M. libycus
from the other two species by certain external characters (hind claws
d&rk= libycus; hind claws p&\e=crassus and shawi), and M. crassus
and M. shawi were distinguished by the size of the auditory bullae
(large in crassus; small in shawi). In addition to the coloration of the
hind claws, the degree of closure of the suprameatal processes was
given as a character separating M. crassus and M. libycus.
Toschi (1954) also recognized three species of the genus Meriones
in the Libyan fauna, M. libycus, M. crassus, and M. shawi, represented
by five subspecies, including the nominate subspecies of all three
species and M. libycus caudatus and M. I. conjalonierii de Beaux. Setzer
(1957), the most recent worker on Libyan mammals, included the
same three species in the Libyan fauna but considered M. crassus
tripolius distinct from M. c. crassus and did not recognize M. libycus
libycus and M. c. crassus as occurring in Libya. In addition, he de-
scribed M.
shawi azizi as a new subspecies from the northern Cyrenai-
can coast. Later, in reviewing the mammals of Egypt, Setzer (1961)
did not consider M. shawi as part of the Egyptian fauna and, further-
more, questioned its validity as a species. He was unable to find any
specific differences between M. shawi and M. libycus and concluded
that the former was probably conspecific with M. libycus farther west
in North Africa. The present author also doubts the validity of the
species M. shawi.
Specimens from coastal Libya, formerly referred to Meriones shawi
shawi and Meriones shawi azizi by Setzer (1957), are now considered
to be referable to Meriones libycus and are now known, respectively, as
Meriones libycus awratus (new subspecies) and Meriones libycus azizi.
However, certain bushy-tailed forms, previously recognized as

Meriones libycus confalonierii de Beaux and Meriones libycus caudatus
Thomas, are sympatric with these coastal subspecies of M. libycus.
It therefore becomes necessary to separate these populations taxo-
nomically. Because the name is older, it is here raised from
caudatus
subspecific status to full species rank. These populations may now be
referred to as Meriones caudatus caudatus and Meriones caudatus
confalonierii.
The jirds of Libya are here considered to be represented by the
following three species and seven subspecies, three of which are here
described as new: Meriones caudatus amplus; Meriones caudatus cau-
datus; Meriones caudatus confalonierii; Meriones caudatus luridus;
Meriones crassus tripolius; Meriones libycus auratus; and Meriones
libycus azizi.
The size and shape of the suprameatal triangle and the amount of
closure of the latter by the enveloping processes of the supraoccipital
and temporal bones varies among the three species ofMeriones and
serves to readily separate them. In M. crassus, the suprameatal tri-
angles are markedly larger than those of M. caudatus and M. libycus
and are open behind rather than completely or partially enclosed by
the supraoccipital and temporal bones. The suprameatal triangles are
of comparable size in M. caudatus and M. libycus, but in M. caudatus
the closure of the suprameatal triangles is complete, whereas in M.
libycus it is usually imperfect.
Individual genetic variation (nongeographic) is relatively high
among populations of jirds in Libya, yet the peculiar physiography
has resulted in the formation of several distinct subspecies. Morpho-
logical features most frequently associated with geographical dis-
tribution include differences in the length of the skull, palate, and
nasals, and changes in the shape and size of the audital and mastoidal
portions of the auditory bullae. In addition to cranial characters, the
size and general form of external features are taxonomically important.
Color, although sometimes highly variable within a given population,
is frequently useful in differentiating the various subspecies.
The subspecies of the several species of Meriones, especially those
of M. more sharply defined morphologically
caudatus, are generally
than those of the various species of Gerbillus. This apparent morpho-
logical segregation in Meriones may be misleading, inasmuch as the
samples representing the subspecies are much smaller than those
available for Gerbillus and may not demonstrate the complete range
of variation for any given character.
Of the three species of Meriones occurring in Libya, only Meriones
caudatus is represented by samples of sufficient size to justify statistical
analysis at the infraspecific level.
Key to the Jirds (Meriones) of Libya
1. Suprameatal triangle large and not enclosed posteriorly by enveloping processes

of temporal and snpraoccipital bones; mastoidal portion of auditory bulla
conspicuously inflated posteriorly and projecting markedly beyond level of
occiput M. crassus
Suprameatal triangle small and enclosed posteriorly by enveloping processes of
temporal and supraoccipital bones; mastoidal portion of auditory bulla not
conspicuously inflated posteriorly and projecting only slightly beyond level
of occiput 2
2. External meatal process of auditory bulla markedly inflated and adnate to
squamous portion of temporal bone; suprameatal triangle completely en-
closed by enveloping processes of the temporal and supraoccipital bones;
tail relatively long and prominently tufted M. caudatus
External meatal process of auditory bulla not markedly inflated and clearly
distinct from squamous portion of temporal bone; suprameatal triangle
imperfectly enclosed by enveloping processes of the temporal and supra-
occipital bones; tail relatively short and without prominent tuft.
M. libycus
Meriones caudatus Thomas
Meriones libycus caudatus Thomas, Ann. Mag. Nat. Hist., ser. 9, vol. 3, p. 267,
March 191'.) (Tamari-Fergian, Tripolitania, Libya).
General distribution of species. Currently known only from
Libya, but range probably includes other portions of North Africa
and adjacent Southwest Asia.
Distribution in Libya. Widespread throughout the desert areas of
Tripolitania and the Fezzan. In Cyrenaica, this species is known only
from the coastal plain and the hamadas surrounding the oasis of
Giarabub.
Meriones caudatus amplus. Fezzan: Oases of Murzuch, Traghen,
Meseguin, El Gatrun, and intervening wadis and hamadas.
Meriones caudatus caudatus. Tripolitania: Coastal plain and in-
terior deserts north of the Hamada de Tinrhert and Gebel es Soda.
Meriones caudatus confalonierii. Cyrenaica and Tripolitania:
Coastal plain and littoral deserts adjoining the Gulf of Sirte.
Meriones caudatus luridus. Cyrenaica: Coastal areas of northern
Cyrenaica and the inland hamadas as far south as Giarabub Oasis.
Published records in Libya. Cyrenaica: El Agheila (de Beaux
[Meriones libycus confalonierii], 1932); El Agheila (Meriones libycus
confalonierii), Giarabub (Meriones libycus caudatus) (Toschi, 1951).
Tripolitania: Ain Hammam and Tamari-Faradie (Toschi [Meriones
schousboei], 1951).
Comparisons. This species differs from Meriones libycus as follows:
Tail longer, with more prominent pencil, and black rather than brown;
auditory bullae noticeably larger and more inflated ventrally and
laterally; lateral meatal process of auditory bulla markedly expanded
schousboeiwas treated as a subspecies of Meriones libycus by de Beaux

(1932) and Chaworth-Musters and Ellerman (1947) and more recently
regarded as a synonym of Meriones libycus libycus by Ellerman and
Morrison-Scott (1951). In the present study, more specimens of M.
caudatus are available from numerous localities throughout Libya, and
the taxonomic position of this jird is better understood. In addition to
the bushy character of the tail, as established by Thomas (1919),
several cranial characters are apparent (see comparisons above) which
clearly establish caudatus as a distinct species from M. libycus of
M.
the Western Desert of Egypt. Its specific status is confirmed by its
being sympatric with populations of M. libycus from several localities
in coastal Libya.
The use of the name M. caudatus for this species can only be con-
sidered provisional at this time. It appears to be the oldest name
available for the Libyan population, but extralimital material (pres-
ently assigned to M. libycus) that I have examined from Algeria
(Beni Ounif) and Iran (Khurasan, Baluchistan, and Kopet Dag
Mountains) indicates that the present species is widespread in dis-
tribution and probably occurs over large portions of both North Africa
and Southwest Asia. It seems almost certain that older names are
available, but until additional material is forthcoming and a thorough
revision of Meriones throughout its entire range can be undertaken,
the name M. caudatus must serve for the Libyan population.
Ecological observations. This species is probably the most
abundant and widely distributed large rodent in Libya. These jirds
seem to prefer arid habitats and reach their greatest numbers in the
deserts of the Saharan interior. They occur sporadically, however, in
the coastal areas, where they are sympatric with Meriones libycus. In
the interior, this species prefers areas of abundant sand and is oc-
casionally taken in areas where large dunes are present; it is never
found associated with mesic habitats.
Throughout the range in Libya, these jirds occur together with
many species of rodents but most frequently are taken with repre-
sentatives of Gerbillus gerbillus, Gerbillus pyramidum, and Gerbillus
amoenus.
Meriones caudatus amplus, new subspecies
Holotype. Adult male, skin and skull, USNM 322681, from El

Gatrun, Fezzan Province, Libya; obtained Jan. 8, 1962, by G. L.
Ranck, original no. 1245.
Specimens examined. Twenty-three, from Fezzan: Meseguin, 8;
Traghen, 2; 28 km E Murzuch, 3; El Gatrun, 10 (1 skin only).
Measurements. Averages and extremes of four adult males and
measurements of two adult females, 322677 and 322679, all from the
type locality, with the measurements of the type in brackets, are,

respectively: Total length 316.8 (302-335), 318,?, [335]; length of
tail 160.8 (151-175), 166, ?, [175]; length of hind foot 38.8 (36-41),
39, 39, [40]; length of ear 21.3 (20-23), 21, 20, [21]; greatest length
of skull 42.2 (41.5-42.8), 42.1, 44.3, [42.8]; length of palate 18 (17.5-
18.5), 18.2, 17.8, [18.5]; length of audital portion of auditory bulla
16.2 (15.6-16.7), 16.4, 16.7, [16.7]; alveolar length of upper molariform
too throw 5.6 (5.5-5.9), 5.5, 5.9, [5.5]; least interorbital breadth 7.6
(7.5-7.8), 7.1, 7.6, [7.5]; length of nasals 15.9 (15.5-16.2), 15.9, 16.7,
[16.2]; breadth of rostrum at level of antorbital foramina 4.1 (4-4.2),
4.1, 4.4, [4.2]; greatestbreadth across zygomatic arches 23 (22.2-23.7),
22.5, 22.3, [23.7].
Diagnosis. rich, lustrous, normally ranging from
Upperparts
Ochraceous-Tawny Buckthorn Brown and with moderate admixture
to
of blackish- tipped hairs; in some paler specimens, the color of the
dorsum approaching Cinnamon-Buff and uniformly colored through-
out; long, silky guard hairs projecting 8 to 10 millimeters beyond
the level of the underlying pelage, particularly on the rump sides and ;
scapular areas lightly washed with gray; distinct whitish and bufFy
patches present above the eyes, between the ears and the eyes, and
below and behind the ears; in the pale color phase, these patches
almost pure white and contrasting markedly with the surrounding
pelage; mystacial, rostral, and circumoral areas ranging from buff to
pure white; eye ring black; pinnae of ears long, sparsely haired, and
in some specimens, almost naked; prominent row of buffy hairs
projecting from the anterior margin of the pinna into the internal
surface; vibrissae long, formed largely of white hairs, and sometimes
extending posteriorly for 25 millimeters beyond the level of the ears;
dorsal surfaces of fore and hind feet pure white; palmar surfaces
naked; ventral surfaces of hind feet densely furred but with promi-
nent hairless areas occupying the proximal one-half of the plantar
surface; both fore and hind feet with five digits with dark-colored
claws; tail exceedingly long for the species, and with a conspicuous
black pencil extending over the distal one-third of the dorsal surface
and continuing onto the ventral surface for perhaps 25 millimeters;
tail unicolorous in the dark color phase, except for some faint
suffusions of black hairs dorsally; tail contrastingly bicolored in the
pale color phase, Cinnamon-Buff dorsally and almost pure white
ventrally; entire underparts usually pure white, but in some speci-
mens, with faint suffusion of buff. Skull: Extremely large, angular,
and massive; zygomata strong and robust; auditory bullae markedly
inflated both ventrally and laterally; lateral meatal expansions
strongly inflated and firmly adpressed to squamous portion of tem-
poral bone; suprameatal triangle small, elliptical in shape, and
completely enclosed by enveloping processes of the temporal and

supraoccipital bones.
Comparisons. From representatives of Meriones caudatus caudatus
from 55 kilometers southwest of Bir Allagh, Tripolitania, the type
and paratypes of M. c. ampins differ in much larger and more robust
crania, particularly the zygomata, larger overall size, and noticeably
lighter dorsal pelage with less suffusion of black. Meriones caudatus
ampins also has proportionately smaller molariform teeth, and the
lateral meatal expansion is less closely adpressed to the squamous
portion of the temporal bone.
Members of this new subspecies can be easily distinguished from
those of Meriones caudatus conjalonierii from coastal Tripolitania
and Cyrenaica by markedly larger and more robust skulls, larger
overall size, and paler dorsal coloration.
Meriones caudatus ampins can be distinguished from Meriones
caudatus Inridus from the vicinity of Giarabub, Cyrenaica, by larger
size of skull and external dimensions, more conspicuous pencil, and
darker (less yellowish) dorsal color, particularly that of the tail.
Remarks. In its much and in external

larger size, both cranially
features, Meriones caudatus ampins is clearly distinct from all other
subspecies of Meriones caudatus. In color, it is closest to Meriones
caudatus caudatus of the Tripolitanian deserts, although it has much
less black on the dorsum. Cranially, this subspecies most nearly
resembles Meriones caudatus luridus from northeastern Cyrenaica.
The latter is also a form of rather large cranial size, but M. c. amplus
is still significantly larger.
Except for a few specimens representing Meriones crassus from Umm
elAbid, Zieghen, and the "Serir" of Murzuch, collected by Edward
Dodson of the Whitaker Expedition to Tripoli in 1901, the present
series constitutes the first record of occurrence of members of this
genus from the Fezzan.
Because of larger size, M. c. amplus might well be regarded as a
distinct species, but because of general resemblance to other sub-
species of M. caudatus in color, body form, general external characters
and in proportions, shape, and configuration of the skull, probably
it represents only an extreme in a clinal gradient.
This subspecies shows no evidence of gene exchange with Meriones
caudatus caudatus to the north. The Gebel es Soda and the barren
areas of the Hamada de Tinrhert and Hamada el Hamra apparently
restrict interbreeding between populations of these two subspecies.
This apparent allopatry has resulted in morphological divergence
in the Fezzanese populations, and this subspecies actually may
represent an incipient species.
In the Fezzan, these large rodents are known from comparatively

few localities. No specimens are available from northern and western
Fezzan, although suitable habitat is widespread, particularly in the
vicinity of the oases of Sebha, Temenhint, Brach, Edri, El Abiad,
Ubari, and Ghat. Future collecting in these areas, however, will
probably reveal the presence of these jirds.
The specimens available at present are almost indistinguishable
morphologically, which suggests a free interchange of genes within
and between populations in the eastern Fezzan. Two distinct color
phases, a dark and a light phase, are present among the specimens
from Meseguin and El Gatrun; some are intermediate in color. Animals
representing all these color types are indistinguishable, however, in
cranial characters. Therefore these wide ranges of color types suggest
a variable genotype for color and do not indicate basic genetic differ-
ences. At present, specimens are too few in number and are from too
few localities to allow any generalizations about the full extent of
variation among the various populations in the Fezzan.
These jirds from the Fezzan are clearly not closely related to any
other subspecies of M. caudatus in Libya and probably have closer
affinities with populations in southern Algeria, Niger, or the northern
Chad.
Ecological observations. These jirds apparently require loose
sand for their burrows. At all collecting sites, areas of pure sand or
small dunes formed the dominant habitats, and specimens were never
taken in areas devoid of sand. At the type locality at El Gatrun, a
good series was obtained from the small dunes and sandy areas inter-
spersed among the date palms. Frequently, these large rodents were
caught in traps set upon sandy elevations having a dense cover of
young palms. Usually in these habitats, large burrows are visible
beneath the palms. The collecting sites near Murzuch, Traghen, and
Mesequin were all characterized by areas of loose sand associated
with densely growing, bushlike date palms.
Apparently they do not occur in the marginal areas of the oases
where areas of loose sand and date palms usually are lacking. These
marginal habitats are typically hamada-like, and if vegetation is
present, it consists of Tamarix and Calligonum growing upon isolated
clay hummocks or sporadic growths of small, succulent plants inter-
spersed on barren playas.
In areas of loose sand, tracks and pathways of these jirds are wide-
spread and usually extensive complexes of burrows are present. On
several occasions, specimens were taken from traps set within or near
entrances to burrows. Only rarely were traps holding full adults found
in position in the trapline. Generally they were found dead several
feet distant from the nearest trap, apparently after having been fatally
struck.
At El Gatrun some of these jirds were purchased from the Arab

inhabitants and had presumably been living commensally with them.
This is not surprising in view of the sandy floors and hastily built
structures in which these people live.
Jirds are primarily nocturnal, but on one occasion at El Gatrun,
an old adult male, 322672, was shot in full daylight while it foraged
near the base of a clump of palms.
Gerbillus pyramidum is commonly associated with M. c. amplus
in these sandy habitats but is always much more abundant. The
smaller Gerbillus gerbillus occurs less frequently with these jirds.
The name amplus, from the Latin meaning large or largest, refers
to the large size of members of this subspecies.
Meriones caudatus caudatus Thomas
Meriones libycus caudatus Thomas, Ann. Mag. Nat. Hist., ser. 9, vol.3, p. 267,
March 1919 (Tamari-Fergian, Tripolitania, Libya).
Specimens examined. Thirty-four, from Tripolitania: 40 km
ENE, Nalut, 2; 5 km N Mizda, 1 ; 55 km SW Bir Allagh, 19; 5 km E
Derg, 6; 2 km SW Hun, 5; Bir Fergian, 1 (skin only).
Measurements. Averages and extremes of five adult males and five
adult females from 55 kilometers southwest of Bir Allagh, are: Total
length 282 (268-298), 276.8 (270-282); length of tail 141.4 (130-151),
138.8 (135-142); length of hind foot 35.8 (34-38), 33.8 (33-35) ; length
of ear 18.4 (17-19), 17.8 (16-19); greatest length of skull 38.7 (37.5-
39.7), 38.3 (37.4-39.5); length of palate 15.6 (15.3-16.1), 15.6 (15.3-
16); length of audital portion of auditory bulla 14.7 (14-15.7), 14.8
(14.3-15.4); alveolar length of upper molariform toothrow 5.3 (4.8-
5.5), 5.3 (5.2-5.5); least interorbital breadth 7 (6.8-7.1), 6.8 (6.5-
7.3); length of nasals 14.7 14.7 (14.4-15); breadth of
(13.8-15.4),
rostrum at level of antorbital foramina 4 (3.9-4.1), 3.9 (3.7-4.1);
greatest breadth across zygomatic arches 20.9 (19.9-21.7), 20.5 (20-21).
Diagnosis. Pelage of dorsum silky and lustrous with long guard
hairs projecting beyond shorter underlying buffy hairs; upperparts
Cinnamon-Buff grading to Pinkish Buff on sides and flanks; all parts
of dorsum strongly suffused with black, being particularly concen-
trated on rump; in most specimens, supraorbital and postauricular
patches prominent, Pale Pinkish Buff, frequently persisting as in-
distinct pale areas anterior to pinnae; eye ring black; circumoral and
mystacial areas, cheeks, and chin pure white, in some specimens, with
faint suffusion of buff; pinna of ear densely furred, same color as
dorsum, with distinct row of buffy hairs along anterior margin; dorsal
surfaces of fore and hind feet usually pure white, but, in some speci-
mens, with slight suffusion of buff; palmar surfaces naked; plantar
surfaces naked proximally, densely furred distally and almost pure
white; fore and hind feet each with five digits with claws, the latter
285-134 O —68 12
;
dark-colored basally and pale-colored distally; vibrissae relatively

long, formed from both light and dark-colored individual hairs, and
extending well beyond the level of the ears tail relatively long, dorsal
;
surface about same color as dorsum, but more heavily suffused with
black, and ventral surface uniformly Pinkish Buff; prominent black
pencil occupies terminal one-third of the dorsal surface of the tail and
extends along the ventral surface one-half this distance. Skull Medium :
in size; auditory bullae markedly inflated both laterally and ventrally

lateral meatal expansion large and applied to squamous part of the
temporal bone; suprameatal triangle small, elliptical, and completely
enclosed by enveloping processes of the temporal and supraoccipital
anterior portion of basioccipital and body of basisphenoid reduced to
narrow rod-shaped structures between the large bullae.
Comparisons. From topotypes of Meriones caudatus luridus, a
single topotype of Meriones caudatus caudatus, 302246, from Bir
Fergian, and near topotypes of M. c. caudatus from near Hun, Derg,
and Bir Allagh, Tripolitania, differ in having dorsal pelage more
heavily suffused with black, shorter tails with more prominent pencils
(black rather than brownish) and being significantly smaller in all
cranial measurements except the length of the upper molariform
toothrow.
When representatives of M. c. caudatus are compared with topotypes
of Meriones caudatus confalonierii, the latter differs in slightly paler
color of thedorsum and slightly smaller size of most cranial characters,
being comparable in least interorbital breadth and width of rostrum.
In general body size animals of the two subspecies are almost in-
distinguishable. Although the above characters are somewhat tenuous,
and M. c. confalonierii is poorly defined morphologically, the dif-
ferences are still sufficient for separating subspecies, and I prefer to
regard M. c. confalonierii as a valid form.
For comparison with M. amplus, see account of that subspecies.
c.
Remarks. A from Bir Fergian, 10 kilo-

single topotype, 302246,
meters south of Socna, Tripolitania, agrees closely in external measure-
ments and color with the original description of Meriones libycus
caudatus Thomas (1919, p. 267). Unfortunately, the skull of this
single topotype was inadvertently lost, and cranial comparisons are
not possible. Cranial measurements of representative series from
nearby localities are decidedly smaller than those of the type specimen.
I have not examined the type specimen of Meriones libycus caudatus
(= Meriones caudatus caudatus), but judging from the literature,

probably it represents an extremely old and outsized male which is
not entirely representative of this subspecies.
The various populations of this subspecies in Tripolitania are rather
uniform in cranial characters, but some local variation is present in
dorsal color. Two specimens from near Nalut, 321833 and 321834,
are the darkest dorsally of any of the Libyan populations of this
subspecies and have more orange on the dorsal surface of the tail.
Specimens from Derg, 321842, and Mizda, 321862, are noticeably
paler dorsally than other representatives of this subspecies.
Judging from the morphological homogeneity within and between
the various populations, these jirds must interbreed freely throughout
their range in Tripolitania. The vast expanses of the Hamada el
Hamra and the rugged escarpments of the Gebel Nefusa have proved
ineffective as deterrents to dispersal of this subspecies. To the north-
east, however, the more humid coastal areas apparently provide
unsuitable habitats, and to the south, the Hamada de Tinrhert and
Gebel es Soda have retarded or entirely prevented gene exchange with
the Fezzanese populations. Specimens are not available from localities
between the ranges of M. c. caudatus and M. c. conjalonierii to the
northeast and M. c. ampins to the south; however, when specimens
do become available from these areas, they probably will show
intergradation between M. c. caudatus and these contiguous subspecies.
Ecological observations. These jirds are unknown from localities
outside Tripolitania but have disjunct distribution throughout the
hamadas and oases of the interior. Then- range includes the inner
portions of the broad coastal plain of northwestern Tripolitania. The
climate and terrain here are not unlike those of the inland deserts
and generally are not typical of the coastal plain.
In Tripolitania, representatives of this subspecies were collected
from a wide variety of habitats as follows: Nalut: Localized, vegetated
dunes supporting Calligonum and bushy, thorny perennials. Numerous
burrows were present throughout the trapline, most of which were
probably made by gerbils. Mizda Occasional mounds and hard eleva-
:
tions in the bottom of a badly eroded wadi. The vegetative cover con-
sisted primarily of two species of thornbush growing among the
hummocks. Bir Allagh: An isolated concentration of sandy-clay
hillocks supporting thornbush in the midst of otherwise barren
hamada. Derg: Margins and bottom of a shallow wadi with occasional
hillocks of sand interspersed with denuded rock-strewn areas. Most of
the trap settings were made near the entrances of active burrows.
Hun: Large, localized permanent mounds supporting almost impene-
trable growths of a thorny perennial in the midst of an extensive
sandy plain.
These rodents are normally nocturnal, but near Hun, a subadult
specimen was caught in full daylight only a few seconds following
the setting of the trap.
At the various collecting sites in Libya, they occurred with Meriones
libycus; Gerbilius aureus; Jaculus jaculus; Gerbillus campestris; Jaculus
;
deserti; Pachyuromys duprasi; Gerbillus gerbillus; Gerbillus pyramidum;

Gerbillus amoenus; Gerbillus eatoni; and Acomys cahirinus.
Meriones caudatus confalonierii de Beaux
Meriones libycus confalonierii de Beaux, Ann. Mus. Stor. Nat. Genova, vol. 55,
p. 384, 1931 (El Agheila, Cyrenaica Province, Libya).
Specimens examined. Seven, from Cyrenaica: 5 km El Agheila, W

5; from Tripolitania: 15 km WNW
Marble Arch, 2.
Measurements. Measurements of an adult male, 302229, and
averages and extremes of four adult females from 5 kilometers west
of El Agheila, are, respectively: Total length 277, 280.5 (277-284);
length of tail 143, 146.5 (145-148) ; length of hind foot 34, 34.5 (34-36)
length of ear 18, 18.3 (18-19); greatest length of skull 36.4, 36.8
(36-37.4); length of palate 15.3, 15.1 (14.2-15.8); length of audital
portion of auditory bulla 13.9, 14.4 (13.7-15); alveolar length of
upper molarifocm toothrow 4.9, 5.1 (5-5.3); least interorbital breadth
length of nasals 13.5, 13.2 (12.7-13.5); breadth of
6.4, 6.6 (6.2-6.8);
rostrum at level of antorbital foramina 3.9, 3.8 (3.7-3.9); greatest
breadth across zygomatic arches 19.3, 19.8 (19-20.4).
Diagnosis. Identical to that of Meriones caudatus caudatus as
given in the preceding account, except for the slightly paler dorsal
pelage with less suffusion of black and the more brownish color of the
pencil. Cranially, confalonierii is smaller than caudatus in almost all
respects but in general salient features of the skull, there are no
significant differences.
Comparisons. This subspecies can be easily distinguished from both
Meriones caudatus amplus of the Fezzan and Meriones caudatus
luridus from northeastern Cyrenaica by its markedly smaller, more
gracile skull and much smaller overall size of body.
For detailed comparison with Meriones caudatus caudatus, see under
"comparisons" in account of that subspecies.
Remarks. Most of the specimens considered here are topotypes of
Meriones caudatus confalonierii and agree rather closely with measure-
ments and descriptions as given by de Beaux (1932) in the original
description.
Setzer (1957) had seven specimens from El Agheila, two of which he
erroneously assigned to Meriones libycus confalonierii (= Meriones
caudatus confalonierii). He also referred a single specimen (skull only,
302311) from Zliten, Tripolitania, to this subspecies. Actually, these
three specimens belong to Meriones libycus auratus (then Meriones
shawi shawi) and are specifically different from the other five specimens
from El Agheila, which were correctly assigned to Meriones libycus
confalonierii. The species M. libycus and M. caudatus thus occur
sympatrically at El Agheila.
This coastal subspecies is known from only the type locality at

El Agheila, Cyrenaica, and from Marble Arch in Tripolitania. Its
range, however, probably includes most of the coastal plain and
transitional desert along the Gulf of Sirte as far east as Benghazi.
The more mesic uplands of the Cyrenaican Plateau and Gebel Achdar
which are interposed between the ranges of this subspecies and that
of Meriones caudatus luridus to the east, provide unsuitable habitats
for these jirds. The coastal plain also is markedly reduced or absent
in many places in northern Cyrenaica and serves to further limit the
extent of habitats suitable for jirds and tends to reduce gene flow be-
tween the two subspecies. There are no physical barriers separating
the range of M. c. confalonierii from that of M. c. caudatus to the south,
but apparently the more arid climate farther inland is unsuitable for
members of this coastal subspecies. Some intergradation doubtless
takes place between these two subspecies somewhere in the transitional
desert south of the Gulf of Sirte.
Ecological observations. This subspecies is apparently confined
to the more humid habitats of the coastal plain and littoral deserts
surrounding the Gulf of Sirte in Cyrenaica and Tripolitania. The two
specimens from near Marble Arch were collected from the northern-
most terminus of the coastal hamada overlooking the coastal plain.
Vegetative cover here was sporadic, localized, and somewhat transi-
tional between that of the more densely vegetated coastal plain and the
much sparser plant cover of the hamadas farther inland. Setzer (1957)
describes the habitat at El Agheila as consisting of sandy areas with a
good vegetative cover. He also observed large, shallow burrows located
under bushes and containing from three to eight entrances. Seeds and
leaves of locally common plants were found in the burrows.
Meriones caudatus luridus, new subspecies
Holotype. Adult male, skin and skull, USNM 325597, from Bahr
el Tubat, 21 km ESE Giarabub, Cyrenaica Province, Libya; obtained
May 29, 1962, by G. L. Ranck, original no. 2187.
Specimens examined. Twenty-nine, from Cyrenaica: 11 km E
Ain el Gazala, 2; 10 km SW Fort Capuzzo, 1 60 km S Bir el Gobi, 17;
;
Giarabub, 2; Bahr el Tubat, 21 km ESE Giarabub, 5; 24 km SSE

