MAMMALS OF THE EASTERN MEDITERRANEAN REGIONj

THEIR ECOLOOf, SYSTSMATICS AND ZOOGEOGRAFHICAL RELATIONSHIPS

Sana Isa Atallah, B,S., M,S.

American University of Beirut, Beirut, Lebanon, 1963

American University of Beirut, Beirut, Lebanon, 1965

A Dissertation

Submitted In Partial Fulfillment of the

Requirements for the Degree of

Doctor of Philosophy

at

The University of Connecticut

1969 Copyright by

SANA ISA ATALLAH

1969 APPROVAL PAGE

Doctor of Philosophy Dissertation

MAMMALS OF THE EASTERN MEDITERRANEAN REGIONj

THEIR ECOLOGY, SYSTEMATICS AND ZOOGEOGRAPHICAL RELATIONSHIPS

Sana Isa Atallah, B.S., M.S.

Major Adviser \a^. V_a $ -g~tcr o

The University of Connecticut 196?

ii ACKNOWLEDGMENTS

The c<- apletj.cn of this work would have not been possible without

the constant assistance and advice of my major advisor. Dr. Ralph M.

Wetzel, I am also greatly indebted to Dr. and Mrs. Robert E. Lewis,

Iowa State University, Ames, Iowa, previously at the Dept, of Biology,

American University of Beirut, Lebanon, for their very kind assistance,

direction and advice while in Lebanon during the years 1963-1966, and

Dr. David L, Harrison, Sevenoaks, Kent, England, for his help in the

field and in identifying and comparing many specimens with material

in his personal collection and at the British Museum (Natural History)

collections,

I am also very grateful to Drs, Ralph M. Wetzel, James A, Slater

and George A, Clark at the University of Connecticut and Dr. Homy

W, Sotzcr at the Smithsonian Institution for their useful suggestions

and cidtical reading of the mar.uyeript.

Thanks are also duo to my parents, Mr. Jacob Qumaioh, Miss Jean

Uridgwood, Mr. Charles Bridgeood, Mr. Richard Biddgwcod, Mr. George

Kassis, Mr, Hanna Qumaioh, Mr. George Khuri and Dr. J. Morton Boyd

fur their valuable assistance in the field; Mrs. Stephana Schaefer,

Miss Susan Kramer and Dr. David Harrison for providing the text

figures; and Miss Kristina Hosko for her help in assembling Appendix

X,

Finally, I wish to exbend my sincere thanks to all my friends

and colleagues for their constant snoouragment and assistance.

iii TABLE OF CONTENTS PAGE

Aclcno'. ledgmenta ;...... lit

Lis b ff tables...... t •«vi

List lit’ illustrations...... vii

Intrci’uction...... -1

Tho E? stem Mediterranean Region...... 12

A. The People and the.Land...... 13

B. Physical Geography...... 17

C. Climate...... 30

D. General Ecology and Fhytogeography...... 37

Materials and Methods...... U?

Species Accounts...... 53

Order Insectivora.5^

Order Chiroptcra...... 88

Order Lagcmorpha,...^x..,)...... 1^42

Order Rodentia...c......

Order Carnivora...... «2/;0

Order Hyracoidea...... «2U5

Order Artiodactyla..2lt6

Domesticated aiid Introduced Mammals.,...... 2149

iv PAGE

Zoog^ .raphlcal Summary...... 2$3

Bibli;. jraphy...... 2^9

Adder* ait to Bibliography...... 280

Appen.. Lx I - List of Measurements...... 281 LIST OF TABLES

TABLE PAGE

1. Checklist of the mammals of tho Eastern 7 Mediterranean Region.

2. Comparison of monthly rainfall and mean 31 Kiaximurrt and minimum temperatures of six localities.

■vi LIST OF ILLUSTRATIONS

IGUR2 PAGE

1. Tli.) Eactai-n Mediterranean Regionj collection 2 and other important localities cited in tlvi text.

2. Schematic representation of regions and sub- 18 regions pertaining to the physical geography of the Eastern Mediterranean Region.

3. Biocli;.iatic zones of tho Eastern Mediterranean 35 Region.

It. Ecological regions of tho Eastern Mediterranean 39 Region,

5. The glans penis of A. Crocidnra russula. SA 60 1153 from Afka, Lebanon, and B, Erinaceus europaeus, SA 1063 from near Damascus, Syria.

6, Head of Nycatalus noctula lebanoticus 129 (HARR. 26,33^0) collected at' Natural Bridge^, Faraya, Lebanon...

7« Postero-lateral view of skull A. Meriones 169 libycus B. Meriones crassus C. Morionos tristrami D, Sekeetamys calurus.

vii INTRODUCTION

The E. •item Mediterranean Region, aa understood here (Fig.l),

represents the smallest natural unit that can be satisfactorily

delineated from tho Middle East ( » Near East), and in essence it

correspond:; to the left horn of the so called "Fertile Crescent"

and the in' ervening Syrian Desert. The Fertile Crescent includes

many centeis of ancient civilizations and extends from the Tigris

and Euphrates River Valleys on the east, westward along the southern

slopes of the Taurus Mountains and southwards along the land bordering the eastern Mediterranean to the Gulf of Aqaba on the Red Sea (Aramco

Handbook, 1968).

The Eastern Mediterranean has been of great zoogeographical

importance since the Miocene, when this land first emerged from under tho Sea of Tethys* Its importance rests on its location^

along the dispersal routes between the two major Old World land mas­

ses: Eurasia and Africa. Consequently, one finds among tho endemic

Palaearctic fauna of this region mammals characteristic of the

Oriental and Ethiopian Zoogeographical Regions.

The objectives of this vork are to provide,as much as possible,

information on the ecology, distribution, systematics and zoogeo­ graphical relationships of mammals presently occurring along the

eastern Mediterranean. Data presented is based on published records, unpublished museum records and original observations accumulated through extensive field work in tho area during tho years 1963-1966 and the summer of 1968,

1 2

EASTERN MEDITERRANEAN REGION 0 ZS SO 15 iqo Km.

s| y„„ • i Jw...* ■'■---' Haita W

H^TAMYK P ',* •N**Wi},?«»1 -r • K»AI . TtbAvivLviv w 0*«O v* *.***» Yafo0 I • *****«*«■•« t*M U • ZlOH i *t.waugnT*«»M f ’ilHOVcFT ^ XUU|«HlH '&tY«kA«%Y / /'• &MI| t OMvtUU vSAh«A e.lM, .W.OftT«AU« . . StOHi, , „ VAlA/*_/ #T»LMil-. Giyr%ti.W * kmKm Y»**r% *”®***S"t8A “E»kY»8 \*.

»Tat»u» \ SyR^N

ShouQAa . EL.-S»fS

• HA«g

• t.u ma«*

• EL-Ou^t'ftft

LiuAtV-# _ S\K

Fig. 1. The Eastern Mediterranean Region; collection sites and other important localities cited in the text. Elevations are in m. 3

Ninety -Tour species and one hundred subspecies of terrestrial, wild mamma}* are recognized from the Eastern Mediterranean Region

(Table l). These figures include only wild mammals occurring at the present time} hoviever thirteen additional species of domes­ ticated anti introduced mammals are briefly mentioned and their impact on the environment discussed,

A brief historical review of the principal publications dealing with the mammals of the Eastern Mediterranean Region is

given below in chronological order,

Pi-e 1600

Conspicuous mammals such as the wild ass, coney, ibex, hyaena, wild boar, leopard, cheetah, etc. are continuously referred to in the Bible and in the works of early travelers and pilgrims

(Aharoni, 1938$ Bodenheimer, I960). Later descriptions of many mammals appeared in the writings of Arab scientists such as Ad-

Damiri (bom 13h9 A.D.) and Al-Qazwini(born 1281-2 A.D.)j tho former in his two volume book "Hayat El-Hayawan *» The Life of

Animals" and the latter in two contributions "Huzhat Ul-Qulub **

Hearts' Delight" and "Aja'eb Ul-Makhloukat - The Wonders of the

Living",

The earliest systematic collections were those of E. Hasselquist and E. Geoffroy St. Hilaire who traveled into southern Palestine from

Sinai and North Africa in the mid and late eighteenth century.

18CO-1900

The foundations of our present knowledge of mammals in the

Eastern Mediterranean Region were laid in this period. Many German and English travelers obtained specimens that were later described a by VJagnor, brants, 0. Thomas and A. Nehring.

Canon 'f. B. Tristram made four trips to Palestine that re­ sulted in h_s major contribution "Fauna and Flora of Western

Palestine" published in 188U, Hovever, he devoted less than 30 pages to his discussion of 113 species, including fossil forms, of domesticated and wild mammals» This work is now outdated and not available except in largo libraries, also many of the species were incorrectly identified.

Other scientists of importance at this time were H. C. Hart,

T. von Heug.lin, E. Rueppoll and C. Cretscchmar; also valuable collections were made by Baron von Schubert and the botanist Kotschy,

1900-1925

Several collections were made during this period especially through the endeavors of British Army Staff such as S, S. Flower,

R. I, Pocock and other collectors such as D. Carruthers, H. Schliiter,

P, A. Buxton, E. Schmitz, S. Merrill and I.Aharoni. These collectors provided European museums with indispensable material, and also published in accounts of their travels valuable information on the habits and habitats of many species of mammals. From these collec­ tions many new species were described by museum personnel at London and Berlin, of these 0. Thomas, A. Nehring, P. Matschie and F. Muller are especially worthy of mention.

I. C. Phillips assisted by W. M, Mann traversed the country south of Mt. Hermon (March-June, 19lll) and collected over 150 small mammals some of which, wore later described by G. M. Allen

(1915) at Harvard. 5

192S-19SO

Many p-iblications appeared in this period, but only a few are noted here.

Tho mu roids of Palestine were described by B. Aharoni (1932) and those of Asia Minor by G, Neuhauser (1936). Both of these studies werj based primarily on collections made by these workers and tho bulk of the material was deposited in the Berlin Zoological

Museum. F, 8. Bodenheimer (1935) reviewed the existing knowledge of the mammals of Palestine and in many ways he brought Tristram's

(I88J4) treatment up to date. D.M.A. Bate (191j5) reported on small mammals obtained from owl pellets at Mt, Lebanon. J. R. Kile man

(I9)j8) published a key to the rodents of soilthwest Asia based on the British Museum (Natural History) collections. In this work he stated "Only from Syria and Palestine is our material not very representative".

1950-to date

The appearance of the "Checklist of Palaearctic and Indian

Mammals, 1758-19U6" by J, R. Ellerman and T. C. S. Morrison-Scott

(1951) provided an excellent summary of the taxonomic status of all species and subspecies of mammals along the eastern Mediterra­ nean and stimulated a tremendous amount of research.

During this period our knowledge of Eastern Mediterranean mam­ mals increased tremendously. Many contributions were made by F. S,

Bodenheimer, A, Zahavi, J, Wahrman, E. Nevo and F. Petter in Israel,

R, E. Lewis, J. II. Lewis and S. I. Atallah in Lebanon, X. Misonne and E. von Lehmann in Syria, S. I. Atallah in Jordan, and D. L.

Harrison throughout the Eastern Mediterranean Region (see biblio­ 6 graphy for details). The most important contribution, however, is that of Harrison (I961|b and 1960a), who summed up many of the published lecords on the mammals of the Arabian Peninsula in two volumes covering all mammalian orders except the Rodentia and

Lagomorpha, Tho last two orders will bo dealt with in a third volume (Harrison, in preparation). Harrison’s work dealing with the remaining orders is an excellent taxonomic account, yet very little information, if any, is available on the habits, habitats and evolutionary trends of the species in question. Also by virtue of the large area covered (almost l/2 the size of continental United

States), Harrison’s account of species and subspecies distribution in any one country is incomplete and is more or less restricted to specimens available to him at tho British Museum (Natural History).

Research on Eastern Mediterranean mammals was enhanced greatly by workers in adjacent areas. Listed below are names of some of these workers and areas of their research (see biblio­ graphy for details):

Iran: D. M. Lay and X, Misonne

Iraq: R. T. Hatt, and D, L. Harrison

Saudi Arabia: D, Vesey-Fitzgerald and D. L. .Harrison

Sinai and Egypt: H. Hoogotraal, K. Massif and H. W. Setzer

Turkey: J. Osborn, H. Kahmann and M. Gaglar 7

Table 1. Checklist of the mammals of the Eastern Mediterranean Region.

Class Mammalia subclass Theria Infraclass Eutheria

Order insectivora - Insectivores

Family Erinaceidae - Hedgehogs Erinaceus europaeus concolor Martin European Hedgehog Hemiechinus auritus calligoni (Satunin) Long-eared Hedgehog aegyptius (Fischer) Paraechinus aethiopicus pectoralis (Heuglin) Ethiopian Hedgehog ludlowi Thomas

Family Soricidae - Shrews Crocidura leucodon .judaica Thomas Bicolor Wiite-toothed Shrew Crocidura lasia Thomas Large Bicolor 'White-toothed Shrew Crocidura russula monacha Thomas Common European White-toothed Shrew Crocidura suaveolens portali Thomas Lesser White-toothed Shrew Suncus etruscus etruscus oavi Savi's Pygmy Shrew

Order Chiroptera - Bats

Suborder Megachiroptera - Fruit-eating Bats Family Pteropidae - Fruit Bats Rhinopoma hardwickei cystops Thomas Lesser Mouse-tailed Eat Rhinopoma microphyllum microphyllum (Bronnich) Greater Mouse-tailed Bat

Family Emballonuridae - Sheath-tailed Bats Taphozous nudiventris nudiventris Cretsschmar Naked-bellied Tomb Bat 8

Table 1. Checklist of the mansnals ....(continued).

Family Nycteridae - Slit-faced Bats Nycteris thebaica thebaica E. Geoffroy Egyptian Slit-faced Bat

Family Rhinolophidae - Horseshoe Bats and Leaf-nosed Bats Rhinoloohus ferrumequinum femur.squ.inum (Schreber) Greater Horseshoe Bat Rhinolophus clivosus clivosus Cretsschmar Cretzschmar*s Horseshoe Bat Rhinolophus hionosideros minimus Heuglin Lesser Horseshoe Bat Rhinolophus euryale judalcus (Anderson & Matschie) Mediterranean Horseshoe Bat Rhinolophus blasii blasii Peters Peters’ Horseshoe Bat Asellia tridens tride'ns (E. Geoffroy) Trident Leaf-nosed Bat

Family Vespertilionidae - Vespertilionid Bats M^’-otis nyotis macrocephalicus Harrison 8c Lewis Greater Mouse-eared Bat .Ryotis blythi omari Thomas Lesser Mouse-eared Bat Myotis emarginatus emarglnatus (E. Geoffroy) Notch-eared Bat I-hrotis caoaccinii bureschi (Heinrich) Long-fingered Bat Myotis nattereri hoveli Harrison Natterer’s Bat Pinistrellus pipistrellus oipistrellus (Schreber) Common Pipistrelle Pioistrellus kuhlii ikhwanius Cheesman 8c Hinton Kuhl’s Pipistrelle Pipistrellus savii caucasicus Satunin Savi’s Pipistrelle Pipistrellus bodenheimeri Harrison Bodenheimer*s Pipistrelle Nyctalus noctula lebanoticus Harrison Common Noctule Eotesicus serotinus serotinus (Schreber) Serotine Bat Eotesicus bottae innesi (Lataste) Botta’s Serotine Bat Qtonycteris hemprichi .jin Cheesman & Hinton Hemprich's Long-eared Bat Plecotus austriacus christiei Gray Grey Long-eared Bat Miniooterus schreibersi pallidus Thomas Schreibers’ Long-winged Bat

Family Molossidae - Free-tailed Bats Tadarida teniotis ruppelli (Temminck) European Free-tailed Bat

Order Lagomorpha - Rabbits and Hares

Family Lepuridae - Hares Leous capensis syriacus Ehrenberg Syrian Hare arabicus Ehrenberg Arabian Hare 9

Table 1. Checklist of the mammals ....(continued).

Order Rodentia - Rodents

Family Sciuridae - Squirrels Sciurus anomalus syriacus Ehrenberg Persian Squirrel

Family Cricetidae - Rats and Mice

Subfamily Cricetinae - Hamsters Cricetulus migratorius cinerascens (Wagner) Grey Hamster Mesocricetus auratus auratus (Waterhouse) Syrian Golden Hamster

Subfamily Microtinae - Voles Arvicola terrestris (Linnaeus) Water Vole Microtus nivalis hemonis Miller Snow Vole ^icrotus socialis (Pallas) Social Vole Microtus guentheri guentheri (Danford and Alston) Gunther's Vole

Subfamily Gerbillinae - Gerbils and Jirds Gerbillus dasyurus dasyurus (Wagner) Wagner * s Gerbil Gerbillus nanus arabium (Thomas) Baluchistan Gerbil Gerbillus henleyi marine (Bonhote) Pygmy Gerbil Gerbillus gerbillus bonhotei Thomas Lesser Egyptian Gerbil Gerbillus allenbyi Thomas Allenby's Gerbil Gerbillus pyramiHum I. Geoffroy Greater Egyptian Gerbil Sekeetamys calurus calurus (Thomas) Bushy-tailed Jird Meriones tristrami tristrami Thomas Tristram's Jird karietini Aharoni Meriones vinogradovi Heptner Vinogradov's Jird Meriones libycus syrius Thomas Libyan Jird Moriones erassue crassus Sundevall Sundevall's Jird Meriones sacramonti Thomas Palestine Jird Psammonys obesus obesus Cretzschmar Fat Sand Rat

Family Spalacidae - Mole Rats Spalax ehrenbergi Nehring Palestine Mole Rat 10

Table 1, Checklist of the mammals ..*«(continued)

Family Muridae - Old World Rats and Mice Apodemus mystacinus mystacinus (Danford & Alston Broad-toothed Field Mouse Apodemus sylivticus tauricus TBarrett-Hamilton) Common Field Mouse Rattus rattus alexandriaus (Desmarest) House Rat i'ruglvoTus (Rafinesque) Rattus norvefpLcus norvepricus (Berkenhout) Brown Rat Mus muscuius nraetextux Brants House Mouse Acomys dimidiatuS'.dimicfiatus (Cretcschmar) Cairo Spiny Mouse Acom^ russatus rassatus (’Vagner) Golden Spiny Mouse Acoinys 'levrisi Atallah Black Desert Spiny Mouse NesoidLa indica bacheri Nehring Short-tailed Bandicoot Rat

Family Gliridae - Dormice Eliomys melanurus (Wagner) S. W. Asian Garden Dormouse Dryonys nitedula"phrygius Thomas Forest Dormouse

Family Dipodiae - Jerboas Jaculus jaculus vocator Thomas Lesser Egyptian Jerboa ScHueteri (Nehring) Allactaga euphratica euph’ratica Thomas Five-toes Jerboa

Family Kystricidae - Porcupines Hystrix indica indica Kerr Indian Crested porcupine

Order Carnivora - carnivores

Family Canidae - Jackals, Wolves and Foxes Canis aureus syriacus Kemprich & Ehrenberg Asiatic Jackal Canis lupus pallipes Sykes Wolf Vulpes vulpes palaestina Thomas Red Fox arablea Thomas Vulpes ruppelli sabaea Pocock Ruppell’s Sand Fox

Family Mustelidae - Weasels, Polecats, Martens, Badgers and Otters Mustela nivalis Linnaeus Weasel II

Table 1. Checklist of the marranals . .•(continued).

Vomela peregusna S7/riaca Pocock Marbled Polecat I'!artss foina syriaca (Nehring) Stone Marten Kellivora capensis (Schreber) Honey Badger Meles neles (.Linnaeus) Badger Lutra lutra siestanica Birula Common River Otter

Family viverridae - Genets and Mongooses Genetta genetta terraesanctae Neumann European Genet Herpestes ichneumon ichneumon (Linnaeus) Ichneumon

Family Hyaenidae - Hyaenas Hyaena hyaena syriaca Matschie Striped Hyaena

Family Felidae - Cats Felis sylvestris tristrami Pocock Wild Cat Felis chaus fufax de Vanto'n Jungle Cat Felis caracal schmitzi Matschie Caracal Lynx Panthera pardus (Linnaeus) Leopard

Order Hyracoidea - Hyraxes and Conies

Family Procaviidae - Hyraxes Procavia capensis s:/riaca (Schreber) Syrian Kyrax

Order Artiodactyla - Even-toes Ungulates

Family Suidae - Pigs Sus scrofa libycus Gray Wild Boar

Family Bovidue - Sheep, Goats and Antelopes Gazella gazella (Pallas) Mountain Gazelle Gazella dorcas (Linnaeus) Boreas Gazelle Gazella subgutterosa narica Thomas Goitered Gazelle Capra ibex nubiana F. Cuvier Ibex THE EASTERN MEDITERRANEAN REGION

12 13

A. THE PEOPLE AMD THE LAND

Tho Eastern Mediterranean Region (Fig. 1) as understood in this paper includes the land bordering the Mediterranean Sea eastern coast between 29° 30* and 36° 00* N. latitude and 3U° l£1 and 39° 30* E. long­ itude, politically comprised of the Republic of Lebanon, the Hashemite

Kingdom of Jordan, the State of Israel, the Gaza Strip (a part of the

United Arab Republic), and adjacent parts of the Syrian Arab Republic.

All localities cited in the text will be referred to one of the following: Lebanon, Syria, Jordan or Palestine (Fig. 1) as described below:

LEBANON:

A political entity that came into existence in 1920 under French mandatory rule. The French had then enlarged the Turkish govemorate of Mount Lebanon to include the country with its artificial boundaries as they are known today. It borders the Mediterranean Sea for some

220 km to the west and is bounded to the north and northeast by Syria and to the south and southeast by Palestine. The average width of the country is not more than 60 km.

Tho country derives its name from its snow topped mountains;

"laban” being the Aramaic word for white, The population was estimated in 1961 at 2,150,526 while land area is about 6,Ii2li sq. km (h,0l5 sq. mi). This makes Lebanon the most densely populated country in the

Middle East with 335 inhabitants per sq. km. The capital is Beirut.

Tho country is known as the Republic of Lebanon, Al-jumhurieh Al-lubnanich (Arabic), or Liban (French). JORDAN:

Tho naniG Jordan uas used since biblical times to refer to the

land bordering the Jordan River. It first came to denote a political

entity in 1921 as a British mandated stato then known as Trans-Jordan.

Tho country gained its independence in 19U6 and following the Palestin

ian War of 19ii8 a small section of Palestine (Cis-Jordan) was annexed

to Trans-Jordan. The two parts (usually referred to as the East and

West Bank) then became knom as the Hashemite Kingdom of Jordan (Al-

Mamlakah Al-Urdinieh Al-Hashemiah in Arabic) until 196? when Israeli

forces took over tho West Bank of Jordan, Jordan, in this paper,

refers to tho East Bank (Trans-Jordan) only.