Giarabub, 2.
Measurements. Measurements of an adult male, 325598, from the

type locality, and the measurements of an adult female, 302317,
from Giarabub, with the measurements of the type in brackets, are,
respectively: Total length 268, 240, [291]; length of tail 131, 123,
[155]; length of hind foot 34, 32, [36]; length of ear 18, 17, [19]; great-
est length of skull 39, 38.6, [40]; length of palate 16.6, 16, [16.3];
length of audital portion of auditory bulla 14.6, 14.5, [15.3]; alveolar
length of upper molariform toothrow 5.1, 5.1, [5.4]; least interorbital

breadth 7.5, 6.4, [7.4]; length of nasals 14.8, 14.5, [15]; breadth of
rostrum at level of antorbital foramina 4, 3.6, [3.6]; greatest breadth
across zygomatic arches 20.6, 20, [20.5].
Diagnosis. Upperparts subdued in color varying in tones of
Ochraceous-Buff, Light Ochraceous-Buff, Warm Buff and Pinkish Buff;
all parts faintly washed with gray; postauricular patches distinct,
almost pure white; region around eyes light gray strongly suffused
with black; cheeks and circumoral areas white; eye ring black; vi-
brissae long, formed of about equal numbers of black and white
hairs; pinna of ear sparsely haired, particularly on medial aspect and
with distinct row of buffy hairs along anterior margin; fore feet
naked ventrally, buffy-white dorsally and bearing five digits with dark-
colored claws; hind feet heavily haired ventrally, except for a promi-
nent hairless area confined to proximal half of plantar surface, white
dorsally and also bearing five digits with claws; tail relatively long,
unicolorous, slightly paler than that of the dorsum, with prominent
black pencil occupying distal one-fourth of dorsal surface and ex-
tending a short distance along ventral surface. Skull: Relatively
largeand robust; zygomata heavy and coarse; anterior palatine fora-
mina narrow and slitlike; auditory bullae large, markedly inflated
both ventrally and laterally, and adnate to squamous portion of
temporal bone; suprameatal triangle large and completely enclosed
by enveloping processes of the supraoccipital and temporal bone.
Comparisons. This new subspecies can be distinguished from both
Meriones caudatus caudatus of the Tripolitanian deserts and Meriones
caudatus confalonierii of the Tripolitanian and Cyrenaican coastal
plains by its larger body size, being larger in all cranial measurements
except breadth of rostrum, alveolar length of upper molariform
toothrow, and zygomatic breadth. In external measurements, Meriones
caudatus luridus is larger in greatest length and length of tail but of
comparable length in the length of hind foot and ear.
Although animals belonging to M. c. luridus are of large size, they
still differ strikingly from M. c. amplus in having smaller and more
gracile crania, particularly the zygomata, and smaller external

measurements.
In its markedly paler, more yellowish dorsal color, particularly
that of the tail, and in the less conspicuous pencil, this subspecies
differs strikingly from all other subspecies of Meriones caudatus.
Remarks. Toschi (1951 and 1954) assigned specimens from the
vicinity of Giarabub to Meriones libycus libycus and Meriones libycus
caudatus. Sympatry of two subspecies of the same species is not in
accordance with modern systematic and evolutionary concepts. In
the present work, however, M. libycus and M. caudatus are considered
as separate species, and the above mentioned sympatry is to be

expected.
I have not examined Toschi's specimens, but judging from his
measurements, they most likely belong to Meriones caudatus luridus
and not to Meriones libycus libycus. This does not preclude the pos-
sibility that the latter species occurs at Giarabub. Representatives
of both species occur sympatrically along the northern Cyrenaican
coast and doubtless are sympatric throughout coastal Egypt.
One specimen of M. c. luridus, 325586, from 11 kilometers east of
Ain el Gazala on the northern Cyrenaican coast is almost identical
in color to topotypical Meriones caudatus confalonierii from near El
Agheila farther west in Libya. This is the only evidence of intergrada-
tion with populations of M. caudatus to the west. The high tablelands
of the Cyrenaican Plateau, the massif of the Gebel Achdar, and the
marked reduction of the coastal plain apparently act as deterrents
to gene flow between members of this subspecies and those farther
west. Meriones caudatus luridus probably has greater affinities with
jirds in western Egypt. Its range probably includes Giarabub Oasis
and all of the inland hamadas south of the Cyrenaican Plateau in
Libya, the Libyan Plateau of extreme northwestern Egypt, and the
extensive low-lying areas of Siwa Oasis, the Qattara Depression, and
the Wadi Natroun in Egypt. The barren vastness of the Serir of
Calanscio and the Sand Sea of Calanscio, in Libya, and the desolate
Western Desert in Egypt apparently limit the southward distribution
of these jirds.
This subspecies is known from comparatively few specimens and
the full extent of its variation is unknown. Two specimens, 302316 and
302317, from Giarabub, however, are noticeably more orangish in

dorsal color and lack the typical yellowish-buff color of the tail. A
large series from south of Bir el Gobi, most of which are subadults,
are constant in showing this pale yellowish dorsal color.
Ecological observations. In Libya, this subspecies is confined
primarily to the hamadas and sandy deserts of the interior, but the
collecting sites 11 kilometers east of Ain el Gazala and 10 kilometers
southwest of Fort Capuzzo are less than 15 kilometers from the
Mediterranean coast. In coastal areas, members of this subspecies
occur together with those of Meriones libycus. The extent of the range
of the latter species farther inland has not yet been ascertained, but
the two species are probably sympatric throughout all of northeastern
Libya and northwestern Egypt.
These jirds have similar habitat preferences to populations of M.
c. caudatus and M. c. confalonierii in western Libya, all having a
decided predilection for sandy areas. The habitats at the following

localities in Cyrenaica are characterized as follows: 11 km E Ain el
Gazala: Series of vegetated sandy-clay hummocks near the roadside,

many of which contained large open burrows. Most burrows were
littered with plant debris and remnants of seeds. This site was situated
on the extreme northern terminus of the coastal hamada and approxi-
mately five kilometers from the Mediterranean Sea. 10 km Fort SW
Capuzzo: A localized pocket of bushes and smaller perennials
surrounded by almost barren hamada. Burrows were widespread in
the coarse, sandy soil. 60 km S Bir el Gobi Localized pockets of vege- :
tation bordering the sandy margins of a small wadi in the midst of

an otherwise featureless and totally barren "serir." 24 km SSE
Giarabub: Residual sandy-clay hummocks and lesser dunes inter-
spersed in areas of loose sand in the bottom of a broad, enclosed
wadi. Bahr el Tubat: Extensive concentrations of large vegetated
dunes. At the latter two collecting sites, Gerbillus gerbillus, Jaculus
jaculus, and Gerbillus campestris were taken along with these jirds.
The subspecies name luridus, from the Latin meaning pale yellow
or yellowish, alludes to the pale-yellow tones of the dorsal pelage.
PMM ?
3 4 ?
2 A ?
Figure 38. — Statistical comparison of greatest length of skull of the subspecies of Meriones
caudatus: A, M. c. amplus; B, M. c. caudaius; C, M. c. confalonierii; D, M. c. luridus.
Meriones crassus tripolius Thomas

Meriones pallidas tripolius Thomas, Ann. Mag. Nat. Hist., ser. 9. vol. 3, p. 265,
March 1919 (Gebel Limhersuk, in the northwest part of the country).
General distribution of species. West Pakistan, Afghanistan,

southern Russian Turkestan, Iran, Iraq, Syria, Lebanon, Jordan,
Israel, Saudi Arabia, Egypt (including Sinai), Libya, Algeria, and
the northern Sudan. The range probably also includes portions of

Mauritania, Niger, and the Chad.
Distribution in Libya. Interior deserts of Tripolitania and the
Fezzan; currently unknown from Cyrenaica, but probably occurs here
also.
Specimens examined. Twenty-four, from Tripolitania: Gebel
Limhersuk, 3 (BM) 5 km S Socna, 4 Bir Fergian, 4 (BM 2 skin only)
; ; ;
;
from Fezzan: Brach, 1 Edri, 1 26 km N Goddua, 1 75 km

; ; Ubari,
; W
4; 55 km SSW Serdeles, 2; 20 km N Ghat, 2; 12 km N Ghat, 2 (one
skull only).
Published records in Libya. Cyrenaica: Sidi Sweya (Thomas,
1902); Tripolitania: Wadi Agarib, Koshebey, Gebel Limhersuk
(Thomas, 1902 and 1919 [Meriones schousboei Loche and Meriones
pallidus tripolius)) Fezzan
; Umm
el Abid, Murzuch, Zieghen (Thomas,
:
1902); Bir el Fatia, Serdeles (Toschi, 1951).

Measurements. Measurements two adult males, 322660 and
of
322662, and one adult female, 322659, from 75 kilometers west of
Ubari, Fezzan, are, respectively: Total length 232, 232, 235; length of
121; length of hind foot 31, 30, 30; length of ear 17, 17,
tail 114, 121,
r^ —^\ l
18; occipitonasal length of skull 36.5, 35.9, 35.5; length of palate 15,
15.2, 15.1; length of anterior palatine foramina 6.1, 5.7, 5.7; length of
audital portion of auditory bulla 14.3, 13.8, 14.4; alveolar length of
upper molariform toothrow 5.3, 5.7, 5.2; least interorbital breadth
length of nasals 13.4, 13.6, 13.6; greatest breadth across
5.7, 5.9, 5.8;
zygomatic arches 19.8, 19.3 19.9; breadth of rostrum at level of ant-
orbital foramina 3.1, 3.4, 3.4.
Diagnosis. Pelage of dorsum fine, silky and lustrous; varying in
color from Light Ochraceous-Buff and Cinnamon-Buff to Pinkish
Cinnamon; most specimens with mild admixtures of brown-tipped
hairs on dorsum, which produce a slightly streaked or marbled ap-
pearance; sides, supraorbital, preauricular, and subauricular regions
with faint suffusions of gray; eye ring black or dark brown; circumoral
and mystacial areas, and conspicuous postauricular patches Pinkish
Buff; pinna of ear densely haired, about same color as dorsum, and
with distinct row of buffy hairs along anterior margin; vibrissae long,
formed of black and white individual hairs; forefeet almost pure
white dorsally, naked ventrally and with five digits bearing pale-
colored claws; hind feet usually white dorsally, with thick covering of
hairs ventrally, except for conspicuous hairless areas on proximal
halves of plantar surfaces, and also with five digits bearing pale-
colored claws; tail relatively short for the species, Pinkish Buff in
color, appearing bicolored owing to slight admixture of black hairs

dorsally, and with distinct brush of black or dark brown hairs distally.
Skull: Relatively small and gracile for the species; auditory bullae
large and bulbous, noticeably inflated ventrally, laterally, and poste-
riorly,but to a lesser extent than in other subspecies; lateral meatal
expansion closely applied to squamous portion of temporal bone;
suprameatal triangle large, triangulariform, and imperfectly closed
posteriorly by enveloping processes of the supraoccipital and temporal
bones.
Comparisons. From representatives of Meriones crassus crassus
Sundevall from the vicinity of Feiran Oasis and St. Catherine's
Monastery, Sinai, Egypt, paratypes of Meriones crassus iripolius from
Tripolitania and representatives of M. c. tripolius from various localities
in the Fezzan darker dorsal color, shorter and less inflated
differ in
auditory bullae, less robust zygomata, and generally much smaller
size of cranium and all external dimensions.
Compared to representatives of Meriones crassus pallidus Bonhote
from the Eastern Desert Governorate, Egypt, M. c. tripolius differs in
the same characters as given for M. c. crassus. Adult specimens of
M. c. asyutensis Setzer are not available for comparison, but judging
from the original description (Setzer, 1961), they are significantly
,
larger than those of M. c. crassus and, by inference, must be markedly

larger than those of M. c. tripolius in every detail.
Meriones crassus can be distinguished easily from both Meriones
libycus and Meriones caudatus by larger and more posteriorly inflated
auditory bullae, larger suprameatal triangles with open posterior
processes rather than closed or closely approximating as in M. libycus
and M. caudatus, shorter tail, and generally paler, more uniform
dorsal color.
Remarks. Setzer (1961) described Meriones crassus perpallidus from
Cairo- Alexandria Road, 4 kilometers from Cairo, Egypt, and stated
that it differed from representatives of Meriones crassus tripolius by
paler dorsal color, larger, more massive skull, wider, shorter rostrum,
and more posteriorly inflated auditory bullae. When paratypes of
these two subspecies are compared, however, M. c. tripolius can be
most easily distinguished from M. c. perpallidus by darker dorsal color,
with stronger suffusion of brownish hairs, and by markedly smaller
size of external dimensions. Paratypes of M. c. tripolius also have
slightly larger and more massive skulls, rather than smaller skulls
as stated by Setzer, and the mastoidal portion of the auditory bulla
is less inflated posteriorly. The breadth and length of the rostra do not
differ appreciably in paratypes of M. c. tripolius and M. c. perpallidus.

When Setzer described M. c. perpallidus, he had only a few specimens
of M. c. tripolius from Tripolitania and apparently did not examine
typical specimens of M. c. tripolius from the Gebel Limhersuk. In the
present work, paratypes of both subspecies are at hand, some additional
characters by which these two subspecies differ are noted, and some of
the differences set forth by Setzer are not apparent.
Thomas (1902) assigned specimens obtained by Edward Dodson in
1901 from several localities in Tripolitania and the Fezzan to Meriones
schousboei Loche. In a later paper (1919), using these same specimens,
he described Meriones pallidus tripolius (= Meriones crassus tripolius)
with the type locality at Gebel Limhersuk, Tripolitania, and named
Meriones libycus caudatus from Tamari-Fergian, Tripolitania. He did
not ascribe a specific range to either species, however, and in no way
indicated to which of the above species the specimens from the
Fezzan might be referred. Many years later, Chaworth-Musters and
Ellerman (1947) assigned these Fezzanese specimens to Meriones
crassus.
Since Thomas described Meriones pallidus trijwlius ( = Meriones
crassus tripolius) in 1919, this subspecies has had an uncertain status.
It was treated as a subspecies of Meriones libycus (Ellerman, 1941)
first
later considered as a subspecies of Meriones crassus (Ellerman, 1947

and Chaworth-Musters and Ellerman, 1947), and finally regarded as
a synonym of Meriones crassus crassus (Ellerman and Morrison-Scott,

1951 and Toschi, 1954). More recently, Setzer (1957) reinstated M. c.
tripolius as a valid subspecies of M. crassus and assigned specimens

from near Socna and Bir Fergian, Tripolitania, to it. He stated that
these jirds from Libya were markedly paler and had smaller skulls
with particularly less inflated posterior portions of the auditory bullae
than typical specimens of M. c. crassus from Sinai. Thomas (1919),
in the original description of Meriones pallidus tripolius, said that it
was very near to Meriones pallidus (= Meriones crassus pallidus), of
the Sudan, but with smaller auditory bullae and slightly darker
dorsal color.
Throughout the known range in Libya, populations of these jirds
are remarkably uniform in cranial characters and dimensions. Jirds
from several localities in the Fezzan clearly belong to this subspecies
but differ from topotypes of M. c. tripolius from farther north in
Tripolitania in having slightly more gracile skulls, more delicate
zygomata, and in being slightly smaller in external dimensions. In
color, all Libyan representatives of this subspecies are inseparable.
Two specimens, a very old male (BM, no. 34.8.2.64) and an old female
(BM, no. 34.8.2.65) from El Golea, Algerian Sahara, are comparable
in color and external measurements to the specimens from Libya
but are noticeably larger cranially. This larger size of the Algerian
specimens may result from increased angularity and thickening of the
skull attributable to extreme old age.
Ecological observations. In Libya, these jirds are represented
by comparatively few specimens from Tripolitania and the Fezzan
and are unknown from Cyrenaica. Several days of intensive trapping
at the oases of Brach and Edri in the Fezzan produced only two
specimens, both from the hamada-like plains several kilometers
distant from the palm groves and sand of the oases proper. Near
Ghat and Ubari they inhabited the margins of shallow wadis that
dissected the desolate, flat hamada. At all these collecting sites, vegeta-
tive cover was reduced to occasional bunches of dry grasses and
scattered acacias. One evening in January, while I camped on the
barren plain 26 kilometers north of Goddua, Fezzan, a single male
specimen, 322693, was captured alive when it approached to within
a few feet of the lantern. Visible vegetation at this campsite was
entirely lacking except for a distant solitary acacia tree. These hardy
rodents seem to prefer the more barren hamadas and sparsely vegetated
wadis located some distance from oases and were never collected
from dunes or sandy areas within palm groves. In the latter habitats,
however, the larger, more bushy-tailed Meriones caudatus is rather
common. Because of their preference for these marginal habitats,
specimens of M. crassus are obtained only rarely and are poorly
represented in collections. The present series probably provides a

very incomplete picture of their actual distribution in Libya. The
jirds obtained by Edward Dodson, plus additional specimens from
Bir el Fatia and Serdeles (Toschi, 1951), those obtained by Setzer
(1957) from near Socna and Bir Fergian, and the present series
collected by the author from the Fezzan constitute the only known
records of occurrence of Meriones crassus in Libya.
Meriones libycus Lichtenstein
Meriones libycus Lichtenstein, Verzeich. Doubl. Zool. Mus. Berlin, p. 5, no. 9,

1823 (Lichtenstein gives the type locality as "e deserto libyco," but Ellerman
and Morrison-Scott [1951:644] limit the type locality to "Near Alexandria.")
General distribution of species. Russian Turkestan, Chinese

Turkestan, Transcaucasia, Kara-Kum, Kizil-Kum, Iran, Baluchistan
(West Pakistan), Afghanistan, Iraq, Syria, Israel, Saudi Arabia,
Sinai, Egypt, Libya, Tunisia, Algeria, and Morocco.
Cyrenaica and Tripolitania.
Meriones libycus azizi. Cyrenaica Coastal areas of extreme northern
:
Cyrenaica.
Meriones libycus auratus. Cyrenaica and Tripolitania: Coastal
plain of Gulf of Sirte and Tripolitania as far inland as the coastal
escarpment.
Published records in Libya. Cyrenaica: Bou Cheifa (Thomas,
[Meriones shawi shawi] 1902) Benghazi, Gheminez (Festa [Meriones
;
guyoni Loche], 1921); Giarabub and vicinity (de Beaux, 1928); Augila
(de Beaux [Meriones guyoni], 1932); Derna (Zavattari, 1934); Merg
( =Barce) (de Beaux, [Meriones shawi shawi] 1938); Benghazi,
Gheminez (Toschi, [Meriones shawi shawi] 1951); Tripolitania:
Wadi Aggar, Wadi Nefed, Tarhuna (Thomas, [Meriones shawi shawi]
1902).
Comparisons. This species can be readily distinguished from
Meriones crassus by coarser, less silky fur markedly smaller and less
;
inflated auditory bullae, and smaller suprameatal triangles with more

closely approximating posterior processes.
From a single specimen (MNHN, no. 704) of Meriones shawi, as
known from Guelt es Stel, Algeria, and two specimens (BM, nos.
60.8.25.12 and 60.8.25.13) identified as Meriones robustus Wagner
(^Meriones shawi [Loche, 1867]) from an unknown locality in Algeria,
representatives of M. libycus from Libya and Egypt are noticeably
smaller cranially and have proportionately larger and more inflated
auditory bullae. In color and external measurements, the specimen
from Guelt es Stel is similar to representatives of M. libycus in Libya.
are subtle, being typical of subspecies and not of species. In my

opinion, specimens from coastal Libya, formerly referred to M. shawl,
actually belong to M. libycus, and if M. shawi is a valid species, its
range is limited to Algeria and areas to the south and west.
This species is represented in Libya by two subspecies, Meriones
libycus auralus and Meriones libycus azizi. As presently understood,
itsrange in Libya is limited to the Mediterranean littoral as far inland
as the coastal escarpment. Apparently the range of the nominate
subspecies does not include Libya. Specimens from Sidi Barrani,
Salum, and Mersa Matruh of western coastal Egypt, although geo-
graphically not far removed from the type locality of M. I. azizi, are
clearly referable to M. I. libycus.
Meriones libycus auratus, new subspecies

Gheminez, Cyrenaica Province, Libya; obtained Oct. 29, 1955, by
H. W. Setzer, original no. 2672.
Specimens examined. Sixteen, from Cyrenaica: Gheminez, 9;
5 km W
El Agheila, 2; from Tripolitania: 12 km Zliten, 3 (1 skullW
only; 1 skin only); 40 km ENE
Nalut, 2.
Measurements. Averages and extremes of four adult males and
three adult females from the type locality, with the measurements of
the type in brackets, are, respectively: Total length 273.5 (269-282),
266.5 (260-273), [269]; length of tail 138 (135-143), 133.5 (132-135),
[137]; length of hind foot 34 (33-35), 32.7 (32-34), [35]; length of ear
16.8 (16-18), 16 (15-17), [16]; greatest length of skull 37.5 (36.5-38.2),
36.5 (36.2-36.8), [37.1]; length of palate 16.5 (16.3-16.7), 16.2 (16.1-
16.3), [16.3]; length of audital portion of auditory bulla 13.3 (12.8-
13.4), 12.6 (12.3-12.8), [13]; alveolar length of upper molariform
toothrow 5.6 (5.4-5.7), 5.5 (5.3-5.6), [5.6]; least interorbital breadth
6.2 (5.7-6.7), 5.9 (5.7-6.2), [6.1]; length of nasals 14.3 (13.7-14.9),
13.7 (13 6-13.9), breadth of rostrum at level of antorbital
[14.9];
foramina 3.6 (3.5-3.8), 3.6 (3.4-3.7), [3.6]; greatest breadth across
zygomatic arches 20.5 (19.8-21.4), 19.9 (19.5-20.3), [20.5].
Diagnosis. Upperparts ranging from Buckthorn Brown to Clay
Color, becoming paler on sides and approaching Cinnamon-Buff; all of
dorsum, including tail, evenly suffused with brownish-tipped hairs;
postauricular patches conspicuous and almost pure white; supraorbital
and preauricular patches conspicuous and almost white with strong
admixture of black-tipped hairs; distinct white patch present on
rostrum immediately behind origin of vibrissae; eye ring black; pinna
of ear relatively short, remarkably uniform in color, but slightly darker
distally and almost devoid of hair, except for a distinct row of pos-
teriorly directed buffy hairs on anterolateral margin; vibrissae fine in
texture, formed of both brown and white hairs, and extending caudad
beyond the level of the ears; scapular regions, upper arms, and pectoral
areas with moderate suffusions of buff forefeet white dorsally, almost
;
naked ventrally, and with five digits bearing claws; hind feet white
dorsally and with five digits bearing dark-colored claws; plantar sur-
faces heavily furred except for a small naked area on the ventral
surface of the heel; tail, dorsally, approaching color of body and
heavily suffused with brown-tipped hairs, and ventrally near Pale
Pinkish Buff; a distinct brownish pencil occupying the distal one-
fourth of the dorsal surface of the tail. Skull: Relatively small in size,
auditory bullae short and well inflated ventrally; anterior palatine
foramina relatively narrow meatal expansion of auditory bullae with
;
abrupt curvature and not applied to the squamous process of the

temporal bone; suprameatal triangle rounded and with posterior
processes imperfectly closed.
Comparisons. The type series of Meriones libycus auratus can be
easily distinguished from near topotypes of Meriones libycus libycus
from the West Desert region of Egypt by markedly smaller and less
robust crania, smaller overall size, especially the small size of the hind
feet, and noticeably paler, more golden dorsal color.
Members Meriones
of this subspecies resemble rather closely those of
libycus azizi from farther north and east in Cyrenaica but differ in
having larger hind feet, wider interorbital breadths and rostra, and
paler, more golden dorsal pelage with more prominent postauricular
and supraorbital patches.
Remarks. Representatives of this subspecies throughout coastal
Libya are remarkably uniform in color and cranial characters. Two
specimens, 321835 and 321836, from 40 kilometers east-northeast of
Nalut on the Tripolitanian coastal plain are almost indistinguishable
from those from the type locality in Cyrenaica, and specimens from
El Agheila on the southern margin of the Gulf of Sirte do not differ
appreciably from topotypes of M. I. auratus. This morphological
homogeneity is the result of constant genetic exchange within and
between populations of these jirds rendered possible by the almost
uninterrupted continuity of suitable habitat of the Cyrenaican and
Tripolitanian coastal plains.
Setzer (1956) referred two specimens from El Agheila and a single
specimen from near Zliten to Meriones libycus confalonierii (= Meri-
ones caudatus confalonierii). These three specimens are specifically
different from the other specimens found in these localities and clearly
belong to Meriones libycus auratus. Regardless of their taxonomic
status at that time this erroneous assignment is unaccountable as the
two forms are easily separable: in the field, by the more bushy tail
with prominent black pencil of M. caudatus and in the laboratory, by
the noticeably larger and more inflated auditory bullae and complete
closure of the suprameatal triangles of M. caudatus.
Ecological observations. These jirds apparently prefer areas of
low elevation on the coastal plain and are unknown from the higher
Saharan interior where they are supplanted by the bushy-tailed
Meriones caudatus.
At Nalut, two specimens, 321835 and 321836, were obtained from
some large dunes near the coastal escarpment. These dunes supported
a sparse vegetative cover of Calligonum and thorny perennials. Two
specimens of M. caudatus and a large series of Gerbillus aureus were
taken from the same trapline. According to Setzer (1957), the habitat
at Gheminez and Zliten consisted of poorly vegetated rocky-clay type
of soil and fallow barley fields.
Although this subspecies is known from comparatively few speci-
mens and has been collected from only four localities, it doubtless
has a wide distribution and is probably sympatric with practically
all other species of coastal rodents throughout the Mediterranean
littoral of Libya. From the vicinity of El Agheila, it has been taken

with Gerbillus eatoni, G. gerbillus, G. pyramidum, G. amoenus, G.
henleyi, Pachyuromys duprasi, Meriones caudatus, and Psammomys
obesus. At Gheminez, Jaculus orientalis, Allactaga, tetradactyla, and
Gerbillus aureus occur with M. I. auratus.
The subspecific name auratus is derived from the Latin "aurum,"
meaning gold or of gold, and has reference to the golden cast of the
dorsal pelage in members of this subspecies.
Meriones libycus azizi Setzer
Meriones shawi azizi Setzer, Proc. Biol. Soc. Wash., vol. 69, pp. 205-206, Dec.
31, 1956 (5 km SE Derna, Cyrenaica, Libya).
Specimens examined. Twenty, from Cyrenaica: 3 km E. Derna,

1; 5 km SE Derna, 2; 7 km E Maraua, 1 2 km N Coefia, 5; 8 km
;
N Benghazi, 2; 20 km E Tobruch, 9.
Measurements. Measurements of two adult females, 302243 and

325604, from 5 km SE Derna and 20 kilometers east of Tobruch
are, respectively: Total length 259, 252; length of tail 131, 123,
length of hind foot 32, 31; length of ear 17, 19; greatest length of
skull 36.8, 36.4; length of palate 16.4, 16.5; length of audital portion
of auditory bulla 12.3, 12.8; alveolar length of upper molariform
toothrow 5.6, 5.7; least interorbital breadth 6.1, 5.7; length of nasals ?,
13.7; breadth of rostrum at level of antorbital foramina 3.2, 3.7;
greatest breadth across zygomatic arches 20.4, 20.6.
Diagnosis. Upperparts ranging from Snuff Brown to Buffy Brown
becoming paler on sides and flanks and approaching Tawny-Olive;
all of dorsum strongly suffused with blackish-brown hairs resulting
285-134 O — 68 13
in a uniformly variegated appearance; circumoral and rostral areas

grading from Pinkish Buff to Cinnamon-Buff; area between ears
and eyes washed with gray and appearing as a faint patch above the
eye; scapular and pectoral areas, chin and upper surfaces of front
legs and feet suffused with varying amounts of Pinkish Buff and
Light Pinkish Cinnamon; forefeet sparsely haired ventrally with
five digits bearing dark-colored claws. Hind feet Pale Pinkish Buff
dorsally, richly haired ventrally except for a bare area near the
heel, and with five digits with dark-colored claws; pinna of ear mod-
erately haired, becoming darker distally and approaching Saccardo's
Umber, with a conspicuous row of buffy hairs along entire antero-
lateral surface; eye ring black; vibrissae formed of both black and
white hairs and projecting backward to the level of the ears; tail
Cinnamon and appearing somewhat bicolored owing to strong ad-
mixtures of blackish hairs dorsally and with a distinct black pencil
confined to the dorsal surface for about the distal one-fourth. Skull:
Small in size; auditory bullae short and expanded ventrally; zygomatic
arches wide; anterior palatine foramina wide; suprameatal triangles
round in shape with posterior processes imperfectly closed.
Comparisons. From near topotypes of Meriones libycus libycus
from various localities in western coastal Egypt, the type specimen
and a paratype of Meriones libycus azizi have significantly smaller
cranial and external measurements and shorter and more inflated
auditory bullae. In color, the two subspecies are quite similar.
For comparison with Meriones libycus auratus, see account of that
subspecies.
Remarks. The specimens from near Coefia and Benghazi are all
immature, but having distinct postauricular spots and slightly

in
paler dorsal color, they suggest intergradation with M. I. auratus
to the south and west.
An adult female, 325604, from 20 kilometers east of Tobruch is
noticeably paler and more grayish in dorsal color and has a more
reduced pencil than the two specimens from the type locality. In
these characters, this specimen resembles neither typical M. I. libycus
nor M. I. azizi, but because of its small cranial and body size and the
geographic position of the collecting locality, it is here included with
the latter subspecies.
With the exception of a single specimen from near Maraua, this
subspeciesis known only from the Mediterranean littoral of northern
Cyrenaica. In these coastal areas, members of this subspecies occur

with Meriones caudatus, Jaculus orientalis, Jaculus deserti, Gerbillus
eatoni, and several species of dipodil gerbils.
Genus Psammomys Cretzschmar

In the past, the taxonomy of the genus Psammomys has been almost
constantly in a state of confusion. Thomas (1902) was the first to
report on representatives of this genus from Libya and described
Psammomys tripolitanus as a new species based on specimens from
Bou Cheifa, Wadi Aggar, and Wadi Cheggar of northern Tripolitania.
He assigned specimens, which were much smaller in size and paler
in color, from Bu Ngem and the Wadi Wagis, Tripolitania, to Psam-
momys roudairei Lataste. The latter species was known from only two
immature types (type specimen and cotype) and had previously been
regarded by Lataste (1887) as a synonym of Psammomys obesus
Cretzschmar. Many years later, Thomas (1925) after examining more
specimens from near the type locality of P. roudairei in Algeria, once
again placed P. roudairei in the synonymy of P. obesus and proposed
the name Psammomys vexillaris for the small, pale-colored specimens
from Bu Ngem and the Wadi Wagis in Libya.
Since that time, P. tripolitanus and P. vexillaris have been variously
regarded as full species (de Beaux, 1932; Zavattari, 1934) or either
one or the other regarded as subspecies of Psammomys obesus (Eller-

man, 1941; Ellerman and Morrison-Scott, 1951; Toschi, 1954).
Ellerman and Morrison-Scott, as well as Toschi, considered P. tri-
politanus as being synonymous with P. o. obesus and accorded P.
vexillaris only subspecific rank under this species. More recently
Setzer (1957), after examining two specimens from near El Agheila
and Gheminez, Cyrenaica, reinstated P. tripolitanus as a subspecies
of P. obesus and regarded P. vexillaris as a full species, based upon a
single specimen from Wadi Bey about 45 kilometers west of Bu Ngem,
Tripolitania. He considered the two forms as specifically distinct on
the basis of the following characters of the skull of P. vexillaris:
Supraorbital bead absent, temporal ridges markedly reduced, and
molars relatively as well as absolutely larger than in P. o. obesus.
In addition to the above characters established by Setzer, this
specimen from near Bu Ngem differs strikingly from topotypes of
Psammomys obesus obesus from the Nile Delta in its much smaller
and more gracile skull, proportionately larger
overall size, less robust
suprameatal triangles, more ventrally inflated auditory bullae, and
less developed parietal crests and postorbital processes. In color and
external characters, P. vexillaris differs markedly in having a much
shorter tail with a grayish rather than blackish
less extensive pencil,
pinnae, greater suffusion white on venter, white rather than
of
strongly buff-colored dorsal surfaces of hind feet, and much paler,
more subdued, dorsal color.
.
I was unsuccessful in obtaining additional specimens of P. vexillaris,

but after reexamination of the single specimen collected by Setzer
from the Wadi Bey, its specific status is unequivocal in my opinion.
Psammomys obesus Cretzschmar
Psammomys obesus Cretzschmar, Riippell Atlas, 58, pi. 22, 1828 (near Alexandria,
Egypt)
General distribution of species. Israel, Jordan, Saudi Arabia,

Egypt, Sudan, Libya, and Algeria.
Cyrenaica and Tripolitania. The range in Libya is doubtless much
wider than the few collecting records indicate.
Psammomys obesus obesus. Cyrenaica: Coastal plain of north-
eastern Cyrenaica.
Psammomys obesus tripolitanus. Cyrenaica and Tripolitania:
Coastal plain and littoral deserts of the Gulf of Sirte.
Published records in Libya. Cyrenaica: Bou Cheifa (Thomas,

1902 [Psammomys tripolitanus]); El Agheila (de Beaux, 1932 [Psam-
Psammomys obesus
I. obesus
2 tripolitanus
Psammomys vexillaris
3. vexillaris
Figure 41. — Distribution of Psammomys obesus and Psammomys vexillaris.

.
momys tripolitanus]) Zauia Mechili (de Beaux, 1938 [Psammomys

;
tripolitanus]); Tripolitania: Wadi Aggar, Wadi Cheggar (Thomas,

1902 [Psammomys tripolitanus]).
Comparisons. Psammomys obesus can be readily distinguished from
Psammomys vexillaris by its darker color and greater size in all di-
mensions.
Remarks. Superficially, members of this genus resemble those of
the genus Merion.es but upon closer scrutiny can be easily identified
by their plain, rather than distinctly grooved, upper incisors and
generally heavier, more angular skulls. Furthermore, representatives
of the genus Psammomys are primarily diurnal, whereas jirds (Meriones)
tend to be almost exclusively nocturnal in their habits.
Specimens of Psammomys obesus obesus from coastal Libya east
of the Cyrenaican Plateau do not differ appreciably from those
representing Psammomys obesus obesus from farther east in Egypt.
The few specimens from the coastal plain along the Gulf of Sirte,
however, differ significantly from representatives of P. o. obesus
from Egypt and are here referred to Psamrnomys obesus tripolitanus.
Ecological observations. In Libya, these large rodents are
limited in distribution to a rather narrow belt of alluvial soils and
abundant vegetation, which occurs along the coastal plain and
occasionally farther inland in portions of the transitional deserts.
At present, specimens are not available from the coastal plain of
Tripolitania, but doubtless this species occurs here also. These rodents
have a similar distribution in Egypt.
All specimens I collected were from the more alluvial soils of the
shoulders of the road. As presently understood, these animals are
exclusively diurnal in their habits, emerging from their burrows
shortly after dawn, remaining active somewhat irregularly throughout
the day, and retiring to their burrows shortly after sunset. They dig
large burrows with conspicuous, well-worn entrances, usually marked
by piles of freshly deposited talus covered with seeds and bits of
local plants. They seem to prefer the loose soils of the elevated roadbed
or nearby large hillocks with a dense vegetative cover.
Frequently, during the daytime, they are seen scurrying across the
road or running about the mounds while foraging. Diurnal traplines
were surprisingly ineffective and only rarely were fully adult animals
caught. On one occasion east of Tobruch, these diurnal traps were left
in position throughout the night and produced three jirds (Meriones)
The occurrence of these jirds in the same trapline suggests that mem-
bers of these two genera occur together in the same burrow complexes.
The jirds, however, forage at night and the sand rats (Psammomys)
feed only during the hours of daylight.
;
Other species of rodents occurring in these more lush habitats of

the Libyan coastal plain include Gerbillus henleyi, Gerbillus campestris,
Gerbillus eatoni, Jaculus orientalis, Spalax ehrenbergi, and Gerbillus
kaiseri.
Psammomys obesus obesus Cretzschmar
Psammomys obesus Cretzschmar, Riippell Atlas, 58, pi. 22, 1828 (near Alexan-
dria, Egypt).
Specimens examined. Sixteen, from Cyrenaica: 20 km E To-

bruch, 13 (1 skin only) ; 5 km W Bardia, 3.
Measurements. Averages and extremes of six adult females and

the measurements of one adult male, 325646, from 20 kilometers east
of Tobruch, are, respectively: Total length 285.8 (271-300), 288;
length of tail 122.5 (110-128), 110; length of hind foot 35.2 (35-36),
38; length of ear 14.8 (14-17), 15; greatest length of skull 42.2 (40.4-
43.5), 40.5; length of auditory bulla 14.3 (13.5-14.6), 13.6; alveolar
length of upper molariform toothrow 7 (6.5-7.6), 6.7; least inter-
orbital breadth 6.7 (6.3-7), 6.8; length of nasals 16.3 (15.3-17.6),
16.3; breadth of rostrum at level of antorbital foramina 4.3 (4-4.6),
4.5; greatest breadth across zygomatic arches 24.9 (23.8-25.5), 23.6;
least breadth between parietal ridges 10.9 (10.5-11.2), 10.5.
Diagnosis. Pelage of dorsum dense and long but in old specimens
heavily worn, particularly in middorsal region where denuded patches
often disclose plumbeous-colored basal portions of hairs; sides
varying from Pale Orange- Yellow to Light Orange- Yellow and
flanks becoming darker in middorsal region approaching Ochraceous-
Tawny; all parts of dorsum interspersed with brownish hairs; thus
entire upperparts formed from a variety of colors and color patterns
resulting in a pronounced variegated appearance; area between the
ears and above the eyes, however, more uniformly colored Cinnamon
or Ochraceous-Tawny circumorbital areas Light Ochraceous-Buff
;
heavily suffused with black; this color pattern extending anteriorly

along the rostrum and posteriorly to the level of the ears circumoral ;
and mystacial areas and dorsal surfaces of fore and hind feet and
legs ranging from Warm Buff to Antimony Yellow, the latter being
more concentrated in the scapular regions, which appear more brightly
colored than the surrounding pelage; ears relatively short, medial
surface of pinna with uniform covering of buffy hairs; anterior margin
of pinna with prominent row of rather stiff hairs, which project
backward into inner surface of the pinna; vibrissae stiff and relatively
short, formed of about equal numbers of black and buff-colored hairs
palmar surfaces of front feet naked and bearing distinct metacarpal
tubercles; four primary digits (2-5) with evenly spaced transverse
ridgesand terminating in well-developed claws; first digit rudimentary
without functional claw; plantar surfaces of hind feet with irregular
covering of hairs; all five digits well defined, sparsely haired, and all
with functional claws; entire underparts strongly suffused with
colors varying from Warm Buff and Ochraceous-Buff to Antimony
Yellow; in all specimens an irregular whitish patch is present in the
pectoral region; tail relatively short and same color as the dorsum
except for some faint admixtures of brownish-colored hairs dorsally;
a prominent pencil, varying in color from Cinnamon-Brown to Prout's
Brown, occupying the distal one-fourth of the dorsal and ventral
surfaces. Skull: Size relatively small for the genus but markedly
angular and robust; zygomata extremely strong and heavy, and in
older specimens, markedly bowed posteriorly; rostrum wide and
heavy; anterior palatine foramina relatively short and broad; posterior
palatine canals indistinct; molariform teeth heavy, grinding surfaces
usually evenly worn; pterygoid processes heavy, with distinctly
knobbed hamulae; auditory bulla small, not markedly inflated
ventrally, but with greatly expanded meatal process, which almost
touches squamous portion of temporal bone; suprameatal triangle
conspicuous, usually completely enclosed by enveloping processes of
temporal and supraoccipital bones; parietals with prominent lateral
ridges extending from lambdoidal ridge to postorbital processes of
frontal bone.
Comparisons. The specimens from near Tobruch and Bardia,
Cyrenaica, differ from representatives of Psammomys obesus nicolli
Thomas from the Nile Delta and from those of Psammomys obesus
terraesanctae Thomas from Palestine in being markedly smaller in all
measurements.
For comparisons of these specimens from Tobruch and Bardia with
representatives of P. o. tripolitanus see account of that subspecies.
,
Remarks. The specimens from Tobruch and Bardia are clearly

referable to Psammomys obesus obesus. Except for their more brilliant
yellowish-orange dorsal color and more worn-looking dorsal pelage,
they are almost indistinguishable from specimens representing P. o.
obesus from Salum, El Alamein, and other coastal localities to the east.
These differences in color are well within the expected range of indi-
vidual variation within a given population.
Setzer (1957) assigned specimens of P. obesus from El Alamein,
Western Desert Governorate, Egypt, to P. o. tripolitanus. In a later
paper, Setzer (1963) regarded these specimens from western Egypt as
intergrades between /'. o. tripolitanus and P. o. obesus but more closely
related to the latter subspecies. The specimens from Tobruch and
Bardia, Cyrenaica, in their slightly smaller size also suggest inter-
gradation with P. o. tripolitanus. It is doubtful that genetic exchange
is common between populations of P. o. tripolitanus and P. o. obesus
because of the reduction or absence of coastal plain in many parts of
northern Cyrenaica.
.
Psammomys obesus tripolitanus Thomas

Psammomys tripolitanus Thomas, Proc. Zool. Soc. London, pt. 2, p. 9, 1902 (Bon
Cheifa, on the coast, Cyrenaica, Libya).
Specimens examined.Six, from Cyrenaica: Gheminez, 1; 20 km

SW Agedabia, 1S Agedabia 3; 5 km
; 10 km El Agheila, 1. W
Measurements. Measurements of an adult female, 325630, from
20 kilometers southwest of Agedabia, are: Total length 300; length of
tail 127; length of hind foot 34; length of ear 14; greatest length of
skull 41.3; length of auditory bulla 14.7; alveolar length of upper

molariform toothrow 7; least interorbital breadth 6.9; length of
nasals 15.7; breadth of rostrum at level of antorbital foramina 4.5;
greatest breadth across zygomatic arches 24.7; least breadth between
parietal ridges 1 1
Diagnosis. Same as for P. o. obesus, except dorsum with stronger

suffusion of brownish hairs, and skull smaller and more gracile.
Comparisons. Compared to representatives of P. o. obesus from
near Tobruch and Bardia, Cyrenaica, and near topotypes of P. o.
obesus from Burg el Arab and El Daba, Western Desert Governorate,
and Damanhur and Hofs, Beheira Province, Egypt, the few specimens
from the coastal plain of the Gulf of Sirte are smaller in overall size
of skull, have wider interorbital breadths, wider rostra, shorter
anterior palatine foramina, and are darker in dorsal color.
Because the range of P. o. obesus is interposed between the ranges
of P. o. tripolitanus and those of Psammomys obesus nicolli from the
Nile Delta and Psammomys obesus terraesanctae from Palestine, critical
comparison with the two latter subspecies was not undertaken, but
from cursory examination both P. o. nicolli and P. o. terraesanctae
appear to be significantly larger both cranially and in external size
and dimensions. Psammomys obesus nicolli is also darker in color,
whereas P. o. terraesanctae is noticeably paler and more uniformly
colored than P. o. tripolitanus.
Remarks. Topotypes of P. o. tripolitanus are not available for study,
but because the specimens from Gheminez and the vicinity of Agedabia
agree closely with the description and measurements of P. o. tripolitanus
as given by Thomas (1902), and because of geographic proximity,
they are here referred to this subspecies.
Psammomys vexillaris vexillaris Thomas

Psammomys vexillaris Thomas, Ann. Mag. Nat. Hist., ser. 9, vol. 16, p. 198,
July 1925 (Bondjem [=Bu Ngem], Tripolitania).
General distribution of species. Libya and Algeria.
Distribution in Libya. At present, this species is known from only
the vicinity of the type locality. The range probably includes most of
the coastal plain and the transitional deserts of Tripolitania.
.
Specimens examined. One, 302254, from the Wadi Bey, 45 km

W Bu Ngem, Tripolitania, Libya.
Published records. Tripolitania: Bu Ngem, Wadi Wagis
(Thomas, 1902 [Psammornys roudairei])
Measurements. The measurements of an old male, 302254, from
near Bu Ngem, are: Total length [224]; length of tail [93]; length of
hind foot 32; length of ear 12; greatest length of skull 35; alveolar
length of upper molariform toothrow 6.1; least interorbital breadth
6.2; length of nasals 13.1; least breadth between parietal ridges 9.5.
Diagnosis. Upperparts varied in color owing to suffusions of various
shades of buff, gray, brown, yellow and orange; a broad, indistinct
Pinkish Cinnamon band extending from the rostrum to the rump;
fine guard hairs projecting beyond the underlying hairs on the dorsum;
sides paler in color than dorsum and with greater admixture of dark
hairs; preauricular and suborbital areas grading from Light Buff to
Warm Brown and strongly suffused with brownish-black hairs; cir-
cumoral and mystacial areas and region between chin and pectoral
girdle white with moderate suffusion of Light Buff; postauricular
patches indistinct, same color as surrounding pelage; pinnae of ears
heavily furred with pale, Light Buff hairs, some of which partially
cover inner aspects of pinnae. Vibrissae relatively short, formed from
both black and white hairs, and extending posteriorly about 8 milli-
meters beyond the ears; scapular areas more brilliantly colored than
surrounding areas and appearing as vaguely defined patches of Warm
Buff and Light Ochraceous-Buff; this same color imparted to the
pectoral region below, but more subdued here; entire underparts
predominantly white, but irregularly interspersed with patches of
Light Buff, Pale Ochraceous-Buff, and Warm Buff; upperparts of
front legs and dorsal surfaces of forefeet Light Buff; palmar surfaces
almost naked; hind feet almost pure white dorsally; sparsely haired
ventrally with conspicuous naked areas on proximal one-half of plantar
surfaces; forelegs with five digits with claws, the first digit being almost
rudimentary; hindlegs with five functional digits bearing well-devel-
oped claws; tail relatively short, Warm Buff, and unicolorous except
for a rather distinct pencil which occupies the distal one-fifth of the
dorsal surface. Noticeably small and gracile; suprameatal
Skull:
triangles relatively large and completely enclosed by enveloping
processes of the temporal and supraoccipital bones; auditory bullae
large and inflated ventrally; postorbital processes and parietal crests
poorly developed; anterior palatine foramina relatively long; molari-
form teeth large and wide; supraorbital bead absent.
Remarks. Originally, Thomas (1925) recognized two subspecies of
Psammornys vexillaris. The range of the nominate subspecies was
thought to include the coastal and interior deserts of Tripolitania,
Libya, and Psammomys vexillaris edusa, with type locality at Mil

Mahases, Chegga, south of Biskra, Algeria, was believed to be confined
to the Algerian Sahara. Ellerman and Morrison-Scott (1951) regarded
P. v. synonym of Psammomys obesus vexillaris, the species
edusa as a
same time being relegated to subspecific rank under
P. vexillaris at the
P. obesus. Psammomys vexillaris is here accorded full specific rank,
based primarily upon its smaller size, both cranially and in external
dimensions, and upon additional characters as given by Thomas
(1925), Setzer (1957), and the present author. Because I have examined
no specimens from Algeria, I cannot comment on the status of P. v.
edusa.
In Libya, this small sand rat is known only from the type and
type series as reported by Thomas (1925) and from a single old male,
302254, obtained in 1955 by Setzer from the Wadi Bey near the type
locality. According to Setzer (1957), this animal was dug from its
burrow which was located in an area of consolidated sand with sparse
vegetation. Myefforts to obtain additional specimens of this rare
sand rat were to no avail.
Family Spalacidae
Genus Spalax Guldenstaedt

Spalax ehrenbergi aegyptiacus Nehring
Spalax aegyptiacus Nehring, S. B. Ges. Nat. Fr., Berlin, p. 180, 1898 (Ramleh,
near Alexandria, Egypt).
General distribution of species. Syria, Israel, Egypt, and Libya.

Distribution in Libya. Cyrenaican Plateau, including the Gebel
Achdar and the coastal plain of Cyrenaica.
Specimens examined. Seven, from Cyrenaica: 5 km Labrag, NW
3; Wadi el Kuf, 13 km WSW
Messa, 1; 8 km N Benghazi, 1; 4 km S
Agedabia, 1; 20 km SW Agedabia, 1.
Published records in Libya. Cyrenaica: Ras el Ferg (Sordelli,
1899); Barce (Ghigi, 1920); Benghazi (Zavattari, 1934; Toschi, 1951).

Measurements. Measurements of two adult females, 325655 and
325656, from 5 kilometers northwest of Labrag and the measure-
ments of an adult female, 325652, from 20 kilometers southwest of
Agedabia, are, respectively: Total length 172, 180, 190; length of tail
15, 15, 16; length of hind foot 21, 22, 23; length of ear 5, 5, 5; con-
dyloincisive length of skull 38.5, 40.7, 39.9; crown length of upper
molariform toothrow 7.4, 7, 7.1; palatilar length 21.8, 22.3, 22.8;
length of anterior palatine foramina 2.9, 3.2, 3.2; greatest breadth
across zygomatic arches 28.8, 29.3, 29.4; least interorbital breadth
6.9, 6.4, 6.7; length of nasals 16.6, 17.8, 17.8; breadth of rostrum at
level of antorbital foramina 7.1, 6.9, 7.4; greatest length of skull
Figure 42. —-Distribution of Spalax ehrenbergi aegyptiacus.
41.2, 43.6, 42.8; length of diastema 14.2, 14.7, 15; basilar length of
skull 34, 35.7, 35.2; greatest breadth of rostrum 7.5, 8, 8.1.
Diagnosis. Most distinctive feature is the featureless, cylindrical
body without a visible tail, and superficially appearing to lack eyes,
ears and nostrils; rostrum furnished with a hardened and slightly
enlarged cartilaginous plate beneath the skin and near the origin of
the vibrissae, which presumably aids in pushing and moving dirt in
the underground tunnels; vibrissae correspondingly small and fine
in texture; pelage short, dense, and silky and of almost uniform color
dorsally, varying in different specimens, however, among shades of
Ochraceous-Tawny, Sayal Brown, and Buckthorn Brown, and ven-
trally somewhat darker owing to less suffusion of these colors. In
older specimens, pelage becoming worn and somewhat darker owing
to greater exposure of the Plumbeous underfur; rostrum, in young
specimens, light gray in color, but, in old specimens, rostral, auricular,
and entire facial areas becoming a much darker gray; eyes not visible
externally, but present as small rudimentary structures in the usual
position in the orbit; pinnae of the ears, on close examination, present
as inconspicuous cartilaginous rings which are obscured by the dense
pelage; forefeet, bearing five digits with claws, remarkably small in

view of the fossorial habits of these rodents; hind feet also relatively
small and with five digits, but with larger claws; fore and hind feet
approaching color of dorsum, but naked ventrally. Skull: Large,
massive, and angular with conspicuous development of the lambdoidal
and sagittal crests; shape of skull almost triangular in outline owing
to the relatively small braincase, the abrupt anterior convergence of
the zygomata and the relatively short and wide rostum; interorbital
breadth markedly constricted; supraoccipital markedly expanded
dorsally, with numerous inflated elevations, and directed a consider-
able distance anteriorly; incisors extremely long, relatively narrow,
and orange in color on their anterior faces molarif orm teeth moderate
;
in size with intricate interwoven patterns in the enamel and dentine,

and with conspicuous styles on the lateral surfaces; infraorbital
foramina large with corresponding reduction in size of the frontal
plate of the premaxillae; dorsal margin of foramen magnum smoothly
rounded; auditory bullae markedly reduced to thick, heavily ossified
structures with prominent external meatal processes and conspicuous,
attenuated styliform processes; anterior palatine foramina markedly
reduced in size; palate with prominent mid ventral rib; posterior
palatine canals small and inconspicuous; pterygoid processes heavy
and with club-shaped hamulae; basioccipital broadly wedge-shaped.
Remarks. Nehring (1897) regarded the mole rats of the Near East
as representing several distinct species. He described Spalax ehrenbergi
from Jaffa, Palestine, Spalax kirgisorum Nehring from northern Syria,
Spalax intermedins Nehring from farther south in Syria, and named
Spalax aegyptiacus Nehring with the type locality at Ramleh, near
Alexandria, Egypt. Later, Mehely (1913) regarded all members of this
group as one species and considered S. kirgisorum and S. aegyptiacus
as varieties of Spalax ehrenbergi. Many years later, Ellerman (1941)
elevated S. kirgisorum to full species status on the basis of page priority
in the original description and relegated £. ehrenbergi and S. aegyptia-
cus to subspecific rank, thus reversing the taxonomic status of S.
ehrenbergi and S. kirgisorum. A short time later, Bate (1945) observed
that, although the description of S. kirgisorum clearly preceded that of
S. ehrenbergi, the latter form is given a prior place in illustrations of
cheek teeth, and is followed by S. kirgisorum. She stated that in works
published prior to 1931, figures are acceptable as valid descriptions,
and therefore the name Spalax ehrenbergi should be regarded as ante-
dating that of Spalax kirgisorum.
More recently, Ellerman and Morrison-Scott (1951) consider S.
ehrenbergi as a valid species but regard S. kirgisorum as a synonym of
S. ehrenbergi ehrenbergi. Thus, the species S. ehrenbergi is currently
composed of two subspecies, S. e. ehrenbergi and S. e. aegyptiacus.
Toschi (1954) assigned specimens from Ras el Ferg, Barce and Beng-
hazi, Cyrenaica Province, Libya, to S. e. aegyptiacus. Although Setzer
(1957) saw evidences of mole rats at Gheminez, Derna, Barce, and
near Agedabia, he obtained no specimens.
In the present study, I obtained seven specimens from four localities
in Cyrenaica. Although they are from widely separated localities which
differ rather sharply in characteristics of soil, vegetative cover, and
climate, they are quite similar morphologically. They differ noticeably,
however, from representatives of S. e. aegyptiacus from Burg el Arab,
Western Desert Governorate, Egypt, in their smaller size of most ex-
ternal and cranial measurements, narrower rostra, which are more
constricted at the level of the infraorbital foramina, more delicate
zygomata, broader frontal plates of the premaxillae, larger infraorbital
foramina, smaller, narrower, and shorter incisors, and more smoothly
rounded dorsal margin of the foramen magnum. In the specimens from
Cyrenaica, the styliform process of the auditory bulla is more attenu-
ated and projects farther medially, and the angular process of the
mandible is smaller and more knobbed.
The above differences would normally be sufficient to warrant con-
sidering these Libyan mole rats as belonging to a distinct subspecies,
but too few specimens are available for accurate designation. Further-
more, they agree rather closely with the cranial measurements given
for Spalax aegyptiacus in the original description.
Despite the foregoing differences, I consider it best, until such
time as additional specimens of mole rats are forthcoming, to refer all
Libyan specimens to S. e. aegyptiacus.
Ecological observations. Mole rats are common on the coastal
plain and uplands of the Cyrenaican Plateau and are relatively
common farther south and west along the coastal plain of the eastern
portion of the Gulf of Sirte. On the Cyrenaican Plateau and the Gebel
Achdar, they seem to prefer loose soils at the margins and bottoms
of the larger valleysand the more fertile soils of the higher tablelands.
Mounds were observed on the rocky slopes north of El Faidia
also
near the highest point of the Cyrenaican Plateau.
These rodents apparently attain the southwesternmost limits of
distribution for the species on the coastal plain south and west of
Agedabia. Currently they are unknown from the Tripolitanian
Mole rats occur inland for varying distances, probably
coastal plain.
depending upon the amount of rainfall for a particular year. While
en route to Gialo in the spring of 1962, we saw fresh diggings 25
kilometers inland from the Mediterranean coast. Setzer (in verbis)
also attests to their presence in these inland localities. It is almost

certain that they do not occur in or near the oases of the Saharan
interior.
These remarkable rodents plug the entrances to their burrows with

soil,and they are rarely observed above ground. On several occasions,
movements of the plug or fresh talus were observed during the day-
time, indicating that they are active both day and night. Specimens
were obtained with the use of Macabee gopher traps but only after
exhaustive efforts because of the unusually hard, resistant nature of
the soil. The small diameter of the entrance of the burrow, which
was almost always difficult to locate, and the insistence of these
mole rats on pushing fresh dirt over the trap, made trapping difficult.
During the rainy season in the spring, the soils of the coastal plain
are relatively soft and easily workable, and diggings of mole rats are
widespread. Accordingly, traps can be more easily set, and the yield
is greater. Later in the summer, after the soil has hardened, the
number of active burrows is greatly reduced, and in some localities
collecting mole rats is much more difficult, if not impossible. At this
time these soils of become almost impervious.
the coastal plain
It is mole rats occupying the more firmly
interesting to note that
packed soils of the Cyrenaican Plateau are generally smaller than
those living in the softer, more sandy soils of the coastal plain. Selec-
tion has apparently favored smaller body size in these areas where
digging is more difficult and a smaller diameter of the burrows is
desirable. The soils of the Cyrenaican Plateau are probably harder
and less sandy than those of the Egyptian littoral farther east, and
this differential in hardness of the soil may account for the differences
in size ofmole rats from the two areas.
The character of the soil seems to be the most critical factor in
the distribution of these burrowing rodents, and vegetative cover and
other physical features of the terrain are of secondary importance.
One specimen, 325652, from near Agedabia was taken from a series
of active burrows in the extremely hard-packed soil near the roadside
on a grassy portion of the coastal plain. These grassy areas, most of
which contained diggings, were interspersed among the rather dense,
shrubby vegetative cover of the coastal plain. Another specimen,
325651, was obtained farther inland where the coastal plain is broad
and featureless and has a more uniform vegetative cover. This speci-
men was obtained during the spring when the recently emerged
grasses were lush and green, and the soil, which normally would have
been almost impervious, was soft and workable. Near Labrag, three
specimens were obtained from the high uplands of the escarpment
which forms the northern terminus of the Cyrenaican Plateau. Even
though fresh diggings were widespread, the soil here was extremely
hard and rocky, and the specimens were obtained only after laborious
efforts. A single specimen, 325657, was obtained from the more alluvial
soils in the bottom of the Wadi el Kuf where the ground was also
nearly impervious.
According to Setzer (1957) and Hartert (1923), the local Arabs

believe that because mole rats lack visible eyes and are apparently
blind, anyone having contact with them will also become blind.
Family Muridae
Genus Rattus Fischer