Jordan is land locked except for a I4 km exit on the Gulf of Aqaba

Red Sea in the south. It is bounded to the north by Syria, to tho

northeast by Iraq, to the south and southeast by Saudi- Arabia and to

tho west by Palestine, The area is about ££,000 square kilometers

and the population may be roughly estimated at 1,000,000. Patai

(19£8) reports that the population of the East Bank was estimated

in 19£8 at 687,000 of which 220,000 were nomads and serai-nomads.

There has been no census of population increase following the 196?

Arab-Israeli War and I have only tentatively placed the population

at one million as a very conservative figure.

Jordan is primarily an agricultural country with over ?£$ of the

people completely dependent on this profession. The country, however,

is very dry and cultivation is nob possible without adequate rain or

irrigation. This has resulted in the government’s extensive support

for irrigation projects, the largest of which is the East Ghor Ganal project,. This canal, almost 7£ km in length, runs parallel to the 1?

Jordan Riv>rr tapping tho waters of the Yarmuk River, a tributary of the Jordan River, and irrigates an area of 120,000 dunums

(30,000 acies),

PALESTINE:

"Palec. ‘/ina", or more correctly Philistina, was the name given by the Greeks to the land of the Philistines who lived along tho southern coast of the eastern Mediterranean. Under tho Roman rule

(about 6? A,D.) Palestine was made a separate province known as

'•Syria Palestina" and is believed to have extended from the lower

Nahr El-Litani (ancient Leontes) Valley in the north to Beersheba in the south and from the coast inland to the desert (Smith, 1897).

Palestine as understood in this paper is bounded to the north by

Lebanon, to the east by the Jordan River, Dead Sea and Wadi Araba, to the northwest by the Mediterranean (290 km of coast line) and to the south and southwest by Sinai and the Gulf of Aqaba, These artificial boundaries were drawn following World War I at the time the British assumed control over tho area.

On May Ilf, 19h8 tho British commissioner of Palestine left the country and on tho same day the State of Israel, a Jewish State, was proclaimed. This lead to the Palestinean War of 19lf0 which was brought to an end in 19lf9 frith a partition of Palestine into three components; Israel, the Gaza Strip controlled by Egypt, and a 3,lf6U sq. km of territory along the west bank of the Jordan River held by the Jordanian Army and later incorporated with the Hashemite Kingdom of Jordan. The area and population of the three components of

Palestine are given below: 16

Coinoonent Area in sq, kra. Population

Israel 12,788 2,179,U9l(estimated 1961)

Gaza 1,280 not available

Jordan (Wee b Bank) 3,I*6U 81^,000(estimated 1958)

Total (Pale..stine ) 17,532 approx. 3,000,000

•

SYRIA:

Syria, more correctly Assyria, was a name originally applied by the Greo’cs to the land north of the Arabian Peninsula bounded by the Mediterranean Sea, Taurus Mountains, Euphrates River, Red Sea and the Isthmus of Suez.- As such this area included the whole Eastern

Mediterranean Region. The present boundaries of Syria were arrived at following t’orld War I and after much dickering between the colonial governments of Britain and France which resulted in the partition of

Syria into Lebanon, Palestine, Jordan and Syria proper.

Today’s Syria (The Syrian Arab Republic; Al~Jumhurieh Al-Arabieh

As-Surieh, in Arabic) occupies an area of 185,180 sq. km (71,1*98 sq. mi). It is bounded by Turkey to the north, Iraq to the east and southeast, Jordan to the south, and Palestine, Lebanon and the Med­ iterranean Sea to the west, Tho population was estimated in 1961 at Li,!?6£,121 of which over 200,000 are nomads (Bedouins), The capital is Damascus,

Only the western two-thirds of Syria are incorporated in this study (3ee Fig.l), B. PHYSICAL GEOGRAPHY

The mout striking feature of the Eastern Mediterranean Region is the great Rift Valley representing the northern end of the Syrian-East-

African Rifb system. This valley is flanked at both sides by mountain ranges extending the whole length of this region from north to south.

The western mountains are separated from the Mediterranean shore by a fertile coastal plain interrupted at various points by spurs of mountains and foothills reaching the sea. Tho eastern mountains, however, terminate at a semi-arid plateau known as the Syrian Desert.

Five zones could thus be recognized: The Coastal Plain, Western

Uplands, The Great Rift Valley, Eastern Uplands, and the Syrian Desert

(Fig. 2). These represent strips of land running parallel to the shore line and can be individually characterized as follows: /For greater detail see Orni and Efrat (1966), Ives (19!?0), Burdon (19^9), and

Poore and Robertson(196U)/:

I, The Coastal Plain:

The Coastal Plain, north of Tartus, Syria, is narrow and inter­ rupted by spurs of foothills from Jabal Ansariyah, while to the south it opens up into the plain of Sahl Akkar which extends southwards to

Tripoli, Lebanon. Farther south the plain narrows and again it is interrupted by rocky spurs from Mount Lebanon. Southwards from Saida the plain widens until it reaches the Lebanese Palestinean border where tho hills of Galilee project to the sea at Ras en Nakura.

Farther south the plain widens again and reaches its greatest width

(IjO-WjO km) in the Gaza Strip. It is, however, interrupted by a spur from Mount Carmel to the north of Haifa.

17 18

EASTON' MEDITERRANEAN REGION 0 AS SO *75 100

i>v ->0

Fig. 2. Schematic representation of regions and subregions pertaining to the physical geography of the Eastern Mediterranean Region. 19

The shore north of Mount Carmel shows an alternation of sandy bays vrith rooky headlands and low cliffs, while to the south sandy beaches predominate. The shore south of the Carmel coast is parall­ eled by a sand dune area which though narrow and discontinuous in the north forms a continuous wide belt from the Judean coast southwards into Sinai.

Cultivation is extensive along the coast especially along the foothills of the Western Uplands whore the heavier terra rossa soil washed from the hills is more suited for agriculture than the red sands of this region. Fertility of the coastal plain is due primarily to the abundance of wadis and rivers that transect this plain draining the Western Uplands and pouring into the sea. The most important rivers in this respect are (from north to south): Nahr El-Kabir,

Nahr Ibrahim, Nahr El-Kelb, Nahr El-Litani, Nahr Hadera and Nahr

Yargon. II. The Western Uplands:

This zone may be divided from north to south into four regions as follows (northernmost region first):

1, The Ansariyah Mountains Region.- These mountains are about

30 km wide, and the highest peak reaches 1562 m. The mountains ter­ minate northwards near Antioch, Turkey, where the Taurus Mountains run perpendicular to them. Southwards these mountains slope gently toward the plain of Sahl Akkar which separates this region from Mount Lebanon.

The western slopes are drained by Nahr Fl-Kabir and many minor rivers leading to the Mediterranean, while the eastern slopes are drained by Nahr El-Assi and its tributaries. 2, Mount Lebanon Region.- This mountain chain, collectively known as Mount Lebanon (or Jabal Lubnan, locally), rises steeply from Sahl

Akkar with the highest peak reaching 3,033 m' (10,131 ft) at Qurnet as- 20

Sawda, the highest peak along the eastern Mediterranean, It is in­ terrupted east of Beirut at Dahr El-Baidar by a low pass (l,5>h0 m),

but quickly rises to a peak at Jabal El-Parouk (1,980 in). It termin­

ates south ards along Nahr El-Litani lower valley where the hills of

Galilee merge into the next region.

The average elevation of this region is over 2,000 m north of Dahr El-Baidar and about l,ii00nito the south of this pass. This region

is also widest at its northern end narrowing to the south. The east­ ern slopes are drained by tributaries of Nahr El-Assi leading north­

wards and Nahr El-Litani leading southwards with a water-shed near

Baalbeck. The western slopes are drained by many rivers of which Nahr Ibrahim, Nahr El-Kelb, Nahr Beirut, Nahr Besri, and Nahr Abou Assouad

are of most importance,

3. The Hill Region,- This region can be further subdivided into

four subregions from north to south as such:

(a) Tine Hills of Galilee: These hills rise from the gorge of Nahr

El-Litani and terminate southwards into the plain of Esdraelon and

the valley of Jerzeel, The hills are divided into two sections, Upper

and Lower Galilee, by the gorge of Nahr Ammud. The Upper Galilee

Hills attain the highest elevation at Jabal Moron (1,203 m), while

the lower hills never exceed 600 m in elevation. These hills are

drained by small streams leading to the Mediterranean on the west and- Lake Hula complex to the east, (b) The Hills of Samaria: These hills occupy a central position in

Palestine. The northern border corresponds with the southern borders

of the Lower Galilee, whore Mount Carmel rises from the bay of Haifa

and extends for some 32 km to the southeast joining the Hills of Menashe

and the Irron Hills, Farther south these join with the main bulk of the

Samarian Hills conveniently separated into two sections corresponding to 21 the Nablus Synclino (Nablus Dotmfold) to the vest and the East Samarian

Upfold to the east. The southern borders of this subregion can not be easily delineated, but geologically one can place these borders just to the north of Jabal Asur where the mentioned folds terminate.

The highest peak of Mt. Carmel reaches 5U6 m, while to the south the elevations are much higher attaining a peak at Jabal Ebal (9h0 m).

The latter mountain is surrounded by many mountains of which Jabal

Shekhem (881 m), Jabal El-Kabir (767 m) and Jabal Hureish (76I4 m) are worthy of mention. Drainage is by many small wadis leading to the

Jordan River on the east and the Mediterranean to the west, with Nahr

Hadera, being the only major river draining the western slopes.

(c) The Hills of Judea: This subregion can be divided into three sections from west to eastj the Foothills bordering the coastal plain, the Hills in the center, and the "Judean Desert" on the eastern slopes.

The central hills rise to a peak in the north (Jabal Asur, 1,016 m) and in the south (Jabal Halhul, 1,019 m) with a low saddle inbetweonn

(800 m), in the center of which lies Jerusalem. The foothills to the vxest average about 10 km in width and form a transitional zone between the hills and the coastal plain with no clear line of demarcation.

The eastern slopes of the hills are steep and the surface is of chalk rocks almost impermeable to water. This results in sudden floods during the rain season and extreme dryness the rest of the year ap­ proaching rigorous desert conditions.

The southern border of this subregion corresponds to a straight line drawn from Beersheba to the Dead Sea.

h. The Negev Region.- This terminology follows the convenience of custom by geographers and does not imply geological or geophysical separation of the Negev from Sinai. 22

Tho Ke;:'ev can bo subdivided into four subregions: the Central

Hills, Beeruheba Foothills, the Paran Plateau, and the Southern Hills. The Central Hills rise south of the Judean Hills along the southern border of tHe Dead Sea and the northeastern border of Uadi Araba and terminate cr.ne 90 km to the southwest into the Sinai Peninsula. The highest elev/ation is at the peak of Har Ramon (1,035 m). Southwards the hills br'eak up and descend into the Paran Plateau, which is inclined eastwards with the elevation decreasing from 600 m on tho

Sinai-Palest/ine borders to 100 m along Wadi Araba. Farther south, near Eilat, She Southern Hills rise to about 900 m and continue southwards into Sinai bordering the Gulf of Aqaba. North of the

Central Hills the Beersheba Foothills extend to about 10 km east of the Gaza Strip descending into the coastal plain,

HI. The Great Rift Valley:

This zone represents the northem end of the Syrian-East-African

Rift System -which extends from near Antioch, Turkey, to Lake Nyasa in

Africa, a distance exceeding 6,500 km (It,000-mi). This zone can be divided into four regions from north to south as such:

1. Nahr El-Assi (Orontes) Valley.- This is the northernmost section of the rift separating the Ansariyah mountains on the west from Ez-Zawiyah mountains and the hills of Homs and Hama on tho east.

The.lower valley, locally known as El-Ghab (also El-Rhab) rarely exceeds

12 km in width. It is continuously flooded with huge areas of swamps and marshes. Cultivation is extensive near Homs and Hama, but restric­ ted to elevated or well drained areas of the lower valley. The upper valley of this river belongs to the next region. 23

2. Tho Eekaa Valley.- This valley separates Mount Lebanon to the west from 1he Antx-Lobanon mountains and Mount Hermon to the east. It is approximately 130 km in length and never exceeds 1^ km in width.

Half-way in the valley lies the city of Baalbeck (ancient Helipolis), situated on a mountain '.trough x;hich rises up to 1,000 m and acts as a water-shed with Nahr El-Assi flowing northwards to lake Homs and Nahr

El-Litani flowing southwards to Lake Qaraoun.- These two lakes mark the borders of the Bekaa, but the rivers finally lead to the Mediter­ ranean. A large swamp, slowly being drained and converted into agricul­ tural land, still persists in the center of the Bekaa, near Ammik.

3. The Jordan Rift Valley,- This region includes the Jordan River valley and the Dead Sea. It extends from the upper Hula Valley in tho north to the southern tip of the Dead Sea. The Hula Valley, formerly occupied by Lake Hula and a complex of swamps, has been recently drained and made suitable for cultivation. The main source of water leading to the Hula Valley is Nahr El-Hasbani, which flaws from Lebanon draining the western slopes of Mount Hermon. The Hula Valley is in turn drained by the Jordan River which continues south via Lake Tiberias to pour into the Dead Sea south of Jericho. The northernmost part of the Jordan

Rift, near Dan, rises some 170 m above the Mediterranean Sea level, while to the south, where the river enters tho Dead Sea, the elevation drops to approximately 1|00 m (l,3l£ ft) below sea level. This repre­ sents the lowest point on earth.

The Jordan River valley is narrow to the north, but widens south­ wards reaching a width of 32 km near Jericho. The river meanders through tliis heavily cultivated valley receiving two major tributar­ ies; Yamuk River south of Tiberias and Zarqa River north of Salt, 2k

The Dead Sea is about 80 km in length and widest north of Ain Gedi

(18 Ion) but narrows to I4 km south of Metsada, where the Lisan Penin­

sula projects inwards dividing the sea into a northem and a southern

basin. The Dead Sea’s ma3.n source of water is River Jordan, also a multitude of wadis lead directly to the sea from the surrounding, mountains, however, of these wadis those draining the mountains of

Moab are the richest in water content, and some flow all year round. The Jordan Rift sends tiro major branches, Harod and Jezreel

Valleys, to the west'which may be considered as part of the rift

system. These valleys are heavily cultivated^rising from the Jordan

Valley to the north of the hills of Samaria and separating these hills from the hills of Galilee to the north. Many other valleys

connect with the rift valley to the east and west, but these are mostly river beds and wadi systems draining the mountains and will be considered part of the Upland Regions,

I4. Wadi Araba Valley,- This represents the southernmost sec­ tion of the Great Rift Valley extending from the southern end of the

Dead Sea to the Gulf of Aqaba on the Red Sea, It is about 165 km in length and never exceeds 32 km in width. The soil is mainly of alluvial sand and gravel resulting from flash floods descending the

steep wadis draining the surrounding uplands. These wadis carry

detritus and due to excessive erosion they build up wide alluvial fans where they enter Wadi Araba, Playa lakes (locally known as Qas, singular Qa) are common to the south, springs and oasis in tho middle, and mudflats along with swamps in the north,

IV. The Eastern Uplands:

This zone extends the whole length of the country bordering the

Giro at Rift Valley to the west and the Syrian Desert to the east. It is divided into seven regions from north to south as such: 2$

1. Eis-Zawiyah Mountain Region.- The northernmost Jabal El-

Wastani occupies a central position between tho Ansariyah Mountains to the west and Jabal Ez-Zawiyah to tho east. It is separated from the former mountains by Hahr El-Assi, while Lake El-Belaa and the

Er-Rouj Valley separates it from Jabal Ez-Zawiyah. Jabal Ez-Zawiyah « borders El-Ghab Valley throughout its length. This mountain is low, with an average elevation of 600 m, but peaks rise up to 939 m, north of Jozef, fill7 m, north of Kafr Aaouaid, and 826 m, south of Kafr Haya,

The western slopes are rather steep while the eastern slopes gradually intergrade with the Syrian Desert.

2. The Hills of Homs and Hama Region.- This region extends from a straight line across the southern border of El-Ghab Valley and Khan

Cheikhoun to Lake Homs in the south. Hahr El-Assi makes a U-turn in this region passing through Homs and Hama. The hills surrounding'the river valley are under extensive cultivation while those farther east are completely barren. The hills are rather low, with only two peaks exceeding 6^0 m in elevation; Jabal Abou Darde (677 m), and near

Arbaine (6$>U m)« 3. Mount Hermon-Anti-Lebanon Mountains Region.- This region ex­ tends 15 km southeast of Lake Homs to the Hula Valley in the south. The mountains are in two blocks, the Anti-Lebanons and Mt. Hermon, separated by a broad pass south of Zabadani. The Anti-Lebanon Moun­ tains average about 1,800 m in altitude with the highest elevation reaching 2,6l6 m at Jabal El-Atneine. Mt. Hermon, on the other hand, represents a single "whale back" ridge rising to 2,8lU m;the highest elevation in this zone. The western slopes are drained by Nahr El-

Assi, Nahr El-Litani, and Nahr El-Hasbani, while the eastern slopes are drained by small streams and wadis leading to the desert steppes. 26

The mountains of this region are sparsely vegetated with the original pine-oak forest restricted to the Zabadani pass drained by Malm Barada which flows eastwards into Damascus. I4. The Basan and Gulan Heights Region,- These hills rise from the foothills 0? Mt, Hermon and extend southwards to the Yamuk River, a tributary of the Jordan River. They abruptly rise from the Hula Valley and Lake Tilerias on the west and slowly intergrade with the Houran

Hills to the east. The northem hills are fairly high with a peak at Tell Ech-Cheikha (1,211 m), while the southern hills are low barely reaching UOO m in altitude.

£. The Ajlun Mountains Region.- This region corresponds to the biblical Gilead Region, bounded by the Yamuk River to the north, the

Jordan Valley to the west. River Zarqa to tho south and the Syrian

Desert to the east. The average width of this region is approximately

60 km, while the highest elevation attained is at Jabal Um-ed-Daraj

(l,2li7 th). Jabal Ajlun itself attains an elevation of 1,127 6, The Hills of Ammon and Moab Region.- This region extends from the Zarqa River to Vadi-El-Hasa along the southern end of the Dead Sea.

These hills correspond and for the most part run parallel to the Judoan

Hills, but in comparison these are much higher and support a greater variety of cultivated and natural vegetation due to a reliable rainfall during the winter. The region is much narrower than the proceeding region attaining an average width of 3£ km. Many streams and wadis drain these hills and lead to the Jordan River or directly to the Dead

Sea. Some of the important wadis are listed below (from north to south):

(a) Wadi Shueib passes through Salt and as it enters tho Jordan

Valley, near Shuna, it becomes known as Wadi Himrin.

(b) Wadi Es-Sir drains the highest peak of the Ammon Hills (Jabal

Dabug, 1,013 m) then joins Wadi Naur foming Wadi En-Nusariyat which 27 enters the Jordan Valley at Wadi Kafrein.

(c) Wadi "irqa Main originates at the hot springs of Ain Ez-Zarqa and drains tho Mountains southwest of Madaba and pours into the Dead Sea.

(d) Wadi El-Kujib is the largest river system in this region. It is

joined by irany side wadis draining the central hills of Moab (eleva­ tion 1,065 m at Shihan).

(e) Wadi El-Karak drains the hills west of Karak (elevation hipest at Tell EL-Meisa, 1,253 ro) and leads to Ghor El-Mazraa at the opening of the El-Lisan Peninsula.

(f) Wadi El-IIasa and Wadi Hudeira drain the southern hills of Moab attaining an elevation of 1,305 m at Jabal Ed-Dahab, the highest peak in this region, both wadis lead to the Dead Sea.

7. The Edom Mountains Region.- This is the largest region in this zone extending from Wadi El-Hasa, south of tho Dead Sea, to tho

Gulf of Aqaba. The mountains rise abruptly from Uadi Araba (100-

200 m) to peaks exceeding 1,600 - 1,700 m and then slope gently to tho east intergrading with the Eastern Plateau and the Southern

Desert. These mountains can be separated into three sections: the

Jebbal Mountains, the Shera Mountains and the Southern Edom Mountains, listed from north to south. This region hardly exceeds 35 hm in width to the north of Maan, but farther south the Southern Edom Mountains penetrate the desert an additional 35-!?0 l

Jabal Baqir (1,592 m), but eastwards where they penetrate the desert 28

Jabal Ram rises up to 1,7514 m«

V. The Syrian Deserts

Tho Syrian Desert occupies a median position in the Saharo-

Sindian Desert Bolt extending from North Africa to India. It is

bounded by the Eastern Uplands to the west, the Euphrates River

Valley, Iraq, to the east, the Nafud Desert, Saudi Arabia, to the

south, and the foothills of the Taurus Mountains, Turkey, to the

north. This great expanse of land can be topographically divided

into three rogions from north to south as follows:

1. The Northern Desert.- This region includes the area north

of the Houran Plain and Jabal Druz, and east of the Eastern Uplands

from Jabal Ez-Zawiyah in the north to the Anti-Lebanon Mountains in

the south. A chain of mountains rising from the foothills of the

Anti-Lebanons extend from Damascus to Palmyra and farther north marking the southern limits of this region.

2. The Eastern Plateau.- This region is comprised of the

area east of the Eastern Uplands from the Houran Plain in the north to

Ras-en-Neqeb in the south. The land is gently uhdulating with wadi

systems draining inland leading to three major depressions: El-Jafr

Depression, Wadi Sirhan, and the Azraq-Shishan Depression, The

desert is primarily of limestone with flints scattered all over

but a small section extending from Azraq-Shishan to Bahr El-Aateibe

in the north is covered by basalt rocks from the size of a pebble to the size of a boulder. These basaltic rocks have resulted from volcanic

outcroppings centered around Jabal Druz (l,G00m),

3. The Southern Desert.- This region includes the desert south

of tho Eastern Plateau primarily of sandstone and granite com­

position. Ecologically, as will be mentioned later, this region

i 29 also includes the Southern Edom Mountains, Uadi Araba and the lover slop'.s of the Eastern Uplands from the Gulan Heights to the Gulf of Aqaba# All wadis load to the Hisma Depression which decends southeasterly into Saudi Arabia running between Jabal Arqa

(1,336 m) and Jabal El-Qannasiya (1,322 m) in the-north and Jabal

Ram (1,7& m) and Jabal Um-Ishrin (1,753 m) in tho south* C. CLIMATE

A Mediterranean type climate predominates over tho Eastern

Mediterranean Region. This is characterized by a mild and moder-.. ately rainy winter and a hot and rainless summer, while spring and autumn are short and indistinguishable. Rainfall is variable and is influenced greatly by latitude, altitude and distance from the

Mediterranean Sea. The northwestern mountains may receive over

1,300 mm of rainfall annually, while parts of the Southern Desert receive as little as 2$ mm of rainfall. This tremendous variation in rainfall, however, is not random as one can quickly observe that rainfall decreases farther inland or southwards and increases at higher altitudes. It also appears that tho amount of rainfall is inversely related to changes in mean temperature, i.e. temperatures increase as one progresses inland or southwards and decrease at higher altitudes. This phenomenon is illustrated in Table 2 where a comparison of rainfall and mean temperatures is provided for six localities representing the Western Uplands (Jerusalem), the East­ ern Uplands (Amman), the Great Rift Valley (Jericho), tho Southern

Desert (Aqaba), and the Eastern Plateau (Mann and Shawmari

Agricultural Research Station).