Rattus norvegicus (Berkenhont)
Mus norvegicus Berkenhout, Outlines N. H. Great Britain and Ireland, vol. 1,
p. 5 (Great Britain).
Remarks. In Libya, the Norway rat is known only from Tripoli

(Toschi, 1951), although probably occurs in most, if not all, of the
it
larger coastal cities and may occur in the wild state in the Cyrenaican
Plateau.
They are thought to be endemic to northern China and Manchuria.
In recent centuries, they have been carried westward as a result of
the increased development of western commerce. Their introduction
into North Africa most likely was through the agency of shipborne
rats in the major ports where ships frequently unloaded their produce.
K^^
.
According to Rode (1948), the Norway rat is abundant in all ports

of North Africa and occurs also in the large villages of the interior.
It is doubtful that they occur in the Saharan interior of Libya. These
rats are known to have a definite predilection for moist habitats and
frequently inhabit stream banks where they demonstrate marked
abilities for swimming and burrowing. Even in the tropics, they sur-
vive only where the habitat has been significantly altered by man.
Moist habitats in some of the larger oases of Cyrenaica and the
Fezzan may provide the ecological requirements of these rats, but
these areas are sporadic and localized. In addition, the barriers
imposed by vast areas of desolate sand and arid desert must be sur-
mounted before they could become established in these interior oases.
In the present study, I collected extensively for several months in
the interior oases and made frequent inquiries regarding the local
rodent fauna. During this period I was unsuccessful in obtaining any
rats (R. rattus or R. norvegicus) and observed no indications of their
presence.
Rattus rattus (Linnaeus)
Mus rattus Linnaeus, Syst. Nat., 10th ed., vol. 1, p. 61 (Sweden).
General distribution of species. Nearly cosmopolitan owing to

its commensal relationship with man; it is said to occur in the wild
state in most of India, West Pakistan, Ceylon, foothills of the Hima-
layan Mountains, Southern China, Burma, Thailand, Laos, Viet Nam,
Malaya, and in the larger islands of the southwestern Pacific, including
the Philippine Islands.
Distribution in Libya. Occurs commensally with man in the
and Derna and apparently
larger coastal cities of Benghazi, Tripoli,
is present in the wild state in parts of coastal northern Cyrenaica.
Specimens examined. Four, from Cyrenaica: 27 km E Apollonia,
3; 12 km NW
Gubba, 1.
Published records in Libya. Cyrenaica: Benghazi (Festa, 1925
[Epimys tectorum Savi]) Derna, Merg ( = Barce) (de Beaux, 1938
;
[Rattus rattus nericola Cabrera]); Tripolitania: Tripoli (Toschi, 1951

[Rattus rattus jrugivorus Rafinesque])
Measurements. Measurements of an adult female, 325658, from
27 kilometers east of Apollonia, are: Total length 343; length of tail
192; length of hind foot 33; length of ear 24; greatest length of skull
39.7; condylobasal length of skull 38.2; alveolar length of upper mo-
lariform toothrow 6.9; greatest breadth across zygomatic arches 19;
least interorbital breadth 5.5; length of nasals 14.5; length of audital
portion of auditory bulla 7; palatal length 21.7; breadth of rostrum
at level of antorbital foramina 4 length of anterior palatine foramina 7.
;
Diagnosis. Upperparts varying among shades of Prout's Brown,

Snuff Brown and Sepia; a large portion of the hairs of the shoulders,
Comparisons. Rattus can be distinguished from Rattus

rattus
norvegicus by more uniformly
the following characters: Longer and
colored tail; one pair of postaxillary, pectoral mammae, rather than
two pairs; smaller front feet; lyre-shaped temporal ridges, as opposed
to parallel temporal ridges; first molar lacking the small accessory
cusp, the latter being present in R. norvegicus.
Remarks. Earlier workers in the systematics of rodents have
applied subspecific names to the three common color types of the
commensal or black These color types are charac-
rat (Rattus rattus).
terized as: black or gray above and below; brown above and dark
gray below; brown above and white below; and correspond, respec-
tively, to the subspecies R. r. rattus (Linnaeus), R. r. alexandrinus
(GeofFroy), and R. r. frugivorus (Rafinesque). In the past, the com-
mensal rats of the larger coastal towns of Libya have been variously
assigned to one or more of these subspecies. More recently, ecological,
behavioral, and genetic studies indicate that these so-called subspecies
do not conform to the modern concept of subspecies and accordingly
should be regarded only as color phases or genetic variants within
the species.
Caslick (1955), in an attempt to resolve some of the relationships of
these color types in Rattus rattus, conducted a series of selective
matings of wild-trapped black rats of mixed origin, and although, in
color, the Utter mates satisfied the requirements of subspecies and
"unclassifiable intergrades" were of rare occurrence, he concluded that
the current forms of R. rattus were best regarded as color phases within
the species.
Setzer (1952) also recognized these same three color types in
commensal rats from the Nile Delta and noted that specimens from a
given locality segregated out as the three expected color types plus
intermediate forms. These intermediate forms, to which he attributed
a single gene character for melanism, were suggestive of intergrades,
but the presence of three distinct subspecies occupying the same en-
vironmental niche (or geographic area) was clearly untenable. Other
workers have frequently reported black rats of different color phases
sharing the same habitat, and the individuals in a given population
that did not conform to one of the so-called subspecies were considered
intergrades.
Johnson (1946) commented on a Utter from the same nest that con-
tained representatives of both the white-belUed and black-beUied color
phases and stated that these color phases had no morphological
and that the ecological differences
distinctions, except those of color,
mentioned by other workers were not evident in the population that
he studied.
CasUck (1955) suggests that the color phases of the commensal rat,
recognized as the subspecies R. r. rattus, R. r. alexandrinus, and R. r.
frugivorus, may be similar to the well-known color phases of the black

bear, red fox, and the gray and fox squirrels.
No differences have been observed in the habits, movements, and
survival differential of R. r. rattus and R. r. alexandrinus, although
R. r. frugivorus has been considered by some to represent a "wild form"
which frequently occurs in the feral state in fields and agricultural
areas. In many cities and villages of Europe, North America, and
North Africa, all three of these color types occur together.
In my opinion, the recognition of rattus, alexandrinus, and frugivorus
as subspecies of R. rattus is of doubtful utility, and I prefer to regard
these so-called subspecies as merely color phases or phenotypic
variants within this widely ranging species.
one subscribes to the older system of applying subspecific names
If
to thecommensal rats of Europe and North Africa, the four specimens
from Cyrenaica clearly represent R. r. frugivorus.
The four specimens from Cyrenaica were trapped in the lush
vegetation bordering small permanent streams. The collecting site
near Gubba is located in a deep canyon of the coastal escarpment and
several kilometers from the nearest human habitation, whereas the
specimens from near Apollonia were taken from an elevated portion of
the coastal plain in traps set only a few hundred meters distant from
human dwellings. These specimens from Cyrenaica constitute the
first records of R. rattus occurring in the wild state in Libya. They are
unknown from the oases of the interior.
Genus Mus Linnaeus

Mus musculus Linnaeus
Mus musculus Linnaeus, Syst. Nat., 10th ed., vol. 1, p. 62, 1758 (Upsala, Sweden).
General distribution of species. Cosmopolitan as a result of

introduction by man.
Distribution in Libya. Occurs as a commensal with man in the
larger coastal citiesand villages of the interior and is quite widely
distributed in the wild state.
Specimens examined. One hundred fifty-eight, from Cyrenaica:
27 km E Apollonia, 11; 11 km SW Susa (= Apollonia), 3; 5 km NW
Labrag, 6; 12 km NW
Gubba, 2; Cyrene, 1; 3 km E Derna, 2; 35
km W
Messa, 1; 10 km SW
El Faidia, 3; 7 km E Maraua, 1; 5 km
W Tocra, 49 (1 skin only); 20 kin SW Tocra, 3; Giarabub, 2; El
Hauuari, Cufra Oasis, 1 El Giof, Cufra Oasis, 12; from Tripolitania:
;
5 km W
Cussabat, 3; 12 km S Chicla, 2; 7 km S El Gheddahia, 1;
20 km E Sirte, 1 (skin only); from Fezzan: Brach, 37 (10 skull only);
Sebha, 1 Goddua, 11 (7 skull only, 2 skin only) Meseguin, 2; Traghen,
; ;
1; 28 km E Murzuch, 1; Murzuch, 1.
204 U.S. NATIONAL MUSEUM BULLETIN" 2 75
Figure 45. — Distribution of Mus musculus. Circles indicate specimens examined; triangle
indicates published record.
Published records in Libya. Cyrenaica: Benghazi, Derna (Klap-

tocz, 1909 [Mus musculus orientalis Cretzschmar]); Benghazi, Ghe-
minez (Festa, 1921 [Mus musculus gentilis Brants]); Benghazi (Har-
tert, 1923); Giarabub (de Beaux, 1928 [Mus musculus musculus and
M. m. orientalis]) Augila, Cufra and Gialo (de Beaux, 1932 [M. m.
;
orientalis}) Derna, Wadi el Kuf (M. m. orientalis) and Merg (Mus

;
musculus brevirostris) (de Beaux, 1938); Tripolitania: Tarhuna

(Thomas, 1902 [M. m. orientalis]); Tripoli (Klaptocz, 1909 [M. m.
Fezzan: Idri, Murzuch (Toschi, 1951 [M. m. gentilis]).
orientalis]);
Measurements. Averages and extremes of 17 males and 8 females
from Brach, Fezzan Province, are, respectively: Total length 164.1
(149-182), 158.6 (156-163); length of tail 82.9 (76-90), 78.3 (75-81);
length of hind foot 19 (18-20), 18.8 (18-19); length of ear 16.6 (14-
21), 15.4 (14-20); greatest length of skull 22 (20.7-23.2), 21.3 (20.7-
22.2); condyloincisive length of skull 21.1 (19.6-22.2), 20.4 (19.8-
21.3); crown length upper molariform toothrow 3.7 (3.3-4), 3.7
of
(3.6-3.9); greatest breadth across zygomatic arches 11.5 (10.9-11.9),
11.3 (10.9-11.6); least interorbital breadth 3.6 (3.5-3.8), 3.7 (3.5-3.8);
length of nasals 8.2 (7.5-8.6), 7.8 (7.1-8.4).
Diagnosis. Two distinct color types of Mus musculus occur in

Libya, a light-bellied form and a dark-bellied form. In the light-
bellied form, the hairs of the venter are either white throughout or
white distally with plumbeous bases. The dark-bellied form has gray
or brown-tipped hairs on the venter, which are always plumbeous
basally. The color of the dorsal and ventral pelage is markedly con-
trasting in the white-bellied form, whereas, in the dark-bellied form,
the color of both surfaces is quite similar. The two forms differ ap-
preciably in the color of the dorsum, that of the dark-bellied form
being markedly and more uniformly darker (ranging from Saccardo's
Umber to Sepia) in the light-bellied form, it is more variable and
;
ranges from Clay Color to Olive Brown and frequently is variegated

owing to suffusions of gray and buff. The dorsal surfaces of the hind
feet are dusky or gray in the dark-bellied form, whereas they are
almost white in the light-bellied form. In both forms, the tails are
naked except for a scanty covering of inconspicouus whitish hairs.
Cranially, the two types do not differ significantly and both have
extremely small, gracile skulls with flattened braincases, short rostra
(more pronounced in the white-bellied form), large anterior palatine
foramina, small posterior palatine canals, relatively small cheek teeth,
broad basioccipitals, and small, bulbous auditory bullae. The "Mus
notch" is pronounced on the upper incisors.
Remarks. The systematic status of the house mouse (Mus
musculus) has never been firmly established. The principal effort to
resolve the nomenclatural relationships of Mus musculus is that of
Schwarz and Schwarz (1943), who considered the wild species, Mus
musculus, to originally consist of four subspecies, M. m. wagneri
Eversmann, M. m. M.
m. manchu Thomas, and
spicilegus Pet6nyi,
M. m. spretus Lataste. According to the above authors, these wild
forms (except for M. m. spretus, which occurs exclusively in the wild
state) have given rise to the various commensal forms of the house
mouse throughout the world. Schwarz and Schwarz contend that most
of these commensal forms, except those of western Europe, Russia,
and Japan, have been derived from M. m. wagneri, which developed
both an eastern and western series of commensals. The commensal
Mus musculus of Libya supposedly arose from the western component
of M. m. wagneri, which had its origin in southwestern Persia ( = Iran)
and from which it spread, by the agency of man and cultivation,
through Iraq, Turkey, Syria, Israel, the Nile Valley, and westward
across North Africa. From Africa these commensal forms have reached
Italy, Spain, and western Europe.
Throughout most of their range, depending upon the degree of
specialization, these commensals form two types, a more specialized
"commensal" from and a more generalized "wild" form. The com-
mensal form, according to Schwarz and Schwarz, has evolved along

specific lines in response to its more intimate association with man
and normally has a longer tail (frequently longer than the length of
the head and body), a darker (more grayish) venter, richer (darker)
shades of color dorsally, a shorter face, and smaller molariform teeth.
In the wild form, the tail is shorter (always shorter than the length
of the head and body), the venter is white, rather than gray, and
forms a sharper contrast with the color of the pelage of the sides and
flanks. These authors contend that, in many areas, both indoor and
outdoor types of house mice are found. The outdoor (feral) types may
represent either wild or commensal types, which are less specialized
than the indoor types.
The house mice of Libya, according to Schwarz and Schwarz, are
represented by two subspecies, M. m. praeiextus Brants and M. m.
brevirostris Waterhouse, representing, respectively, the wild and
commensal forms.
Toschi (1954, pp. 264-265) summarized the work of all previous
workers on Libyan mammals and listed four subspecies of Mus mus-
culus as occurring in Libya. These included M. m. musculus, M. m.
brevirostris, M. m. gentilis (=M. m. praetextus), and M. m. orientalis.
Both M. m. gentilis and M. m. musculus were listed from Benghazi
and apparently represented, respectively, the feral and commensal
forms of M. musculus. These subspecies mentioned by Toschi were
based on specimens from widley scattered localities in Libya, and
many were assigned to a given subspecies without regard to distribu-
tional or taxonomic concepts.
In the present study, many more specimens are available from
widely scattered localities in Libya, and two distinct color types are
discernible. Those from all interior localities represent the light-bellied
form and are white ventrally with the individual hairs white through-
out or plumbeous-colored basally. Dorsally, the coloration varies
from extremely pale individuals to quite dark-colored forms. In all
specimens from the interior of Libya, the dorsum and venter are
markedly contrasted in color. Specimens from the coastal plain near
Tocra in Cyrenaica typify the dark-bellied form in which the venter
is gray or brown and thus not contrasting with the color of the dorsum.
Light-bellied individuals also occur near Tocra but are less abundant.
Setzer (1957) obtained both feral and commensal house mice in
Libya and was able easily to distinguish two types on the basis of
their dorsal and ventral coloration. He found, based on color, that
the commensal kind agreed with the description of the feral form
given by Schwarz and Schwarz, and that the feral kind agreed with
the description of the commensal form as given by the above authors.
He further stated that these color types did not represent subspecies
as understood by modern taxonomists.
All Libyan specimens I collectedwere taken in the wild or feral

state and, except for those from the Cyrenaican coastal plain, conform
in the pattern of their coloration to the wild form M. m. praetextus
as established by Schwarz and Schwarz. The dark-bellied specimens
from the coastal plain clearly represent the commensal form, M. m.
brevirostris. These coastal specimens have shorter rostra than those
from other localities, which is a character used by Schwarz and
Schwarz to separate M. m. brevirostris from M. m. praetextus, but
they lack the long tail which is reputed to be a character of the
commensal form. Furthermore, both color types occur together on
the coastal plain near Tocra, and neither show commensal tendencies.
To me, these different color types are only color variants of the same
subspecies. On the coastal plain near Tocra, and doubtless in many
other localities in Libya, both color types (or subspecies) occur
together; this clearly violates the concept that "no two subspecies
occupy the same geographic range." For these reasons, I prefer to
assign all of these mice occurring in Libya to the species AIus musculus
and suggest that the different color types be interpreted as various
stages in commensalism rather than as distinct subspecies as used
in the context of modern taxonomists.
Ecological observations. In Libya, house mice are widely
distributed and occur in a wide variety of habitats. Frequently,
they act as strict commensals sharing the same dwellings as the
local people. At some localities, they inhabit gardens and palm groves
of the larger villages and oases. In many areas, they exist in the wild
or feral state, completely divorced from the nearest influence of man.
The largest series were taken on the coastal plain or from the larger
interior oases. At Brach, in the Fezzan, house mice are abundant
in the interior of the oasis where slightly elevated patches of Phrag-
mites are interspersed among areas of open water. They are common
inhabitants of the dense pockets of sedges and other mesophytic
plants that encircle the saline lakes at El Giof and El Hauuari at
Cufra Oasis. On the coastal plain near Tocra, Cyrenaica, a large
series was taken from the dense, brush-covered coastal plain near
the sea. In other parts of Cyrenaica, they occur in the dense, meso-
phytic plant cover bordering small permanent streams or inhabit
localized mesic pockets in the coastal escarpment. They were occa-
sionally trapped in the chaparral vegetation of the higher portions
of the Cyrenaican Plateau.
Relatively high humidity, mesic conditions, and dense plant cover
are the features common to all the above habitats. Only rarely are
these commensal mice taken from dry situations, and they never
inhabit soils formed exclusively of sand. Perhaps these dry habitats
are marginal and the collection of house mice from them was entirely
fortuitous.
It is interesting to note that apparently house mice are absent from

several of the large oases of southern Cyrenaica, such as Tazerbo and
Bzema. Suitable habitat occurs in these isolated oases, but, unlike
Cufra Oasis, they are far removed from the caravan routes and have
relatively few outside visitors; thus, the introduction of commensals
has probably never taken place.
Genus Acomys Geoffroy

Acomys cahirinus viator Thomas
Acomys viator Thomas, Proc. Zool. Soc. London, pt. 2, p. 10, 1902 (Wadi Sultan,
near Socna, Tripolitania, Libya).
General distribution of species. Libya, Egypt, and Algeria, and

range probably includes additional countries and provinces of the
Sahara.
Distribution in Libya. This species is known from only the type
and from the oases of El
locality near Socna, Tripolitania Province,
Barcat, Fezzan Province, and El Giof (Cufra Oasis), Cyrenaica
Province.
;
Specimens examined. Twenty-three, from Cyrenaica: El Giof,

Cufra Oasis, 19; from Tripolitania: 5 km S Socna, 1; Bir Fergian,
10 km S Socna, 1 (skin only); from Fezzan: El Barcat, 2.
adult females from El Giof, Cufra Oasis, are, respectively: Total
length 202 (195-209), 203.1 (185-229) length of tail 104.3 (100-107),
;
104.3 (99-115); length of hind foot 19.4 (19-20), 19.4 (19-20) length ;
of ear 19.6 (19-20), 19 (19-19); greatest length of skull 29.3 (28.4-

30.1), 28.7 (27.3-30); condyloincisive length of skull 27.3 (26.7-28),
26.5 (25-27.6); crown length of upper molariform toothrow 4.2 (4.1-
4.3), 4.2 (4-4.5); breadth of rostrum at level of antorbital foramina
3.2 (3.1-3.3), 3.2 (3.1-3.3); least interorbital breadth 4.8 (4.7-5), 4.8
(4.6-5); greatest breadth across zygomatic arches 14.4 (14-14.9), 14
(13.3-14.7); median length of nasals 10.5 (10.1-10.8), 10.3 (9.7-10.6).
Diagnosis. Upperparts ranging from Smoke Gray on scapular
areas and sides to Deep Grayish Olive on middorsal region and rump
pelage of rump coarse and hispid owing to individual hairs modified
into stiff bristles and spines; total dorsum mildly suffused with tones
of Light Drab; base of pinna surrounded by band of distinct buffy-
gray hairs; pinna large, sparsely haired, and ranging in color from
Deep Grayish Olive to Dark Grayish Olive; chin, mystacial, and
circumoral areas, forelegs, hindlegs, dorsal surfaces of fore and hind
feet, and entire underparts pure white; ventral surfaces of fore and
hind feet naked, with distinct palmar and plantar pads, and bearing
five digits with claws; vibrissae relatively long, fine in texture, and
formed largely of grayish hairs; tail fusiform, conspicuously bi-
colored, Clove Brown above and Deep Olive-Buff below, and composed
throughout its entire length of successive, imbricated segments, from
the bases of which arise short, stiff, white hairs. Skull: Medium in
expanded dorsally; anterior palatine foramina
size; parietals slightly
long and wide; molariform teeth relatively small and toothrow short;
pterygoid fossae broad and long and with prominent elliptical fo-
ramina on laterocaudal margins; auditory bullae small and noticeably
inflated ventrally; pterygoid processes convergent anteriorly and
forming a distinct rib separating the halves of the pterygoid fossae
and extending to the level of the palate.
Comparisons. From specimens representing Acomys cahirinus
cahirinus (Geoffroy) from various localities both east and west of the
Nile River, Egypt, specimens from El Giof, Cufra Oasis, differ in
slightly more robust skulls, particularly the rostrum and zygomata,
slightly wider braincases, more expanded parietals, and a larger fora-
men on the laterocaudal portion of the pterygoid fossa. The foramen
magnum, in specimens from El Giof, is more rounded dorsally and
extends farther dorsally into the supraoccipital bone. In external

measurements the populations from Egypt and Libya are comparable,
but, in color, the Libyan specimens are more uniform, whereas the
Egyptian specimens show a wide range of color, divisible generally
into two distinct color phases.
The Libyan specimens differ significantly from representatives of
Acomys dimidiatus dimidiatus (Cretzschmar) form Feiran Oasis, Sinai,
in their smaller, more gracile skulls, more flattened braincases, and
narrower anterior palatine foramina. The foramen magnum, in the
Libyan specimens, is much more rounded dorsally and the foramina in
the pterygoid fossae are appreciably larger. In color, they are much
darker, gray as opposed to buff, and more uniformly colored.
In cranial measurements, specimens from Cufra resemble those of
Acomys dimidiatus hunteri de Winton from the Eastern Desert
Governorate of Egypt and the Sudan, but differ markedly from the
latter in their darker color, smaller size, prominently rounded dorsal
surface of foramen magnum (as opposed to markedly flattened or
truncated in A. c. hunteri), and shorter anterior palatine foramina.
Remarks. The series of spiny mice from Cufra Oasis is here referred
to Acomys cahirinus viator but with reservations. In color, specimens
from Cufra closely resemble two near topotypes of A. c. viator from
near Socna and Bir Fergian, Tripolitania. They are slightly smaller in
overall length but are of comparable size in all other external dimen-
sions. Unfortunately, the skull of the adult specimen from Bir Fergian
was lost, and the other specimen from near Socna is extremely young;
thus, cranial comparisons are not possible. Compared to measurements
of the type specimen of A. c. viator, as given by Thomas (1902) in the
original description, specimens from Cufra are slightly smaller in
greatest length of skull and have slightly wider interorbital breadths.
It seems unlikely that the same subspecies would occur in such
widely separated geographic areas in Libya,such as Socna, El
Barcat, and Cufra, particularly since suitable habitat is sporadic and
localized. The referral of these specimens to A. c. viator must therefore
be considered provisional until such time as additional specimens of
A. c. viator are forthcoming and permit more accurate comparisons.
Spiny mice from Cufra represent geographic isolates, clearly distinct
morphologically from all other populations to the east, and probably
will prove also to be subspecifically distinct from A. c. viator.
In the past, A. cahirinus and A. dimidiatus have been variously
regarded as either distinct species or the latter treated as a subspecies
of A. cahirinus. Ellerman and Morrison-Scott (1951) regarded the
smaller A. cahirinus purely as a commensal form and relegated the
largerand exclusively wild form (A. dimidiatus) to subspecific status
under A. cahirinus. Setzer (1959) recognized two species of spiny
mice as occurring in Egypt and Sinai; he considered A. cahirinus
as a strict commensal but regarded it as a monotypic species with a

range confined to the Nile Valley and distinguishable from A. dimi-
diatus by more flattened braincase, more rounded foramen magnum,
more nearly vertical (less recurved) upper incisors, and broader and
less elongated pterygoid fossae.
The above differences are apparent when spiny mice from Sinai and
Baluchistan, Iran, are compared to those from the Nile Delta, but
specimens from Libya do not show these same differences, and there
is too much overlap of characters among populations of these mice
from Libya and the Nile Valley to suggest specific differences. I agree
with Setzer that the "cahirinus group" is divisible into two species,
A. cahirinus and A. dimidiatus, but extend the range of A. cahirinus
to include Libya, as well as the Nile Valley of Egypt, and limit the
range of A. dimidiatus to areas to the south, east, and north. For the
present, until a thorough revisionary work of this poorly understood
genus has been undertaken, I prefer to include the spiny mice of
Libya and the commensal forms of the Nile Valley of Egypt within
a single species, Acomys cahirinus. Commensalism is unknown in the
Libyan populations, but this is not a valid criterion for specific
designation and could have arisen independently in the populations
of spiny mice in the Nile Valley.
Cranially, two specimens from El Barcat in the extreme south-
western Fezzan resemble those from Cufra but differ in having more
flattened and wider braincases, shorter anterior palatine foramina,
wider rostra, and shorter nasals. In color, they more closely resemble
the two specimens from Socna but are slightly more grayish dorsally
and are darker on the ventral surface of the tail. It is apparent that
these two specimens from the Fezzan represent a population of spiny
mice differing significantly from others in Libya, For the present,
however, until larger series are obtained from the vicinity of El Barcat
and Ghat, in the Fezzan, and more topotypes of A. c. viator become
available from Tripolitania, these two specimens are provisionally
referred to A. c. viator largely on the grounds of geographic occurrence.
I have not examined specimens of Acomys cahirinus seurati Heim de
Balsac, the type locality of which is Iniker, in the Ahaggar Mountains

of southern Algeria, but the spiny mice from the Fezzan may represent
this subspecies.
Ecological observations. Spiny mice were obtained from areas
with dead fronds and debris, near the bases of
of loose sand, littered
date palms.
According to Thomas (1902, p. 11), spiny mice from the Wadi
Sultan near Socna resembled in color "the small blackish stones
which lie about among the Soda Mountains." This statement sug-
gests that the habitat in the Wadi Sultan was primarily of a rocky
character.
It is possible that these mice occur more abundantly in rocky

environments and that the oases may represent marginal habitats.
Rocky areas are not uncommon in Libya, but owing to their seemingly
desolate and barren appearance, trapping efforts in them are minimal
and may account for the sporadic representation of this species in
Libya.
According to Setzer this species is almost exclusively
(1959),
commensal throughout range in the Nile Valley and Delta. In
its
Libya, spiny mice apparently have not developed this intimate

relationship with man, although frequently gerbils (Gerbillus) and
jirds (Meriones) were purchased from the local inhabitants of the
oases and had presumably been living commensally with them.
Family Gliridae
Genus Eliomys Wagner
Eliomys quercinus (Linnaeus)
Mus quercinus Linnaeus, Syst. Nat., 12th ed., vol. 1, p. 84, 1766 (Germany).
General distribution of Europe, western U.S.S.R.,

species.
Sicily, Corsica, Turkey, Saudi Arabia,
Sardinia, Balearic Islands,
Syria, Israel, Sinai, and North Africa, including Egypt, Libya,
Algeria, Morocco, and Spanish Sahara (Rio de Oro).
Distribution in Libya. Northern Cyrenaica, northwestern Tripol-
itania,and the Fezzan.
Eliomys quercinus cyrenaicus. Cyrenaica: Cyrenaican Plateau
and adjacent coastal plain.
Eliomys quercinus denticulatus. Fezzan: Large oases and adjoining
areas.
Eliomys quercinus tunetae. Tripolitania: Tripolitanian Gebel.
Published records in Libya. Cyrenaica: Gheminez (Festa,
1921); Benghazi (Hartert, 1923); Tripolitania: Gharian (Klaptocz,
1909 [Eliomys munbyanus tunetae Thomas]); "Gebel Tripolitano"
(Toschi, 1951 [Eliomys munbyanus munbyanus Pomel]).
Remarks. Owing the relatively few specimens available to
to
workers in the past, a variety of names has been applied to the
dormice of western Libya and Algeria, and the systematic position
of this group has never been firmly established. Thomas (1903)
described Eliomys lerotinus tunetae from Karouana, Tunisia, and
stated that it might prove to grade into the earlier described Eliomys
munbyanus (Pomel) of Morocco and western Algeria. A few years
later, Klaptocz (1909) confirmed Thomas' suspicion and regarded
E. I. tunetae as a subspecies of E. munbianus {=E. munbyanus),
based on specimens from Gharian, Tripolitania. These specimens
Eliomys quercinus
I. cyrenaicus
2. denticulotus
3. tunetoe
Figure 47. — Distribution of the subspecies of Eliomys quercinus. Circles indicate specimens
examined; triangle indicates published record.
from northern Tripolitania constituted the first record of occurrence

of dormice in Libya.
Since that time, E. m. tunetae has been regarded as a subspecies
of E. lerotinus (Lataste) (Zavattari, 1934), regarded as a subspecies
of E. munbyanus (Ellerman, 1941), or considered as a pure synonym
of E. m. munbyanus (Cabrera, 1932; and Allen, 1939). Toschi (1951)
was in agreement with Cabrera and Allen and assigned specimens
from the Tripolitanian Gebel to E. m. munbyanus. Ellerman and
Morrison-Scott (1951) included all dormice of the genus Eliomys,
exclusive of Eliomys melanurus (Wagner) of Egypt and the Middle
East, as a single species, Eliomys quercinus, and regarded E. m.
tunetae as a synonym of Eliomys quercinus munbyanus. Toschi (1954)
and Setzer (1957), although they examined no actual specimens,
considered the dormice of western Tripolitania to represent E. q.
munbyanus.
The systematica of the dormice of Cyrenaica have undergone less
confusion in the literature. Specimens from Gheminez were originally
described as a full species, Eliomys cyrenaicus (Festa, 1921). Later
E. cyrenaicus was relegated to subspecific rank under Eliomys lerotinus
(Zavattari, 1934), regarded as a subspecies of E. munbyanus (Allen,

1939), reinstated as a full species, E. cyrenaicus (Ellerman, 1941),
and more recently regarded as a subspecies of E. quercinus (Ellerman
and Morrison-Scott, 1951; Toschi, 1954; Setzer, 1957).
The most recent attempt to clarify the nomenclature of dormice
of the genus Eliomys is that of Niethammer (1959), in which he
united all members of the genus into a single species, Eliomys quercinus,
which he divided into the "quercinus," "lusitanicus," and "melanurus"
groups, based entirely upon the pattern of coloration on the ventral
surface of the tail. He recognized two subspecies of E. quercinus
in Libya, E. q. cyrenaicus of northern Cyrenaica, and E. q. tunetae
of northwestern Tripolitania, the latter again being considered as
distinct from E. q. munbyanus. These two subspecies, in addition
to E. q. melanurus of coastal Egypt, Sinai, and the Middle East,
constitute the "melanurus" group. The "quercinus" group is distrib-
uted over most of Europe, western U.S.S.R., and extends into
North Africa including Morocco and western Algeria. The range of
the "lusitanicus" group includes southern Spain and Italy and
several of the larger islands in the Mediterranean.
In the present work, I have followed Neithammer's classification
by including all Libyan dormice within the "melanurus" group but
have extended appreciably the range of E. quercinus to include
the Fezzan.
Eliomys quercinus cyrenaicus Festa
Eliomys cyrenaicus Festa, Boll. Mus. Zool. Anat. Comp. Univ. Torino, vol. 36,
no. 740, p. 4, December 1921 (Gheminez, Cyrenaica).
Specimens examined. One, from 5 km SE Derna, Cyrenaica

and skeleton).
(skin, skull
Measurements. The measurements of the above specimen, a
subadult female, 302274, are: Total length 211; length of tail 100;
length of hind foot 27; length of ear 25; greatest length of skull
31.9; condyloincisive length of skull 29; length of audital portion of
auditory bulla 9.9; crown length of upper molariform toothrow 5.4;
least interorbital breadth 4.2; greatest breadth across zygomatic
arches 17.9; length of nasals 11.4; breadth of braincase 14. G.
Diagnosis. Upperparts Avellaneous with strong suffusion of gray,
becoming paler, more grayish, on sides; dorsal portion of rostrum
and interauricular area grading from Buckthorn Brown to Tawny-
Olive; cheeks, mystacial, subauricular, and scapular areas Pale
Pinkish Buff; area between eye and origin of vibrissae, entire eye
ring, and a broad band passing from eye to base of the pinna of the
ear, black with mild suffusion of gray; postauricular patches indis-
tinct; pinna of ear moderate in size, sparsely haired, approaching
the color of the pelage of the dorsum, and with a conspicuous tuft
of buffy hairs on antero ventral margin; vibrissae relatively long

with white and black individual hairs; fore and hind feet relatively
small with four and five functional digits respectively, white above
and naked below, with prominent palmar and plantar pads; tail
medium in length and with dense covering of grayish-brown hairs
proximally and of black hairs distally both above and below, the
latter color present throughout terminal four-fifths of the length.
Skull: Small and gracile; rostrum narrow and attenuated; inter-
orbital region markedly constricted; parietals noticeably inflated;
interparietal expanded laterally; zygomata bowing slightly laterally;
auditory bullae conspicuously inflated ventrally; basioccipital broad
and triangular; pterygoid processes fragile; hamular processes applied
to anteromedial surfaces of the auditory bullae; pterygoid region
with numerous large foramina; molariform teeth large, with two
prominent cusps on the labial surfaces; anterior palatine foramina
large and laterally expanded posteriorly; mandible with a small
oval foramen in the angular process.
Comparisons. From Eliomys quercinus melanurus Wagner as known
from the vicinity of St. Catherine's Monastery, Sinai, the specimen,
302274, from five kilometers southeast of Derna, differs in much smaller
size, both cranially and in external dimensions, relatively larger and
wider molariform teeth, smaller size of the auditory bullae, and
smaller size of the foramen on the angular process of the mandible.
In color, this specimen from Cyrenaica closely resembles representa-
tives of E. q. melanurus but is slightly darker dorsally. The pinnae
of the ears of the specimen from Derna are much smaller and in this
respect differ strikingly from those of the representatives of E. q.
melanurus.
The specimen from Derna differs markedly from a representative
(BM, no. 22.5.30.45) of Eliomys quercinus munbyanus from Tagouidert
(Haha), Morocco, in having more subdued dorsal color, ventral sur-
face of the tail black rather than white, much larger molariform teeth,
and a smaller foramen in the angular process of the lower jaw.

According to Festa (1921), Eliomys cyrenaicus (= Eliomys quercinus
cyrenaicus) differs from Eliomys lerotinus tunetae (= Eliomys quercinus
tunetae) primarily by larger body and larger auditory bullae.
For comparison of this specimen from Derna with those representing
Eliomys quercinus denticulatus from the Fezzan, see the account of
E. q. denticulatus.
Remarks. This specimen from near Derna, a specimen from Beng-
hazi (Hartert, 1923), and the type series of E. q. cyrenaicus from
Gheminez constitute the only records of occurrence of dormice in
Cyrenaica. Dormice are unknown from farther south in the interior
of Cyrenaica, although suitable habitat is present in the larger oases,
;
such as Gialo, Tazerbo, and Cufra. Also they apparently do not occur
in the desert regions of Egypt farther east. The desolate areas of the
Serir of Calanscio and the immense dunes of the Sand Sea of Calanscio
in Libya and comparable areas of desolation in the Western Desert
of Egypt apparently have proven to be insurmountable barriers to
the dispersal of dormice, both at present and in the past.
The exact type locality of E. cyrenaicus is in doubt. Festa (1921)
did not state whether the type series was taken from the local gardens
of Gheminez or from the nearby Gebel. A dried up dormouse was
found in the gardens of Benghazi by Hartert (1923), and since these
gardens are also common at Gheminez, it seems most likely that the
specimens came from within, or very near, the village. Setzer (1957)
considered it most likely that the type series was collected from the
massif some 40 kilometers east of Gheminez, particularly since dor-
mice are unknown to the local residents of Gheminez.
According to Setzer (1957), the specimen from near Derna was
live-trapped from the upper slopes of a large wadi near the brink of
the coastal escarpment. The vegetative cover at this site consisted of
bushlike chaparral, and the dormouse was taken from a trap set near
the base of a large evergreen bush.
In the present work, traps were frequently set throughout these
chaparral habitats on the Cyrenaican Plateau, but we were unsuccess-
ful in obtaining additional dormice and observed no indications of their
presence.
Eliomys quercinus denticulatus, new subspecies
Holotype. Adult female, skin and skull, USNM 322757, from El

Gatrun, Fezzan Province, Libya, obtained Jan. 11, 1962, by G. L.
Specimens examined. Six, from Fezzan: Temenhint, 3 (immature);
Goddua, 1 (skin, skull and skeleton) 28 km E Murzuch 1 (immature)
;
El Gatrun, 1 (skin, skull and skeleton).

Measurements. Measurements of an adult male from Goddua,
322758, and of the type specimen (in brackets) from El Gatrun, are
respectively: Total length 200, [213]; length of tail 96, [115]; length
of hind foot 27, [25] length of ear 24, [24] greatest length of skull 30.7,
;
;
[31.2]; condyloincisive length of skull 27.7, [28.3]; length of audital

portion of auditory bulla 10.1, [10.5]; crown length of upper molari-
form toothrow 4.7, [4.5]; least interorbital breadth 4.3, [4.4]; greatest
breadth across zygomatic arches 18.1, [17.6]; length of nasals 10.6,
[11.2]; breadth of braincase 14.8, [14.7].
Diagnosis. Upperparts Avellaneous, heavily washed with gray,
becoming paler on sides and approaching Light Grayish Olive; dorsal
part of rostrum and interauricular area more brightly colored than
dorsum near Light Pinkish Cinnamon and Cinnamon-Buff; cheeks,
mystacial, subauricular, and scapular areas nearly pure white and

sharply contrasting with black areas around eyes and base of ear;
postauricular patches inconspicuous and Pale Pinkish Cinnamon
strongly suffused with gray; pinna of ear moderate in size, sparsely
haired, approaching color of the dorsum, and with a conspicuous tuft
of buffy hairs on the anteroventral margin; vibrissae relatively long
with white and black individual hairs; fore and hind feet relatively
small with four and five functional digits respectively, white above
and naked below with prominent palmar and plantar pads, tail relative-
ly long with three distinct zones of color on both surfaces; a proximal
region of grayish-brown, a more extensive middle region of black, and
a terminal tuft of white. Skull: Small and gracile; rostrum narrow
and attenuated; interorbital region markedly constricted; parietals
prominently domed; zygomata bowing slightly laterally; auditory
bullae conspicuously inflated ventrally; basioccipital broad and tri-
angular; pterygiod region with numerous large foramina; molariform
teeth small, with two prominent cusps on the labial surfaces; anterior
palatine foramina large and laterally expanded posteriorly; mandible
with large oval foramen in the angular process.
Comparisons. From representatives of Eliomys quercinus melanurus
Wagner from the Gebel Musa and near St. Catherine's Monastery,
Sinai, the type specimen of Eliomys quercinus denticulatus from El
Gatrun, Fezzan, differs in markedly smaller size, particularly the ears
which are strikingly smaller; more domed braincase; much shorter
molariform too throw and smaller individual teeth; longer inter-
parietal and a more truncated dorsal margin of the foramen magnum;
dorsum slightly darker with greater suffusion of light brownish hairs,
especially in the interauricular and rostral regions; postauricular
patches more conspicuous; tail white terminally, rather than black,
and darker proximally owing to a greater duffusion of brown. In E. q.
denticulatus, there is a more gradual transition from the brownish-
gray color of the base of the tail to the solid black of the middle and
distal portions, whereas in E. q. melanurus the grayish color of the
base of the transforms abruptly into black.
tail
Compared to a representative of Eliomys quercinus cyrenaicus Festa

from five kilometers southeast of Derna, Cyrenaica, the type specimen
of Eliomys quercinus denticulatus from El Gatrun differs in having
skull smaller, interparietal longer, braincase more domed, rostrum
narrower and more attenuated, anterior palatine foramina slightly
wider, foramen in the angular process of the mandible larger, molar-
iform too throws markedly shorter, and with smaller individual teeth.
These two subspecies are comparable in external dimensions except
for the longer tail of E. q. denticulatus, which is white terminally,
rather than black as in E. q. cyrenaicus. They are also similar in dorsal
285-134 O— 68 15
color, but E. q. denticulatus is slightly paler in the interauricular and

rostral areas and has less suffusion of buff on the cheeks and scapular
areas.
The type specimen of E. q. denticulatus from El Gatrun can be
readily distinguished from a representative of Eliomys quercinus
munbyanus from Tagouidert (Haha), Morocco, by much more sub-
dued dorsal color (Avellaneous as opposed to Tawny or Kusset) and
paler facial region with less black enveloping the base of the pinna
of the ear. In E. q. denticulatus the ventral surface of the tail is black
except for a terminal white whereas in E. q. munbyanus it is
tuft,
white throughout its entire length. The type specimen of E. q. dentic-
ulatus from El Gatrun is smaller in all external dimensions, except
the length of the and has a more domed braincase, a narrower
tail,
and more attenuated rostrum, slightly more laterally and ventrally

inflated auditory bullae, and shorter molariform toothrows with
smaller individual teeth.
From a specimen of Eliomys quercinus tunetae from Bone, northern
Algeria, the specimen of E. q. denticulatus is more subdued in dorsal
color and is paler in the facial region. Cranial comparisons with this
specimen of E. q. tunetae from Bone, Algeria, are not possible, but
compared with measurements of the type specimen of Eliomys lerotinus
tunetae (= Eliomys munbyanus tunetae) from Karouana, Tunisia, as
given by Thomas (1903) in the original description, the type specimen
of E. q. denticulatus from El Gatrun is slightly smaller in size of skull
and has a markedly shorter molariform toothrow. The slightly smaller
cranial size of the type specimen of E. q. denticulatus is further indi-
cated by comparison of its cranial measurements with those of a
specimen representing E. q. tunetae from Gherran ( = Gharian),
Tripolitania, as reported by Klaptocz (1909).
Remarks. The few specimens available at the present time probably
do not demonstrate the full range of variation in the Fezzanese popu-
lation, but because they differ strikingly in morphological characters
from all other subspecies of E. quercinus, and owing to their isolated
geographic position, they warrant designation as a new subspecies.
The most distinguishing features of this new subspecies of dormouse
are the small molariform teeth, the relatively long tail with the white
tip,and the generally smaller size. The immature specimens from
Temenhint and near Murzuch show these same characters.
These specimens from the Fezzan represent the first records of
dormice from the Saharan interior of Libya. The specimen from El
Gatrun is perhaps the southernmost record for Eliomys quercinus in
the interior of North Africa and closely approaches the southern
limits of distribution for the species on the African continent.
The presence of dormice in the Fezzan is inexplicable in view of

the present physiography and climate of Libya. The various popula-
tions within the Fezzan are separated from neighboring populations
by vast areas of unsuitable habitat, and the Fezzanese population,
as a whole, is geographically and physiographically isolated from
populations of dormice on the Tripolitanian Gebel nearer the coast.
These dormice of the Fezzan probably represent relicts of a former
period in Libya when a more humid climate prevailed and habitats
suitable for dormice were more widespread and thus permitted their
southward dispersal.
Ecological observations. In the Fezzan, dormice seem to be
confined largely to the environs of the oases. At Temenhint and
28 kilometers east of Murzuch, young dormice were taken, along with
gerbils and jerboas, from areas of loose sand covered with fronds at
the bases of young, unpruned date palms, as well as from sandy-clay
soils at the bases of hardened mounds which supported sparse growths
of tamarix. At Goddua, an adult dormouse was trapped from the
midst of a group of young palms into which dunes of considerable
size were encroaching. The specimen from El Gatrun was obtained
from the edges of a large clump of tamarix located several kilometers
from the palm groves of the oasis. In the tamarix zone, an individual
clump is usually separated from others by several hundred meters of
barren sand alternating with hard-packed clay. It is possible that in
this type of habitat dormice are confined to single clumps and utilize
tamarix as their sole source of food. A species of dipodil (Gerbillus
amoenus) was the only other species of rodent obtained from these
outlying concentrations of tamarix.
All specimens were obtained during the winter months when night-
time temperatures frequently dropped below 30° F. These low tempera-
tures may account for the sporadic representation of dormice in the
Fezzan, as they are known to hibernate during prolonged cold periods.
The name denticulatus from the Latin meaning small teeth, refers
,
to the diminutive size of the molariform teeth in members of this

subspecies.
Eliotnys quercinus lunetae Thomas
Eliomys lerotinus tunetae Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 2 p. 495,
May 1903 (Karouana, Tunis).
Remarks. In Libya, the range of this subspecies is apparently
confined to the Tripolitanian Gebel. Klaptocz (1909) assigned speci-
mens from Gharian to Eliomys munbianus (munbyanus) tunetae, and
later,Toschi (1951) referred a specimen from the "Gebel tripolitano"
to Eliomys munbyanus munbyanus. More recently, Neithammer (1959)
reinstated E. m. tunetae as a subspecies of the widely ranging Eliomys
quercinus.
I have not examined these specimens from the Tripolitanian Gebel,
but because the descriptions of these specimens given by Klaptocz

and Toschi agree quite closely with the original description of Eliomys
lerotinus tunetae (= Eliomys quercinus tunetae), and on account of
geographic proximity, they are here referred to E. q. tunetae.
Family Dipodidae
Genus Jaculus Erxleben
Jerboas in Libya show varying amounts of geographic variation

depending upon the species. Jaculus jaculus has undergone the
greatest amount of differentiation associated with geographic dis-
tribution. The other two species, occurring in Libya, Jaculus orientalis
and Jaculus deserti, are less variable morphologically. The latter two
species are represented by samples of relatively small size and, in
some cases, the full range of variation for any given character may
not be accurate. In jerboas, as with gerbils, geographic variation is
usually not manifested by ordinary meristic and quantitative char-
acters, such as external measurements and cranial indices, but is
indicated by more subtle, qualitative differences, which do not
ordinarily lend themselves to measurement or statistical analysis.
The most important of these characters associated with geographic
variation include differences in color, shape and configuration of the
auditory bullae, angularity and massiveness of the skull and zygo-
mata, and changes in the degree of doming of the braincase.
Key to the Jerboas of Libya
1. Ears usually more than 40 mm; four toes on hind foot; rudimentary premolar
present in upper jaw Allactaga tetradactyla
Ears usually less than 40 mm; three toes on hind foot; rudimentary premolar
lacking in upper jaw 2
2. Body size large, hind foot greater than 70 mm; baculum with pair of curved
terminal spines Jaculus orientalis
Body size small, hind foot less than 70 mm; baculum lacking pair of terminal
spines 3
3. Dorsal color dark; two foramina on angular process of mandible; sole of hind
foot and metatarsal area suffused with brownish hairs J. deserti
Dorsal color pale; a single foramen on angular process of mandible; sole of

hind foot and metatarsal area white or buff and lacking suffusion of brownish
hairs J. jaculus
Jaculus deserti (Loche)
Dipus deserti Loche, Explor. Alger. 100, 1867 (Ouargla district, northern Al-
gerian Sahara).
Jaculus jaculus deserti Ellerman, J. R., and T. C. S. Morrison-Scott, Checklist
of Palaearctic and Indian Mammals, 1758-1946. Brit. Mus. (Nat. Hist.),
p. 539, 1951.
General distribution of species. Arabia, Iraq, Israel, Sinai,

Egypt, Libya, and Algeria.
Distribution in Egypt. Sinai, Eastern Desert, and coastal environs
west of the Nile.
Distribution of the subspecies in Egypt.
Jaculus deserti schluteri (Nehring). Sinai: St. Catherine's Monastery
Area; Wadi Raha; Feiran Oasis; Eastern desert governorate:
Cairo-Suez Road; Cairo Area; Gebel el Ahmar; Wadi Ghuweibba;
beginning of Wadi el Asyuti; Digla, near Cairo; Ain Sukhna.
Distribution in Libya. Coastal areas of the Gulf of Sirte, the
northeastern Libyan coast, and the great hamadas of central Tripoli-
tania and western Cyrenaica.
Jaculus deserti favillus. Cyrenaica and Tripolitania: Littoral
deserts near the Gulf of Sirte and northern Cyrenaican coast.
Jaculus deserti Juscipes. Tripolitania: Interior and coastal hamadas.
Jaculus deserti rarus. Cyrenaica: Extreme southeastern Cyrenaica.
Jaculus deserti vastus. Cyrenaica: Interior hamadas of central-
western Cyrenaica and the Gebel el Harug el Asued.
Distribution in Algeria. Northern Sahara, including the High
Plateaus and the Saharan Atlas.
Distribution of the subspecies in Algeria.
Jaculus deserti deserti. Vicinity of Ouargla between the Grand
Erg Oriental and the Grand Erg Occidental, and Biskra in the
Saharan Atlas.
Jaculus deserti
subspecies by its proportionately larger auditory bullae and generally

smaller size of most cranial and external measurements (Ellerman
and Morrison-Scott, 1951; and Toschi, 1951). At this time, com-
paratively few areas in Libya were represented by specimens, and
the above assignments were based largely on geographic occurrence
with little regard for taxonomic relationships. Zavattari (1934, p. 896)
recorded both J. j. deserti and J. j. jaculus from El Agheila on the
Cyrenaican Coast. The occurrence of these two subspecies in the
same geographic area clearly violates the concept of subspeciation.
Setzer (1957), the most recent worker in Libyan mammals, had no
specimens of the lesser jerboa available and accordingly was unable
to investigate this problem. In the present study, adequate series
are available from widely scattered localities and thus provide an
opportunity to clarify the taxonomic status of the small jerboas
of Libya.
No topotypes of J. j. deserti are available for study, but Loche
(1867, pp. 100-101), in the original description, described this jerboa
as follows: "Head short, terminated by a small muzzle; eyes black
and shining, ears medium and taking their origin from a white patch;
pelage of the upper parts a pale dun color, slightly tinted with black
on the rump, resulting from the blackish tips of the otherwise dun-
colored hairs; hairs of the sides of the body and flanks white, ter-
minated by tawny and brownish hairs; face, outer parts of shoulders,
thighs and ventral surface of tail uniformly dun-colored; posterior
surfaces of hind limbs, pencil and plantar surfaces of the hind feet a
tawny-brownish color; upper eyelids, sides of the muzzle, all of the
front limbs, all underparts and inner surfaces of the hind feet are
white: a line of this same color extends ventrally along the tail to
the terminal brush, which is brown and white; nails white; total
length 310 mm., length of tail 170 mm." Loche (1867) indicated
that Dipus deserti (= Jaculus jaculus deserti) was a miniature repro-
duction of Dipus gerboa Olivier ( = Jaculus orientalis).
Representatives of J. j. deserti from Biskra, Algeria, agree closely
with the above description and are found to differ from typical J. j.
jaculus in a number of additional characters, particularly in having
brownish rather than white hairs in the soles of the feet, much darker
dorsal color, and two foramina in the angular process of the lower jaw,
instead of one as in J. j. jaculus. Furthermore, in several localities in
Libya, jerboas of the deserti type and those typical of J. j. jaculus
occur sympatrically and show no evidences of interbreeding. The
present work indicates that J. j. deserti is sufficiently distinct to
warrant reinstatement as a full species, and because of priority (1867),
assumes precedence as the nominate form. The extralimital forms
Jaculus loftusi vocator Thomas, 1921 (type locality: Sohar, Muscat,
Arabia) Dipus schluteri Nehring, 1901 (type locality: Jaffa, Palestine);

;
and Dipus loftusi Blanford, 1875 (type locality: Mohumrah, Iraq)

were referred to Jaculus jaculus by Ellerman and Morrison-Scott
(1951, p. 539). I have, however, found that these forms definitely have
characters which ally them to the "deserti" type and consider them as
subspecies of J. deserti rather than J. jaculus. Henceforth, these forms
should be known as Jaculus deserti vocator Thomas, Jaculus deserti
schluteri (Nehring), and Jaculus deserti loftusi (Blanford).
Although intergradation is not demonstrable between populations of
Jaculus deserti favillus and Jaculus deserti juscipes in Libya, the ranges
of these two subspecies are not separated by physical barriers. In the
future, when more adequate series become available, this intergrada-
tion will probably become apparent, and clinal gradients will be
demonstrable within these two widely distributed populations. Farther
east, however, the Nile River appears to serve as an effective barrier
for the exchange of genes between J. d. javillus and Jaculus deserti
schluteri.
Jaculus deserti, in the dark color of dorsum and soles of hind feet,
resembles Jaculus blanfordi (Murray), whose range lies far to the east
and north. This similarity in color is probably the result of conver-
gence or a response to a similar color of substrate. Vast areas of gray
and brown hamadas are of common occurrence in the ranges of both of
these jerboas. Morphologically, J. blanfordi is more closely related to
the larger Egyptian jerboa (J. orientalis).
Ecological observations. Four distinct populations comprise
this species in Libya. Jaculus deserti favillus is apparently confined to
the more humid Mediterranean coast and adjacent areas of low
elevation. Jaculus deserti vastus, J. d. rarus and J. d. fuscipes prefer
the more arid deserts of the interior. In general, this species of jerboa
is more confined to the "hamadas" or great "pebble deserts" of the
Libyan hinterland. In the interior hamadas, vegetation may be entirely

wanting or confined to narrow strips along the margins of wadis, the
brinks of escarpments, or the fringes of inland "play as." The larger
wadis sometimes support stands of acacia of appreciable size, but these
are of rare and local occurrence.
Near the coastal plain at Marble Arch, J. jaculus and J. deserti occur
together. In other coastal areas the large Egyptian jerboa (J. orientalis)
also occurs sympatrically with these smaller species.
Jaculus deserti favillus Setarer
Jaculus jaculus favillus Sctzer, Proc. Biol. Soc. Washington, vol. 68, p. 184, Dec. 31,
1955 (Bir Bosslanga; Salum, Western Desert Governorate, Egypt).
Specimens examined. Two, from Cyrenaica: 10 km SW Fort

Capuzzo, 1; from Tripolitania: 15 km WNW
Marble Arch, 1.
Measurements. Measurements of the above specimens, an adult

male, 325806, and an adult female, 325819, are, respectively: Total
length 300, 312; length of tail 186, 195; length of hind foot 63, 60;
length of ear 24, 23; greatest length of skull 34.7, 33.8; condyloincisive
length of skull 30.5, 30.6; crown length of upper molariform toothrow
5.2, 5.5; greatest breadth across zygomatic processes 23.5, 23.2; least
interorbital breadth 13, 12.8; length of nasals 13.2, 12.5; breadth of
rostrum at level of antorbital foramina 5.5, 4.5; greatest breadth of
braincase 19.1, 18.5; greatest breadth across antorbital processes 24.4,
24.1.
Diagnosis. Upperparts, including dorsum of tail, Avellaneous
becoming darker posteriorly owing to greater admixture of brown-
tipped hairs; interorbital and interauricular areas Cinnamon-Buff;
mystacial, suborbital, postauricular, and pectoral areas Light Buff*;
pinna of ears badly disfigured by ants, but remnants Clay Color; in
both specimens Plumbeous underfur exposed in large eroded patches
on dorsum behind ears; underparts white grading to Light Buff on
lateral margins; lateral margins of rump creamy and washed with
flecks of gray; prominent whitish bands on the posterolateral areas
of the rump converge medially onto the tail; fore and hind feet
white dorsally, forefeet naked ventrally; soles of hind feet heavily
furred ventrally with strong admixture of Light Pinkish Cinnamon;
tail prominently bicolored and pinnate, Avellaneous dorsally and
Pale Pinkish Buff ventrally; pinna Bister proximally and white

distally; vibrissae profuse and dark basally. Skull: Large for the
species; auditory bullae large and markedly inflated; zygomata
heavy, relatively flattened in profile; angular process of mandibles
fragile and with two distinct foramina of unequal size on anterior
margin.
Fromnear topotypes of the nominate subspecies from Biskra,
Algeria, these two specimens from Libya are larger in all external
measurements and cranial characters, being of comparable size only
in the length of the ears.
These two specimens differ from Jaculus deserti schliiteri as known
from Feiran Oasis, Sinai, Egypt, as follows: Size: Smaller in all
external measurements, especially in length of hind foot and length
of ear. Skull: Smaller in greatest length, condyloincisive length, and
breadth of rostrum at level of antorbital foramina; larger in length
of molariform toothrow, least interorbital breadth, and size and
degree of inflation of auditory bullae. Color: Slightly paler and less
variegated dorsum, supraorbital and rostral areas more uniformly
colored Cinnamon Buff, and soles of feet with less suffusion of dark
hairs.
The specimens from Libya, although clearly referable to J. d.
favillus, differfrom topotypes of the latter in having slightly shorter

ears, paler color of dorsum, longer molariform toothrows, greater
least interorbital breadth, and the two foramina of the angular
process larger, more distinct, and more uniform in size.
For comparisons with J. d. fuscipes, J. d. rarus, and J. d. vastus,
see accounts of those subspecies.
Remarks. This subspecies was originally assigned to J. jaculus by
Setzer (1955) and considered by him to represent an eastward ex-
tension of the Libyan fauna into Egypt. The present study indicates,
however, that this subspecies has a much wider distribution in Libya
and Egypt than was formerly supposed and shows characters allying
it with J. deserti rather than to J. jaculus.
The specimen from Fort Capuzzo was taken alive while making
a nighttime check of a trapline which was set in a localized pocket
of vegetation in a sandy depression in the otherwise denuded coastal
plain. The plants forming these sporadic "pockets" of vegetation
were composed of juniperous species and other woody shrubs.
The habitat at Marble Arch is similar to that of the coastal plain
at FortCapuzzo but with more widespread areas of rocky hamada.
Jaculus deserti
favillus apparently representspopulation of
a
jerboas of large size restricted to the more humid environs of the
coastal plain and low-lying areas of the Libyan and Egyptian littoral
deserts, and it is genetically distinct from those of the Saharan
interior. Setzer (1958) suspected that the range of this subspecies
was restricted to the humid areas of the Mediterranean coastline.
Jaculus deserti fuscipes, new subspecies
Holotype. Adult female, skin and skull, USNM 322761, from