Rainfall is more or less restricted to the months of November through April, with occasional- showers possible during October and

May. Rain, however, is not evenly distributed over the six month winter period, but instead limited to 30-60 rainy days. For example

30 31

Localities Jerusalem Amman Jericho Shavmari Mann Aqaba Months 166 .0-* 6b.0 39.0 1372 9.1} 5.3 January U.6-12.7** 3.9-12.5 8.7-19.8 2.3-H4.6 2.3-2J4.8 9.5-22.14 1214.0 60.0 31.0 8.9 2.9 3.9 February 5.0-114.0 14.5-H4.0 9.14-20.9 U.0-17.1 3.2-16.2 10.2-23.0 97.0 hit .0 17.0 I4.I 0.3 14.3 March 6.2-15.8 6.2-17.2 11.2-25.3 6.3-21.1- 5.7-19.9 12.9-27.1 314.0 13.0 9.0 23.5 6.8 3.6 April 9.1l-20.8 9.6-22.7 114.9-29.7 10.7-28.3 9.1-2li.l 16.7-31.U • 7.0 5.0 3.0 16.5 0.3 0.0 May 12.U-25.7 13.7-28.0 18.2-314.0 12.7-30.8 13.0-28.9 20.1.-35.7 0.0 0.0 0.0 0.0 0.0 0.0 June 15.5-28.2 16.3-30.8 22.0-38.U 17.0-36.3 16.2-33.5 23.9-39.9 0.0 0.0 0.0 0.0 0.0 0.0 July 17.1-29.5 18.14-32.0 2U.3-39.1 18.14-37.2 17.U-314.9 25.1-39.8 0.0 0.0 0.0 0.0 0.0 0.0 August I8.O-3O.I4 18.6-32.6 28.2-39.l4 18.2-37.8 17.14-3U.6 25.5-140.5 1.0 1.0 0.0 0.0 0.0 0.0 September 16.14-28.3 16.3-30.5 22.1-36.8 15.2-3U.2 H4.7-32.U 22.7-36.8 8.0 I4.O 3.0 10 .U 2.9 0.1 October H4.2-26.0 13.9-37.5 20.0-33.9 12.14-31.0 11.8-28.6 19.8-314.3 79.0 32.0 - 20.0 13.3 1.7 2.5 November 10.0-19.5 9.8-20.8 lii.3-29.0 7.14-23.1 7.14-21.6 15.9-39.5 109.0 143.0 30.0 16.14 6.7 8.8 December 6.2-114.5 5.1j-U.7 10.1-21.5 14.14-19.9 3.9-17.1 n.7-2l4.3

Table 2. Comparison of monthly rainfall and mean maximum and minimum temperatures of six localities. -H-Rainfall in mm; Min-Max mean temperatures in C° Data obtained from Jordan Tourism Authority, Amman, Jordan. 32

Amman receives 279«l| mm of rainfall (average for the years 1923-

19h7), but this rain is received in 3^4 rainy days with a maximum

daily fall of 78,7 mm (Hemsley & George, 1966), This phenomenon

is of tremendous importance especially in the desert -where rainfall

is less than $0 mm, evaporation is high, and a certain amount of

surplus viator must be made available to the plants for some time

to be able to survive.

Heavy rainfall causes extensive erosion,and the accumulation of

the eroded material in wadis allows many plants to thrive most of the

year, Playa lakes (Qas) which are common throughout the Syrian Desert,

are formed by surplus water drained by many wadis leading to a single

basin as well as a rise in ground water level (Hemsley and George,

1966).

Snowfall is restricted to tho Uplands north of the Dead Sea

Basin, It is, however, rather infrequent south of Mt, Lebanon to

the west and Mt. Hermon to the east. Jerusalem, for example, re­

ceives an average of two days of snowfall while Mt. Lebanon and

Mt. Hermon are usually snow covered from December through May with small patches of snow surviving the whole year. Hail is

frequent along the coast, the uplands north of the hills of Moab

and Judea, and the Rift Valley north of Tiberias.

Dewfall and relative humidity are highest near the shorei de­ creasing inland. Winds are westerlies, but occasionally when a low barometric pressure center develops over Egypt or Libya,easterlies blow over the eastern Mediterranean bringing fine dust and high

temperaturesj these winds are called ’’Khamsin". 33

Tho relationship of rainfall to evaporation determines tho amount of water available or retained within the ecosystem.

Evaporation rate has been operationally correlated with temper­ ature by Long (195?) for 2h stations in Jordan and Palestine, using the foil Giving fonnula of Emberger's:

Q X 1,000 (M + m) (M - m)/2

whore P is the annual average rainfall of the site

M is the average maximum temperature of the hottest

month

m is the average minimum temperature of the coldest

month

(All temperatures in absolute values)

He then plotted the values of Q against m and divided the studied areas into four bioclimatological zones (Figure 3).. Al­ though additional factors help determine evaporation rate, these zones are here adopted for the area of the present study:

1. Sub-humid Mediterranean Bioclimate.- This zone includes the coastal plain, north of Gaza, the Western Uplands north of the central hills of

Judea, the Rift Valley north of Tiberias, and the high peaks of the western Uplands north of Salt. Annual rainfall typically exceeds

£00 mm,

2. Semi-arid Mediterranean Bioclimate.- This zone extends from the central hills of the Negev northwards along the eastern slopes of the Western Uplands to Tiberias. It also includes the lower slopes of the Eastern Uplands north of Wadi El-Mujib and tho crests of the

Jebbal and Shera mountains farther south. Annual rainfall 300-500 mm. 3h

3. Arid Mediterrancan Bioclimate.- This zone is mostly a transition zone between the Seini-arid zone and the following, Saharan-Mediterran­ ean, zone. It is comprised of the middle Jordan Valley, tho foothills of the Eastern Uplands, the Northern Desert as well as the Houran

Plain and Jabal Druz of the Eastern Plateau. Annual rainfall about

1^0-300 mm.

It. Saharan Mediterranean Bioclimatc,- This zone includes the Eastern

Plateau south of the Jabal Druz area, the Southern Desert, Wadi Araba, the lower Jordan Valley, the Dead Sea basin, and the Negev excluding tho foothills of Beersheba and the central hills. Annual rainfall less than l£0 mm.

Climatic changes during Pleistocene times are of importance in zoogeographic studies. Four glacial periods dominated the climate in the Northern Hemisphere during the Pleistocene, During each glacia­ tion the polar ice cap extended southwards to nearly £o°N latitude reaching southern England and central Germany in Eurasia. No part of the Eastern Mediterranean Region was glaciated and there is no evidence that any of the high mountains in this region supported a permanent ice cap. Southward extension of the polar ice cap, however, resulted in a southern displacement of climatic belts. During glacial periods in the north the Eastern Mediterranean Region had increased rainfall, which have been called ,,pluvial,, periods. Glacial or pluvial periods alternated with warm periods called "interglacial" and "interpluvial" periods respectively. Pluvial periods allowed northern biotas (of the Euro-Siberian and Irano-Tauranian Rogions) to extend southwards along tho mountains of the Eastern and Western

Uplands as well as the principle drainage systems such as the Rift

Valley and various wadi systems leading southwards and eastwards 35

aloo Sv\b - V\unV»^

ScxW qo « B \oel»ot\

2o

7o em’\ - IVi

lo - KrtcH He^V^raA^o

20 : SHvueb » Z«rc^c\ S<\K<\ran H^AilefrOwACan lo ; <*[\n.rc.<\ •||,-HS • Ma.ixn "BxocVvvnaVvt Zoa« • El 7«fr * A^tok ■.i—. j... 0 " -j______i_ —1------>------1------A- +2 -*5 4? +6 •+T 48 4^ +\0 4 1\ 4ii m

Fig, 3* Bioclimtic zones of the Eastern Mediterranean Region, (Q= quotient pulviothermique of Emberger; m = mean of minimum temperatures of the coldest month). Modified from Long (1957), 36

into the desert. These northern biotas subsequently were forced to retreat during the dry and warm interpluvial periods, except as relicts on mountain tops and in lush valleys.

Less drastic changes in climate continued to occur during Recent times as evidenced from writings such as those of the Bible as well as archeological digs which have revealed cities and villages in some presently uninhabitable places. The desertification of the

Eastern Mediterranean Region, however, may not be entirely due to climatic changes alone as man himself has misused and deprived the envircment of its natural biota in his search for food and shelter for thousands of years. The deforestation of the Eastern Uplands and a good portion of the Western Uplands is a good example of man's destruction of his habitat. D. GENERAL ECOLOGY AND PlfiTOGEOGRAPIIY

The flora of the Eastern Mediterranean Region, and in particular

that of Palestine and Jordan, has been dealt with in great detail by many authors (Eig, 1931-1932; Zohary, 19h7 and 1962; Boyko, 19$h}

Feinburn and Zohary, 1955; Zohary and Orshan, 1956; Orshan and

Zohary, 1962; Poore and Robertson, 196li; and others). These studies

show that tho Eastern Mediterranean falls at the meeting point of three phytogeographical regions: the Mediterranean, the Irano-

Tauranian and tho Saharo-SIndian. Moreover, along with the flora representative of these three regions a few species characteristic

of the more northem Euro-Siberian Region and the southern Sudano-

Decanian Region can also be found. The numerical species proportions of these five phytogeographical regions in Palestine are listed below as an example for the whole area:

Mediterranean element 863 species

Irano-Tauranian element 309

Saharo-Sindian element 299

Euro-Siberian element 15

Sudano-Decanian element 20

^Tabulated here are species restricted to one region. For

example none of tho 36? species shared by the Mediteixanean

and the Irano-Tauranian Regions are included. (Extracted

from Zohary, 1962)

37 38

Elements of the Euro-Siberian Region are scattered throughout

the Eastern Mediterranean, vhile those of the Sudano-Eecanian Region

are confined to local enclaves in tho lower Rift Valley. The three principal yhytogeographical regions, however» are restricted to

particular belts representing three ecological regions: the

Mediterranean Region, the Steppe Region, and the Desert Region

(Fig. U). These are described below:

I. The Mediterranean Region:

This is comprised of the Coastal Plain north of Gaza, except for a stretch of sand dune area bordering the sea south of Haifa, the Western Uplands from the crests of tho Judean Hills northwards, the Rift Valley north of Lake Tiberias, and the Eastern Uplands from the peaks of the Jebbal and Shera Mountains northwards to the Ez-Zaw­ iyah Mountains.

This region corresponds to the Sub-Humid and Semi-Arid Mediter­ ranean Bioclimatic Zones where rainfall exceeds 300 mm. The soil is. typically of the terra-rossa variety, however, rendzina and alluvial soils are also common. The flora is typical of the Mediter­ ranean Phytogeographical Region.

The climax vegetation is evergreen sclerophyllous forest and maquis. The name "mquis" has been defined by Zohary (1962:03) as a term conventionally denoting "Mediterranean woodland dominated by sclerophyllous evergreen low trees and shrubs up to height of k m. or so..."*

Mediterranean maquis thus appear to represent a forest pre-climax which "under favorable ocologic conditions and in the absence of human interference, will develop into forest dominated by one or 39

CAST CRN MtOITCRRANCAN REGION

’•** Jti*»*»**,*'> 25 50 IS '00

OSO Mediterranean

fga Hn Desert

• v‘

.>•«•••*• It

Fig. I(. Ecological regions of the Eastern Mediterranean Region. Arrow indicates desert penetration along the coastal sand dune area of southern Palestine. ho more tree species".

Plant associations in this region can best be discussed under tvro headings as follows:

A. Mediterranean Forest and Maquis

Cedar forests once common in the Lebanon Mountains have disap­ peared in recent years except for isolated stands on mountain tops exceeding 1,900 m near Barouk, Becharre and Jaj. Pine and oak woodlands occupy a lower altitudinal vegetation zone extending from

sea level to mountain peaks. These woodlands have also been reduced

to isolated stands in recent times by man's attempt to expand avail­

able cultivated areas and his need for fuel.

Pine forests, typically confined to rendzina soils, extend from

sea level to altitudes exceeding 1,200 m in Syria and Lebanon. Tho

Aleppo pine, Pinus halepensis, is tho dominant species, but farther north Pinus brutia and Pinus uinea become equally abundant. Pine

stands are usually associated with an undercover of maquis, espec­ ially Quercus calliprinos, Pistacia (3 species), Arbutus andrachne and many other species of oak and a ground cover of low shrubs typical of the Mediterranean garigue and batha formations (these formations are defined on p.Ij2). Stands of pine mixed with

Junlperus phoenicea, J. oxycedrus and Cupressus sempervlrens can also be found scattered throughout the Western Uplands north of

Galilee•

Oak forests are found throughout the Upland Regions from Hebron and Petra northwards, but with different associations at different altitudes and localities. Deciduous oak forests arc common on the

Hills of Galilee (association of Quercus ithaburensis and Styrax officinalis), tho western slopes of the Ajlun Mountains, Basan and the Zabadani saddle (association of Q, ithaburensis and Pistacia atlantica) and the Sharon Plain where remnants of pure stands of

0 ithaburensis have been preserved until today. These forests are restricted to altitudes not exceeding $00 m. Evergreen oak forests and maquis are much more widely spread representing the commonest plant formations in this region. The dominant species, Quercus calliprinos, can be found in association with Juniperus spp. at altitudes exceeding 1,000 m and with Pistacia sp. at lower al­ titudes but never below 200 m. The Quercus-Pistacia association includes a variety of Mediterranean trees and shrubs such as

Laurus nobilis, Arbutus andrachne, Hhamnus alatemus, Styrax officinalis, Cercis siliquastrub, Rhamnus palaestina and Crataegus azarolus (the first three species are evergreen,the rest deciduous).

The Quercus-Juniperus association includes Crataegus azarolos,

Pistacia palaestina or P, atlantica and Rhamnus palaestina. A ground cover of Mediterranean batha and garigue is often associ­ ated with both of these associations.

Typical maquis communities can be found along the foothills of the Western Uplands where the main tree is a carob, Ceratonia siliqua. Undisturbed communities usually contain carob associated with Pistacia lentiscus accompanied with a dense ground cover of

Mediterranean garigue spec5.es. This formation extends onto consol­ idated sand dune areas along the Sharon Plain, while to the east along tho foothills of the lower Galilee and Samaria it extends into the Steppe Region and becomes associated with many species of the

Irano-Tauranian communities. h2 River valleys are usually heavily cultivated vhile river banks are dominated by Populus spp., Platans orientalis. Eucalyptus spp, and Nerium olgaonder,

B, Mediterranean Garigue and Batha

The communities considered here arc typically of dwarf shrub associations. "Garigue" is a term used to denote sclerophyllous shrub associations lying on or near the ground, usually less than

1,5 m in height, while "Batha" refers to shrub associations not exceeding 0.5 m in height. The leading plants of the garigue com­ munities are Calycotome villosa, Cistus villosus, Cistus salvifolius.

Salvia triloba, Satureja thymbra and Phlomis viscosa, while those of the batha communities are Poterium spinosum. Thymus capitatus, Fumana sp., Teucrium polium, Ballota undulata. Salvia spp., Echinops viscosa,

Origanum maru and others. These communities are also associated with scattered tree species representing the woodland climax towards which the batha or garigue sere is progressing. This is not always true as these formations may also represent a stage in woodland destruc­ tion ultimately leading to bare ground.

Mediterranean batha and garigue associations are common on mountain slopes extending into the Steppe Region where they are joined by several Irano-Tauranian species. These communities are of tremendous importance as safeguards against soil erosion, also they support an abundance of herbaceous winter annuals which bloom six months of the year providing enough food for livestock and wild gi'azing mammals. It is regretable that many villagers and bedouins rely solely on Poterium spinosum, the dominant species of the batha communities, and many other species as fuel for fii'e. This usually results in extensive erosion and bare ground in the cleared areas.

II. The Steppe Region

This region forms a belt around the Mediterranean Region (Fig.h) and in many respects it represents an "ecotone" between the Mediter­ ranean and Desert Regions. Geographically, this region extends from the central hills of the Negev and the Beersheba Foothills northwards along the eastern slopes of the Western Uplands to the upper Jordan

Valley, southwards along the western slopes of the Eastern Uplands to the Ez-Zawiyah Mountains increasing gradually in width from south to north.

This region corresponds to the Arid Mediterranean Bioclimatic

Zone receiving approximately 150-300 mm of rainfall. The flora is typically of the Irano-Tauranian Phytogeographical Region corre­ lated locally to edaphic conditions. The predominant habitats are rocky hills, gray limestone steppe soil and alluvial or loess soil.

The climax vegetation may have been a "Tall Grass Steppe"

(Poore and Robertson, 196ii:l8). At present the vegetation consists of a brush steppe, a grass steppe and a forest steppe. Cultivation is intensive in many sections of this region, especially in Syria and northern Jordan.

A. Forest Steppe

The dominant and only tree, Pistacla atlantica, forms open forests on the central hills of the Negev, eastern slopes of the

Ajlun Mountains, .the Anti-Lebanon Mountains and on certain ridges of the Northern Desert, The trees are of considerable age and young seedlings are never allowed to grow due to intensive grazing. Thus unless certain measures are taken this vegetation type is bound to disappear. The ground cover is a scrub of Irano-Tauranian species such as

Artemisia herba-alba and Noaea Mucronata and some Mediterranean species.

D. Brush Steppe

A brushwood association dominated by Ziaynhus lotus is found along the eastern slopes of the Western Uplands facing the middle

Jordan Valley, while farther south an association dominated by

Retama raetum and Rhus tripartita becomes more abundant especially on rocky hills. The commonest brush association of this region, however, is dominated by the sagebrush, Artemisia herba-alba. This association is found on the gray and yellow limestone steppe soils of the rolling hills and valleys, associated with other shrub species such as Ononis natrix, Noaea mucronata and Echinops spp. along with a ground cover of Poa sinaica and Carex pachystylis.

C. Grass Steppe

Grass steppe are found on loess silty soils, rocky hills and hill tops. The characteristic dominants are Poa sinaica and Carex pachystylis.

Ill, The Desert Region

This is chiefly comprised of the Negev except for the Central

Hills and the Beersheba Foothills, Wadi Araba, Dead Sea Basin, the

Judean Desert, the Eastern Plateau, the Southern Desert and a strip of sand dime area from Haifa south-ward to Sinai*

The region corresponds to the Saharan Mediterranean Bioclimatic

Zone where rainfall is usually less than 15>0 mm. The flora is typi­ cally of low halophytic orxerophytic shrubs many of which have leaves thickened or modified into thorns and spines. The majority of the snecies are confined to the Saharo-Sindian Phytogeographical Region. Zohary (1962:128-9)- distinguishes three types of deserts in the

Middle East based on vegetation as follows:

(1) 'Main deserts" where vegetation is sparse but maintained by rainfall, '.'his habitat extends into the Steppe Region.

(2) "runoff deserts" whore vegetation is restricted to depres­ sions due to accumulation of rain. None of the plant species is a true desert species.

(3) "absolute deserts" where no vegetation is po;- :ible.

All throe types can be found in the geographical .a studied.

The distribution of these three types, however, depends solely on local physiographic features which makes a comprehensive treatment of the Desert Region using this system rather futile. Poore and

Robertson (19614:17-18) divide the desert into two sections on geological grounds, as follows:

A. The Basalt and Limestone Desert

This desert is the typical formation of the Eastern Plateau and ’• eastern." Negev. The ground is covered by a layer of flints except for the northern section which is covered by basalt rocks of varying sizes resulting from lava flows originating at Jabal

Druz and other mountains in that area. Flint deserts in North

Africa, the Middle East and Arabia have been termed "hammadas" defined as "slightly rolling gravelly desert plains whose surfaces are strewn with vari-sized stone fragments and pebbles" (Evenari and Orshansky, 19l48:l).

Vegetation is abundant in wadis, sparse elsewhere. Ii6

B, The Sandstone and Granite Desert

This formation includes the Southern Desert, the Rift Valley from the loi-er Jordan Valley southwards, the Judean Desert, the southern cc.ist of Palestine and eastern Negev, The relief is rugged with mountains rising to over 1,£00 m. This desert consists of sandstone and granite with localized areas of blown sand and very few stretches of hammadas.

Vegetation dominated by Anabasis, Acacia and Haloxylon is abundant in wadis and on the mountains. Enclaves of Sudano-

Decanian vegetation are found at several localities in the Rift

Valley,

The principal plant communities and habitats of the Desert

Region are:

1, Grasslands,- This type is found between the scattered basalt rocks of the lava beds as well as on alluvial fans or large wadis, e.g. Wadi El-Mu.jib and Wadi El-Hasa. The dominant species are

Poa sinaica and the sedge, Carex pachystylis, Also, Bromus, an annual grass, can be found locally in depressions within the hammada,

2, Salt flats,- This type is found near the Azraq pools, the Dead

Sea and Wadi Araba, The dominant species, Nitraria rotusa, is usually associated with Aelurops littoralis, Tamarix passerinoides and Prosopis sp, (Dead Sea). A similar community found on coarse alluvium soils, common in Wadi Araba, is dominated by Salvadora persica. Near Azraq Nitraria communities are found on mounds scattered throughout the salt flats. The mounds result from the accumulation of brovm silt at the bases of these shrubs,

3, Mud flats.- These are usually devoid of vegetation except along the odge vhero a sparse cover of Anabasis sp, and some annual

grasses can be found. These flats will be covered with water for

a short period in the rainy season.

I4. Hranmadas.- The surface is covered with cherts of varying sizes

and vegetation is sparse on hills becoming more abundant in depres­

sions. The depressions are dominated by Artemisia sp,, Phlomis sp.,

Anabasis avbiculata and many annual grasses, while the hammada in

between is either bare or sparsely vegetated with annual grasses and widely scattered shrubs. In the proximity of sandstone the hammada may be covered with a thin layer of blown sand; vegetation is usually more abundant. In central and southwestern Negev some depressions

in the hammada may contain Acacretun fortilis along with Anabasis

artlculata.

5. Wadi spreads.- Flat silty wadi spreads common in the limestone and basalt desert are dominated by Artemisia herba-alba and Sledlitzia rosmarinifolla. Sandy and gravelly wadis in the limestone desert are of tall shrub such as Retama raetum and Atriplex halimus and a sparse shrub vegetation such as Traganum nudatum, Haloxylon spp. and

Zygophyllum dumosum. In the sandstone and granite desert the char­ acteristic dominants ox' the wadis are Haloxylon spp. and Anabasis articulata.

6, Blown sand.- This type is abundant in the sandstone and granite desert especially Wadi.Araba and the Southern Desert. The dominant plants are Haloxylon persicum and/or H. salicornicum on coarse sand and Traganum nudatum on fine sand (locally Tamarix sp, may become equally abundant in both cases). Steep hill slopes covered with sand are .usually devoid of vegetation.

7. Dare rocks.- These are scattered throughout the desert and can be granite, sandstone, limestone, chalk, conglomerate or basalt.

This habitat is devoid of rooted vegetation.

8. Coastal sand dunes.- The commonest plant association here is i Artemisia monosperma - Arisfcida scomria. The sand is fairly mobile.

9. Reed swamps.- This type is confined to the Azraq oasis. Phragmites sp. and Juncus sp. are the dominant plants.

10. Tropical or Sudano-Eecanian enclaves.- These are widely spread in Vadi Araba and the Dead Sea Basin. The principal plant associa­ tion is dominated by Acacia, 3 species, and a shrub layer of Anabasis articulata, Haloxylon salicornicum and many other Saharo-Sindian and

Tropical dwarf shrubs. Another association typical of this type is the Zizyphus sninachristi - Balantes aegyptiaca association, which also includes two Acacia species.

11. Human habitations.- This includes villages and scattered houses and research stations in the desert MATERIALS AND METHODS

This rfcudy is based on examination of somo 3,000 specimens, more than cv.a half of which I have collected myself on the following occasions:

1. 1963-1965. Master's thesis research at the American Univer­ sity of Bail Tit, Beirut, Lebanon. Also employed as assistant to Dr*

R. E. Lewis, Curator, AUB-Museum of Natural History* Survey of the mammals of Lebanon and short collecting trips in Syria and Jordan.

2. April-May, 1966. Mammalogist on the International Biological

Program- International Jordan Expedition* Survey of the mammals of the Azraq Desert Basin in northeastern Jordan.

3. June, 1966, Survey of the mammals of El-Jafr Basin, southern

Jordan, by request from Director, Bedouin Settlement Pilot Project,

Ministry of Agriculture, Amman, Jordan.

U. July-August, 1968. Field work in Lebanon and Jordan support­ ed by University of Connecticut, N3F travel grant (6b4j306X) for graduate studies In evolutionary and enviromental biology.

The specimens examined are either in ray private collection (SA) or were made available for study through the courtesies of Dr. H. W,

Setzer, U. S. National Museum (USM), Washington, D. C ., Drs. S.

Anderson and K. Koopman, American Museum of Natural History (AMNH),

New York, Dr. J. C. Moore, Field Museum of Natural History (FMNH),

Chicago, Dr. G. B. Corbet, British Museum.(EM), London, Drs. R. E.

Lewis (PEL) and J. Buchard, American University of Beirut-Museum of Natural History (AUB), Beirut, Drs. J. Oppermann and R. Angermann,

Zoological Museum of the Humboldt University (BZM), Berlin, Dr. E von Lehmann, Museum Alexander Koenig (AKM), Bonn, the late Dr. E, Mohr,

Hamburg University Museum (HUM), Hamburg, Dr. B. Lawrence and Mr. C.

Mack, Museum of Comparative Zoology (MCZ), Cambridge and Dr. D, L.

Harrison (DLH), Private collection (HA.RR), Sevenoaks, Kent, England.

(In parentheses are abbreviations as used in the text.)

The following is a list of abbreviations and definitions of cranial and external measurements used in this study:

TL => Total Length, from tip of nose to tip of tail, omitting

terminal hair.

T “ Tail, from dorsal base of tail to its tip omitting terminal

hair.

HB » Head and Body Length, from tip of nose to base of tail.

HF « Hind foot, from tip of longest claw to extremity of heel.

Measurements obtained from specimens at HI or DIH collections

do not include the claw.

E « Ear Length (from notch), from tip of pinna to lowest part of

auricle.

FA =* Fore Arm, from elbow joint to vrrist joint.

TR ** Tragus, the greatest length from base to tip

Cranial Measurements

GLS a Greatest Length of Skull, from most projecting point of

incisors to most projecting region at postei’ior extermity,

regardless of structure under question. 51

CBL Condylobaaal Lenrrth, frcr.i most projecting point of

ii.cisors to extremity of occipital condyle,

ZB » Zy■ .omatic Breadth, taken across greatest breadth of

zygomatic arches,

BB ** Br eadth of Bralncase, taken across iridest point of braincaso,

IG ** In'..ororbital Constriction, taken across narrowest breadth

interorbital region,

UM “ Upper Molars, from front of first upper molar to end of

crown of last upper molar. Taken only for rodents, lag-

morphs and hyracoids,

C-M3 " Maxillary Toothrow Length, taken from anterior of canine to

end of crown of last upper molars,

LM “ Lower Molars, from front of first lower molar to end of

crown of last lower molar. Corresponds to UM measurement,

C~m3 «* Mandibular Toothrow Length, taken from anterior of canine

to end of crown of last lower molar. Corresponds to C-M3

measurement,

M ® Mandible, from tip of incisor teeth to extremity of

mandibular condyle,

TB *» Tympanic Bullae, taken across the greatest diagonal length

from apex to anterior of mastoid portion of bullae.

Means and extremes of most of these measurements are listed in a systematic order in Appendix I, Standard deviations are given when two populations are compared.

Capitalized color terms mentioned in the text follow the system of Maerz and Paul (1950), 52

Few special Arabic and Hebrew terms used in conjunction with names of localities, worthy of mention, are defined below:

Ain “ Spring, a small source of water.

Bir 0 Well, sometimes spelled Beer.

Hammada 0 Flint desert (defined in text).

Jabal “ Mountain*

Jebal " Mountains,

Mogharet " Cavo,

Nahr “ River.

Qa » Playa lake, mudflat

Has » Tip; Ifead. i.e. source of a river, or a head-land.

Rosh in Hebrew.

Sahl " Plain

Wadi “ Valley, typically used for ephemeral stream beds, however,

it has been erroneously applied to permanent streams, e.g.

Wadi El-Mujob. Wadi El-Hasa, etc. SPECIES ACCOUNTS

53 This section treats 70 species belonging to the orders

Insectivora, Chiroptera, Lagcraorpha and Rodentia in some detail,

while the remaining 2h species belonging to the ordersCarnivora,

Hyacoidea and Artiodactyla are listed and a separate key is

provided for all species in each order.

Orders, families and genera arc arranged in phylogenetic

order according to Simpson (19W). Discussions under these headings

pertain to the systematic and zoogeographic problems associated with

these taxa in the Middle East. Also keys to families, subfamilies,

genera and species are provided under the appropriate heading.

Original citation and type species for each genus are taken from

Ellerman and Morrison-Scott (19^1).

Original citation, type locality, common names (English and

ArhlJc names only) and distribution are provided for each species

followed by some systematic remarks regarding the subspecies status

of Eastern Mediterranean populations.

Subspecies are treated in detail under the following headings:

Synonymy: All synonymies listed by Ellerman and Morrison-Scott

(19!?1) are omitted. Those listed in the text are either new or

published following the appearance of Ellerman and Morrison-Scott*3

Checklist. The authority and cate of publication of each proposed synonymy are indicated in parahthesis folloidng the citation. Type mater .alt If examined,all data pertaining to number of specimens, date, sex, locality, collector and the whereabouts

of these spBcimens will be provided} if no^then only the typo locality is listed.

General dlugribution: A very general statement followed by a paragraph or two listing all records from the Eastern Mediterranean

Region,

Specimens examined: Localities, numbers of specimens and collections whore these specimens are housed are provided here for all specimens examined.

Identification: Only few diagnostic characteristics are listed} more measurements are available in Appendix I,

Biology: A summary of most published data and my.personal field observations on the habits and habitat of tho subspecies. Order INSECTIVORA

Tills order includes a stock of primitive mammals which gave rise to tho diverse assembly of therian mammals as well as a group of modern highly specialized descendants. The geologic range of tho order extends from Upper Cretaceous to Recent.

Of the three Palaearctic insectivore families only two, Sori- cidae and Erinaceidae, are represented in the geographical area covered. Seetzen (l85h), however, reports on the occurrence of a talpid. Taina europea, from Lebanon. This report is questionable inasmuch as this boreal species has not been taken for over a century.

Following is a KEf to Eastern Mediterranean families:

1. Large forms, CBL over bO mm, HF over 2£ mm; zygomatic arch

present; eyes large and conspicuous; pelage very coarse with

spines covering most of the dorsal surface. (Hedgehogs),..,

...... Erinaceidae

2. Small forms, CBL less than 35 mm, HF less than 2h mm;

zygomatic arch absent; eyes small and inconspicuous; pelage

short, fine and dense. (Shrews)...,,...,...... Soricidae

Family ERINACEIDAE Bonaparte, 1833

Hedgehogs

This family is widely, distributed throughout the Ethiopian

Region (not found on Madagascar), the Oriental Region except for

Ceylon, and the Palaearctic Region except for Japan and the cold

56 S7 temperate coniferous forests above 60° N latitude.

Tx

Cabrera, 1925 a strictly Oriental subfamily including four genera

of hairy hedgehogs (moon rats), and Erinaceinac Gill, 1872 with

a distribution approaching that of the family.

There aopears to be no general agreement on the number of

genera belonging to tho subfamily Erinaceinac. Anderson and Jones

(1967) following Simoson (l9)-t5) list five genera, while Ellerman

and Morrison-Scott (1951) list three, Ellerman and Morrison-Scott

considered the two African genera Atelerix Pomol, I8I4O and Aethechinus

Thomas, 1918 as synonymous and accorded Atelerix only subgenoric rank

under Erlnaceus. This synonmy appears to be justified as the two

African genera are nonotypic and do not show sufficient structural

differentiations from Erlnaceus to warrant their recognition as

separate genera (Ellerman and Morrison-Scott, 1951 and Ellerman &

Korrison-Scott and Hayman, 1953)* Miller (1912) and Bobrinski et al

(1965) take the extreme position of recognizing only the nominal genus

as valid, however, this over simplification has been Condemned by many authors.

Three genera, Erlnaceus, Hemiechinus and Paraechinus, are pert­

inent to this paper. Geological records and the present distribution

of these three genera suggests a north African origin for the genus

Paraechinus, an east Asiatic origin for Hemiechinus and a European

origin for Erlnaceus. Thus, it is not surprising to find all three

genera in sympatry in the eastern Mediterranean as the area falls

along the dispersal routes of all three forms. Erlnaceus reaches the southernmost limit of its distribution in southern Palestine where Bodenheimer (1958) recorded a few "stragglers" near Gaza. 53

Eeserts seen to bo the primary barrier for this form, which is a typical brush and cultivated land inhabitant, On the other hand

Paraechinus and to some extent Fomicchinus are adapted for desert conditions. They seem to have dispersed along the Saharo-Sindian desert belt from opposite directions and at present their distri- i bution overlaps only in the center of this belt (Egypt to Afghan­ istan).

It is believed that hedgehogs (no particular species) were regarded as bad omens in ancient Egypt (Bodenheimer, I960). Today, however, hedgehogs are sought by many people in the Middle East for their so called "medicinal potentialities". Cooked hedgehogs es­ pecially their liver is supposed to be the cure for many diseases of which arthritis and rheumatism are tho most frequently indicated.

Bodouins look on hedgehogs strictly as a food item. They are usually fried in oil or barbecued along with other desert rodents such as jerboas and gerbils. The use of hedgehogs for foodis in no way:restric­ ted to the Middle East. Gypsies and Spaniards especially on Ibiza, one of the Balearic Islands off tho coast of Spain are known to rear, pickle and store Erinaceus algirus in barrels (Corbet, 1966).

All hedgehogs lack a baculum. The semi-pendulous penis (Fig.^B) has been examined in all three genera. The glans penis is identical in shape in the two genera Hemiechinus and Erinaceus, but is much larger in the latter. Paraechinus, however, lacks the lateral folds very prominent in the glans of the. two former genei’a. This close association of Hemiechinus and Erinaceus and their distinct separation from Paraechinus will also be noted in the following KEY to Eastern genera of hedgehogs 59

1. Spines parted anteriorly with a conspicuous mid-dorsal bare

patchj pterygoid fossa inflated, connected to tympanic cavity

dorsallyj four sacral vertebrae (Gupta, 1965)...... Paraechinus

Spines without an anterior mid-dorsal bare patch or vnth an

inconspicuous slit-like bare patch; pterygoid fossa shallow,

not connected to tympanic cavity; three sacral vertebrae...2

2. Largo form, CBL more than 50 mm, HF over 36 mm, ears short

less than 32 mm...... Erinaceus

Small, CBL less than 1;8 mm, HF less than 35 mm; ears long

always exceeding 35 mm...... Hemiechinus

Genus Erinaceus Linnaeus,

1758* Erinaceus Linnaeus, Syst. Nat., 10th ed., p. 52.

Type species: Erinaceus europaeus linnaeus

Two subgenera, Atelerix and Erinaceus, are recognised. The subgenus Atelerix is primarily African in distribution except for

Erinaceus (Atelerix) algirus which extends into Spain and south­ eastern France. The subgenus Erinaceus is highly eurytropic. Ognev

(1920) recognized ten species and twenty subspecies under the subgenus

Erinaceus in the Palaearctic Region, while Ellerman and Morrison-Scott

(1951) recognized 18 subspecies and only two species, E. europaeus and E. dauuricus. Tho latter form reported fron northern China,

Mongolia, and Transbaikalia has been referred to genus Hemiechinus 60

1.0 rum

I______I 1.0 mm

Fig. 5. The glans penis of A. Crocidura russula. SA 1153 from Afka, Lebanon, and B. Erlnaceus europaeus. SA 1063 from near Damascus, Syria. 61

(Allen, 193<3J Bobrinski ot al, 19Wi) but there still is no

general consensus in regard to its systematic status.

Erinaceus europaeus Linnaeus

17£8. Erlnaceus europaeus Linnaeus, Syst. Nat., 10th ed.,

p. 52.

Type Locality: Wamlingbo, south Gothland Island, Sweden.

Common names: European Hedgehog, Common Hedgehog, Kunfuth, Konfud,

Kababet Chouk, Khlund. (The last four names are classic and collo­ quial Arabic names and are used for no particular genus or species

of hedgehogs.)

Distribution: Europe, southern USSR from Ukraine to tho Pacific, north and northwestern China, northwestern Iran, Iraq, Turkey and

Eastern Mediterranean.

The characters used in distinguishing subspecies are based primarily on the color of the pelage, especially the fur on the underside, as well as skull dimensions. These characteristics, however, are highly variable even within local populations as is evident in collections from Lebanon and Syria. This may account for tho great amount of confusion in the literature regarding the subspecific status of Eastern Mediterranean hedgehogs. Barret-

Hamilton (1900) tentatively placed four specimens from Mount

Lebanon (no exact locality) under E. e. concolor. He also described, in the sane paper, E« europaeus roumanlcus as a subspecies restrict­ ed to Roumania and adjacent lands west of the Black Sea and the Sea of Marmara. Subsequently Thomas (1918b) raised roumanlcus to speci­ fic status and described E. roumanlcus sacer from Palestine and re­ ferred the four Lebanese specimens to it. Specimens from Lebanon and 62

Syria can individually satisfy criteria used in distinguishing both

E. e. concolor and K. e. roumanicus. However, in considering dis­ tribution patterns of these subspecies I am inclined to refer all

Eastern Mediterranean populations to the following subspecies:

Erinaceus europaeus concolor Martin

1838, Erinaceus concolor Martin, Proc. Zool. Soc. Lend, , p, 103*

1918. Erineeus roumanicus sacer Thomas, Ann, Mag, Nat’,-Hist'.,2(9) :212,

Near Jerusalem, Palestine, (Harrison, 1961>b)

1951, Erinaceus europaeus concolor Martin, Elleman and Morrison-

Scott, Checklist, p. 20,

Holotype: H-l Collection no, J>J>.12.2lj.83 collected by Mr, Keith Abbott

near Trebizond, Turkey, No data on sex and date of

collection.

General distribution: Lebanon, Palestine, Jordan, Syria, Iraq, western

Turkey and Transbaikalia,

Harrison (1961jb) recorded this form from near Natanya, Tivon, near Lod, Beit Dagan and near Migdal Ha'Emeq in Palestine, He erroneously quoted Misonne (195>7) as reporting this species from

Jabal Abiad, Syria, Misonne only mentioned that it is common in

Syria, Barret-Hamilton (1900) described four specimens collected by Mr, Baroody on Mount Lebanon,

Thomas (1918b) reported on a specimen from- Jerusalem, Palestine in­ dicating tliis to bo the southernmost limit of the species, but Bodonhei- mer (1958) showed that some reach farther south along the coast to Gaza,

Other Records,- Bodenheimer (1920 and 1935) Tristram (1866a and I88I4), 63

Specimens examined: 31« LKBANON (3 BM no exact locality): Beirut

(1 USNM; 2 AUB; 1 SA); Baalbek (2 SA); Dier Zahrani (1 SA); near

Junieh (1 SA); Nabi Chite (l SA); Ras Baalbek (3 SA). SYRIA:

Damascus (10 SA). JORDAN: Amman (1 SA). PALESTINE: Bethlehem

(l SA),' Jerusalem (li BM).

Identification: E. £. concolor approximates the size of Paraechinus aethiopicus, but is larger than Hemiechinus auritus (see measurements

Appendix 1.). A median spineless patch at the anterior margin of the spines on the dorsal surface is diagnostic for this form as well as P. aethiopicus, however, it is narrower (width less than

$ mm) in E. europaeus. Ear length usually does not exceed 32 mm in E. europaeus but usually over 3^ mm in the two other species

H, auritus and P. aethiopicus.

Biology: E. e. concolor is common along the coastal plain of

Lebanon and northern Palestine. It has successfully invaded tho hills of Galilee and Judea in Palestine, but does not ascend the

Lebanon mountains beyond 1,000 m. It is common in the Bekaa

Valley, the vicinity of Damascus and the Jordan uplands. The south­ ern limits of its distribution may be indicated by a line joining

Gaza on the Mediterranean to Tafila in Jordan. Obviously this boreal woodland species has been capable of adapting to steppe and seniarid conditions, but has failed to invade the extreme arid con­ ditions in southern Palestine, Jordan and the Syrian Desert.

The European hedgehog is strictly nocturnal in habits. If surprised at night it first'runs for cover, but when approached it stops and rolls up in a ball exposing the spines on the back and flanks only. This behavior is an excellent protection against 6h normal predators such as wild cats, foxes, jackals and large owls.

However, this also accounts for the large number of dead specimens seen along highways. One early morning in August, 1968 I counted ill I’oad kills along the highway between Baalbek and Has Baalbek, a distance of approximately 38 km. These hedgehogs do not dig their own burrows but seem to take shelter among vegetation, especially along tho bases of shrubs, or in stone walls. Near Bethlehem,

Palestine, I have seen fresh tracks of a hedgehog leading into a deserted fox den.

These hedgehogs are omnivorous in their diet, more so than other Eastern Mediterranean species. In tho laboratory they were found to relish eggs (boiled or raw), bread, apples, figs, pears, cheese, crackers, hamburger, dead mice, birds and lizards. The stomach contents of a specimen caught feeding on a dead lizard,

Agama stellio (Linn,), contained feathers, exoskeletons of beetles and grasshoppers as well as some unidentifiable vegetable matter.

They make loud growling sounds while feeding and will occasionally produce a sharp squeak. They display an aggressive behavior when confined to a cage, a captive adult male was found to jump upwards, with its spines erect, a distance over cm in an attempt to hit any object held directly above it.

E. exxropaeus is known to hibernate in Europe and the more northern sections of its range during winter months (Corbet, 1966),

However, no records of such behavior are reported from the Middle

East. Mating and reproduction' has been studied by many European scholars of which the work of Herter (1938) is outstanding. The 6$ fine anatomy of tho semi-pendulous penis, which has not. been illustrated before is shorn in Fig. 5>B. The urethra opens in a depression at the tip of a medial projection. This projection has two prominent lateral projections on either side of its base. The glans average 1)4 mm in length o-nd 6 mm in width in six specimens collected near Damascus,

Syria.

Genus Hemiechinus Fitzinger,

1866. Hemiechinus Fitzinger, S. Bi, Akad, Hiss, Wien,

Type Species: Krinaceus Plo ';is Sundevall = Hemiechinus auritus

aegyptius (Fi; ir)

Sixteen species have been referred to this genus, however, fourteen of them were subsequently reduced to subspecific level under two species complexes, H. auritus and H. megalotis.

Of these two species only H. auritus is found along the Eastern

Mediterranean.

Hemiechinus auritus (Gmelin)

1770, Erinaceus auritus Gmelin, Nov. Comment. Aced. Sci. Petrop.,

Type locality: Astrakhan, southwestern USSR.

Common names: Long eared Hedgehog. (Arabic names as for Erinaceus

europoaus)

Distribution: Cyrenaica, Egypt, Sinai, Palestine, Jordan, Syria,

Turkey, Cyprus, Iraq, Iran, ^Afghanistan, Caucasus, Kazakstan, western

Siberia, Mongolia, Turkistan, northern India and W. Pakistan.

Identification: This species lacks the mid-anterodorsal spineless patch characteristic of Paraechinus and Erinaceus, It is smaller in all cranial measurements than either genus in this area (Appendix 66

I). Ears long averaging over 35 nun in length. Pterygoid fossa shallow and does not connect to tho tympanic cavity.

Biology: H. auritus appears to inhabit extreme desert conditions as in the Gobi Desert of Mongolia (Kowalski, 1963) and the Rajastan

Desert in India (Prakash, 1963b and 19614b), semi-arid steppes as in most of the Middle Eastern countries and cultivated land especially along river valleys that transect the Saharo-Sindian desert bolt, such as the Nile Delta (Hoogstraal, 1962) and the Euphrates-Tigris

River system (Harrison, 196!|b). This species exhibits a step further toward desert adaptation than Erinaceus europaeus, and in a way it occupies an intermediate biotope between this species and the highly adapted desert species Paraechinus aethiopicus. Thus along the eastern Mediterranean H. auritus overlaps in distribution with E. europaeus in the north and northwest and with P. aethiopicus in the south and southeast.

Most of our information on the biology of this form comes from studies with H. a. collaris in the Rajastan Desert. The breeding season begins in May (Nader, 1968) and continues through October, but most births occur in July, August, and September (Prakash, 1955a and I960), One to six spineless, young/are usually bom; and their eyes stay closed for 21 days (Gupta and Sharma, 1961). The glans penis was examined in only one young specimen collected near Amman,

Jordan. It resembles that of E.europaeus in all respects except size.