7 km S El Gheddahia, Tripolitania Province, Libya; obtained Dec. 3,
1961, by G. L. Ranck, original no. 909.
Specimens examined. Five, from Tripolitania: 20 km E Rumia,
1; 7 km S El Gheddahia, 3; 55 km SW Bir Allagh. 1.
Measurements. Averages and extremes of three females from the
type locality, with the measurements of the type in brackets, are,
respectively: Total length 287 (285-290), [290]; length of tail 180.7
(173-192), [192]; length of hind foot 61.7 (61-62), [62]; length of ear
23.7 (23-24), [24]; greatest length of skull 34 (33.7-34.4), [34]; con-
dyloincisive length of skull 30.5 (30.2-30.7), [30.2]; crown length of
upper molariform toothrow 5.3 (5.3-5.3), [5.3]; greatest breadth
across zygomatic processes 22.7 (21.5-23.5), [23.5]; least interorbital
breadth 12.4 (12.2-12.6), [12.6]; length of nasals 12.0 (11.7-12.4), [11.8];
breadth of rostrum at level of antorbital foramina 5 (4.6-5.2), [5.2];
greatest breadth of braincase 18.2 (17.6-18.6), [18.4]; greatest breadth

across antorbital processes 23.1 (23-23.1), [23.1].
Diagnosis. Upperparts Avellaneous grading into Cinnamon-Buff;
supraorbital, interauricular and rostral regions and all parts of dorsum
heavily suffused with dark being particularly concentrated on
hairs,
lateral margins of rump and extending onto flank; sides and scapular
region with strong admixture of brownish hairs; entire dorsum thus
appearing streaked or variegated in color; mystacial, eircumorbital,
and small patches near anterior margin of ear Pale Pinkish Buff;
entire underparts white, becoming more buffy on ventrolateral sur-
faces; pinna of ear dark (Drab) on distal margins and anteriorly grad-
ing to same color as dorsum of body with a thin row of buffy hairs;
vibrissae long, almost as long as body, and dark basally; fore and
hind feet white dorsally, the former naked ventrally and bearing five
claws; soles of hind feet heavily furred with markedly contrasting
inner hairs of Mummy Brown and bearing three claws; tail indistinctly
bicolored Avellaneous, dorsally, and ventrally, Light Buff, the latter
color becoming whiter distally; distal portion of tail distinctly pinnate
and bicolored brownish-black proximally and white distally. Skull:
Medium, robust, and compacted; incisors large and heavy; upper
molariform toothrow long; rostum wide and short; nasals short;
zygomata heavy; braincase moderately vaulted; condyloincisive
length long; auditory bullae large and markedly inflated; mandible
heavy and angular process of mandible with two distinct foramina of
nearly equal size on ventrolateral margin.
Comparisons. Members of this subspecies resemble near topotypes
of Jaculus deserti deserti from Biskra, Algeria, but differ in their
darker color of dorsum and soles of the hind feet, larger hind feet,
larger and more distinct foramina on the angular process of lower
jaw and slightly larger size of skull.
From specimens of Jaculus deserti schliiteri as known from Feiran
Oasis, Sinai Governorate, and Wadi Ghuweibba, Eastern Desert
Governorate, Egypt, Jaculus deserti Juscipes differs in slightly darker
color of dorsum and soles of hind feet, smaller body size, markedly
smaller hind feet and ears, shorter skull, smaller and less inflated au-
ditory bullae, foramina of angular process larger and more uniform
in size, more vaulted braincase, longer molariform toothrow, larger
interorbital breadth, shorter nasals, and slightly smaller breadth of
braincase.
Compared with specimens representing Jaculus deserti javillus from
20 miles west of Sidi Barrani, Western Desert Governorate, Egypt,
this subspecies differs in smaller size of all external measurements,
being especially smaller in length of ears and hind feet. In color,
J. d. fuscipes is slightly darker dorsally with a more prominent ad-

mixture of black hairs on lateral margins of rump and darker hairs
on soles of hind feet. Cranially, these two subspecies are quite similar,
but J. d. fuscipes can be distinguished by its longer molariform tooth-
row, slightly more vaulted skull, more robust zygomata, and larger
and more distinct foramina on the angular process.
Jaculus deserti fuscipes differs from Jaculus deserti vastus in markedly
smaller size of external and cranial measurements except length of ear,
condyloincisive length of skull, length of molariform toothrow, and
greatest breadth of braincase. Jaculus deserti vastus is paler and more
uniform in color of dorsum with less admixture of dark hairs, lacking
blackish hairs on lateral margins of the rump, and is more buffy
ventrally.
Remarks. This subspecies of jerboa can be distinguished from all
others in Tripoli tarda by its markedly darker dorsal color, darker
hairs on the sole of the hind foot, more compact skull, the presence
of two distinct foramina in the angular process, larger and more
greatly inflated auditory bullae, and proportionately wider upper
incisors.
The range of Jaculus deserti fuscipes includes the extensive hamadas
of central and northern Tripolitania. Throughout these "pebble
deserts," vegetation occurs sporadically in localized concentrations
along the margins of dry watercourses and other low-lying areas.
These jerboas probably have a much wider distribution in Libya than
is currently supposed.
The above specimens were all obtained from these interior hamadas.
The type locality, although near the Mediterranean coast, is typical
"hamada" desert and has no physical relationship to the nearby
coastal plain. Fat-tailed sand rats {Pachyuromys duprasi) also show
a decided preference for this rocky, upland type of habitat and were
obtained at Rumia and El Gheddahia along with jerboas.
The specimens from near Bir Allagh and Rumia are darker dorsally
and have a greater suffusion of dark hairs on the soles and metatarsal
areas of hind feet than the specimens from the type locality but are
definitely referable to this new subspecies.
The subspecific name fuscipes refers to the dark-colored soles of
the hind feet.
Jaculus deserti rarus, new subspecies

319781, from Ain
Zueia, Gebel Uweinat, Cyrenaica Province, Libya; obtained Mar. 26,
1961, by H. W. Setzer, original no. 2987.
Specimen examined. The type only.
Measurements. The type specimen measures: Total length 296;
length of tail 184; length of hind foot 61; length of ear 24; greatest
length of skull 33; condyloincisive length of skull 29.2; crown length

of upper molariform too throw 4.9; least interorbital breadth 11.6;
length of nasals 12.1; breadth of rostrum at level of antorbital foram-
ina 5.1; greatest breadth of braincase 16.9; greatest breadth across
antorbital processes 22.
Diagnosis. Upperparts Cinnamon-Buff grading to Light Ochraceous
Buff in interauricular and supraorbital regions; rump decidedly darker
owing to greater admixture of brown-tipped hairs; plumbeous under-
fur exposed in many areas of dorsum; rostral, pectoral, cheek, flank,
and mystacial areas, and entire venter Pale Ochraceous-Buff; lateral
areas washed with darker color; fore and hind feet sparsely haired
and white dorsally; ventral surface of hind feet with long tufts of gray
hairs; tail indistinctly bicolored, Cinnamon-Buff dorsally, Light Buff
ventrally and with a prominent pinna, black proximally and white
distally. Skull; Small, zygomata delicate, auditory bullae small and
little inflated, rostrum narrow, foramen magnum small, molariform
toothrow short, and angular process of mandible with two distinct

foramina on its anteromedial surface.
Remarks. This subspecies differs from all other subspecies of
Jaculus deserti in the smaller size of all cranial and external measure-
ments. The type specimen of J. d. varus resembles the type series of
Jaculus jaculus cufrensis in body size, greatest length of skull, and
condyloincisive length. In crown length of upper molariform toothrow,
least interorbital breadth, length of nasals, breadth of rostrum at
and degree of inflation of auditory
level of antorbital foramina, size
bullae, and in external measurements, it is closer to Jaculus jaculus
butleri Thomas. The above are only superficial resemblances, however,
inasmuch as specimen has two foramina in the angular process of
this
the mandible, brownish hairs on the soles of the hind feet, and markedly
darker dorsal coloration with greater suffusion of brownish hairs on
the rump and sides. The latter are all diagnostic characteristics of
Jaculus deserti and, in the aggregate, are unknown in Jaculus jaculus.
This single specimen was obtained from the barren hamada at the
base of the Gebel near Ain Zueia by Henry W. Setzer, while en route
to the Tibesti Mountains in the spring and early summer of 1961.
It was obtained only after three successive nights of continuous trap-
ping throughout an area of abundant diggings. Efforts to obtain ad-
ditional specimens by "night hunting" were without effect.
The extensive sand seas to the north have apparently isolated this
subspecies from the ranges of all other populations of Jaculus deserti.
The small size and other cranial differences of this subspecies may
have arisen through the agency of genetic drift and consequent rapid
genetic fixation in an effectively small, isolated population of jerboas.
The term rams is from the Latin meaning scarce or scattered and
has reference to the apparent scarcity of jerboas of this subspecies.
Jaculus deserti vastus, new subspecies

Wadi er Rueis, Gebel el Harug el Asued, 340 km WNW
Tazerbo
Oasis, Cyrenaica Province, Libya; obtained Apr. 29, 1962, by G. L.
Specimens examined. The type only.
Measurements. Measurements of the type specimen are as follows:
Total length 311, length of tail 203; length of hind foot 67; length
of ear 24; greatest length of skull 34.3; condyloincisive length of
skull 30.3; crown length of upper molariform toothrow 5.3; greatest
breadth across zygomatic processes 23.6; least interorbital breadth
12.6; length of nasals 12.3; breadth of rostrum at level of antorbital
foramina 5.2; greatest breadth of braincase 18.2; greatest breadth
across antorbital processes 23.2.
Diagnosis. Upperparts Light Pinkish Cinnamon grading to Pale
Pinkish Cinnamon on sides, flanks, pectoral regions, and rostral
areas; all upperparts with strong suffusion of brownish-tipped hairs
more concentrated on rump; mystacial, circumorbital, subauricular
areas, forearms and entire venter, Pale Pinkish Buff; vibrissae long
and darkly colored basally; pinnae of ears sparsely haired and white
dorsally; forefeet naked ventrally and bearing five claws; hind feet
heavily furred ventrally, bearing three claws, and sole with strong
suffusion of Mummy Brown hairs. Tail indistinctly bicolored Pinkish
Buff dorsally and Light Buff ventrally and with terminal pinna which
is Clove Brown proximally and white distally. Skull: Large, auditory
bullae large and markedly inflated; zygomata robust; incisors large

and heavy; braincase slightly vaulted; ramus of mandible heavily
fenestrated and angular process with two distinct oval foramina of
unequal size.
Comparisons. From representatives of J. d. deserti of Algeria,

this specimen from the Wadi er Rueis differs in its much larger size,
particularly in the length of the hind feet and tail.
Jaculus deserti vastus differs from representatives of Jaculus deserti
favillus from 20 miles west of Sidi Barrani, Western Desert Gover-
norate, Egypt, in slightly longer tail, much larger hind feet, shorter
ears, longer molariform toothrow, larger and more uniform foramina
on angular process, paler, less variegated dorsal pelage, and more
buffy venter.
This specimen closely resembles representatives of Jaculus deserti
from Feiran Oasis, Sinai Governorate, and Wadi Ghuweibba,
schliiteri
Eastern Desert Governorate, Egypt, but differs in smaller body size,
slightly shorter length of tail, hind feet, and ears, longer molariform
toothrow, wider interorbital breadth, shorter nasals, slightly paler,
less variegated color of dorsum and more buffy venter.
For comparison with Jaculus deserti fuscipes, see account of that

subspecies.
Remarks. Jaculus deserti vastus can easily be distinguished from all
other small Libyan jerboas by its much larger body size, markedly
longer tail and hind feet, and much greater suffusion of buffy hairs
on the forelegs, sides, and venter.
Although specimen doubtless belongs to the "deserti" group
this
of jerboas, theabove differences are of sufficient magnitude to warrant
the designation of a new species. Owing to lack of adequate material
it is deemed better, however, to limit the category to subspecific
rather than specific rank.
The type specimen is the only record of occurrence for this subspecies
in Libya. The above specimen was obtained from the rocky, denuded
margins of the Wadi er Rueis at a point where it emerges from the
higher parts of the Gebel el Harug el Asued (Black Gebel). Volcanic
extrusions are widespread in this area and in many places encroach
upon the Wadi. Vegetative cover is very limited and is composed
primarily of acacias, woody shrubs, and hardy grasses.
This large jerboa probably has a much wider distribution in Libya
than the present study would indicate, and its range doubtless in-
cludes the entire massif of the "Black Gebel" and the low-lying pe-
ripheral areas wherever suitable habitat occurs.
Jerboas inhabiting the Gebel el Harug el Asued are effectively
isolated from the ranges of other subspecies of Jaculus deserti in
Libya, Gene exchange with these other populations is accordingly of
infrequent occurrence and genetic fixation and differentiation have
thus progressed rapidly. Owing to the remoteness and consequent
inaccessibility of central Cyrenaica, additional specimens of this
unique jerboa will probably long be wanting.
The name vastus, from the Latin meaning empty and waste, alludes
to the remote regions inhabited by jerboas of this subspecies.
Jaculus jaculus (Linnaeus)
M us jaculus Linnaeus, Syst. Nat., 10th ed. p. 63, 1758 ("In Arabia, Calmukia";
Giza Pyramids, Egypt [G. Allen, 1939]).
General distribution of species. Iraq, Syria, Lebanon, Israel,

Jordan, Saudi Arabia, North Africa southward through the Sahara
including Sudan, Chad, Niger, Mauritania, and Spanish Sahara.
Distribution in Libya. Widespread through the coastal and in-
terior areas of Cyrenaica, Tripolitania, and the Fezzan.
Jaculus jaculus arenaceous. Fezzan: Oases and connecting wadis.
Jaculus jaculus collinsi. Cyrenaica: Gialo and Tazerbo and the
intervening Serir of Calanscio.
Jaculus jaculus cufrensis. Cyrenaica: Environs of Cufra Oasis.

Jaculus jaculus tripolitanicus. Tripolitania: Gefara plain.
Jaculus jaculus whitchurchi. Cyrenaica: Coastal plain of Gulf of
Sirte,and Giarabub Oasis.
Published records in Libya. Cyrenaica: Giarabub (de Beaux,
1928); El Agheila, Cufra (de Beaux, 1932); Giarabub (Zavattari,
1937); Zauia Mechili (de Beaux, 1938); Giarabub, Gialo, Marada,
Tazerbo (Toschi, 1951); Tripolitania: Socna, "AttickLoumonileh",
"Oumsinerma" (Thomas, 1902); Tripoli (Klaptocz, 1909); Sirte
(Zavattari, 1937); Tripolitania settentrionale (Toschi, 1951); Fezzan:
Murzuch, Serdeles, Bir el Fatima (Toschi, 1951).
Remarks. The present study reveals that jerboas of this species
are an assemblage of rather localized populations showing consider-
able differentiation resulting from relatively long periods of isolation
of allopatric populations and ensuing genetic fixation. Local differ-
entiation of these jerboas is demonstrable in several areas in Libya.
These areas may be delineated as the oases and serirs of central and
southern Cyrenaica, the oases and intervening wadis of the Fezzan,
and the littoral deserts along the Gulf of Sirte in Tripolitania. This
latter region is divisible into two distinct areas the broad coastal —
Jaculus jaculus
I arenaceous
2 collinsi
3 cufrensis
4 tripolitanicus
5 whitchurchi
Figure 50. — Distribution of the subspecies of Jaculus jaculus.

plain between the Mediterranean Sea and the coastal escarpment and
the transitional desert farther inland.
The Sand Sea of Rebianna serves as an effective barrier to gene flow
between jerboas inhabiting the oases of Cufra and Tazerbo, and the
Sand Sea of Calanscio prevents contact between these populations
and those of the nominate subspecies to the north. Genetic interchange
between animals from Gialo and those of coastal Libya is of doubtful
occurrence owing to extensive sandy plains and dunes interposed
between their ranges. Some genetic exchange occurs between coastal
and interior populations in the region of the oases of Socna, Hun and
Uaddan, but generally, the Fezzanese jerboas have maintained their
genetic integrity. In northern Tripolitania, the coastal escarpment
effectively separates the jerboas of the coastal plain from those of the
"high deserts" of the interior.
Ecological observations. Jaculus jaculus inhabits both the
littoral desertsand the interior oases of Libya. On the coastal plain
they are common in the sandy hummocks and large coastal dunes, and
farther inland are confined to the sandy margins of the oases where
vegetation occurs sparingly for several kilometers beyond the palm
groves of the oasis proper. This peripheral vegetation, composed of
Calligonum, Tamarix, and thorny perennials, is interspersed with
barren areas of sandy, rock-strewn plains. Large, solitary hummocks
are present in these outlying areas and provide suitable sites for
burrow construction above the water table.
Occasionally, jerboas are taken from sandy areas at the bases of date
palms and tamarix in the interior of the oasis and occur here with
Gerbillus gerbillus, Gerbillus pyramidum, and Gerbillus amoenus.
They are of rare occurrence in rocky deserts and playas far removed
from oases. In these latter areas, J. jaculus is supplanted by the
darker colored Jaculus deserti. These two species are known to occur
sympatrically along a narrow zone of contact on the inner margins of
the coastal plain and probably occur together in other parts of Libya.
Jerboas normally begin foraging shortly after dusk, but when
cold temperatures prevail, they emerge from their burrows later in
the night. In Libya, prolonged periods of cold are uncommon, hence
it is doubtful if jerboas hibernate there. In a few instances, jerboas
were taken when temperatures fell below 30° F.
Jerboas are difficult to catch by standard trapping methods, and
accordingly trap yield is usually low, thus rendering an incorrect
estimate of population size. Jerboas are probably more abundant in
Libya than trapping success would indicate.
The pale, ochraceous color of these jerboas is undoubtedly an
adaptive response to the prevailing color of the soil on and in which
they live. Jaculus jaculus is markedly paler in dorsal color than J.
deserti, which inhabits the grayish brown soils of the hamadas.
285-134 O —68 16
;
Jaculus jaculus arenaceous, new subspecies
Holotype. Adult male, skin and skull, USNM 322812, from Edri,
Fezzan Province, Libya; obtained Feb. 9, 1962, by G. L. Ranck,
original no. 1624.
Specimens examined. Seventy-one, from Tripolitania: 2 km SW
Hun, 5; from Fezzan: Brach, 1; Edri, 23 (2 skeletons); Temenhint
Oasis, 30 km NE Sebha, 3 km N Sebha, 1 3 km
; 4 Sebha, 2
; NW
Sebha, 3; 7 km SW Sebha,1; El Abiad, 60 km SW Sebha, 9; 75 km
W Ubari, 2; Murzuch, 5; 55 km SSW Serdeles, 7; El Gatrun, 3;
12 km N Ghat, 3; Ghat, 4.
type in brackets, are, respectively: Total length 289 (279-307), 299.4
(288-308), [307]; length of tail 178.4 (160-200), 182 (172-189), [202];
length of hind foot 63.2 (61-66), 63.3 (62-65), [65]; length of ear 21.9
(21-23), 21.8 (21-22), [23]; greatest length of skull 34.1 (33.5-34.7),
34.3 (32.9-34.8), [34.7]; condyloincisive length of skull 29.9 (28.9-
30.8), 30 (28.9-31.4), [30.6]; crown length of upper molariform
zygomatic processes 22.1 (21.3-23.6), 22.3 (21-22.8), [22.3]; least
interorbital breadth 12.4 (11.8-12.8), 12.4 (11.8-13.2), [12]; length
of nasals 12.6 (12-13.4), 12.8 (11.7-14), [14]; breadth of rostrum at
level of antorbital foramina 5 (4.8-5.2), 4.9 (4.7-5.3), [5] greatest
breadth of braincase 18.1 (17.3-18.7), 18 (17.5-18.7), [18]; greatest
breadth across antorbital processes 22.7 (22-23.4), 22.8 (21.7-23.7),
[22.3.]
Diagnosis. Upperparts Light Pinkish Cinnamon becoming darker
posteriorly with strong admixture of brownish-tipped hairs causing a
markedly variegated appearance; rostral, mystacial, circumorbital,
postauricular, pectoral, and lateral areas Pale Ochraceous-Buff;
entire underparts, dorsal surface of fore and hind feet, and terminal
portion of tail white; forefeet naked ventrally and bearing five claws;
hind feet relatively large, heavily furred ventrally, artificially stained
Pale Yellow-Orange and with three toes; tail indistinctly bicolored
Pale Yellow-Orange dorsally and Light Buff ventrally, and ter-
minating in a distinct transversely bicolored pinna, Bister proximally
and white distally; pinna of ear Clay Color, sparsely haired with
fringe of buff-colored hairs on anteromedial margin. Skull: Large,
auditory bullae moderately large and inflated; zygomata convergent
anteriorly; pterygoid processes slightly divergent; nasals relatively
long; molariform toothrow relatively long.
Comparisons. Jaculus jaculus arenaceous from the nominate
differs
subspecies in greater overall length, less compacted skull, slightly
less vaulted braincase, more inflated bullae, larger infraorbital
foramina, longer and more widely flaring nasals, wider rostrum,

greater breadth of rostrum at level of antorbital foramina, and longer
molariform toothrow. In color, J. j. arenaceous differs from topotypes
of Jaculus jaculus jaculus in more uniform orangish dorsal coloration,
less suffusion of dark hairs on sides, darker pinna of ear, and vibrissae
of more uniform color.
Compared to the type and type series of Jaculus jaculus whitchurchi,
J. j. arenaceous differs in greater overall length of skull and body,
smaller hind feet, less vaulted braincase, more robust zygomata,
larger, infraorbital foramina, larger foramen on anterior fossa of
angular process and larger size in all other cranial features. In color,
jerboas from Edri are darker, more variegated and far less uniformly
colored dorsally, have more admixture of dark hairs on the sides, and
have a more extensive buffy patch anterior to dark portion of pinna of
the tail.
From the type and type series of Jaculus jaculus collinsi, J. j.
arenaceous can be differentiated by more massive skull, relatively

its
narrower rostrum, larger lachrymals, slightly larger and more inflated

auditory bullae, more tenuous pterygoid processes, larger size of all
cranial measurements (except greatest breadth across zygomatic
processes and greatest breadth of braincase), and markedly larger hind
feet. Jerboas from Edri appear darker and more variegated in color
owing to greater suffusion of brownish hairs on dorsum, rump, and
sides. An old male, 322768, from Temenhint Oasis, 30 kilometers north-
east of Sebha, is markedly more orangish (ochraceous) in color of
dorsum and probably represents an aberrant specimen or an extreme
in pelage color incident to old age.
This subspecies can be easily separated from J. j. sefrius Thomas
and Hinton from Ain Sefra, southwestern Algeria, by its significantly
smaller size of external and cranial measurements.
A single specimen (BM, 12.11.14.53) of J. j. centralis Thomas and
Hinton from 85 kilometers south of El Golea, central Algerian Sahara,
somewhat resembles J. j. arenaceous in color, but in external measure-
ments and cranial characters it is much smaller.
For comparisons with Jaculus jaculus tripolitanicus and Jaculus
jaculus cufrensis, see accounts of those subspecies.
Remarks. Specimens from 2 kilometers southwest of Hun are the
only known representatives of J. j. arenaceous from Tripolitania
Province and are intergrades with J. j. whitchurchi. In length of upper
molariform toothrow, greatest breadth across zygomatic processes,
greatest breadth across antorbital processes, and length of the hind
foot they resemble the latter subspecies, but in the majority of cranial
and external measurements, they are closer to J. j. arenaceous to
which they are here referred. The vast hamadas to the north and east
of Hun, Socna, and Uaddan provide favorable habitat for jerboas and
accordingly allow for genetic exchange between coastal populations
and those in the interior.
In Fezzanese populations of jerboas, genetic uniformity has been
maintained owing to the continuity of suitable habitat and conesquent
free genetic interchange. Numerous dry watercourses such as the Wadi
es Sciati, north and west of Sebha, the Wadi Tenezoft, north of Ghat,
and the broad hamadas Unking together the oases of Tesaua, Murzuch,
Traghen, Umm el Araneb, Meseguin, Zuila, and Tmessa provide suit-
able habitat and insure the maintenance of genetic integrity within and
between populations of jerboas in these far-reaching areas. These
hamadas and wadis allow jerboas to circumvent the desolate "ramleh"
or sandy areas. El Gatrun, however, unlike the aforementioned oases,
is effectively isolated by the great sandy areas of the Idehan Murzuch,
and three specimens from here show slight differences from populations
elsewhere in the Fezzan. Until such time as adequate series become
available, I prefer to include jerboas from El Gatrun within the sub-
species J. j. arenaceous.
In the Fezzan, jerboas were obtained from a wide variety of habitats,
ranging from practically barren hamadas to the bleak margins of sand
seas. In these habitats, plants occur sparingly and are represented by
relatively few species, one kind usually being dominant. In most
instances, jerboas were not found in the oases proper but in the more
open, outlying areas.
The name arenaceous, meaning of sand or sandy, refers to the
sandy color of the pelage of these jerboas.
Jaculus jaculus collinsi, new subspecies

Tazerbo Oasis, Cyrenaica Province, Libya; obtained Apr. 15, 1962,
by G. L. Kanck, original no. 2039.
Specimens examined. Twenty, from Cyrenaica: Gialo Oasis,
5; Bir el Harasc, 1 Bir bu Zarregh, 1 Tazerbo Oasis, 11 El Gezira, 2.
; ; ;
Measurements. The averages and extremes of five males and six

females from the type locality, with the measurements of the type in
brackets, are, respectively: Total length 292 (282-307), 294.2 (275-
313), [293]; length of tail 182.2 (175-192), 183.3 (166-197), [182];
length of hind foot 61.6 (61-63), 61 (58-63), [62]; length of ear from
notch 22.2 (21-23), 21.7 (21-23), [23]; greatest length of skull 33.6
(32.5-34.5), 33.3 (32.3-34.3), [34.4]; condyloincisive length of skull
29.9 (28.6-30.9), 29.6 (28.7-34.3), [30.7]; crown length of upper
molariform toothrow 5.2 (5-5.3), 5.2 (4.9-5.4), [5.3]; greatest breadth
across zygomatic processes 22.2 (21.6-23.4), 21.8 (21.4-22.9), [21.7];
least interorbital breadth 12.2 (11.8-12.3), 12 (11.5-12.6), [12]; length
of nasals 12.2 (11.6-12.8), 12.3 (11.6-12.8), [12.6]; breadth of rostrum

at level of antorbital foramina 5 (4.7-5.3), 5 (4.7-5.1), [5.1]; greatest
breadth of braincase, 18.2 (17.6-18.6), 18 (17.5-18.4), [18.4]; greatest
breadth across antorbital processes 22.7 (22.3-23.1), 22.5 (21.7-23.6),
[23.1].
Diagnosis. Upperparts Light Ochraceous-Buff with suffusion of
darker hairs on rump; all hairs of dorsum Plumbeous basally; rostral
circumorbital, mystacial, subauricular, interorbital, and pectoral
regions Pale Ochraceous-Salmon; pinna of ear sparsely haired and
Clay Color; underparts, and dorsum of fore and hind feet
flanks,
white, grading to Light Buff in some specimens; lateral vibrissae
long and white; median vibrissae smaller and darker; hind feet
heavily furred ventrally and Pale Pinkish Buff; palmar surfaces of
front feet naked; tail Pinkish Buff dorsally, Light Buff ventrally,
with prominent bicolored terminal pinna, Warm Sepia proximally
and white distally; encroachment of buffy areas of the sides onto
dorsum imparting a decidedly pale aspect to all of the specimens.
Skull: Triangular; size medium; molariform teeth medium; auditory
bullae moderately inflated; infraorbital foramina large; braincase
relatively flattened, rostrum heavy; zygomata convergent anteriorly;
pterygoid fossae large and partially enclosed ventrally by the palatines;
pterygoid processes long and attenuated; basisphenoids, alisphenoids,
presphenoids, and orbitosphenoids markedly reduced and restricted
to median plane; lachrymals appearing as a posterior process of the
maxillary and situated dorsad of the winglike antorbital processes;
lateral margins of basioccipital convergent anteriorly; jugular proc-
esses small; distinct foramen located on ventroanterior margin of
condyloid process and anterior margin of angular process; coronoid
process markedly reduced.
Comparisons. From the type and the type series of Jaculus jaculus
cufrensis,specimens of J. j. collinsi differ in larger size of most cranial
characters, being smaller only in the least interorbital breadth and of
comparable size in the greatest breadth across antorbital processes.
Members of this subspecies also have larger molariform teeth, heavier
zygomata, narrower rostra, and less compact skulls. Jaculus jaculus
collinsi is paler in color owing to encroachment of the buffy areas of
the sides onto the dorsum, less suffusion of dark hairs on the flanks,
and the general overall Light Ochraceous-Buff of the back.
Jaculus jaculus collinsi appears to be most closely related to J. j.
jaculus but can be distinguished by longer molariform toothrow, less
inflated auditory bullae, larger infraorbital foramina, heavier rostrum,
and less vaulted braincase. In color, J. j. collinsi is markedly paler,
has greater suffusion of white or cream-colored hairs on the dorsum
of the tail, and has less admixture of dark hairs on sides and flanks.
Jaculus jaculus collinsi differs from the type and the type series of
J. j. whitchurchi in larger body size, longer tail, longer ears, more
variegated color of dorsum, paler flanks and lateral regions, smaller
lachrymals, wider rostrum, less vaulted braincase, heavier zygomata,
and generally larger cranial measurements.
For comparison with Jaculus jaculus arenaceous, see account of
that subspecies.
Animals from Gialo are noticeably paler and smaller in size than
those from the type locality which lies farther to the south, suggesting
intergradation with Jaculus jaculus whitchurchi, whose geographic
range is to the north. This similarity in color could best be inter-
preted as a response to the paler colored soils nearer the coast, inas-
much as extensive areas of sand and barren hamada occur between
Agedabia and Gialo. It is doubtful if genetic exchange is of common
occurrence across this barrier.
Some were obtained from a large concentration
of the type series
of sparsely vegetated dunes associated with date palms, and the re-
mainder were taken from among large clay hummocks at some distance
from the oasis proper. The latter area supported sparse growths of
Tamarix and Calligonum.
The subspecific name collinsi is proposed in recognition of Alan
C. Collins, who made possible the trip to the Tibesti Mountains during
the early spring and summer of 1961.
Jaculus jaculus cufrensis, new subspecies
Holotype. Adult female, skin and skull, USNM 325785, from El