There are many conflicting records of the feeding habits of this species. Most workers seem-to indicate that it is predominantly carnivorous but will occasionally feed on seeds and other vegetable 67 matter (McCann, 1937; Krishna and Prakash, 1956; Prakash, 1959, 1963a and 1961ja).’,rear Amman, Jordan I captured a young male H. a. calligoni which was feeding on a jird, Merioncs tristrami Thomas, caught in one of my traps. Stomach contents suggested a strictly animal diet as tho stomach was packed with insects, fur and skeletal and skin remains of an agamid lizard, Agama ruderata Olivier, Cannibalistic behavior has been also reported in this species in Iraq (Nader, 1968) and India

(Prakash, 1953 and 1955b)j however, in both of these cases tho animals were confined to a cage with insufficient food.

Long-eared hedgehogs are strictly nocturnal. They are surpris­ ingly fast runners^ Prakash (19514) clocked their maximum speed at

2$ inches per second. They construct a simple burrow which lies about 20 cm below the surface and is usually less than 50 cm in length (Krishna and Prakash, 1955), The burrow is usually construc­ ted at the base of a shrub or a rock to give it additional protection.

Systematic remarks: Two subspecies are recognized along the Eastern

Mediterranean: H. a. aegyptius from southern Palestine, Sinai and eastern Eygpt and H* a, calligoni inhabiting the steppes bordering the Syrian Desert from Aleppo, Syria to Amman, Jordan and penetrates much farther north through Turkey to the Caucasus, The populations of these two subspecies are widely separated from each other by the

Rift Valley and tho Western Uplands from Judea to Lebanon,

Hemiechinus auritus aegyptius (E, Geoffroy) l0O3« Erinaceus aegyptius Geoffroy, Cat. Mus. N. H. Paris, p. 1:6,

1876. Erinaceus syriacus Wood, Bible Animals, p. 83. Palestine,

(Harrison, 19614b) 4 68

19^1. Hemicchinus aurltus an^n3tins Fisher; Ellerman and Morrison-

Scott, Checklist, p. 2l|.

Type material: Egypt (no exact locality). Not located.

General distribution: Egypt east of the Nile Delta, Sinai and

Palestine.

Harrison (196!jb)re corded this form from Beersheba, 2? km N.

Beersheba, 27 km E, Beersheba and near Rishon le Zion in Palestine, while Bodenheimer (1958) reported it to bo common along the coastal plain from Tel-Aviv southward to Gaza and El Arish.

Other records.- Bodenheimer (1935 and I960)*

Specimens examined: 2. PALESTINE: Jaffa (1 Ei); near Lod (1 Ei).

Identification: H. a. aegyptius is identical to H. a. calligoni in all respects except for its slightly yellowish belly and an over­ all light coloration. The relationship between these two subspecies is poorly known. Most workers assign their specimens to either of these forms solely on geographical grounds.

Hemiechinus auritus calligoni (Satunin)

1901. Erinaceus calligoni Satunin, Port. Obsch. Est. Kazan, no.

192 p. 2 and P.Z.S., 1901, 2:281i.

1951 • Hemiechinus auritus calligoni Satunin; Ellerman and Morrison-

Scotfr Checklist, p. 25.

Lectotype: Aralyk, about 26.5 km south of Erivan, eastern Anatolia,

Turkey; not examined.

General distribution: Transcaucasia, Turkey, Syria, Iraq, Iran.and

Jordan. 69

Trouossart and Kollman (1923) report this hedgehog from Djeround and Ataibe in Syria. Harrison (I96ljb) recorded it from near Aleppo and Karyatein in Syria and Amman, Jordan, and Lehmann (1966a) from-

Damascus, Syria.

Specimens examined: 7* SYRIA: Aleppo (2 AUB); Karyatein (1 EM)j

JORDAN: Jerash (2 SA)j near Amman (1 SAJ 1 BM).

Identification: Same as aegyptius except for its pure -white belly and slightly darker coloration due to an extention of the median black band on the spines of the dorsal surface.

Genus Paraechinus Trouessart

1879» Paraechinus Trouessart, Rev, Zool. Paris, 7i2lj2.

Type species: Erinaceus micropus Blyth

Three species are recognized. P, micropus from India and W.

Pakistan, P. hyponelas from central Asia and southern Arabia, and

P, aethiopicus with a wide distribution extending from Morocco through north Africa and most of the Arabian Peninsula to Iraq

(Ellerman & Morrison-Scott, 195>l). P. aethiopicus is a highly adapted desert species, while micropus and to a greater extent hypomelas are primarily steppe and dry rocky mountain slopes inhab­ itants . On morphological grounds micropus and aethiopicus appear to be closely related, and as the two species are nllopatric it has been suggested (Ellerman and Morrison-Scott, 1951:27) that mi cmpps is only a subspecies of P. aethiopicus.

Of the three species only P. aethiopicus is found along the

Eastern Mediterranean* 70

Paraechinus aethiopicus (Ehrenberg)

1833. Erinaceus aethionicns Ehrenberg in Hemprich and Ehrenberg,

symb. Phys. Mamin., 2:sig. k, recto (footnote).

Type locality: Dongola Desert, Sudan.

Common names: Ethiopian Hedgehog. (Arabic names as for Erinaceus

europaeus)

Distribution: Morocco, Algeria, Libya, Sudan, Ethiopia, Egypt,

Sinai, Palestine, Jordan, Arabian Peninsula and Iraq.

Identification: A prominent mid-anterodorsal spineless patch (width

10 mm, length 20 mm) is characteristic of this form. Pterygoid fossae highly inflated and connected with the tympanic cavity dorsallyj

shallow with no connection to tympanic cavity in all other Eastern

Mediterranean hedgehogs. Ears long, usually over h2 mm in length.

Biology: The function of the bare patch on the crown of the head

is unknown. I believe, however, it may be of importance as a stri-

dulatory organ used as a warning device or in claiming territory.

A stridulatory organ has been shorn to occur in the Tenericidae (Gould

and Eisenberg, 1966), an insectivore family resticted to Madagascar,

and in no other mammal. In this group the spines are arranged in 2 bands in a localized area (1 cm ) on the back and the sound is pro­ duced when these bands are rubbed against each other. The spines on either side of the bare patch of P. aethiopicus as well as Erinaceus europaeus are close enough to touch each other and will produce an audible sound if rubbed together on a freshly killed specimen.

Ethiopian hedgehogs are capable of survival under extreme desert conditions. Their distribution in tho Middle East, however, indicates 71

a preference within the desert to oases and vegetated wadis where food

and water are more abundant. Stomach analysis of a specimen taken at

Azraq-Shishan, a desert oasis, revealed a strictly carnivorous diet.

More than of the contents appeared to be frogs (probably Rana

ridibunda Pallas,the only ranid that has been reported from this

locality) while the rest consisted of arthropods, particularly insects.

Another specimen taken near El-Jafr in a vegetated valley had almost

an empty stomach with very few insect remains,

Tho penis is tube shaped, pointed distally and without any

prominent lateral projections as in E. europaeus or H, auritus,

Systematic remarks: Ellerman and Morrison-Scott (1951) recognized

nine subspecies. Six of these P. a, dorsalis (Anderson and de Winton)

from southern Arabia, P, a. ludlowi Thomas from western Iraq, P, a,

oniscus Thomas from Aden, P, a, pectoralis (Heuglin) from Jordan,

P, a. deserti (Loche) from the Sahara, and the nominal race from

Sudan and southeastern Egypt, wore originally described as full spec­

ies, The remaining three forms were described as subspecies, albior

Pocock and albatus Thomas belong to P, dorsalis and blancalis Thomas

to P. deserti, Hoogstraal (1962) following Setzer (1957a) revived this

old classification, but Harrison (l96Ub:23) presents the following argu­ ment: "Quite a good case has been advanced by Setzer (1957a) for maintain­

ing the Arabian foras as a distinct species on morphological grounds, par­

ticularly because of the cranial differences from true aethiopicus.

However, tho Arabian forms are nowhere sympatric with typical aethiopicus

and there are many important similarities between them, especially when 72

all the Arabian races are considered. It therefore seems reasonable

at present to regard tho Arabian forms as well differentiated subspec­

ies of aethiopicus.11 I have examined specimens of P. a. dorsalis, P.

a. ludlowi, P. a. pectoralis and P. a, aethiopicus (SA, BM and FHNH

collections) and found no characters that could justify elevating any

of the throe former subspecies to a specific status. I am able to

show, however, contrary to my previous statement (Atallah, 1967c),

that P. a. dorsalis is a junior synonym of P. a. pectoralis (see details

under subspecies discussion).

Two subspecies are recognized in the geographical area covered,

these are: P, a, pectoralis and P. a. ludlowi.

Paraechinus aethiopicus pectoralis (Heuglin)

1861. Hemiechinus pectoralis Heuglin, Nova Acta Loop, Carol,, 29:22.

1901. Erinaceus dorsalis Anderson & de Winton, Ann, Mag. Nat, Hist.,

7:h2, Hadramut, southern Arabia. NEW SYNONYMY'

19^1. Paraechinus aethiopicus pectoralis Heuglin ■; Ellerman and

Morrison-Scott, Checklist, p, 27,

Holotype: Heuglin (l86l) does not indicate where the type was

deposited. I have looked through the collections of the

Berlin Zoological Museum, but was unable to locate it.

The single specimen on which this name was based,

however, was obtained from Petra, Jordan.

General distribution: Sinai, southern Palestine, Jordan and Saudi

Arabia, Harrison (196Lib) reported on specimens from Wadi Araba 'and Beer-;_>

sheba: in:Palestine,'--Tristram (l88]j) .and Bodenheimer (1953)- reported it as common in Southern Palestine. Atallah (1967c) 73 recorded a specimen from El-Jafr, Jordan,

Specimens examined: JORDAN: El-Jafr (2 SA), 16 specimens of dorsalis

(here a synonym of pectoralis) from Sinai and the Arabian Peninsula in collections of BM (lO'-Holotype), FMNH (£), and AUB (l).

Identification: Spines are double banded (white base-black band - i white band - black band - white tip), however, the white tip is reduced to absent along the medial surface. This results in the appearance of a dark median stripe. Head with a prominent dark brown medial band separating the white fur posterior to the eyes on either side.

Systematic remarks: I had maintained in a previous publication

(Atallah, 1967c) that pectoralis was "definitely different" from dorsalis because it lacked the prominent dark median stripe of tho latter. This was based on a comparison of one specimen from El-Jafr

(practically a topotype for pectoralis), Jordan to a dorsalis from

Rafha, Saudi Arabia. However, when long series of dorsalis were examined the variations in the degree of prominence of the dark median stripe were immediately noticed. Specimens from Sinai, re­ ferred previously to P. a. dorsalis (Wassif and Hoogstraal, 195>h), have a very faint median stripe identical to the Jordanian specimens.

On the other hand specimens from northeastern Saudi .Arabia (Rafha specimen for example) show some intergradation with P. a. ludlowi which results in the increase of prominance of the dorsal stripe.

They do not, however, reach ludlowi color pattern where the spines on the lateral surfaces are completely bleached while the dorsal spines are predominantly black tipped. Paraechinus aethionicus ludlowi Thomas

1919. paraechinus ludlowi Thomas, J. Bombay Nat. Hist, Soc., 26:7h8.

19!?1. Paraechinus aethiopicus ludlovd Thomas J Ellerman and Morrison-

Scott, Checklist, p. 28.

Holotype: BM collection no, 19.3.1.14. collected at Hit along the

Euphrates, Iraq on August 8, 1918,

General distribution: Iraq west of Bagdad and Jordan east of Amman.

Atallah (1967a) recorded two specimens from Azraq-Shishan,

Jordan. Lewis, Lewis and Harrison (1969) had erroneously referred

a specimen of P. a. pectoralis from Rafha, Saudi Arabia to this fom.

Specimens examined: JORDAN: Azraq-Shishan (1 SA),

Identification: The spines on the lateral surface are completely yellowish-white, lacking the subterminal dark band present in pectoralis. This results in the appearance of a black median stripe flanked on either side by a white band. Fur on head with a continuous * white band posterior to the eyes transected by an irregular thin greyish brown band extending from the bare patch on the cream of the head to tho tip of tho nose. This band widens anteriorly to cover the whole face from tho eyes forward.

Family SORICIDAE Gray, 1821

Shrews

This family is widely distributed throughout the northern hemisphere except for the cold polar areas above 75’° N latitude and southern hemisphere except for the Australian Region 'and south of the tropics in South America, 7?

Two genera, Suncus and Crocidura, and five species are recorded from the Eastern Mediterranean* All of these species are of a more northern origin and must have spread southwards during pluvial periods of the Pleistocene.

In general, shrews are restricted to wet habitats such as water courses and heavily vegetated mountain slopes. Their presence at an oasis (Atallah, 1967a) separated by a long expanse of desert from the next habitable place can only be explained if one is allowed to postulate that during pluvial periods the Mediterranean fauna and flora was able to extend much deeper into the arid zone. Tho with­ drawal of the biota during dry inter-pluvial* would then allow mesic species to remain only in areas with an ample supply of water such as oases, and would result in the present pattern of distribution.

All soricids lack a baculum. The glans penis is tube shaped with the urethral opening situated in a. depression at the tip guard­ ed by a number (usually two) of tubercles. The exterior of the glans is either grooved or covered with tiny spicules (Martin, 1967). The glans penis of the two Eastern Mediterranean genera, Suncus and

Crocidura, are surprisingly similar (Vinogradov, 1958} Bobrinskii et al, 1965). This indicates a close relationship between these genera and would further implement Ellerman and Morrison-Scott's

(l95l> p. 61;-65) statement: "The retention of the genus Suncus, based on species which have an extra small upper unicuspid tooth, is largely a matter of convenience, strictly speaking, it is not more than a subgenus of Crocidura." 76

Following is a KEY to Eastern Mediterranean genera of shrews:

30 (rarely 32) teeth, U upper unlcusplds...... Suncus

28 teeth, 3 upper unicuspids...... Crocidura

Genus Crocidura V.raglor

1832. Crocidura Wagler, Isis, p. 2?£.

Type species: Crocidura leucodon (Hermann)

The nine Palaearctic species of this, genus (Elleman and

Morrison-Scott, 19^1), can be subdivided into north African (3)

and Eurasian (6) species. Four species of the latter group have

been recorded from western Europe and they all reach their

southernmost limit of distribution along the Eastern Mediterranean.

Two species, C. russula and C. suave ole ns, however, v:ere able to

penetrate much farther south into the African continent, but

they did so across the Strait of Gibraltar between Spain and

Morocco on the other side of the Mediterranean (Setzer, I960).

This appears to be the case as both species are absent (C, russula)

or rare (C. sauveolens) in southern Palestine and Jordan, and common

in Algeria, Morocco and Spain.

Crocidura leucodon, C, lasia and C. suaveolens are extremely rare

along the eastern Mediterranean. This is indicated in collections

as not more than half a dozen specimens of any of these three species have been taken from this area to date. C, russula on the other

hand is very abundant and occupies a wide range of habitats includ­

ing semi-arid steppes, Tho scarcity of the first three cited spec­

ies may be attributed to the general tendancy of animals to become

less abundant along the fringes of their ranges, A stronger argument, 77 however, may be in favor of the competitive exclusion principle where the high adaptive potential of C. russula has allowed it to compete satisfactorily with the other three species in the area.

Crooidurans appear to have been much more abundant throughout the Pleistocene of Europe and the Eastern Mediterranean (Haas, 1966;

Kurten, 1968). Three of the four presently described species of western Europe and the Eastern Mediterranean are knows as fossils since the last glacial period (100,000 years), while the fourth species, C. lasia, has no fossil record.

Following is a KEY to Eastern Mediteranean species of

Crocidura:

1. Lino of demarcation between dorsal and ventral surfaces color

sharp; tail short, less than 36/5 of TL...... 2

Line of demarcation not sharply defined; tail long, more than

36:5 of TL...... 3

2. Large, CBL more than 22.0 mm, HF more than 13.0 mm, TL usually

over 120 mm...... C. lasia

Smaller, CBL less than 21.5 mm, HF less than 12,5 mm, TL less

than 115 mm..C. 3cucodon

3. Medium size, TL more than 100 mm, CBL more than 18,0 mm......

...... C. russula

Small, TL less than 100 mm, CBL less than 17.5 mm..,......

...... C. suaveolens

Crocidura leucodon (Hermann)

1780, Sorex leucodon Hermann,- in Zimmermann, Geogr. Gesch,, 2:382.

Type locality: Vicinity of Strasbourg, Bas Rhin, Eastern France.

Common names: Bicolor VJhite-toothed Shrew, Salita (Arabic). 78

Distributi< n: Europe from France eastwards into western and southwestern

USSR and northwards to Central Siberia, Turkey, Iran, Lebanon, and

Palestine.

Harrison (1963b) observed that tho type specimen of Crocidura russula .judcdca described from Palestine actually belongs to the

Crocidura leucodon group. Thus, he subsequently transferred the form judaica to the latter species hitherto unknoim from the Eastern

Mediterranean. This population represents the southernmost record of the species,

Crocidura leucodon Judaica Thomas

1919. Crocidura russula Judaica Thomas, Ann. Mag. Nat.-Hist.^3(9):32,

1963. Crocidura leucodon Judaica Thomas; Harrison, Bull.

Res. Council Israel, 11B(I4):177-183.

Holotype: BM collection no. 18.8.1.3 An adult male collected by Capt,

0, C. Shortridge near Jerusalem, Palestine, in April, 1918.

Distribution: Palestine and Lebanon.

Thomas (1919a) described two specimens taken near Jerusalem,

Palestine, while Harrison (196Ub) recorded it from Carmel Caves (20 km

S. of Haifa), Mishmar Ha’Emeq and Nahr Rubin in Palestine.

Atallah and Harrison (1967) reported on the first record of this species in Lebanon, where tiro specimens were taken.

Specimens examined: 16. LEBANON: 10 km NE Rayak, Bekaa Valley (1 SA)j

S. slope Mt. Sanino (l SA)j Ajaltoun (1 SA). PALESTINE: Near Jerusalem

(2 BM). 3,1 specimens at USNM from Turkey. 79

Identification: Distinctly bicolored, grey dorsura and white belly,

with sharp line of demarcation along the flanks. In this respect it

resembles C. lasia but the latter form is much larger (Appendix I).

It overlaps in all external and cranial measurements with C. russula,

however the ratio of tail length to total length is usually less than

36^ in leucodon and always greater in russula.

Biology: Bicolor white-toothed shrews favor extremely wet habitats

with a dense planb cover. On Mt. Sanine (elev. 2000 m), Lebanon,

I caught one specimen at the base of a shrub, Berberis cretica L.,

which was abundant at this location. However, at the Bekaa Valley

(elev. 050 m), near Rayak, Lebanon, I trapped another in a grassy field.

In both of these cases the shrews were caught at burro:-: entrances

belonging to Microtus guentheri, a highly colonial vole. It appears

that these shrews do not dig their own burrows instead they rely on

rodent burrow systems. Nothing is known of this shrew's mode of life,

Crocidura lasia Thomas

1906, Crocidura leucodon lasia Thomas, Ann. Mag. Nat; ..Hist.*; 17(7):!»l6,

1991. Crocidura lasiura lasia Thomas^ Ellerman and Morrison-Scott,

Checklist, p. 81:,

Holotype: BM collection no. 6.3.6,10, an adult female collected on

November 20, 1905, from Scalita, near Trebizond, Turkey.

Distribution: Turkey and Lebanon.

Harrison (I96hb) reported on one specimen from Shemlan, Lebanon,

at the BM. Atallah and Harrison (1967) recorded it from near Mar

Abda, Nahr El-Kelb, Lebanon, 60

Specimens examined: h« LRHAl-ION: Near Mar Abda, Nahr El-Kelb (1 SAJ 1

HARR) j Shemlan (1 BM); ?. km V, An jar (1 SA).

Systematic remarks: The form lasia is here elevated to a specific status for tho first time.

Lay (1967) presents an excellent discussion of the status of this form. He convincingly illustrates that lasia does not belong to the C. lasiura group as maintained by Ellerman and Morrison-Scott

(19^1), Harrison (l96ljb) and Atallah and Harrison (196?). Lay concludes that Thomas (1906) was correct in placing lasia as a subspecies of C, leucodon. The specimens from Lebanon, however, are found to occur sympatrically with C. leucodon judaica. Thus lasia cannot be considered as a subspecies of C. leucodon, instead it should be accorded full specific status as maintained here.

Identification: This is the largest shrew along tho Eastern Medi­ terranean with CBL exceeding 22.0 mm; CBL in all other species never reaches 21.5. It resembles C. leucodon in being sharply bicolored, with a white belly, and short tail. However, these species differ in all measurements, C. lasia having larger dimensions.

Biology: This is a rare species as only four specimens have been taken since 1906. The three specimens collected in Lebanon (2 SA;

1 HARR) were trapped around dusk, two in a thick hedge along a stream and one within a Microtus guentheri colony in an apple orchard.

Crocidura russula (Hermann)

1780. Sorex russulus Hermann, in Zimmermann, Geogr, Gesch., 2:382,

Typo locality: Near Strasbourg, Bas Rhin, Eastern France.

Common names: Common European White-toothed Shrew, Salita (Arabic).

Distribution: Northwestern Africa (Morocco and Algeria), Europe except 81

for the British Isles and Scandinavian countries,southwestern USSR,

Turkey, Syria, Lebanon, Palestine, Jordan, Iraq, Iran, Afghanistan,

Kashmir, China and Japan.