Giof, Cufra Oasis, Cyrenaica Province, Libya; obtained Apr. 2, 1962>
Specimens examined. Seventeen, from Cyrenaica: El Giof, Cufra
Oasis, 15 (1 skull only); El Hauuari, Cufra Oasis, 2.
Measurements. The averages and extremes of five males and
seven females from the type locality, with the measurements of the
type in brackets, are, respectively: Total length 279.8 (273-285),
282 (270-297), [290]; length of tail 172.8 (164-178), 174.9 (165-184),
[180]; length of hind foot 63.6 (61-65), 61.4 (60-64), [60]; length
of ear 21.2 (20-22), 21 (20-22), [20]; greatest length of skull 33.2
(33-33.6), 32.7 (32.2-33.5), [32.7]; condyloincisive length of skull
29.3 (28.8-29.8), 28.7 (27.8-29.7), [28.9]; crown length of upper molar-
iform toothrow 5.1 (4.8-5.4), 4.9 (4.7-5.1), [5.1]; greatest breadth
across zygomatic processes 21.8 (21.1-22.7), 21.8 (21-22.7), [22.7];
least interorbital breadth 12.4 (12.1-12.9), 12.2 (12-12.5), [12.3];
length of nasals 12.1 (11.8-12.5), 11.6 (11.2-11.9), [11.7]; breadth
of rostrum at level of antorbital foramina 4.8 (4.6-5), 4.8 (4.4-5.4),
[4.8]; greatest breadth of braincase 17.6 (17.2-18.1), 17.7 (17.4-18.2),
[17.8]; greatest breadth across antorbital processes 22.3 (22-23.5),

22.1 (21-22.8), [22.7].
Diagnosis. Upperparts Cinnamon-Buff, more uniform anteriorly
and becoming darker posteriorly owing to a slight admixture of
brownish-tipped hairs; entire underparts, postauricular regions, and
flanks whitish, the latter with infusions of brownish hairs imparting
a streaked appearance; cheeks, mystacial areas, circumorbital regions,
and pectoral areas Pale Pinkish Buff; anterior margins of pinna of ear
with row of Pinkish-Buff hairs; remainder of pinna sparsely haired
and Clay Color; tail and hind feet Pale Pinkish Buff dorsally; hind
feet heavily furred, frequently Ochraceous-Salmon resulting from
artificial staining by the substratum; tail, dorsally, approximating
color of back but becoming lighter, more Pinkish-Buff, posteriorly
and terminating in a distinct tuft or pinna of Bister Brown proximally
and white distally; brown portion of the pinna becoming indistinct
ventrally; tail, ventrally, Light Pinkish Cinnamon becoming lighter
distally. In some specimens, the pinnate tail is stained a deep Ochra-
ceous-Orange owing to prolonged contact with the reddish-orange
substratum. Skull: Small; molariform teeth relatively small; auditory
bullae moderately inflated; rostrum moderately narrow; zygomata
convergent anteriorly; malar bone expanded laterally forming a wing-
like antorbital process; lachrymals distinct and appearing as posterior
appendages of the maxillary; infraorbital foramina large; braincase
moderately vaulted; upper and lower incisors relatively short and
narrow; angular process of mandible with posterolaterally directed
process and possessing a well-defined foramen on its anterior margin,
coronoid process small and reduced to a secondary process.
Comparisons. From topotypes of Jaculus jaculus jaculus, J. j.
cufrensis differs in markedly smaller cranial measurements and gen-
erally smaller body size. The auditory bullae are less inflated in
J. j. and the skull appears less massive in all general aspects.
cufrensis,
In color, J. j. cufrensis is lighter and more uniform in dorsal coloration
(Cinnamon-Buff as opposed to Tawny-Olive), has paler colored
vibrissae, less suffusion of dark hairs on lateral body surface, flanks
white or creamy as opposed to buffy or tan, and much less admixture
of dark hairs on the dorsum.
When compared with Jaculus jaculus butleri Thomas from the
Sudan, J. j. cufrensis is similar in greatest length of skull, condy-
loincisive length and least interorbital breadth but is significantly
smaller in zygomatic breadth, length of nasals, size of infraorbital
foramina and breadtli of rostrum. Jaculus jaculus cufrensis has a
slightly more vaulted skull, greater flaring of the nasals anteriorly
and more ventrally inflated auditory bullae. In color, J. j. cufrensis
differs from J. j. butleri in having less admixture of dark hairs on the
dorsum, lighter coloration of the dark portion of the pinna of the

tail, and much lighter, more ochraceous color of the dorsum.
In smaller size of almost all cranial and external measurements,

this subspecies is readily separable from Jaculus jaculus arenaceous.
For comparison with Jaculus jaculus collinsi, see account of that
subspecies.
Remarks. Specimens were taken only from extensive areas of
"hamada" or pebble desert supporting a scanty vegetative cover.
While foraging at night, these jerboas seem to prefer the rather barren
areas peripheral to the oases proper. The conspicuous absence of bur-
rows in these areas suggests, however, that the animals actually
dwell and construct burrows in the sandy areas nearer the oases.
The name cujrensis is proposed in reference to the oasis of Cufra in
southern Cyrenaica.
Jaculus jaculus tripolitanicus, new subspecies

325773, from 25
km N Gharian, Tripolitania Province, Libya; obtained Mar. 6, 1962,
Specimens examined. Five, from Tripolitania: 25 km N Gharian,
4;40kmENE Nalut, 1.
Measurements. The measurements of an adult female, 325771,

and the averages and extremes of three adult males, 325770, 325772,
and 325773, from the type locality, with the measurements of the
type in brackets, are, respectively: Total length 302, 298.3 (290-300),
[300]; length of tail 193, 181.7 (174-184), [184]; length of hind foot 66,
66.3 (61-65), [61]; length of ear 23, 21.7 (21-22), [22]; greatest length
of skull 36.4, 34.9 (34.9-34.9), [34.9]; condyloincisive length of skull
31.9, 30.4 (30.2-30.6), [30.2]; crown length of upper molariform tooth-
row 5.3, 5.5 (5.4-5.7), [5.7]; greatest breadth across zygomatic proc-
esses 22.5, 22.6 (21.4-24.1), [24.1]; least interorbital breadth 12.5,
12.6 (12.3-12.9), [12.6]; length of nasals 13.4, 12.4 (11.7-13), [12.5];
breadth of rostrum at level of antorbital foramina 5.2, 4.8 (4.7-4.8),
[4.8]; greatest breadth of braincase 19.2, 18.7 (18.2-19.1), [18.9];
greatest breadth across antorbital processes 23.7, 23.1 (22-23.8),
[23.8].
Diagnosis. Upperparts, subauricular region, and flanks Cinnamon-
Buff with strong admixture of brown-tipped hairs, more concentrated
on rump; lateral areas Pinkish Buff with suffusion of dark hairs;
overall dorsal coloration a uniform blending of light and dark hairs;
rostral, supra- and infraorbital, mystacial, and postauricular areas
Pale Pinkish Buff; pinna of ear sparsely haired with fringe of buffy
hairs along anteromedial margins and otherwise Clay Color basally
and Tawny-Olive terminally; fore and hind feet white dorsally; fore-
feet naked vent-rally with five distinct claws; hind feet large and heavily
furred ventrally, Pinkish-Buff with three distinct claws; tail indis-
tinctly bicolored Cinnamon-Buff dorsally and Pale Pinkish Buff ven-

trally; prominent terminal pinna transversely bicolored Bone Brown
proximally and white distally; dark portion of pinna of tail becoming
white ventrally. Skull: Large and massive; cheek teeth heavy;
zygomata heavy; braincase flattened; auditory bullae moderately in-
flatedand large; anterior palatine foramina short and wide; lachrymals
large; foramen on anterior margin of angular process markedly
enlarged.
Comparisons. Ja cuius jaculus tripolitanicus differs from the type
and type series of Jaculus jaculus whitchurchi in larger, more massive
skull, relatively, as well as actually, larger, more inflated, auditory
bullae, heavier zygomata, wider anterior palatine foramina, larger in-
fraorbital foramina, markedly less inflated braincase, and larger
cranial and external measurements. In color, J. j. tripolitanicus has
a much stronger and more uniform suffusion of dark-tipped hairs on
the dorsum, a greater admixture of brownish hairs on flanks, and a
more sharply bicolored tail with a much darker anterior portion of the
pinna.
Compared with the type and type series of Jaculus jaculus arena-
ceous, J. j. tripolitanicus is larger and more massive in almost all
cranial measurements, has more inflated auditory bullae, markedly

wider anterior palatine foramina, more divergent pterygoid processes,
larger lachrymals, more robust zygomata, and larger, more oval
foramina on anterior fossa of angular process of mandible. In color,
J. j. tripolitanicus has less admixture of dark-tipped hairs on sides
and dorsum, more suffusion of buffy hairs on flanks, a more prom-
inently bicolored tail, and a darker anterior portion of the pinna.
From topotypical Jaculus jaculus jaculus, specimens from Gharian
are larger in all cranial measurements and have larger more inflated
auditory bullae, a larger foramen magnum, more divergent pterygoid
processes, more robust zygomata, less vaulted braincases, larger
lachrymals, and more widely flaring nasals. Jaculus jaculus tripol-
itanicus also differs from the nominate subspecies in paler dorsal
coloration, owing to less admixture of dark-tipped hairs, greater
suffusion of buffy hairs on sides and pectoral regions, and more
uniformly colored vibrissae.
In their generally large overall size and massive skulls, members
of this subspecies resemble representatives of Jaculus jaculus sejrius
from western Algeria but differ in having significantly longer hind
feet, much larger ears, and longer molariform toothrows. The simi-
larities probably do not indicate close taxonomic relationship, how-
ever, because populations of Jaculus jaculus centralis, a subspecies
markedly smaller in size than either J. j. tripolitanicus or J. j. sefrius,

are interposed between their ranges.
Remarks. Jaculus jaculus tripolitanicus is larger cranially and has
larger hind feet than any other subspecies of Jaculus jaculus in Libya.
A specimen from 40 kilometers east-northeast of Nalut,
single
Tripolitania, differsfrom the type series of J. j. tripolitanicus in
more uniform color of dorsum with more yellowish coat, less admix-
ture of dark-tipped hairs, and slightly larger hind feet. Cranially,
this specimen does not differ significantly from animals from Gharian
and is accordingly referred to J. j. tripolitanicus.
This large subspecies is apparently confined to the more humid

environments of the Gefara Plain north of the Gebel Nefusa.
Both the type series and the specimen from 40 kilometers east-
northeast of Nalut were taken from localized concentrations of
vegetated dunes near the escarpment of the coastal plain. Fresh
burrows were abundant in the sand throughout the traplines, sug-
gesting a high population density. Gerbillus aureus and Meriones
caudatus were also collected from this same habitat.
The name tripolitanicus refers to the range ascribed to this sub-
species in northern Tripolitania Province.
Jaculus jaculus whitchurchi, new subspecies
Holotype. Adult male, skin and skull, USNM 325807, from 10

km S Agedabia, Cyrenaica Province, Libya; obtained June 18, 1962,
Specimens examined. Thirty-one, from Tripolitania: 12 km W
Sirte, 3; 15 km WNW
Marble Arch, 1; from Cyrenaica: 10 km
S Agedabia, 17; Giarabub, 8; 24 km SSE Giarabub, 1; Bahr el Tubat,
21 km ESE Giarabub, 1.
Measurements. The averages and extremes of 12 males and 3

females from the type locality, with the measurements of the type in
brackets, are, respectively: Total length 287.6 (280-296), 300.3
(294-305), [282]; length of tail 178.6 (174-182), 186.3 (184-190),
[177]; length of hind foot, 62.3 (59-64), 62.7 (60-65), [62]; length of
ear 21.4 (19-23), 21.7 (20-23), [21]; greatest length of skull 33.2
(31.8-34.5), 34.1 (33-34.8), [32.8]; condyloincisive length of skull
29.4 (28.5-30.4), 29.9 (29.3-31.1), [28.7]; crown length of upper
molariform toothrow 5.1 (4.7-5.4), 5.2 (5-5.4), [5]; greatest breadth
across zygomatic processes 21.7 (20-22.5), 23 (22.5-23.8), [21.9];
least interorbital breadth 12.4 (11.8-13), 12.6 (12.4-13), [12.4]; length
of rostrum at level of antorbital foramina 4.8 (4.5-5.3), 4.8 (4.6-5),
[4.8]; greatest breadth of braincase 18 (17.1-18.9), 17.8 (17.5-18.2),
[17.1]; greatest breadth across antorbital processes 22.4 (21.3-23.8),
22.8 (22.6-23), [22.3].
Diagnosis. Upperparts Cinnamon-Buff becoming darker on rump

owing to admixture of brownish hairs; rostral, mystacial, circum-
orbital, pectoral, and postauricular areas Light Buff; flanks and
lateral areas with slight admixture of brownish hairs; entire under-
pays, dorsum of front and hind feet and terminal portion of tail
white; pinna of ear sparsely furred and Clay Color; hind feet,ventrally,
heavily furred and frequently stained Pale Orange-Yellow; tail in-
distinctly bicolored, Pale Yellow-Orange dorsally and Light Buff
ventrally; pinnate tail sharply demarcated into a Bister anterior
portion and white posterior portion. Skull: Small; triangular in shape;
braincase prominently vaulted; zygomata convergent anteriorly;
auditory bullae moderately inflated; lachrymals relatively large;
rostrum narrow; zygomata relatively fragile; foramen on anterior
margin of angular process enlarged longitudinally; coronoid process
markedly reduced.
Comparisons. When compared with topotypes of Jaculus jaculus
jaculus, J. j. body size, shorter tail,
whitchurchi differs in smaller
shorter hind foot, markedly shorter ears, and generally smaller size
of cranium. Specimens of J. j. whitchurchi are paler and more uniformly
colored dorsally and have less suffusion of brown on the sides and
flanks.
For comparisons with J. j. tripolitanicus , J. j. collinsi, and J. j.
arenaceous, see accounts of those subspecies.

Remarks. This new subspecies of jerboa is largely restricted to
the Libyan littoral desert along the Gulf of Sirte. Specimens from
Giarabub, however, are referable to this subspecies and occur more
than 200 kilometers from the coast. Apparently, the vast hamada
north and east of the Sand Sea of Calanscio provides suitable habitat
for jerboasand allows for gene flow between these coastal and interior
populations. North and east of the Gulf of Sirte, dispersal is rendered
impossible by the escarpments of the Cyrenaican Plateau and Gebel
el Achdar. Widespread areas of sand between Gialo Oasis and the
coast also preclude interbreeding between J. j. whitchurchi and J. j.

collinsi.
The type series was obtained from a complex of heavily vegetated
coastal dunes about one-half kilometer from the Mediterranean coast.
Specimens from Sirte and Marble Arch were collected from the
alkaline-clay soils of the coastal plain and farther removed from water.
A specimen, 325802, from Giarabub, Cyrenaica, is markedly paler
and more uniformly colored dorsally than any other jerboa examined
from Libya. This particular specimen is an old female and probably
represents an extreme color variation incident to age.
J. j. whitchurchi is proposed in recognition of Lt. Colonel Thomas
Whitchurch, former Commanding Officer of the 64th Engineering
;
Battalion, Wheelus Air Force Base, Tripoli, who provided support

essential to the field work in Libya.
Jaculus orientalis orientalis Erxleben
Jaculus orientalis Erxleben, Syst. Regni Animalis, p. 404, 1777 (Egypt, in the
mountains separating Egypt from Arabia).
General distribution of species. Egypt (including Sinai),

Libya, Algeria, Tunisia, and Morocco.
Distribution in Libya. Coastal areas of Cyrenaica and Tripolitania
and inland to Augila (Gialo Oasis).
Specimens examined. Twenty-five, from Cyrenaica: 12 km NW
Gubba, 1 5 km; NW
Labrag, 1 2 km ; W
Tocra, 1 (skull only) 2 km N ;
Coefia, 2; 8 km N Benghazi, 1 Gheminez, 18 (1 skin only; 1 skeleton)

;
from Tripolitania: 20 km E Rumia, 1.

Published records in Libya. Cyrenaica: Sidi Faradie (Thomas,
1902); Gheminez (Festa, 1921) Mechili (Festa, 1925); Agedabia, El
Agheila, Augila (de Beaux, 1932); Zauia Mechili (de Beaux, 1938);
Tripolitania: Tripoli (Klaptocz, 1909); Tripolitania (Toschi, 1951).
Measurements. The averages and extremes of 14 adult males and
4 adult females from Gheminez, Cyrenaica Province, and the measure-
Jaculus orientalis orientalis
Figure 51. — Distribution of Jaculus orientalis orientalis.

ments of an adult male, 325835, from 20 kilometers east of Rumia,

Tripolitania Province, are, respectively; Total length 370.7 (354-382),
361 (354-372), 362; length of tail 222.1 (204-230), 215.5 (214-217),
210; length of hind foot 76.5 (75-78), 73.5 (71-76) 79; length of ear
from notch 32.8 (30-36), 31.5 (30-35) 30; greatest length of skull 38.4
(36.6-39.4), 37.9 (37.2-39.2), 38.1; condyloincisive length of skull 35
(33.7-36.1), 34.8 (34.6-35.1), 35.1; crown length of upper molariform
toothrow 5.9 (5.5-6.1), 6.1 (6.1-6.2), 6.5; least interorbital breadth
14.1 (13.6-15.1), 14.2 (13.8-14.6), 14.4; length of nasals 14.8 (13.9-
15.6), 14.5; breadth of rostrum at level of ant-
13.8 (13.6-14.2),
orbitalforamina 5.8 (5.4-6.4), 5.5 (5.2-5.9), 6.3; greatest breadth of
braincase 20.4 (19.4-21.2), 20.4 (19.8-21.1), 20.8; greatest breadth
across antorbital processes 27.7 (25.4-29.1), 27.5 (27.2-28), 28.1.
Diagnosis. Upperparts and flanks Clay Color with strong suffusion
of dark hairs resulting in a streaked or marbled appearance; this effect
more prominent on rump; sides, ventrolateral and scapular areas
Pinkish Buff with strong admixture of brownish-black hairs; under-
pays and forearms white but suffused with Pale Pinkish Buff; sub-
auricular, postauricular, and postorbital areas Cinnamon-Buff grading
to Pale Pinkish Buff in circumorbital and mystacial areas; all areas,
except those which are white, with strong admixture of black hairs;
pinnae of ears sparsely haired and Sepia on internal and lateral
aspects, Tawny Olive on medial margin; vibrissae long, proximally
dark in color (Fuscous) and white distally; forelegs and hindlegs
sparsely haired and whitish dorsally; fore feet naked ventrally and
bearing five claws; hind feet bearing three claws, heavily furred
ventrally with dark hairs (Clove Brown) concentrated on sole and
extending along ventral surface of metatarsus; digital areas Pale
Pinkish Buff; tail indistinctly bicolored Clay Color dorsally and Light
Buff ventrally with terminal pinna, transversely bicolored, Fuscous
Black proximally and white distally. Skull: Large and robust; brain-
case slightly vaulted; molariform teeth large; zygomata heavy and
bearing large, winglike antorbital processes; auditory bullae mod-
erately large; anterior palatine foramina expanded posteriorly and
laterally; nasals flaring anteriorly; rostrum relatively wide and heavy;
angular process of mandible with large, single oval foramen on its
anterolateral surface.
Comparisons. From Jaculus orientalis orientalis, as known from
Mersa Matruh, Western Desert Governorate, Egypt, the series from
Gheminez differs in having slightly larger size of most external and
cranial measurements but in being smaller or of comparable size in
length of ears and breadth of rostrum at level of antorbital foramina.
Jerboas from Gheminez also have slightly larger lachrymals, and the
anterior palatine foramina are less expanded posteriorly. The two
246 U.S. NATIONAL MUSEUM BULLETIN" 2 75
populations agree closely in color, but specimens from Gheminez

have slightly less admixture of dark hairs on the rump.
Compared to a near topotype (BM, no. 12.6.12.170), of Jaculus
mauritanicus Duvernoy from Guelt es Stel, Central Plateau,
orientalis
Algeria,and a specimen (BM, no. 25.6.3.8) representing J. o. mauri-
tanicus from near Sfax, Tunisia, the Libyan specimens are more
uniformly colored dorsally, have less suffusion of Ochraceous-Salmon
on the sides and flanks, are somewhat smaller in total length and
length of tail, have narrower and longer rostra, and are slightly
smaller in the length of skull.
Jaculus orientalis, in Egypt and Libya, although resembling Jaculus
blanfordi (Murray) in some superficial characters, differs markedly in
all of the salient features. Specimens of the latter species as known
from Majan (50°02' E, 32°35' N), Dasht-i-lut, and near Jangal,

Khurasan Province, Iran, differ from those of J. orientalis from
Gheminez in smaller size of every cranial and external measurement.
Jaculus blanfordi also differs from J. orientalis in darker, more uni-
form dorsal color, more distinctly bicolored tail, less buffy sides
more heavily washed with gray, smaller terminal spines on baculum,
darker vibrissae, whiter underparts (as opposed to Pale Pinkish Buff),
proportionately larger front feet and claws, and darker, almost black,
proximal portion of pinna of tail. Cranially, J. blanfordi has a more
flattened skull, proportionately more fragile zygomata, smaller molari-
form teeth, relatively larger and more inflated auditory bullae, and
two, rather than one, distinct, oval foramina in the angular process
of the mandible.
Remarks. An adult male, 325835, from 20 kilometers east of Rumia,
Tripolitania Province, differs significantly from the series from Mersa
Matruh and other specimens from Libya, in that it is much darker
dorsally with markedly stronger admixture of dark hairs, resulting in
a streaked or marbled appearance. The sides and pectoral areas are
more heavily suffused with Ochraceous-Salmon. Cranially, this speci-
men from Tripolitania differs from typical Jaculus orientalis orientalis
in having a wider rostrum, smaller lachrymals, narrower anterior
palatine foramina, larger foramen in angular process, longer cheek
teeth with more laterally directed occlusal surfaces, and lateral
margins of antorbital processes more abruptly curved.
In the above characters, this specimen from near Rumia shows
intergradation with Jaculus orientalis mauritanicus, whose range in-
cludes northern Algeria and Morocco. No topotypical specimens of
the latter subspecies are available for study, but Loche (1867, pp.
98-99) gives the following description: "The jerboa of Mauritania
isvery large, very powerful and darker in color (more russet) than
the jerboas of Tripoli. The underparts, the throat, and the extremities
.
dorsum is more streaked or marbled.

are light yellow or russet, and the
The brownish rust above with some blackish hairs at the tip
tail is
(6 or 7 cm)." Jaculus o. mauritanicus apparently also has a shorter,

wider muzzle, a larger head, smaller ears in proportion to the body,
and generally a larger, more robust body. The jerboa from Rumia
shows some of these characters, particularly in width or rostrum,
streaked and darker dorsum and proportionately smaller ears, but in
general body size and in most cranial characters, it is closer to the
nominate subspecies and is here so referred. The referral of this
specimen from Tripolitania to J. o. orientalis is, however, provisional
until such time as adequate series of these large jerboas become
available from western Libya and permit a more thorough study.
The type locality of J. o. orientalis is not firmly fixed but is believed
to be somewhere in the mountains separating Egypt from Arabia
and thus probably near Sinai. Only one subadult specimen is available
from Sinai, however, and thus valid comparisons with jerboas from
Libya are not possible. It is doubtful if the large jerboas inhabiting
the coastal areas of Sinai, Egypt, and Libya are all of the same
subspecies. Setzer (1957) also questioned the occurrence of a single
subspecies over such a wide geographical area.
Local differences in pelage color of these large jerboas are notice-
able throughout their range in Cyrenaica. These differences are
probably attributable to differences in age, sex, or seasonal change.
Much larger series are required to determine the exact taxonomic
significance of color in these large jerboas.
Ecological observations. In Libya, these large jerboas are
confined almost exclusively to the more humid environment of the
coastal plain. Occasionally, specimens were taken from hamadas
farther inland, and on one occasion, in Tripolitania, one was collected
from the brink of the coastal escarpment. This species was recorded
by de Beaux (1932) from Augila, Cyrenaica Province, at a distance
somewhat removed from the coastal plain. The elevation at Augila,
however, is relatively low and accordingly the general habitat here
is similar to that of the coastal plain.
These jerboas are difficult to catch by standard trapping procedures
and trap yields are usually low. Setzer (1957) was able to obtain
the series from Gheminez only by digging the animals from their
burrows. At this time, he also obtained several specimens of the
four-toed jerboa (Allactaga tetrad a cti/la)
On two occasions, adult specimens were picked up some distance
from a trapline. Apparently, these animals had been struck on the
head by a trap but were able to move some distance before dying.
Museum special traps are not of sufficient size to immediately kill
and hold animals of this size.
On several occasions these jerboas were observed hopping about

the dense vegetation of the Cyrenaican coastal plain near Tobruch
shortly before dusk. They appear to be most active immediately
following dusk.
Genus Allactaga Cuvier
Allactaga tetradactyla (Liechtenstein)
Dipus tetradactylus Lichtenstein, Verz. Doubl. Mus. Berlin, p. 2, 1823 (near

Alexandria, Egypt).
General distribution of species. Coastal hamadas of Egypt

and eastern Libya. In Egypt, this species is known from the type
locality, Mersa Matruh, Burg el Arab, Bahig, Sidi Barrani, and
Mariut (El Amiriya).
Distribution in Libya. Known only from Gheminez, El Agheila,
and Zauia Mechili.
Specimens examined. Three, all from Gheminez, Cyrenaica
Province.
Published records in Libya. Cyrenaica: Mechili (Festa,

1925); El Agheila (de Beaux, 1932); Zauia Mechili (de Beaux, 1938).
Measurements. The measurements of two adult males, 302294
and 302295, and an adult female, 302293, from the above locality,
are, respectively: Total length ?, 295, 293, length of tail ?, 179, 176;
length of hind foot 57, 59, 58; length of ear 42, 44, 43; greatest length
of skull 32, 32.3, 31.6; condyloincisive length of skull 29.4, 29.9,
29.1; crown length of upper molariform toothrow 5.9, 6, 5.9; greatest
breadth across anterior zygomatic processes 22.2, 22.5, 21.5; least
interorbital breadth 9.8, 9.8, 10.2; length of nasals 11.4, ?, 11.1;
breadth of rostrum at level of antorbital foramina 5.5, 5.6, 5.7;
greatest breadth of braincase 18.9, 19, 18.9.
Diagnosis. Dorsum generally dark (Drab) becoming lighter
(Cinnamon) on rump; entire dorsum strongly suffused with grayish
hairs and appearing uniformly variegated, the latter becoming more
pronounced on sides owing to a blending of white and blackish hairs;
flanks, preauricular, subauricular, and pectoral areas Cinnamon-
Buff; distinct Light Buff patch between eye and base of ear; mystacial,
suborbital, and circumoral areas Pale Ochraceous-Salmon; vibrissae
relatively short and uniformly colored Mummy Brown; ears very large;
base of pinna Pinkish Cinnamon grading to Drab on sparsely haired
distal and medial portion; internal face of pinna lacking hair and Dark
Olive; forefeet sparsely haired and white dorsally, heavily furred
ventrally with sole and metatarsal area Bister and bearing three
primary digits with claws and one rudimentary digit and claw applied
to metatarsus; tail Sayal Brown dorsally, becoming slightly lighter
ventrally and with terminal pinna, transversely bicolored dark brown
proximally and white distally; entire underparts white. Skull: Small
and compact; braincase, particularly the parietals, markedly inflated,
sloping markedly anteriorly from coronal suture to front margins of
nasals; medial portions of frontals forming a distinct depression in
median line; squamous portion of temporal projecting markedly laterad
thus rendering a "bowed" appearance to zygomata; zygomata rela-
tively fragile and markedly convergent anteriorly; antorbital processes
reduced; lachrymals relatively large; infraorbital foramina relatively
large; rostrum wide; upper incisors small and short; upper molars
large and directed laterad; small rudimentary premolar applied to
anterior surface of first upper molar; anterior palatine foramina large
and markedly expanded posteriorly; pterygoid hamulae heavy and
knobbed; posterior palatine canal large and elongated; palate heavy,
with distinct spine directed caudad; auditory bullae markedly small
and but little inflated; basioccipital wide, occiptal condyles and
jugular processes large; mandible large and bearing a deep fossa on
285-134 O— 68 17
ventral margin of coronoid process; angular process reduced and with a

single trianguliform foramen on an tero ventral margin; molariform
teeth large, lower incisors thin and attenuated.
Comparisons. Specimens from Gheminez are slightly larger
cranially and in external body measurements than specimens from
Burg el Arab, Western Desert Governorate, Egypt. In color, animals
from Libya are more uniform and less streaked dorsally, have more
darkly colored soles of the hind feet and metatarsal areas, have a more
prominent buffy patch above the eye, and are less suffused with buff
on the sides and pectoral areas.
Remarks. The above differences are of insufficient magnitude
to be of taxonomic significance and probably fall within the range of
variation for the species. In view of the wide geographic range of
Allactaga tetradactyla, it has undergone comparatively little local
differentiation. Setzer (1958) referred to this genetic stability and
was also unable to demonstrate significant variation either between
or within populations of this kind of jerboa. Apparently, suitable
habitat occurs uninterruptedly throughout its range and, as a con-
sequence, there has been little interruption of gene flow. When
adequate series of A. tetradactyla become available, however, some
variation undoubtedly will be demonstrable, and this species will
prove to be polytypic.
Ellerman and Morrison-Scott (1951) did not include Libya within
the range of A. tetradactyla. Apparently, they were unaware of the
collections reported by Festa (1925) and de Beaux (1932) from Zauia
Mechili and El Agheila. The present study contributes but little to
the knowledge of the distribution of this kind of jerboa. The few
collecting localities in Cyrenaica do suggest a limited occurrence
in Libya but probably do not provide an accurate distributional
picture. As with most uncommon species, the few collecting sites
indicate a fortuitous meeting of collector and animal and do not
render an accurate estimate of the population size.
Ecological observations. The four-toed jerboa inhabits the
hamadas of the Mediterranean littoral. These coastal deserts are
relatively free of sand and are composed of boulder-strewn plains
where vegetation grows sparingly or is frequently wanting. These
jerboas apparently dislike habitats of a sandy character and accord-
ingly are unknown from the Saharan interior. The collecting locality
at Zauia Mechili is still well within the zone of coastal hamadas
and has no real relationship to the Sahara proper.
Setzer (1957) obtained the small series from Gheminez by digging
the animals from their burrows but was unable to obtain specimens
by trapping. Collecting efforts by the author were to no avail.
Family Hystricidae
Genus Hystrix Linnaeus