All item Meditein'anean shrews belonging to this species were

referred to C. r. .judaica Thomas (Thomas, 1919aj Bodenheimer, 1935

and 1958), until Harrison (1963b) shoved that the type of .judaica

actually belongs to the C. leucodon group. Subsequently, Harrison

(I961|b) referred all his specimens of this species from Lebanon and

Israel to the race C. r. monacha Thomas, described from Scalita,

Turkey,

Recently Richter (1966) and Lehmann (1966a) have independently

elevated the form C. r. guldenstaedti (Pallas), known from the Cau­

casian mountains just to the north of the range of C. r. monacha

(Ognev, 1928j Ellerman and Morrison-Scott, 1951), to a full specific

status and referred specimens from Turkey, Syria, Israel, Greece

and Crete to it. There appears to be, in my opinion, no justification

by these authors for such a systematic rearrangement. Lehmann (1965

and 1966a) assigned all his specimens from Turkey to C. g. guldenstaedti

giving no reasons for elevating guldenstaedti to a specific level nor any

discussion of the status of C. r. monacha previously described from that

country. Richter (1966), on the other hand, maintained that monacha

may prove to be a good ^subspecies of C. guldenstaedti or just a

synonym to the nominate form of this species. He does not, however,

give any satisfactory explanation for recognising C. guldenstaedti

as a distinct species. I have examined over 150 specimens of C. russula from southwestern Europe, Turkey, Iran, Lebanon and Palestine (SA, BM,

BZM and FMNH collections) and found no justification for distinguishing 02 the form f^ldonstacdti as a distinct species. I also believe that

C, r. monacha is perfectly valid on account of its shorter tail and in general smaller external measurements when compared to C. r. guldenstaedti. Thus as understood here C. r. guldenstaedti repre­ sents a northern population restricted to the Caucasus mountains while C. r. monacha represents a southern population extending from the Anatolian Plateau, Turkey southwards along the Mediterranean to Jordan and Palestine,

Crocidura russula monacha Thomas

1906. Crocidura russula monacha Thomas, Ann. Mag. Ikt.Huh, 17(7)':U71,

Holotype: BM collection no. 6,3.6.13, An adult female collected

Dec. 3, 190£, at Scalita, south of Trebizond, Turkey (elev. 700-

1000 m).

Distribution: Turkey, Syria, Lebanon, Palestine and Jordan,

Harrison (1963b) reported this species from Beit Oren, Haifa, Dan,

Beit Heir, Hulata, Wadi Tira, Beit Hananya and Carmel Caves in Palestine, while Allen (1915) recorded it from Rashaya, Baniyas, Amik and Aithenit in Lebanon. Harrison (l96l|b) added the following collection sites: Beit

Yannani, Palestine and Kammacha, Lebanon. Atallah (1967a) reported on one specimen from the Azraq-Shihan desert oasis, the first record of this species from a desert habitat.

Specimens examined: 118. LEBANON: Nahr El-Kelb, near Mar Abda (IJ4 SA)j

10 km N. E. Rayak, Bekaa (l SA); Natural Bridge, Faraya (2 SA); Ras

El-As si (1 SA); Afka (3 SA); El-Rumeileh, 7 kmS.W. Laklouk (3 SA); 2 km

W. Anjar (l SA). PALESTINE: Bethlehem (9 SA); Beit-Sahur (3 SA); h km N. V/. Beit Fajjar (5 SA). JORDAN: Azraq-Shishan (1 SA). TURKEY:

(Ml BZM; 21; USNM). 83

Identification: This is a medium sized shrew resembling £. leucodon in size, but differs from this form by having a longer (more than 36^ of TL) tail and an indistinct lino of demarcation between tho dorsal and ventral surfaces. The overall coloration varies from light brown to dark grey.

Biology: This shrew is common throughout the Mediterranean Region.

It is usually found near streams, rivers, lakes, swamps and other moist habitats. Though specimens can be caught any time of the day and night, they seen to show a peak of activity around 1700 - 2000 hours. They do not construct burrow systems but instead they take refuge in rock piles, terrace walls and hedges. They feed primarily on snails, earthworms, insects and other arthropods.

Pregnant shrews were caught in March (3 females with 14,14 and 3 embryos), April (one with U embryos) and June (one with 3 embryos), while three specimens caught in July were lactating. T of the pregnant females (June and late March) were also lacta’ at the time. This suggests that these shrews breed continuous).^ from March through July, Kahman & Kinlechner (195>9) noted that in Sardinia this species breeds from March to September with litter sizes from

2-6. Adult male testes are green colored (I can't find any reference pertaining to the significance of this coloration, consistent in all specimens examined) measuring 5x3 mm.

Crocidura suaveolens (Pallas)

1811. Sorex suaveolens Pallas, Zoogr. Ross. As., 1:133, pi. 9 Fig. 2.

Type locality: Khersones, Crimea, USSR.

Common names: Lesser White-toothed Shrew, Salita (Arabic) Distributi* u vSouthem Europe front Spain to Poland, central and southern

Russia to r .istern Siberia, Mongolia, China, Afghanistan, Iran, Turkey,

Eastern Met terranean, Iraq and varius localities in northeast and

west Africa,

All s; ocincns from the Eastern Mediterranean are referred to the » following i -.tbspecies: •

Crocidura suaveolens portali Thomas

1920, Crocidura portali Thomas, Ann. Mag. Nqt.Hist., £(9).*119,

193$* Suncus tristrami Bodenheimer, Animal Life in Palestine, p. 95.

NEW SYNONYMY

1951. Crocidura suaveolens portali Thomas; Ellerman and Morrison-"

Scott, Checklist, p, 77*

Holotype: BM collection no. 19.il.11.9 from Ramleh, Palestine.

General distribution: Palestine, Lebanon, and Sinai.

Harrison (1956b) recorded this species from Iraqi Kurdistan

based on a damaged specimen, but he has since decided that this

specimen represents C, russula (Pers. Comm.). Bodenheimer (1958)

reported seeing two specimens one from the Negev and one from Dan,

Palestine, while Bate (19ii5) reported this species from owl pellets

at Becharre and Jaj, Lebanon. Wassif (I95iia), Setzer (1957a) and

Hoogstraal (1963) have reported and discussed the existence of one

specimen collected in southern Sinai.

Specimens examined: Only the holotype.

Identification: This is tho smallest white-toothed shrew in this

region viith CBL less than 17.5 mm. '

Biology: Nothing is known of the ecology of this species in the Eastern

Mediterranean. Its presence in the Negev and Sinai, however, indicates

a high tolerance for arid conditions. 8$

Genus Suncus Ehrenberg

1833. Suncus Ehrenberg,in Hemprich and Ehrenberg, Symb. Phys. Mamm.,

2:k,

Type specif s: Suncus sacer Ehrenberg e Suncus murinus sacer Ehrenberg.

Two species, Suncus etxmscus and Suncus murinus, are reported from tho Middle East, but only the former species has been collected from the geographical area covered. The house shrew, S. murinus, is widely distributed throughout the Old World tropics and has also been reported from the Arabian Peninsula, Egypt, Iraq and Afghanistan.

It is believed, however, that the occurrence of this shrew outside the tropics has been greatly enhanced by man (Cheesman, 1920; Ellerman and Morrison-Scott, 1951J Harrison, 19614b), especially because of its highly commensal habits. If it has not been already introduced, it could be in any one of the major ports on the Mediterranean or the Gulf of Aqaba. Actually, in 1965 I was told by natives at

Aqaba, Jordan of the occurrence of "Mice with pointed snout", but never had the opportunity to trap for them.

Suncus etruscus (Savi)

1822. Sorex etruscus Savi, Nuovo Giom, de Letterati, Pisa, 1:60.

1951. Suncus etruscus Savi£ Ellerman and Morrison-Scott, Checklist, p»68.

Type locality: Pisa, Italy.

Common names: SaviT3 Pygmy Shrew, Dwarf Shrew, Salita (Classical

Arabic used for shrew, no particular species).

Distribution: Southern Europe, southwestern USSR, Turkey, Lebanon,

Palestine, Iraq, Iran, PeninsularIndia, Burma, Ceylon

and the following countries in Africa: Algeria,

Nigeria, Kenya, Tanzania, Republic of South Africa. 86

The peculiar discontinuous distribution of this species may be attributed to tho difficulties encountered in collecting such a minute form which can escape from almost all kinds of traps normally used by mammalogists.

The spotty distribution of this species in the Middle East makes the question of subspecies a most formidable one. Thus most workers in the Middle East, including myself, have evaded the question by referring their specimens to the nominal subspecies. I am afraid I cannot improve on the situation at this time and will follow the general trend by assigning all Eastern Mediterranean specimens to the nominate form.

Suncus etruscus etruscus (Savi)

Synonymy under species.

Type material: From Pisa, Italy. Not examined.

General distribution: Southern Europe, southwestern USSR, Iran, Turkey

Iraq, Aden, Palestine, Lebanon, Jordan, Algeria, Morocco and Nigeria.

Harrison (196)48) recorded this form from Tiberias, Dan, Rehoboth,

Hulata, near Tel Hassan, 3 km W. of Wadi Zikhron}Ya1aqov and Beit Yanani in Palestine. Tristram (l88!t) reported a specimen identified as Sorex pygmaeus from near Mar Saba, Palestine which Bodenheimer (1958) had reasons to believe that it represents S. etruscus. Atallah (196?a) recorded the species for the first time from Jordan at Azraq-Shishan oasis,

PEL gave mo a specimen collected at Nahr Beirut, Lebanon, a new record for this country.

Specimens examined: JORDAN: Azraq-Shishan (2 SA)j LEBANON: Nahr

Beirut, near Hazmiyeh (l SA). I also had a chance to examine DIH 87 collection;* from the localities listed above.

Identification: This species is believed to be the smallest mammal in the world, TL less than 80 mm, CBL less than 13.3 mm (other mea­ surements a Appendix I).

Biology: Nothing is known aboxit the biology of this species; collection localities indicate its preference to mesic habitats.

The stomach of the two specimens from Jordan contained only exoskel­ etons of b. etles.

Systematic remarks: Though my specimens from Jordan are close in their measurements to the rest of the specimens from the Eastern

Mediterranean they differ drastically in coloration. Jordanian specimens unlike Palestinian and Lebanese specimens have a light colored pelagae and a definite bicolored tail. They could prove to represent a well defined population, however, we know little of tho amount of variation possible in this species. Order CHIROPTERA

The 27 species of bats reported from the Eastern Mediterranean

Region (Table l) represent seven families and constitute almost ?$% of the mammalian fauna. Following is a KEY to the seven families represented: ^Harrison (19£6c) provides a Key to the bats of the

Arabian Peninsula, but he includes only 20 of the species considered here_.7

1. CIS over LiO mmj FA greater than Q$ mm; Tail rudimentary (Suborder

Kegachiroptera)...... PTEROPIDAE

CIS less than 3I4 mmj FA less than 83 mm; Tail well developed (Sub­

order Microchiroptera)...... 2

2. A considerable amount of the tail free of the interfemoral mem­

brane (protruding over 20 mm)...... 3

Tail entirely or almost entirely enclosed within the interfemoral

membrane...... £

3. One upper premolar- ; snout with a rudimentary noseleaf; free

portion of tail at least three times the enclosed part......

...... RHINOPQMATIDAE

Two upper premolars; snout normal, without a rudimentary noseleaf;

free portion of tail equal or slightly longer than the enclosed

part...... ,U

I4. Three lower incisors; free terminal portion of tail emerges from

tho hind margin of the interfemoral membrane...... MOLOSSIDAE

83 89

Two lower incisors; free terminal portion of tail emerges upwards

from near the middle of the interfemoral membrane....,......

...... EMBALLOIIURIDAE

Face nor.rial, unadorned by noseleaves of any kind......

...... VESPERTILIOHIDAE

Face with noseleaves...... 6

6, Face with a deep central slit housing fleshy noseleaves; incisors

2/3...... NYCTERIDAE

Face without a deep central slit, membranous noseleaves present;

incisors 1/2...... RHINOLOPHIDAE

Khafash, Wat-Wat and Tayr-el-layl are three Arabic names used for bats with no particular reference to any one species.

Suborder Kegachiroptera

This suborder is more or less confined to the Ethiopian and

Oriental Regions except for a few species which extend farther north into the Palaearctic Region. One such species is Rousettus aegyp- tiacus which extends into the Mediterranean Region reaching its northern limit in southern Turkey (Kosswig, 19$$; Eisentraut, 19^9}

Kahmann & Caglar, I960; Osborn, 1963 and Lehmann, 1966a). This species belonging to the family Pteropidae is the only representa­ tive of this suborder reported from the Eastern Mediterranean

Region.

Family PTEROPIDAE Gray, 1821

Fruit-eating Bats

This is tho only family included in the suborder Kegachiroptera.

Koopman and Cockrum (196?) noted two main evolutionary trends within this family: a trend to shorten the jaws resulting in more powerful 90 jaws adapts l for chewing hard fruits and a trend to lengthen the jaws as an daptation for nectar-feeding. Rousettus is a typical example of ..he former trend.

Genus Rousettus Gray

1821. Rou.v:btus Gray, London Medical Repository,

Type species: Pteropus aegyptiacus E. Geoffrey

Dental formula: 2-1-3-2/2-1-3-3 53 3U

This genus includes eight species. Three species are reported from Africa (Hayman, 1967), of these only R. aegyptiacus is known to extend northwards into the Eastern Mediterranean Region.

Rousettus aegyptiacus (E. Geoffroy)

1810. Pteropus egyptiacus Geoffroy, Ann. Mus. II. N. Paris, l£:96.

Corrected to aegyptiacus in 1818, Description de'l'Egypte H.

N., 2:131:, pi. 3, fig. 2.

1951. Rousettus aegyptiacus Geoffrey} Ellerman and Morrison-Scott,

Checklist, p. 92.

Type locality: Great Pyramid, Giza, Egypt.

Common names: Egyptian Fruit Bat.

Distribution: Africa from the Capo Province northward to Senegal,

Egypt, Syria, Palestine, Jordan, Lebanon, Cyprus and Turkey. Also

Arabia, Iran, Pakistan and Western India if R. arabicus is to be con­ sidered as a subspecies of this form, as suggested by Eisentraut (19^9)•

Rousettus aegyptiacus aegyptiacus (E. Geoffrey)

Synonymy under species.

Type material: Mot examined.

General distribution: As for the species except for the four localities listed for R. a. arabicus. 91

This species has been reported as very corrmon in Palestine

(Tristram, 1881|j Bodenheimer, 1935 and 1958; Anon, 19i{6j Dor, 19h7j

Harrison, 1961(b) where it has been recorded from Dan, Mt. Carmel,

Wadi Kurn, Jericho, Jerusalem, Rehoboth, Me'arath Hateamin, Carmel

Caves, Herzliya Cave and near Hatruv. In Lebanon it has been reported by Eisentraut (1959), Lewis and Harrison (1962) and Harrison

(I961|b) from Beirut, Antelias, Amchite, Jamhour and Mt. Lebanon.

Specimens examined: 31. LEBANON: Mt. Lebanon (1 BM); Antelias (1 BM);

Beirut (1 SAJ 3 AUB); Ras Beirut (h SA); Mogharet Saleh, near Amchite

(15 SA) Roman Aqaduct, 2 km E. Hasmich (2 AUB); Cave, J4 km S. E. Beit

Meri (li AUB). PALESTINE: (l BZM).

Identification: This is the largest bat found along the Eastern Med­ iterranean. FA is over 89 mm in length and GLS over 1(0 mm, while all other bats are smaller with FA less than 83 mm and GLS less than 3h mm.

Morover, this is the only bat with a rudimentary tail.

Biology: Egyptian fruit bats were found to become active shortly after dusk. Leaving their roosts in moist caves, they swarm around fruit trees on which they feed. They have been reported to feed on date, fig, carob, sycamore, mulberry and azedarach trees (Anderson, 1902;

Bodenheimer, 1935; Lewis & Harrison, 1962; Harrison 1961(b). They normally consume only the fruits, however Lewis and Harrison (1962) recorded these bats feeding on leaves as well, especially of tho ornamental fig, Ficus religiosa L. I have observed these bats feed­ ing on a variety of fruit trees including apricot, peach and apple.

Those observations wore made at Antelias, Junieh, Tripoli and Saida in Lebanon 92

Tho a- undanco and feeding habits of these bats causes a tremendous (’.image to fruit trees grown for human consumption.

Farmers awai o of the bats1 activity have resorted to either placing a thick net ling around unripe fruits, customarily seen in date palm plantations, or seeking bat colonies and destroying then (Lewis and

Harrison, IS'62; Harrison l

Sound perception, temperature regulation and parturition has been studied in some detail by Kulzer (I960, 1963 & 1966). His studies confirm initial observations made in the field suggesting that these bats can rely solely on auditory orientation in complete darkness. He also confirms that they do not hibernate. When the temperature drops they tend to become inactive "but not torpid or lethargic,"

The number of young is usually one, but two are not uncommon. Four nursing females collected in mid-August, 1968, were found carry­ ing only one young each, while of six pregnancies checked three were found carrying twins. In Lebanon young are usually bom in the months of June through August. Anderson (1902) reported young bom in

February and March in Egypt, while Flower (1932) found that in the laboratory young were bom every month of the year, suggesting no fixed breeding season.

These bats do not appear to have long migrations. The reduc­ tion in the size of colonies in Lebanon during the winter has been attributed to dispersal (Lewis and Harrison, 1962). An altitudinal migration, however, can also be postulated to explain this sudden reduction in colony size. Egyptian fruit bats may thus leave their 93

cold cavern-, in the mountains and descend to the Coastal Plain or

the Rift Valley where warmth prevails most of the winter months.

Suborder Microchiroptera-

This luborder includes the bulk of the species representing

l£ living families. Many are insectivorous, but fish-feeding, nectar­

feeding, blood-sucking and other modes of feeding are also represented

in this suborder. Six families are reported from the Eastern Mediter­

ranean Region, all of which are primarily insectivorous.

Family RHINOPOM/VTIDAE Dobson, 1827

Mouse-tailed Bats

This primitive monotypic family ranges from Spanish Sahara east­ ward across north and central Africa to Egypt, Sudan and Ethiopia

into southern Asia from Arabia to Sumatra.

Genus Rhinopoma E. Geoffrey

1818. Rhinopoma Geoffrey, Descriptions do I’Egypte, 2:113.

Type species: Vespertilio jiicrophyllus Brunnich

Dental formula: 1-1-1-3/2-1-2-3 ".28

This genus includes two species, both of which are reported from the Eastern Mediterranean Region. These species have also been reported in sympatry in India /~Kha,]'uria, 1953 and Prakash, 1963a; only if R. kinneari is to be considered a subspecies of R. microphyllum as suggested by Aellen (195977* Afghanistan (Gaisler et al, 1968 and

Keyer-Oehmo, 1965), Iran (Lay, 196?), Arabian Peninsula (Harrison,

1961|b) and Egypt (Hoogstraal, 1962) usually inhabiting the same caves and sharing the same roost. 9)4

Most Id records of Rhinopoma from the Middle East were referred

to R. micro ’lyllum, a species rarely found by recent workers. This

probably re ulted from the failure of earlier workers to differentiate

between R. i; icrophyllum and R. hardwickei. Following is a KEY to these

two species:

FA usually more than 62 mmj tail always shorter than FA; CBL over 18.5

mm...... R. microphyllum

FA usually less than 60 mmj tail longer than FA; CBL less than 17.£ mm

...... «...R. hardwickei

Rhinopoma microphyllum (Brunnich)

1782. Vespertilio microphyllus Brunnich, Dyrenes Hist., 1:£0, pi. 6,

Figs. I-I4.

19^1 Rliinoporaa microphyllum Brunnich; Elleman and Morrison-Scott,

Checklist, p. 102.

Type locality: Egypt and Arabia.

Common names: Larger Mouse-tailed Bat or Rat-tailed Bat.

Distribution: Nigeria, Egypt, Sudan, Palestine, Lebanon, Iran,

Afghanistan and India.

All specimens from the Eastern Mediterranean Region are referred

to the nominate subspecies.

Rhinopoma microphyllum microphyllum (Brunn.)

Synonyray under species.

Type material: Not examined.

General distribution: Throughout the species range exclusive of India.

Tristram (1866a and I86I4) and Bodenheimer (1933> and 1958) report­ ed this species as common in Palestine. However, since this is not 95 tho case it is probable that their records were based on misidentif- ication of 1. hardwickei. Dor (I9h7) recorded it from owl pellets in Palest!'.', while Harrison (196I»b) reported specimens taken at Dan,

Jerusalem, .iinereth and Mi. Quarantania, near Jericho. Harrison

(1963a) noted the occurrence of this species in Lebanon on the basis of single specimen collected in a cave near Nahr El-Litani.

Specimens examined: 1. PALESTINE: Mt. Quarantania, near Jericho

(1 BM).

Identification: Tail shorter than forearm. Larger than R. hardwickei in all measurements except tail length.

Biology: Individuals of this species are reported Inhabiting dry open caves and rock crevices roosting together with the more common,

5* hardwickei. Gaisler et al (1968), however, reported that in

Afghanistan both species are equally common and though they were found together, frequently R. microphyllum prevails in one system of caves and R. hardwickei in another,

Yo'ong are born in June (Asdell, 1961:),

Rhinopoma hardwickei Gray

1831. Rhinopoma hardwickei Gray, Zool. Misc., p. 37.

Type locality: India,

Common names: Lesser Mouse-tailed Bat.

Distribution: North Africa, Palestine, Jordan, Iraq, Arabia, Iran,

Afghanistan, India, Kashmir, Burma and Thailand,

This species is represented in the Eastern Mediterranean by the following subspecies:

Rhinopoma hardwickei cystops Thomas 96

1903. Rhj or)om cystops Thomas, Ann. Mag. Nat.Hist., 11(7):1j96.

1913. Rh:i 'cporaa estops arablum Thomas, Ann. Mag. Nat.Hist.,

12(i)589» (Harrison, 1961

1951. Rhi ..opoma hardwickei cystops Thomas; Ellerman and Morrison-

Sco^ t. Checklist, p. 102. *

Holotypo: HM collection no, 2,1,17,2, an adult female obtained from

Luxor, Egypt ^

General distribution: North Africa, Arabia, Palestine, Jordan and

Iraq,

Harrison (1959a and 196hb) recorded this species from Tiberias,

Dan, Arbel, 30 km S, Ain Ck5di, Wadi Amud, Wadi Dalam, Ras en Nakura,

Bir Hindis, Wadi Zalam, Hanita, Ain Gedi and Mt, Quarantania (also

Thomas, 1913) in Palestine,

Tristram's and Bodenheimer*s records of R. microphyllum

probably represent this species.

Specimens examined: 23. JORDAN: Jerash (17 SA). PALESTINE: Ain

Fashkha (3 SA); Mt. Quarantania (3 BM).

Identification: This species is smaller than R. microphyllum.

CBL never exceeds 17.5 mm. FA usually smaller than tail, never

over 60 mm.

Biology: These bats are abundant among the ruins of Jerash with

colonies of varying sizes, 25-200 individuals. I have also observed

them along the western bank of tho Bead Sea where they are common

in open caves, (Qumran Caves), each cave with 5-15 individuals.

No other species of bat was found sharing the same cave or complex

of ruins with this species. 97

Collc< ';ion dates and observations in the field suggest that these bats do not hibernate, but remain active all year round.

I have observed an active colony near Qumran on three consecutive days during December and found that none of the bats left the cave during four to five hours of observation after dusk. This may i suggest that in this species a period of reduced activity replaces hibernation.

Nothing is known of the breeding habits of this species in the

Middle East, Prakash (I960) noted that in India young are born in June and July.

Family EMBALLONORIDAE Dobson, 1872

Sheath-tailed Bats

This family includes a dozen genera, primarily restricted to the New and Old World tropics, with only Taphozous represented along the Eastern Mediterranean.