Hystrix cristata Linnaeus
Hystrix cristata Linnaeus, Syst. Nat. 10th ed., vol. 1, p. 56, 1758 (near Rome,
Italy).
Remarks. I have not examined any Libyan specimens of the

porcupine, and judging from the few records available in the litera-
ture, they occur only sparingly in the mountainous areas near the
coast. They are apparently unknown from the deserts and oases
of the interior.
Figure S3. — Known localities of occurrence of Hystrix cristata.
While we were camped in the Gebel es Soda of southern Tripoli-

tania, a local bedouin mentioned having frequently seen porcupines
in the surrounding mountains, but I observed no signs of their presence
here or elsewhere in Libjâ. Setzer (1957) found spines of the porcupine
at Wadi Ahmar, 40 kilometers southeast of Benghazi, in Cyrenaica,
but obtained no specimens.
Records of porcupines from Gharian (Klaptocz, 1909) and Barce

(Festa, 1921) indicate that they occur on the Cyrenaican Plateau,
as well as the Tripolitanian Gebel, and probably are commonest in
these mountainous regions of Libya. The broad, chaparral-covered
valleys of the Cyrenaican Plateau and the rocky canyons of the
coastal escarpment in Cyrenaica and Tripolitania probably provide
the most suitable habitats for porcupines. A specimen from Derna
in northern Cyrenaica (de Beaux, 1938) constitutes the only known
record from the coastal plain proper, but the coastal escarpment near
Derna approaches near to the sea, almost obliterating the coastal
plain, and probably this specimen was taken from the escarpment
rather than from the coastal plain.
Family Ctenodactylidae
Genus Ctenodactylus Gray

Ctenodactylus gundi (Rothman)
Mus gundi Rothman, Schloezer's Briefwechsel, p. 339, 1776 (Gharian, 80 km S

Tripoli, Libya).
General distribution of the species. Libya, Tunisia, Algeria,

and Morocco.
Distribution in Libya. Mountainous areas and the larger wadis
of northern Tripolitania.
Ctenodactylus gundi gundi. Tripolitania: The Gebel Nefusa,
Gebel Tigrinna, Gebel Jefren, and other parts of the Tripolitanian
Gebel.
Ctenodactylus gundi vali. Tripolitania: The transitional desert
between the Gebel es Soda and the Gulf of Sirte.
Published records in Libya. Tripolitania: Gharian (Yarrel,
1831; Klaptocz, 1909; Toschi, 1951); Wadi Titti, Bu Ngem (Thomas,
1902); Wadi Soffegin at Orfella (Toschi, 1951).
Remarks. In the past, the gundis of Libya have been variously
regarded as two distinct species, Ctenodactylus gundi and Ctenodactylus
vali (Thomas, 1902; Klaptocz, 1909; Zavattari, 1934; Toschi, 1951;
Petter, 1961), or considered as one species, C. gundi, with C. vali
treated as a subspecies (Ellerman and Morrison-Scott, 1951; Toschi,
1954; Setzer, 1957).
Petter (1961) regarded C. vali and C. gundi as separate species but
used sympatry as his primary argument in establishing their specific
identity. He apparently misinterpreted Toschi (1951) and presumed
that the collecting site of C. vali in the Wadi Soffegin and the collecting
gundi on the Gebel Nefusa were part of the same plateau
sites of C.
complex. Actually, the ranges of C. vali and C
gundi are geographi-
*,
such as the Wadi Bey, Wadi Soffegin, and Wadi Titti located farther
to the southeast.
Ctenodactylus gundi gundi (Rothman)
Mus gundi Rothman, Schloezer's Briefwechsel, p. 339, 1776 (Gharian, 80 km S

Tripoli, Libya).
Specimens examined. Twenty-nine, from Tripolitania: 5 km W

Cussabat, 24 (1 skeleton); 20 km E Rumia, 5.
Measurements. Averages and extremes of five adult males and
seven adult females from 5 kilometers west of Cussabat, are, respec-
tively: Total length 221.4 (200-240), 232 (220-245); length of tail
22.8 (20-25), 24.3 (20-30); length of hind foot 40.6 (38-44), 37.6
(35-44); length of ear 18.8 (18-20), 19.3 (18-20); greatest length of
skull 48.3 (47-49.3), 48.3 (46.4-51.4); basilar length of skull 36.8
(35.4-38.1), 37.7 (36.5-39.7); greatest breadth across zygomatic arches
32.2 (30.7-33.4), 31.8 (30.3-33.2); length of nasals 19.4 (18.4-20.7),
20.3 (18.5-21.6); greatest breadth of nasals 6.2 (5.8-6.5), 6.5 (6.4-6.7);
least interorbital breadth 12.1 (11.8-12.6), 12.3 (11.5-12.6); breadth
of interparietal 12 (11.4-12.6), 11.7 (11.4-12.3); length of diastema
12 (11.3-12.5), 12.1 (11.6-12.8); length of anterior palatine foramina
7.8 (7.4-8.2), 8 (7.7-8.4); crown length of upper molariform toothrow
9.5 (8.4-9.9), 9.7 (9.4-10.2); oblique length of auditory bulla 16.8
(16-17.6), 16.9 (16.2-17.5); height of auditory bullae 14.4 (13.7-15.3),
14 (13.6-14.5); jugular breadth 19.5 (18.8-20.6), 19.2 (18.1-20).
Diagnosis. Entire pelage silky and lustrous, Cinnamon-Buff dor-
sally with moderate suffusion of brown on middorsal and facial re-
gions, and Pinkish Buff ventrally with frequent exposure of the
Plumbeous underfur; long brown guard hairs of the dorsum extending
noticeably beyond level of underlying pelage; circumoral area and
chin slightly paler than surrounding pelage and approaching Light
Ochraceous-Buff; dorsal surfaces of fore and hind feet Light Buff
with mild suffusion of Warm Buff; ventral surfaces of fore and hind
feet black and naked, with prominent metacarpal and metatarsal
tubercles, and with thickened pads at the bases of the short black
claws; each appendage with four functional digits; each digit of the
hind foot enveloped terminally by stiff, bristle-like hairs, some of
which are markedly hardened and resemble claws; vibrissae long,
black, and extending posteriorly beyond level of the ears; pinna of
ear small and inconspicuous, round in shape and with black inner
surface; outer rim of pinna approaching Light Buff and contrasting
markedly with the darker color of the surrounding pelage; tail
markedly short and uniformly colored Warm Buff. Skull: Large,
massive, and extremely angular with heavy parietal ridges and zygo-
mata; nasals wide and slightly flaring anteriorly; lachrymals and dor-
sal portion of the jugals greatly expanded, forming a large portion of

the wide antorbital plate; maxillary component of the antorbital plate
narrow, owing to the extremely large size of the infraorbital foramen;
parietal ridge terminating in a conspicuous flattened protuberance on
posterior margin of the orbit; interparietal large and broadly elliptical;
auditory bullae heavily ossified and irregular in shape; audital portion
of bulla mildly inflated and meatal process heavy and projecting con-
spicuously laterad mastoidal portion of bulla moderately inflated and
;
extending posterior to occiput; portion of bulla above meatus ex-

panded into a rounded prominence between the mastoidal portion of
the bulla and the squamous portion of the temporal bone; jugular
processes large with distal portions directed anteromedially; basioc-
cipital broadly wedge-shaped and with faint median rib; pterygoid
processes rounded on medial aspect and hamulae applied to the styli-
form processes of the auditory bullae; basisphenoid present as a dis-
tinct rod-shaped structure between the pterygoid processes, which
become progressively narrower anteriorly; pterygoid fossae large,
partially enclosed ventrally by the palatine bones, and with large
foramina on their posterolateral portions; mandibular fossae large and
deeply grooved; upper molariform teeth with shallow kidney -shaped
crowns; anterior palatine foramina extremely large and bowed lat-
erally; upper incisors relatively small, with a decided notch distally
and with plain outer faces; mandible coarse and angular and with a
distinct medial curvature; lower molariform teeth kidney-shaped and
with conspicuous styles on medial surfaces; coronoid process reduced
to a slight elevation of the ramus; angular process small and atten-
uated; lower incisors small and distinctly chisel-shaped.
Comparisons. Specimens representing Ctenodactylus gundi gundi
from near Rumia and Cussabat on the Gebel Nefusa, northwestern
Tripolitania, differ strikingly from a single to po type of Ctenodactylus
gundi vali from the Wadi Bey, northwest of Bu Ngem, Tripolitania,
in having smaller and less inflated mastoidal portions of the auditory
bullae, narrower rostra, larger and wider infraorbital foramina, wider
interorbital breadths, and heavier angular process of the mandible.
The skull of the specimen from Bu Ngem is badly broken, and, conse-
quently, most cranial measurements could not be taken. By inference,
however, C. g. gundi would seem to be slightly larger in most all of
the standard cranial measurements. Thomas (1902), in the original
description of Ctenodactylus vali, gives the following characters by
which vali differs from C. g. gundi: Auditory bullae larger; nasals
longer and narrower; antorbital projections more delicate; inter-
parietal narrower, molars smaller and more delicate (shorter antero-
posteriorly), and the last molar less distinctly L-shaped than in
C. g. gundi. In external dimensions, representatives of C. g. gundi
from northwestern Tripolitania are generally larger than the specimen

of C. g. vali from the Wadi Bey, the two forms being closest in the
length of the hind foot and ear. Except for total length, the external
measurements of the type specimen of C. g. vali, as given by Thomas,
are also significantly smaller. In Ctenodactylus gundi vali, the pelage
is longer, silkier, slightly paler in color, and more evenly suffused
with brown hairs. The venter is paler than in C. g. gundi and has
less suffusion of buff or tan. The pinna of the ear in C. g. vali is brown
on its inner surface rather than black as in C. g. gundi, has a thicker
covering of hair, a more prominent tuft of buffy hairs on the anterior
margin, and the color of the hair on its outer surface is similar to
that of the dorsum and not markedly contrasting in color, as in
C. g. gundi.
A
specimen, 122110 (sex unknown), from Gafsa, Tunisia, agrees
rather closely in cranial features and measurements with those of
C. g. gundi from Tripolitania but is noticeably wider in interorbital
breadth. External measurements of this Tunisian specimen are not
available, but judging from the relative size of the prepared skin,
it appears to be comparable in size to typical C. g. gundi from Libya.
In color, it is somewhat darker dorsally, paler ventrally, and has more
admixture of yellow on the dorsal surfaces of the feet. These few
differences are not sufficient for separation at even the subspecific
level, and this specimen is here referred to C. g. gundi.
Kemarks. The five specimens from the Gebel Nefusa, 20 kilo-
meters east of Rumia, Tripolitania, are near topotypes of Ctenodactylus
gundi gundi, and in addition to the large series of gundis from 5 kilo-
meters west of Cussabat, Tripolitania, represent the first records of
this subspecies in northwestern Tripolitania since those reported
from the type locality (Klaptocz, 1909).
The large series of gundis from near Cussabat probably contains
the greatest number of specimens (24) ever collected from a single
locality in Libya and in number probably exceeds all those previously
collected in Libya.
While I was at Sebha Oasis in the Fezzan in 1962, an engineer
employed by U.S.O.M. (United States Overseas Mission) mentioned
having seen, earlier that year, "wild guinea pigs" in the rocky out-
croppings near the village of Cussabat in northern Tripolitania. His
description of them, although quite fragmentary and incomplete,
suggested characters of the gundi, particularly as guinea pigs were
not known to occur in Libya, at least not in the wild state. This brief
discussion at Sebha prompted me, later that year, to search the vicinity
of Cussabat and led eventually to the discovery of this large popula-
tion of gundis.
The collecting site on the Gebel near Cussabat is located approxi-

mately 110 kilometers northeast of the type locality of C. g. gundi at
Gharian and 135 kilometers northeast of the collecting locality near
Rumia and is part of the same escarpment which sharply delineates
the inner margin of the Tripolitanian coastal plain. The escarpment at
Cussabat is neither as extensive nor as high as the gebel at Gharian
and Rumia, but the character of the rocky outcroppings and coarse
talus are similar in the two areas. Gundis from Cussabat and those
from Rumia differ more than would be expected, however, in view
of the similarity in terrain, the ecological continuity of the escarpment,
and the relatively short distance separating them.
Specimens from Cussabat and Rumia are almost indistinguishable
in the general shape and configuration of the skull, and both clearly
represent C. g. gundi. In cranial and external measurements, however,
those from near Cussabat are slightly larger in total length and length
of ears, least interorbital breadth, and length of anterior palatine
foramina. They are slightly smaller in length of tail and hind foot,
width of interparietal, height of auditory bullae, and jugular breadth.
In specimens from Cussabat, the angular process of the mandible is
heavier (less attenuated), the pelage is more worn, shorter, thinner,
less fluffy and silky and has less suffusion of brown hairs in the mid-
dorsal region. Ventrally, they are less buffy, being grayish rather
than tawny, and have darker tails distally. The specimens from near
Rumia were obtained in early March and presumably were in winter
pelage, and those from Cussabat were taken in late June and con-
sequently were in summer pelage. The differences in color and luster
of the fur between specimens from these two areas may thus reflect
seasonal characteristics of the pelage and may not indicate actual
differences.
This subspecies attains its easternmost limits of distribution in
northwestern Libya where it is known from only the vicinity of the
type locality on the Gebel Nefusa and from near Cussabat. Its range,
however, doubtless includes all of the coastal gebel as far east as the
western margins of the Gulf of Sirte, south of which this subspecies
probably intergrades with C. g. vali. The western limits of its range
are unknown, but this subspecies probably occurs widely throughout
the mountains and escarpments of Tunisia and the eastern portions
of the AtlasMountains in Algeria. Gundis are unknown from the
Hamada el Hamra and the Hamada de Tinrhert of the Tripolitanian
interior. Apparently suitable habitat does not occur in Libya this
far inland.
Ecological observations. Near Rumia, gundis were occupying
and interstices of large rocks which together form the face
fissures
of the coastal escarpment. All specimens were collected near the

brink of this escarpment where it becomes less abrupt and transforms
into the high rolling uplands of the Gebel Nefusa. The coastal escarp-
ment is varied in its degree of steepness and surface configuration.
Frequently well-defined terraces exist on its slopes, into which have
accumulated irregular masses of boulders and smaller rocks, all of
which provide suitable habitat for the gundi. The less precipitous
slopes of the larger wadis, which dissect the Gebel Nefusa and descend
through the coastal escarpment onto the coastal lowlands, have ir-
regular cliffs, rocky outcrops and boulder-strewn areas, which are ideal
for the habitat requirements of the gundi. The fact that specimens
were collected and observed only in the higher portions of the gebel
near Rumia is a direct result of my efforts having been concentrated
there and does not indicate that gundis are most abundant at higher
elevations or that they do not occur at lower elevations.
Near Cussabat, gundis were abundant in the rocky hillsides of the
easternmost limits of the Gebel Nefusa. The escarpment here is lower
in elevation and more gradual in slope, and rocky areas are more
sporadic and localized. A large series of gundis was obtained from the
ledges and rocky outcroppings near the valley floor. Vegetative cover
was denser here than at Rumia and consisted primarily of clumps of
perennials scattered among the rocks and cliffs.
Vegetative cover among the rocks and boulder fields, although never
dense, is surprisingly uniform. Several species of succulent, herbaceous
perennials are widespread in these rocky areas, but grasses are almost
always the dominant type of vegetation. Freshly cut stems and leafy
portions of these plants, mingled with fecal droppings, were frequently
observed on rocky ledges and beneath boulders and indicated the
presence of gundis in a given area.
Hunting these animals with a shotgun proved to be the most effec-
tive method. They are usually extremely wary and appear for only
short periods of time in the sunlight or in the shaded recesses of large
rocks, rocky ledges, and projections. On one occasion on the gebel near
Rumia, three gundis were shot in rapid succession, each one appearing
on the same rocky ledge as its predecessor after the latter had fallen
dead into a nearby crevice. Presumably these rodents are exclusively
diurnal and reach a peak in their activity during the hours of mid-
morning have warmed the ledges and feeding
after the sun's rays
stations. They seem have a secondary surge of activity in the late
to
afternoon after the heat of the day has subsided. In the middle of the
day, when maximum temperatures prevail, they remain in the sanc-
tuary of their nests in crevices and fissures of the rocky outcroppings.
Even though the gundis near Rumia were collected in late winter
(early March), daytime temperatures reached 98° F and, at night,
did not drop below 45° F. At Cussabat, which we

visited during early
summer (late June), the maximum daytime
temperatures were even
higher and exceeded 100° F, and at night the minimum temperature
was 57° F. The relative humidity, for the two days spent at Cussabat,
was 72 and 70 percent respectively. To what extent these high tem-
peratures and the high humidity influenced the behavior of the gundis
is not known, but certainly the high daytime temperatures induced
them to remain more inactive during the middle of the day.

Gundis do not share these rocky habitats with any other species of
large rodent, but near Rumia, several specimens of Gerbillus campestris
were taken from traps set among rocky outcrops known to contain
gundis, and at Cussabat, an elephant shrew (Elephantulus Thomas and
Schwann) was obtained from a rocky surface over which gundis
frequently travelled.
A skull of a young female, 325855, from near Cussabat, shows two
distinct bregmatic bones in the middorsal region near the fronto-
parietal suture.
In general appearance, habits and habitat preferences, the gundi
resembles the pikas of the genus Ochotona, which are entirely unrelated
and confined to temperate Asia and North America.
Ctenodactylus gundi vali Thomas

Ctenodactyhis valiThomas, Proc. Zool. Soc. London, pt. 2, p. 11, October 1902
(Wadi Bey, northwest of Bu Ngem, Libya).
Specimens examined. One, from Wadi Bey, 45 km W Bu Ngem,

Tripolitania.
Measurements. The external measurements of the above specimen,
an adult female, 302296, are: Total length 178; length of tail ?; length
of hind foot 38; length of ear 19. No cranial measurements are avail-
able.
Diagnosis. Entire pelage extremely long, fluffy and silky, Light
Pinkish Cinnamon dorsally, uniformly suffused with brown and black-
tipped hairs, and Light Buff ventrally with faint admixture of Warm
Buff; characters of the facial region, fore and hind feet, and ear
essentially as in C. g. gundi, but hind feet more nearly white dorsally
and color of outer margin of pinna of ear darker and more nearly
approaching that of the dorsal pelage. Skull: Similar in general shape
and configuration to that of C. g. gundi but with markedly larger and
more inflated mastoidal and audita! portions of the auditory bulla,
less flaring nasals anteriorly, and smaller infraorbital foramina.
Comparisons. For comparisons with Ctenodactylus gundi gundi, see
account of that subspecies.
Remarks. Thomas, in 1902, described Ctenodactylus vali from the
Wadi Bey near Bu Ngem, Tripolitania, and examined specimens from
the Wadi Titti, east of Socna. More recently, Toschi (1951) recorded a
specimen from the Wadi SofFegin and assigned it to C. g. vali. Setzer
(1957) recorded the most recent specimen of C. g. vali taken from the
type locality at the Wadi Bey. These few specimens constitute the
onlyknown records of this subspecies in Libya.
The Wadi Bey is located about 300 kilometers southeast of the type
locality of Ctenodactylus gundi gundi, but the range of C. g. vali
probably includes most of the arid transitional desert between the
Gebel es Soda and the coast. Most of the broad wadis in this region,
which drain northward into the Gulf of Sirte, provide suitable habitat.
The drier climate of these interior localities and the irregular scarps
which are present on the margins of these wadis differ markedly from
the more humid climate of the Gebel Nefusa and its higher and more
imposing escarpments. The ranges of C. g. gundi and C. g. vali are
thus physiographically and climatically distinct.
Gundis are unknown from the Gebel es Soda and the Gebel el Harug
el Asued, both of which appear to have an abundance of suitable
habitat. The Gebel es Soda is also only a short distance south of the
Wadi Titti, a previous collecting site for the gundi. In the future, gundis
will probably be found to occur in both of these areas.
Setzer (1957) stated that, during his visit to the Wadi Bey in 1955,
his guideinformed him that in the past numerous gundis had inhabited
the "broken rock scarps" but owing to a severe drought the vegetation
had dried up and most of the gundis had died.
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U.S. NATIONAL MUSEUM BULLETIN 275, PLATE 1
•=/«
Mi T*«?.J*
Coastal plain near Tocra showing typical habitat of Microtus mustersi and Mus musculus,
May 1962.
Broad coastal plain 20 kilometers east of Tobruch, Cyrenaica. The densely vegetated
hummocks provide ideal habitat for the large Egyptian jerboa (Jaculus orientalis) and a
small gerbil {Gerbillus kenleyi), June 1962.
285-134 0—68 18
U.S. NATIONAL MUSEUM BULLETIN 275. PLATE 2
5ush-covered coastal plain near Tocra, Cyrenaica. Gerbillus campestris is the dominant
rodent in this type of habitat, May 1962.
Coastal plain near Benghazi, Cyrenaica, showing characteristic hillocks supporting dense
vegetative cover interspersed among bare, saline areas. The large Egyptian jerboa (Jaculus
orientalis) and the Libyan jird (Meriones libycus) occur in this type of habitat, May 1962.
:
»<PV
Fresh talus and burrow entrances of the sand rat {Psammomys obesus) in a large hummock
on the coastal plain near Agedabia, Cyrenaica, May 1962.
iP-^" --* *Q4 : r'r*
Bouldcr-strcwn slopes such as these of the escarpment near Gharian provide ideal habitat
for gundis (Ctenodactylus gundi), March 1962.
U.S. NATIONAL MUSEUM BULLETIN 275, PLATE 4
Rocky outcroppings and large boulders of the escarpment 5 kilometers west of Cussabat,
Tripolitania, where a large series of gundis (Ctenodactylus) was obtained in June 1962.
Hamada (pebble desert), acacias, and escarpment approximately 100 kilometers west-
southwest of Ubari, Fezzan. This "pebble desert" is the most characteristic type of
terrain in the Libyan Sahara. Rodents occur sparingly in these "hamada" deserts and are
usually represented by Meriones crassus or the ubiquitous Gerbillus campestris, December
1961.
Bir bu Zarregh located near the "Cufra Track" in the desolate Sand Sea of Calanscio,
Cyrenaica. Vegetation at this remote site consists of only three stunted date palms, a
castor bean tree, and a eucalyptus tree. In spite of the isolated location of Bir bu Zarregh,
jerboas (Jaculus jaculus) and gerbils (Gerbillus gerbillus) were found to occur there,
March 1962.
JH^-fT-
. a-
> ;•*».
J*
:
'**Wlk* ':
. U.-
Mounds of the mole-rat (Spalax ehrenbergi) on rocky slope north of El Faidia near the
highest point of the Cyrenaican Plateau, May 1962.
Rock-strewn plain of the Gebel Nefusa, 12 kilometers south of Chicla, Tripolitania. The
fat-tailed sand rat (Pachyuromys duprasi) prefers these areas where the surface is covered
with coarse pebbles and the vegetative cover is sparse and localized, May 1962.
Large mound with dense cover of Calligonum near the oasis of Tazerbo, Cyrenaica. These
outlying hummocks of the Saharan oases provide shelter and sustenance for several
species of desert rodents, especially for jerboas (Jaculus jaculus) and gerbils (Gerbillus
gerbillus), April 1962.

Vegetated dunes and sandy-clay hummocks supporting dense growths of Tamarix and
Calligonum surrounding Tazerbo Oasis, Cyrenaica. These peripheral zones are inhabited
by large numbers of jerboas (Jaculus jaculus) and gerbils {Gerbillus gerbillus), April 1962.
w.-;-M
Dense pocket of Phragmites growing in an area of abundant fresh water near the oasis of
Brach, Fezzan. A large series of Mus musculus was collected from the above habitat,
February 1962.
Outlying tamarix near the oasis of El Gatrun, Fezzan. This peripheral zone of tamarix is
typical of most of the larger oases in the Libyan Sahara and provides suitable habitats
for several species of Saharan rodents, January 1962.
*&t
Date palms interspersed with sandy-clay hummocks supporting tamarix at El Abiad Oasis,
Fezzan. The large gerbil, Gerbillus pyramidum, is abundant in the sandy areas around
the palm trees where the dense, basal fronds afford shelter and the fallen dates provide a
convenient source of food, December 1961.
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SMITHSONIAN

The Rodents of Libya

Division of Mammals, U.S. National Museum

SMITHSONIAN INSTITUTION PRESS

U.S. GOVERNMENT PRINTING OFFICE

For sale by the Superintendentof Documents, U.S. Government Printing Office

History), London; and Dr. Francis Petter, Curator of Mammals,

aid reading of the manuscript.

ing staff members of the Department of Zoology and Entomology,

Libya is a vast expanse of semiarid and arid desert of variable

Longitudinally, Libya lies between 9° and 25° east longitude. This

is entirely lacking. Where

pressions of the hamadas, or occurs as narrow fringes bordering the

larger wadis (dry watercourses). Vegetation also occurs sparingly

elevations and greater amounts of rainfall, the Tripolitanian Gebel

One to 14 days were spent at eacli collecting site. The majority of

gauge shotgun, using number 9 shot or a .32-caliber auxiliary barrel

and again in the spring of 1961, while on a trans-Saharan trip to the

When specimens become available from more localities on the coastal

Physical Description of Libya

Libya is divisible into several rather broad physiographic elements.

Figure 1. — Physical features of Libya.

The Coastal Plain

The Libyan coastal plain varies in width from a few hundred

owing to the encroachment of the escarpments of the Cyrenaican

The Coastal Escarpment

A broad belt of pre-Saharan desert is typical of vast areas of Libya

Saharan desert of the interior. These transitional deserts are generally

graphic features with their deep canyons, boulder-strewn slopes and

The overall coloration of thesehamadas is generally gray, but

Occasionally, bedrock is covered with a veneer of sand and gravel

western margins of the Sand Sea of Calanscio to the eastern limits

In some areas of Libya, the surface layer is composed of extremely

The depressions associated with Giarabub Oasis and Bahr el Tubat

Mountainous Areas of the Interior

The Gebel es Soda and the Gebel el Harug

are mountains of this type. Both have been regarded

The Cyrenaican Plateau

rainfall, fresh water accumulates in the bottom of the valley to a

The Tripolitanian Gebel

The Gebel Nefusa of northwestern Tripolitania is less clearly de-

South of the rugged coastal escarpment the interior of the Gebel

The Eocene transgression outcrops widely north and east of the

east-northeast west-southwest anticline, which is connected in the

southwest to another anticlinal zone near Ghat. This anticline extends

Near Murzuch the Cretaceous Nubian Sandstone outcrops, forming

Volcanic activity occurred in three regions of Libya: the Gebel

Arid or Desert Type

high. Relative humidity as low as 2 percent, with temperatures of

Semiarid or Steppe Type

A region of low latitude steppe climate characteristically bounds

beltbetween it and the humid climate of the Mediterranean littoral.

On the coastal plain, rainfall generally approaches 15 to 25 inches.

evaporated, and Mediterranean climate of Libya can be described as

On the Cyrenaican Plateau and Tripolitanian Gebel, the summers

Additional Meteorological Data

Meteorological data obtained during field work in 1961 and 1962

coastal plains, were, respectively: 113.7° F (108°-120° F), 62° F (53°

General Features of the Saharan Flora