Genus Taphozous E, Geoffroy l8l8, Taphozous Geoffroy, Description do 1» Egypte, 2:113.

Type species: Taphozous perforatus Geoffroy.

Dental formula: 1-1-2-3/2-1-2-3 a 30.

Taphozous nudiventris is tho only species reported from tho geographical area covered. This species has been designated as the type species for Liponycteris Thomas, 1922, given subgeneric rank by Simpson (19^2).

Taphozous nudiventris Cretzschmar

1826. Taphozous nudiventris Cretzschmar, in Ruppell’s Atlas zu der

Reise im nordlichen Afrika, Saugethiere, p. 70, fig. 27b. 98

Typo loca!t oy: Giza Pyramid, Egypt.

Coirmon n-rr. :j: Nakod-bellied Tomb Bat, Nakcd-rumpod Bat.

Distributi. a: Africa, from Senegal and Tanzania northward and

eastward t< Arabia, Palestine, Iraq, Iran and Afghanistan (Meyor-

Oehme, i960).

Taphozous nudiventris nudiventris Cretzschmar

Synonymy under species.

Type: Not examined.

General distribution: Egypt, Sudan and Palestine.

Tristram (1866a and 1881:) recorded ’’iramense numbers in ravines

in Galilee”. Allen (1915) reported on specimens from Jericho,

Harrison (I961jb) from north of Haifa, Lake Tiberias and Wadi Amud,

and Bodenheimer (1958) and Dor (l9l»7) from no exact locality in

Palestine. Specimens examined: 3. PALESTINE: Lake Tiberias (3 BM),

Identification: Free portion of tail emerges from near the middle

of the intorfcmoral membrane. Rump naked. FA 6)4-80 mm, GI5 26-29 mm.

Biology: These bats are highly colonial, inhabiting rock crevices in

arid and semi-arid areas. Harrison (196ijb). had reasons to bolievo

that in Iraq they migrate southward during winter months. This

sort of migration, however, would probably be unnecessary in Pales­

tine as these bats are restricted to the southern Rift Valley where winter is mild.

Family NYCTERIDAE Dobson, 18?5

Slit-faced Bats

This monotypic family-is primarily confined to the Ethiopian Region and. tho Malaysian Subregion (as outlined by Chasen, 19l|0). 99

Genu.*3 Nycteris Cuvier and Geoffroy

179$• Nyc .eris Cuvier and Geoffroy, Mag. Encyclop., 2:186.

Type speci s: Vespertilio hlspidus schreber

Dental for .ula: 2-1-1-3/3-1-2-3 ■ 32.

This .jenus includes a dozen species, eleven African (Hayman,

1967) and one Malaysian (Chasen, 19h0). All African species are restricted to Africa south of the Tropic of Cancer except for one species, N. thebaica, which extends northward along the western coast of Africa to Morocco and along the eastern coast to Egypt,

Arabia (western coast), Sinai, Palestine, and tho Island of Corfu.

Hycteris thebaica E. Geoffroy, 1818

1818. Hycteris thebaicus Geoffroy, Description de 1'Egypte, 2:119*

pi. 1, no. 2.

195>1. Hycteris thebaica Geoffroy; Ellerman and Morrison-Scott,

Checklist, p. 107.

Type locality: Egypt.

Common names: Egyptian Slit-faced Bat.

Distribution: Africa south of the Tropic of Cancer, Morocco, Egypt,

Arabia, Sinai, Palestine and the Island of Corfu, Greece.

Most nycterids are tree dwellers and solitary. However, this species is basically a semi-gregarious cave dweller (Rosevear, 196^).

Such habits make thebaica easier to procure than other nycterids, many of which are known only from few specimens. At least five subspecies are recognized, but only the nominate form is represen­ ted in the geographical area covered by this study. 100

Nyeterls thebaica thebaica E. Geoffroy

Synonymy in.der species.

Type material: Not examined.

General distribution: Egypt, Sinai, Palestine, northwestern Arabia and Corfu, Greece.

The only record of this species is from Beit Shean, Palestine

(Aharoni, 19l»h), Bodenheimer (19!?0) and Harrison (196lib) repeated this record.

Anderson (1902) reported this species as common in Sinai.

This, however, does not appear to be the case as only one specimen is available from this area in all museum collections examined.

Specimens examined: None from the geographical area covered.

Sinai: (l BM).

Identification: Face with a deep central slit. FA IjO-^O mm,

GLS 16-18 mm. Last caudal vertebra T-shaped.

Biology: In Egypt this bat is known tb frequent ruins in large numbers (Anderson, 1902). Nothing is known of its habits.

Family RHINOLOPIIIDAE Bell, 1836

Horseshoe Bats and Leaf-nosed Bats

This family is widely distributed throughout Africa, southern

Europe, southern Asia from Turkey and the Arabian Peninsula east­ ward to Japan and southward to Malaysia and into northeastern

Australia. Two subfamilies are recognized (Koopman and Cockrum,

1967), these are: 101

Rhinolophin s, represented In tho Eastern Mediterranean by five

species of r-hinolophus, and Hipposiderinae, represented by

Asellia trii'ens. The latter subfamily has been given full

family posiMon by many authors (Simpson, 19h$i Hayman, 1967;

Ellerman & Morrison-Scott, 1991 and others). Following is a

KEY to the two genera considered in this paper:

1. Premolars 2/3} superior border of noseleaf forming one vertical

projection.,...... Rhinolophus

2. Premolars 1/2; superior border of noseleaf bearing three prom­

inent vertical projections ...... Asellia

Genus Rhinolophus Lacepedo

1799. Rhinolophus Lacepede, Tabl. Hamm., p. 19.

Type species: Vespertilio forrum-equinum Schreber

Dental formula: 1-1-2-3/2-1-3-3 **32.

Following is a KEY to five species recorded from the area of - studyp

1. Large FA more than Sh mm, GLS more than 23 mm..R. ferrumequinum

Smaller, FA less than 92 mm, GLS less than 21.9 mm...2

2. Small, FA less than 37 mm, GLS less than 16 mm...,..,.,..,,,...

...... R. hipposideros

Medium-sized, FA more than bh mm, GLS more than 17 mm...... 3

3. Connecting process of sella pointed...... It

Connecting process of sella blunt...... R. clivosus

I:, Sella with parallel sides, connecting process of sella acutely

pointed projecting more than sella itself

euryale 102

Sella dth cuneate sides, connecting process of sella less

prominent than sella..*...... *...... R. blasii

Rhinolophus ferrumequinum (Schreber)

177U* Vesi'-ortillo ferrum-equinum Schreber, Saugeth., l:17ll> pi. 62.

1951. Rhinolophus ferrumequinum Schreberj Ellerman and Morrison-

Scott, Checklist, p. 111.

Type locality: France.

Common names: Greater Horseshoe Bat.

Distribution: Southwestern Europe, southern USSR, Japan, Korea,

China, Nepal, Kashmir, Afghanistan, Iran, Iraq, Turkey, Eastern

Mediterranean, Morocco and Algeria,

The relationship of Rhinolophus ferrumequinum to Rhinolophus clivosus is not clearly understood. R. clivosus is more or less restricted in its distribution to the African continent while R. ferrumequinum is primarily a Palaaarctic species. Thus in many ways these two species are complementary to each other in their distribution. Both species, however, have been reported from

Morocco, .Algeria and Palestine, but even here they have never been recorded in sympatry (in one cave). This has led

Harrison (1959b) to reduce clivosus to subspecific rank under

5* forrumequinum. However in a later paper Harrison (I96ljb) considered both species valid. 102a

Only t-j.0 nominato race is recorded from tho Eastern

Mediterranean*

Rhinolophus fcrrumaquinum ferrumequinum (Schreber)

Synonymy under species.

Type material: Hot examined.

General distribution: Southwestern Europe, Turkey, southwestern

Russia, Eastern Mediterranean, Algeria and Morocco.

This species has been reported as common in Palestine

(Tristram, l866annd I88I4 ; and Bodenheimer, 1920, 1935 and 1958),

Lebanon (Lewis and Harrison, 1962) and Jordan (Harrison, 1959a).

Harrison (1961tb) listed many localities from Palestine of which

the following localities: Tiberias, Haifa, Dan, Me’arath

Hateamim and Jerusalem are not listed in "specimens examined"

below.

Specimens examined: LEBANON: Tripoli (2 BM)j Kogharet Saleh, near Amchite (6 AUB; h SA)j Anjar (2 SA)j Roman Aquaduct, 2 km

E, Haznieh (22 AUB)j Dier Mar Maroun, J? km S. Hermel (1 AUB)j

Afka (1 REL). JORDAN: Jerash (U3 SA). 103

Identificai on: This is tho largest horseshoe bat found along

the Eastern I-'editerranean, FA 53-63 mm and GLS22.5-2)j,5 mm.

Biology: G eater horseshoe bats are common throughout the

Mediterrane m Region. They are found in large colonies during the

summer months roosting in moist dark caves and among ruins, es-

pecially vh m these lead to dark underground tunnels. The rest of

the year only a few individuals can be found hibernating in any

one cave. It Is obvious that the bulk of these bats migrate

during the winter. No one has as yet attempted to study migra­

tion routes, duration of hibernation or reproduction of this

species in the Middle East. Ransomo (1968) presented an excell­

ent study of this species in southern England,and Asdell (196U)

summarized all.reproductive data available for this species in

Europe, I found juveniles abundant during July and August which may suggest that young are born in May and June. Hibernating bats were collected in November, December and January.

Large colonies of R. ferrumequinum usually include a few

R. euryale sharing the same roost. Flight is graceful with quick wing strokes.

Rhinolophus clivosus Cretzschmar

1828. Rhinolophus clivosus Cretzschmar, in Ruppell, Atlas Reise

Nordl. Africa, Saugeth., p. Ii7.

Type locality: Kuwaileh, Saudi Arabia. (27° ^9* N, 35° 30* E)

Common names: Cretzschmar’s Horseshoe Bat.

Distribution: Africa except for tho forest belt from tho Congo westward, southwestern Arabia, Sinai and Palestine. Also south­

eastern Russia and northern Afghanistan if R. bocharicus is to

be considered a ^subspecies of this form as suggested by Aellen(1959)• Rhinolophus clivosus clivosus Cretzschmar

Synonymy a above

Typo mator al: Not examined.

General di- bribution: Northwestern Saudi Arabia, southen\ Palestine and Sinai.

Trist:, an (l866a) reported this species from near Lake Tiberias but later in his final work (Tristram, 1801;) did not treat the species as a member of the Palestinian fauna. Harrison (1961jb) recorded two specimens collected at Revivim and Abde in tho Negev, the only authentic record of the species from Palestine.

Identification: This is a medium sized bat slightly larger than

R. euryale and R. blasii, but smaller than R. ferrumequinum.

FA l;5>~f)0 mm and GLS 19.£t21.0 mm. Connecting process is blunt resembling R. ferrumequinum; pointed in both R. euryale and R. blasii.

Biology: Nothing is know; of the habits and habitat of this species in southern Palestine, Hoogstraal (1962) reported col­ lecting specimens in store-houses, stone huts and hillside, caves at El-Arish and Feiran Oasis in Sinai, Two subspecies R, c. acrotis and R. c. brachygnathus vrere reported either hanging singly or in numbers up to £0 in buildings (Sanborn and Hoogstraal,

1953 and Harrison 1961;b) or in large colonies in desert caves

(Hoogstraal, 1962).

Rhinolophus hipposideros (Bechstein)

1800, Vespertilio hipposideros Bechstein, in Pennant, Uebers,

Vierf , Thicre., 2:629. 105

Type locality: France.

Common name,;: Lesser Horseshoe Bat.

Distribution: Europe, southwestern USSR, Kashmir, Afghanistan,

Tran, Iraq, Turkey, Eastern Mediterranean Region, Sinai, Saudi

Arabia, Sudan, Eritrea and Morocco.

The following is the only subspecies represented in this region:

Rhinolophus hipposideros minimus Heuglin l86l« Rhinolophus minimus Heuglin, Nova Acta Leop. Carol. 29, 8:6,

1951* Rhinolophus hipposideros minimus Heuglin; Ellerman and

Morrison-Scott, Checklist, p. 116.

Typo material: Hot examined. From Keren Eritrea,

General Distribution: Portugal, Spain, France, Switzerland, Italy,

Turkey (Strinati, 1959), Lebanon, Palestine, Saudi Arabia (Morrison-

Scott, 1939), Sinai, Sudan, Eritrea and all large islands in the

Mediterranean Sea,

Bodenheimer (1958) reported this species from Herzlia, Beit

Arava and Sheikh Abreik in Palestine, Lewis and Harrison (1962) from Marjayoun and Beit ed Din iri Lebanon, and Harrison (19614b) from Machghara, Lebanon and Jerusalem, Dan, Beit Guvrin and Ain

Yahav in Palestine.

Specimens examined: 5. LEBANON: Beit ed Din (1 AUB); 1 km N. W.

Ain Anoub (2 AUB). PALESTINE: Jerusalem (2 BM). 106

Identifica’ .on: This is tho smallest horseshoe bat along the Eastern

Kediterran< n« FA less than 37 mm and GLS less than 16 mm, while all other ; .ecies have FA over Wi nim and GLS more than 17 mm.

Biology: r). ds species is rare throughout the Mediterranean

Region occu pying caves, deserted buildings and vmderground tunnels.

Not more t’um one or two specimens have been found at any one locality.

Asdell (l?6U) observed that usually a single young is bom in June or July. Young individuals reach sexual maturity at one year and mate in the fall.

Rhinolophus euryale Blasius

1853. Rhinolophus euryale Blasius, Arch. Naturgesch. 19, l:h9.

Type locality: Milan, Italy.

Common names: Mediterranean Horseshoe Bat,

Distribution: Portugal, Spain, Franco, Italy, Sardinia, Austria,

Yogoslavia, Greece, Turkey, Iran, Caucasus, Russian Turkestan, Iraq,

Syria, Lebanon, Palestine, Jordan Egypt, Algeria, Tunisia and

Morocco, (Almost circum-Mcditerranean).

The relationship of this species to R. meheyli is still uncertain. Sanborn and Hoogstraal (1955) and Hoogstraal (1962) refer all specimens from Egypt to R. meheyli instead of R, euryale as reported by Anderson (1902) and Flower (1932). Harrison (I961tb) examined a few specimens from Egypt but found them identical to

R. euryale judaicus from Palestine,

Only R. e. judaicus has been reported from the geographical area studied. 107

Rhi; »lophus euryale judaicus (Anderson and Matschie)

190b. Rurv 'lus judaicus Anderson and Matschie, S. B. Gesch,

Natv. -f. Fr. Berlin, p, 80.

1951. Rhir. ■loohus euryale judaicus Anderson and Matschie;

Elle ,vman and Morrison-Scott, Cliecklist, p, 120.

Holotype: BZM collection no. 12.978. A female collected at Cave of

Adullan, near Jerusalem, Palestine. I fruitlessly tried to locate this cave by asking officials and people around Jerusalem, How­ ever, recently I ran across a description of this cave (Tristram,

1881) which gives tho Arabic name for the cave as "Mogharet

Xhureitun". I am very familiar with this cave located about

If km southeast of Beit Sahur (31° bO* N and 35° l£' E) and about

10 km southeast of Jerusalem.

Genor-al distributions Syria, Lebanon, Palestine and Egypt (see discussion under species).

This species has been reported from Aleppo (Vettstein, 1913) and near Damascus (Trouessart and Kollmann, 1923) in Syria, Tiberias

(Tristram, 188b), Jerusalem and Herzlia (Bodenheimer, 1950 and

Harrison, 196bb) in Palestine and .Tyre (Tristram, 188b), Ras en

Nakura, near Araya and Roman Aqueduct, 10 km E. of Beirut (Harrison,

196bb) in Lebanon,

Other records.- Bodenheimer (1920 and 1935) and Lewis and

Harrison (1962) from Palestine and Lebanon respectively.

Specimens examined: 30. LEBANON: Cave, near Roman Aqueduct, 6 km E. Beirut (16 AUB); Mogharet Saleh, near Amchite (7AUB; 2 SA)* 108

PALESTINE: (1 FMNH)j Jerusalem (3 HM)j Ramleh (2 BM).

JORDAN: J..- ash (l SA).

Identifica! on: This species is slightly larger than R. blasii with which t may be easily confused. However R. euryale has longer eara (19~23 nm) and GI5 usually over 19»0 mm. Sella with parallel sj ,23 and connecting process sharply pointed.

Biology: Tils species is less common than R. ferrumequinum with which it ur. nlly roosts, Imiiiatures wera collected in August, also female.j collected at this time had well developed mammae which would suggest that young are born in the spring. Dr, Lewis

(bars. Comm,) collected a few hibernating bats in December and

January hanging solitarily in crevices at Mogharet Saleh, near

Amchite, Lebanon,

Rhinolophus blasii Peters

10^7. Rhinolophus clivosus Blasius, (Nec Cretzschmar), Saugeth,

Deutschlands, p, 33. •

1866, Rhinolophus blasii Peters, Mbcr. Preuss, Akad, Wiss,, p» 17*

New name for R, clivosus Blasius not R. clivosus Cretzschmar,

Type locality: Italy,

Common names: Peters' Horseshoe Bat.

Distribution: Italy, Greece, Turkey (Zimmemann, 1953), southwestern

USSR, Cyprus, Crete, Iran, Afghanistan, Syria, Palestine, southwestern

Arabia, Transvaal, Rhodesia, Malawi, Congo, Eritrea and Morocco.

Only tho nominate form is recorded from the geographical area covered. 109 Rhinolophus blasii blasii Peters

Synonymy as above.

Type material; Not examined.

General distribution; As for the species excluding all African

localities except Morocco.

Tristram (188U) reported this species as "abundant about

Jerusalem, Bethlehem and Hebron" in Palestine. Bodenheimer (1958)

reported it from Herzlia and Sheikh Abreik and Harrison (196Iib)

added Jenin and 2 mi S. of Jenin in Palestine. Wettstein (1913)

cited the first record of this species in Syria at Aleppo. Boden­ heimer (1920 and 1935) listed this species as a member of the

Palestinian fauna but did not cite any particular locality.

Dor (I9h7) recorded one specimen from pellets of Tyto alba

(Scopoli) in Palestine.

Specimens examined; l6, PALESTINE; (l FMNH)j Jenin (5 BM); Gave of

Adullam, 10 km southeast of Jerusalem ( 1 SA} 3 BM)} Jerusalem

(U BM)} Near Jerusalem (2 AMNH).

Identification; This is a medium-sized horseshoe bat, FA 145-U7.5 mm

and GIS19.0-19.5 mm. Sella cuneate or wedge-shaped. Connecting

process pointed. No marked contrast between crown areas of anterior

and posterior lower molars.

Biology; Nothing is known of the biology of this species. At the

Cave of Adullam I found this species hanging solitary or in couples

along with Plecotus austriacus. no

Gonus Asellia Gray

1838» Asel ia Gray, Mag. Zool. Bofc., 2ih93*

Typa Specie ': Rhinolophus. trldena E, Geoffroy,

Dental fom-la: 1-1-1-3/2-1-2-3 - 28,

Harris-n (I96l>) recognized Asellia patrizii do Beaux,

1931 described from Ethiopia, however, Hayman (196?) reduced it to a suhspecific status under A. tridens,tho only species included in this genus.

Asellia tridens (E. Geoffroy, 1813)

1813. Rhinolophus tridens Geoffroy, Ann, Mus. H. N. Parris, 20:26£,

1951, Asellia tridens Geoffroy; Ellerman and Morrison-Scott,

Checklist, p. 130.

Type locality: Egypt,

Common names: Trident Leaf-nosed Bat,

Systematic remarks: Two subspecies are reported from the Middle

East. The nominate from Gambia, northeastern Africa and Sinai

(Hayman, 196?)j and i. t. murraiana (J. Anderson, 1881) from India,

Pakistan, Afghanistan, Iran and Iraq (Aellen, 1959} Hatt, 1959}

Siddiqi, 1961} Harrison, 196Ub; and Lay, 1967), A. t, murralana from Iraq and Central is much greyer in coloration and slightly larger in size than the nominate form, FA U9-58 mm in murraiana and ltb-53 mm in tho nominate. The Eastern Mediterranean Region falls on the fringe of the distribution of both of these sub­ species, Harrison (1957 and 196ljb) after studying many examples of both of these subspecies concluded that "the two races intermediate in Palestine and N. V/. Arabia". Atallah and Harrison Ill

(1967) note 1 tho first record of this species in Syria and indicated that their ••pocimons show intergradation between the two subspecies mentioned above. The Eastern Mediterranean populations, however, show a greater similarity to tho nominate subspecies and will be treated here as belonging to it,

Asellia tridens tridens (E. Geoffroy)

Synonymy under species

Type material: Not examined.

General Distribution: Syria, Palestine, Sinai and Northern Arabia.

Tristram (188U), Aharoni (1930) and Bodenheimer (1935 and 1958) recorded this species from the Dead Sea Basin. Bodenheimer (1958)and

Harrison (19614b) added the following localities in Palestine: Jerusalem,

Hezme, Jaffa, Tel Aviv and Nahal Amram, 10 km N. of Eilat,

Other records,- Bodenheimer (1920) and Harrison (1957).

Specimens examined: I4. SYRIA: Palmyra (1 HARR; 2 AUB),

PALESTINE: near Jerusalem(l BM), Identification: Noseleaf bears three vertical projections.

FA I4I1-5U ran. Only one upper premolar (two in Rhinolophus).

Biology: This species is more or less restricted to the Desert

Region, It is highly colonial occupying caves, ruins and under­ ground water tunnels, Harrison (1956b and 19614b) and Lay (1967) maintained that these bats leave their summer retreats during winter months but return to the same grounds - every summerv Many— of the caves occupied by these bats during the summer are reported ideal for hibernation (Harrison, 1956b).

Weber (1955) reported on the high temperature tolerance of this species. No data is available on the reproductive biology of this species in the Middle East. 112

Family VESP2RTILI0NIDAE GRAY, 1021

Vespertilionid Bats

This .1 daily attains a world vido distribution except for

some distnnt islands in the south Pacific and tho polar ico caps.

Six subfami lies are recognized, but only two of these are re­

presented 3n the Eastern Mediterranean Region; the nominate sub­

family represented by six genera and the monotypic subfamily

MLniopterirae by the nominato genus. Following is a.KEY to these

seven genera:

1. Second phalanx of third finger about three times as long as the

first phalanx. (Subfamily Miniopterinae)...... Miniopterus

Second phalanx of third finger less than three times as long as

tho first phalanx, usually equal or subequal in length to it.

(Subfamily Vespertilioninae)...... 2

2. Upper jaw with-three premolars j tragus- long and sharply point­

ed, ...... tlyotiso

Upper jaw with one or two premolarsj tragus usually short and

rounded at tip, but if long and pointed then ears are over 35

mm in length...... 3

3. Two upper incisors on each side...... lj

One upper incisor on each side...... Otonycteris

U. Ears very large exceeding 3h mm in length, joined across the

forehead...... Plecotus

Ears not very large and widely separated from each other....,5

5. Medium sized bats, FA 1j2-56 ...... 6

•anall sized bats, FA 30-38 mm...... ,,,.,...... Pipistrellus 113 6, Four cl ockteeth behind the canine in each upper jaw; tragus

rounded distally not expanded into a club-shaped extremity..*..

...... Eptesicus

Five cheekteeth behind the canine in each upper jaw; tragus ex­

panded distally into a club-shaped extremity...... Nyctaius $ “ I- -ir* - ~ r a

Genus Myotis Kaup

1829. Myotis Kaup, Skizz. Europ. Thierwelt, 1:106.

Type species: Vespertilio myotis Eorkhausen,

Dental formula: 2-1-3-3/3-1-3-3 0 38.

This is the most widely spread genus of bats achieving a dis­

tribution similar to that of the family. It is represented along

the Eastern Mediterranean by five European species. All of these

species are restricted to the Mediterranean Region reaching their

southernmost limit of distribution in Palestine.

Tristram (1881:) erroneously reported II. daubentoni and M. mystacinus from the Eastern Mediterranean Region. Vespertilio murinus reported by Tristram (188U) and Bodenheimer (1920 and 1935)

refer to Myotis sp.(vide Bodenheimer, 1958).

Following is a KEY to the five species represented in the

geographical area covered:

1. Large, FA more than 55 mm, GISovor 21 mm...... 2

Small, FA less than U5 mm, GISless than 17 mm...... 3

2. GLS over 2I4 mmj FA usually more than 61: mm...... «U. myotis

GLS less than 23.5j FA less than 63.5...... M. blythi

3. Outer edge of leg (tibia) hairy; ear less than ll: mm

...... »M. capaccinii

Outer edge of .leg not hairy; ear more than ll: mm.1; llli

In Interf ioral membrano with a dense fringe of hair; BB more than

7.3 iitti, outer border of ear not notched...... M. nattereri

Interf floral membrane with only scattered hairs along the edge;

BB les.i than 7.3; outer border of ear notched....H. emarginatus

Kyotis myotis (Borkhausen)

1797. Vcsj.arbilio myotis Borkhausen, Deutsche Fauna, 1:80.

Typo locality: Thuringia, Germany.

Common names: Gx*eater Mouse-eared Bat.

Distribution: Southern Europe, Turkey, Eastern Mediterranean Region and southeastern Russia,

The conflicting views regarding the relationship of this species to M. blythi and the subspecific status of Eastern Mediterranean

M. myotis are adequately reviewed by Harrison and Lewis (1961).

Harrison and Lewis (loc. cit.) described M. m. macrocephalicus from Lebanon and referred all Middle Eastern populations to it.

Myotis myotis macrocephalicus Harrison and Lewis

1961, Myotis myotis macrocephalicus Hariris on and Lewis, J. Mamma}..,

1»2(3):373.

Holotypo: REL collection no. V-6oli8 deposited at the BM. An adult male collected on August 3ii, i960 at a cave (Mogharet Saleh) 2 km E. of Amchite, Lebanon.

General distribution: Lebanon, Palestine, Syria and Turkey,

A paratypo is represented by one specimen from Toll Kalakh, Syria, the only authentic record from this country. Lewis and Harrison (1962) reported on specimens from a cave near Halba, 2km E, of Amchite and the ruins of Baalbek in Lebanon. Atallah (1969b) recorded specimens taken at a cave near Anjar (8^0 m), Lebanon. Theodore and Moscona (195k) 115

listed tho species as one of the hosts examined for parasites in

Palestine, but do not list collecting sites. For additional

records see discussion under the genus heading.

Specimens examined: 76. LEBANON: Mogharet Saleh, 2 km E.Amchite

(8 SA; 30 AUBj h BM)j Cave near Anjar (2h SA); Cave near Halba

(7 AUB); Ruins of Baalbek (2 AUB). SYRIA: Tell kalakh (l BM).

Identification: This is the largest mouse-eared bat in the

region, FA 58-72 mm, GIS2U,5-27.0 mm. It differs from the nominal

race by its paler coloration and larger external and cranial

measurements (see Harrison and Lewis, 1961: Tables 1 & 2),

Biology: This is a gregarious cave duelling species. At Mogharet

Saleh these bats were found to roost along with M. blythi in deep

inaccessible fissures at the terminus of the cave, while at Anjar

225 bats roosted in a shallow (75 cm high) 2-ij m dark extension

along the side of the cave. At the latter locality these bats were found to emerge shortly after dark in great numbers. Some

circled over the cave but most of the bats headed toward the spring,

50-100 m from the cave, for a drink. Bats entering the cave were

found to hang singly or in small numbers in small cracks on the walls

of the cave. Close observation of those bats showed that they

retreat to these cracks with their prey, usually moths, and feed

on them while they hang.

Theodor and Moscona (195^) maintained that a definite antagon­

ism exists between M. myotis and Rhinolophus spp., as all

Rhinolophus spp. "were observed to leave a cave in May when Myotis myotis began to appear". This does not appear to be tho case in 116

Lebanon as in all caves visited both Myotis myotis and at least two species of Rhinolophus were collected,

Asdell (1961i) maintained that mating takes place usually in the fall and again in the spring, A single young is born in June, attaining sexual maturity in one year.

Greater mouse-eared bats were found to leave their roosts by mid-October and return by the end of March, No data are available on wintering areas of this species. In a few instances one or two individuals were found to remain through the winter in the summer range (Lewis and Harrison, 1962),

Myotis blythi (Tomes)

1857« Vespertilio blythii Tomes, P,Z.S,, p, 53

Type locality: Nasriabad, Rajputana, India,

Common names: Lesser Mouse-eared Bat,

Distribution: Southern Europe (along the Mediterranean Sea),

Morocco, Algeria, Tunisia, Turkey, Eastern Mediterranean Region,

Iraq, Iran, southern Russia, Afghanistan, (Zimmemann, 1956),

Kashmir, India and eastern China,

Eastern Mediterranean populations have been referred to tho following ^subspecies (Harrison and Lewis, 1961):

Myotis blythi omari Thomas, 1 6

1906. Myotis myotis omari Thomas, Proc, 'Zool, Soc, Lohd.,2:521,

1961, Myotis blythi omari Thomas; Harrison and Lewis, J, Mamm.,

ij2(3):378.

Holotype:'- BM collection no, 5.10.1nlh. collected at Derbent, 50 miles west of. Isfahan, 6,500 ft, Iran, 117 General dis! ribution: Iran, Iraq, Syria, Palestine, Lebanon and

Turkey,

Harrison and Lewis (1961) reported on specimens from Tell

Kalakh, Syria and 2 km E.Amchite, Lebanon, while Lewis and Harrison

(1962) recorded specimens from the Natural Bridge, 7 km S. E. Faraya, f Lebanon, Hr i’rison (l96hb) noted the occurrence of this species in

Palestine, uo exact locality.

Other -records.- Atallah (1969b) and Theodor and Moscona (19510*

Specimens examined: 30, LEBANON: Mogharet Saleh, 2 km E.Amchite

(2U SA; 6 AU3).

Identification: This species is slightly smaller than the precoeding species with which it is sympatric throughout its range. Fa less than

36,5 mm and GISless than 23,5 mm. Also ears smaller than M. nyotis, over 27 mm in the latter and less than 25 mm in M, blythi. For comparison with M. blythi oxygnathus, the southern European and

North African race, see Harrison and Lewis (1961).

Biology: This species resembles M. myotls in its habits and habitats.

Both species roost together, but in any one cave one species predom­ inates .

Nyotis emarginatus (E, Geoffrey)

I806, Vespertllio emarginatus Geoffroy, Ann. Hus. N. H. Paris, 8:198,

1951. Kyotis emarginatus Geoffroyj Ellorman and Morrison-Scott,

Checklist, p. lltl.

Type locality: Charlemont, Givet, Ardennes, Franco.

Common names: Geoffrey’s Bat, Notch-eared Bat.

Distribution: Holland, France, Germany, Switzerland, Italy, Hungary,

Greece, Bulgaria, Poland, southern USSR, Afghanistan, Iran, Palestine, 118

Lebanon, Morocco and Algeria.

Only the nominate form is pertinent to this study,

Nyotis emarginatus emarginatus (E, Geoffroy)

Synonymy under species.

Type material: Not examined.

General distribution: European and North African range of the species

and Palestine,

Tristram (1881|) reported on specimens from Tyro, Lebanon, and

Mt. Carmel, Palestine, Bodenheimer (1920, 1935 and 1958) listed the

species as a member of the Palestinian fauna based on Tristram's records and a specimen obtained by M, Dor, Lewis and Harrison

(1962) reported on a specimen taken near Amchite, but Atallah (1969b) found this specimen to represent M. capaccinii instead, Harrison

(196Ijb) recorded this species from Nahal Oren, Mt. Carmel, and

Jebal in Palestine.

Specimens examined: 2h» PALESTINE. Jebal (2 BM)j Mt, Carmel

(22 HARR).

Identification: This is a small myotine bat characterized by a

conspicuous notch on the posterior border of the oar. BB less

than 7.3 mm and CBL II4,0-15.5 mm. Outer edge of tibia not hairy as in M. capaccinii and interfemoral membrane without a dense fringe of hair as in M. nattereri.

Biology: Harrison (l96Ub) records a maternity colony of more than two hundred individuals at Mt, Carmel in April, 1962. A single embryo was found in each of 21 females examined. Young are born in June and become sexually mature in their second summer

(Asdell, 196h). 119

I-Iyotis capaccinii (Bonaparte)

1837• Vost> .'.•'tillo capaccinj.! Bonaparte, Faun, Ital., I, fuse, 20,

Type locality: Sicily, Italy,

Common nam'' : Long-fingered Bat,

Distributic, : Southern Europe, southwestern USSR, Iran, Turkey

Lebanon, Poostine, Morocco and Algeria,

Atallah (196%) and Hair is on (lS?6^b) referred their specimens from Lebanon and Palestine to M, c, bureschi described from Bulgaria,

Hyotis capaclnii bureschi (Heinrich)

1936, Leuconoe capaclnii bureschi Heinrich, Mitt, Naturu, Inst,

Sofia, 9:38.

19!?1. Hyotis capaclnii bureschi Heinrich; Ellorman and Horrison-

Scott, Checklist, p. ll»3,

Holot;/pe: BZM no, 312 an adult male collected at Karamlcr, Strandja-

Balkan, (800 feet), Bulgaria, on July 2, 193!?,

General distribution: Bulgaria, Turkey, Lebanon and Palestine,

Bodenheimer (19!?8) reported the species from Palestine based on few specimens obtained by M, Bor, no exact locality or date,

Harrison (196ljb) found large colonies at the Carmel Caves and near

Tiberias representing the first authentic record of the species,

Atallah (1969b) obtained a few specimens at Mogharet Saleh, near

Amchite, Lebanon.

Spocimens examined: 1J?. LEBANON: Mogharet Saleh, near Amchite

(h SA; 1 AU3). PALESTINE: Carmel Caves, S. of Haifa (10 HARR).

Identification: Outer edge of leg densely covered with hair.

Ear, smaller than in H. nattereri and M. emarginatus, less than

Ik mm in length. Feet largo, more than half the length of the 120 tibia.

Biology: I! (.'risen (196hb) found these bats in large maternity

colonies at the Carmel Caves about mid-April. All females

were progn? t, a single embryo each, irith embiyos fairly well

developed. Atallah (I96?b) found four specimens occupying a deep

crevice ale g the entrance to Mogharet Saleh, Lebanon, roosting

together vi h M, nattereri. Two males examined had well developed

testes sugg •sting a fall mating season. The glans penis and baculum were, illustrated by Atallah (1969b, Fig. 1-c and Fig.v2)V'.

Hyotis nattereri (Kuhl)

1818. Vespertllio nattereri Kuhl, Ann. Wetterau Ges, Naturk,,

h, Ij33,

Type locality: Hanau, Hessen, Germany.

Common names: Natterer’s Bat.

Distribution: Europe, southern USSR, Korea, Japan, China, Iran

(Harrison, 1963), Turkey, Lebanon, Palestine and Morocco.

Harrison (I96ha) described the x'acc M. n. hoveli from Palestine.

Atallah (1969b) referred Lebanese specimens to this subspecies but maintained that '‘further collections in Lebanon may shot/

intennodiation between hoveli in the south and the typical race,

12* H* nattereri, in the north."

Myotls nattereri hoveli Harrison

196!i. Kyotis nattereri hoveli Harrison, Z. Saugetierk., 29(1):f>8.

Holotype: HARR collection no. 11.3393, an adult femalo obtained on

April 30, 1961, at Aqua Bella, near Jerusalem, Palestine,

General distribution: i'alestine and Lebanon. 121

Bodcnh.imor (1953) maintained that this species is quite common in Galilee, Horzlia and Aqua Bella referring his specimens to the nominate fo'.t. Harrison (l96Ub) referred specimens from Hatruv,

Mt, Carmel nd Herzlia in Palestine to hoveli, Atallah (1969b) reported on the first record of this species from Lebanon, collected i at Mogharet Saleh, near Amchite,

Specimens econined; lit, LEBANON: Mogharet Saleh, near Amchite

(It SA), PALESTINE: Aqua Bella, near Jerusalem (10 HARR),

Identification: Ears average longer than H, ccpaccinii and

M, emarginatus, 15-18.5 mm, Interfemoral membrane with a dense fringe of hair between the calcar and the tail, Braincase over

7,3 mm in width.

Biology: Harrison (1961tb) found a maternity colony at Aqua Bella on April 30, 1961. Pregnant females carried only one young.

Asdell (196U) reports that young are born late in Juno or early

July in Europe. Judging from the size of embryos examined by

Hairrison (loc. cit,) in April young would bo bom in late Hay or early June in Palestine,

Three females and one male were found occupying a deep crevice along the entrance to Mogharet Saleh, near Amchite, Lebanon, roosting with M. capaccinii. They started leaving the roost at

7:25 p.m. The baculum of this form has been illustrated by

Atallah (1969b, Fig. 1-d).

Genus Plpistrollus Kaup

1029. Plpistrollus Kaup, Skizz. Europ. Thierw., 1:98.

Type species: Vespertilio pipistrellus Schrebcr.

Eental formula: 2-l~2~3/3-l-2~3 13 3h. Occasionally in P, savii the small upper premolar is missing. 122

Four .v^ocles are recorded from the Eastern Mediterranean.

Three of th ;se are typically European species reaching their southernmovv; limit of distribution in Palestine, ■while tin fourth, P. . odanheimeri, is an endemic Arabian desert species.

Following i a KEY for identifying these four species of Eastern « Mediterranean pipistrelles.

1. First upper incisor unicuspidj ving membrane with a well defined

Tfhito border..P. kuhl-H

First upper incisor bicuspid} wing membrane dark throughout..,,,

...... 2

2. First upper premolar displaced internally, not visible in later­

al view} color of dorsal surface pale buffy brotm, belly whitish

...... 3

First upper premolar slightly displaced, easily visible in a

lateral view} color of dorsal surface dark brown, belly light

brown...... ,P, pipistrellus

3. GLS'more than 13.0 mm} FA more than 31.£ mm....,...... P. savii

CIS loss than 12.0 mm} FA less than 31.5 mm......

...... P. bodenhetmeri

Pipistrellus pipistrellus (Schrebcr)

177U. Vespertilio pipistrellus Schrebcr, Saugeth,, I, pi.

(text p. 167).

195l• Pipistrellus pipistrellus Schrebcr} Elloman and Morrison-

Scott, Checklist, p. I6I1.

Type locality: Franco,

Common names: Common pipistrelle. 123

Distributie i- Europe, southueatom USSR northwards to Moscow,

Korea, Jap-! ■, Taiwan, Kashmir, Afghanistan, Iran, Turkey, Lebanon and Morocco

Only t ) nominate form is pertinent to this study,

Plpisi nllus pj.plstrollus pipistrellus (Schreber)

Synonymy ur 'er species.

Type mateid. 1: Not examined.

General die'ribution: Europe, Turkey, Iran and Lebanon,

Lewis nd Harrison (1962) recorded two specimens from

Lebanon (exact localities listed below), the first from the

Arabian Peninsula,

Soecimens examined: 2, LEBANON: Machghara (1 AUB)j Ammik,

Bekaa Volley (l BM),

Identification: This species resembles P. bodenheimeri in its small dimensions, FA 27-31 mm and GLS 11-12,5 mm. However, they differ in coloration, P, pipistrellus with a dark black dorsum and P, bodenheimeri irith a pale buffy dorsum. Also, in the for­ mer species the first upper premolar is but little displaced medially, while it is displaced so completely in the latter to bo not visible in a side view. First upper incisor bicuspid.

Biology: Nothing is known of the habits and habitats of this species in the Eastern Mediterranean Region due to its scarcity,

Asdell (19614) summarized all studies dealing with reproductive biology of this species in Europe, These studies show that a single young is usually born around July after a gestation period of lih days. Young males and females reach maturity when two years old and mate in the fall. 12U Pipistrellus Icuhlii (Kxihl)

1019. Vesp rtilio leutilii Kuhl, Ann. VJetterau. Ges. Naturk., !j, 2:199.

Typo locnli y: Trieste, Italian-Yugoslavian border.

Common name, : Kuhl*a pipistrelle.

Distributio;Circtun-Hcditerrancan northwaixis into Germany and eastwards to southwestern USSR, Afghanistan, Kashmir, Pakistan,

Iran and southwards to southern Arabia and the Cape Province in

Africa,

Lewis and Harrison (1962) showed that intergradation between

P. k. ikhwanius Cheesman and Hinton and P. k. kuhlii takes place in Lebanon, but they maintained that on the ibole Lebanese specimens "are decidedly paler and less rufous brown than the typical race”. They also maintained that though young specimens of P, k. ikhwanius are appreciably darker than the adults, these are still paler than the typical race. Harrison (196Ub) followed this for Palestine, Lebanon and Cyprus, referring all populations from the Arabian Peninsula to P. k. ikhwanius.

Pipistrellus kuhlii ikhauanius Cheesman and Hinton

192I4. Pipistrellus kuhli ikhwanius Cheesman and Hinton, Ann. Hag,

Hat. Hist., Ib(9):$k9.

Holotype: BM collection no. 2li.8.2.1., an adult male collected at

Hufuf, Hasa, Saudi Arabia on December 7, 1923.

General distribution: Saudi Arabia, Yemen (Sanborn and Hoogstraal,

1953)> Oman, Kuwait, Bahrain, Iraq, Iron, Turkey, Syria, Lebanon,

Jordan, Palestine and Sinai,

Tristram (1866a and 1881:), Bodenheimor (1920, 1931? and 195>0), 12$ Dor (I9li7) .nd Harrison (19^Ub) recorded this species from tho following I'.calities in Palestine: Ayelet Hashkhar, Dafne,

Don, Rehbot>i, Mt, Carmel, near Haifa, Wadi Tira, near Tivon, Hulata,

Wadi Anud, icar Eeorsheba, Arbel, Beit Cren, Kishon, Kefar Ruppin,

Lake Tiberias, Plain of Acre, Kefar Monachem, Tel Aviv, Tantuza,

Bethlehem and Cave of Adullam, Trouessart and Kollnan (1923) and Harrison (l96Ub) reported on specimens from Djeroud and Damascus,

Syria, Harrison (196Iib) and Atallah (1966 and 196?a) recorded the species from Sanour and the Azraq Basin in Jordan, Allen

(191$), Stencel (1961), Lewis and Harrison (1962) and Harrison

(19614b) reported on spocimens from Chtura, Beirut, Halba, Ehden,

Baalbek, Amchite, Zahle, Barja, Saida, Machghara and Tyre in

Lebanon,

Specimens examined: 36 JORDAN: Azraq ed Druz (3 SA)$

Azraq-Shishan (3 SA). PALESTINEj Beit-Sahur (6 SA)j Tiberias

(1 BZM). LEBANON: Beirut (20 AUB; 3 SA)j Ajaltoun (1 SA).

SYRIA: Damascus (1 BM),

Identification: Wing membrane with a r.’ell defined white border.

First upper incisor unicuspid, bicuspid in all three other

Eastern Mediterranean species. Size larger than P. pipistrellus and P. bodenheimeri, FA 32-37 mm and GIB 12,5-lii.O mm.

Biology: Natural history of this species .in Lebanon has been studied in some detail by Stencel (1961), This species was found common in Lebanon roosting in large colonies in buildings around cities and villages along the coast up to an altitude of 1^00 m. Blanford (1888-1091) maintained that two young are usually born at a time. Harrison (I961jb) noted this in a specimen collected in April at Dan, Palestine, Kales collected 126 in tho fall had enlarged testes suggesting a fall mating season.

Young arc b .-u during late April and Hay (Lewis and Harris on, 1962),

Kuhl1s bats vere found to leave their roosts at dusk and fly straight and high to their feeding area, where they tend to fly low nd erratically. They are active all year round and can bo seen feeding on mild winter days. A reduced activity period seems to replace hibernation.

Pipistrellus savii (Bonaparte)

1837. Vespertilio savii Bonaparte, Faun. Ital., Itfasc. 20.

Type locality: Pisa, Italy.

Common names: Savi’s Pipistrelle,

Distribution: Southern Europe, southwestern USSR eastwards to

Siberia, Hongolia, Turkey, Lebanon, Algeria and Morocco.

Harrison (1961) referred the three known examples of this species fren tho Eastern Mediterranean Region to the race P,

£. caucasicus.

Pipistrellus savii caucasicus (Satunin)

1901, Vesperugo (Vesperus) caucasicus Satunin, Zool. Anz., 2)j:Ij62.

19!?1. Pipistrellus savii caucasicus Satunin j Ellerman and Morrison-

Scott, Checklist, p. 170

Type material: Not examined, collected at Tiflis, Caucasus,

General distribution: Caucasus, Crimea and Turkestan in Russia,

Turkey and Lebanon. £Known only from Ainab, Lebanon (Harrison, 1961)7.

Specimens examined: 3. LEBANON: Ainab (3 BM).

Identification: A medium sized pipistrelle, Fa 31.£-33.£ mm and

GL$13.0-l)i,0 mm. Color of back pale brown, belly whitish. First 127 upper promo'! displaced internally from tho tooth row or com­ pletely absc it.

Biology: Collection sites indicate that this species is a high altitn, a form. Nothing is knotm of its habits in tho

Middle East *

Finistrollus bodenheimeri Harrison

I960. Pipl- trollus bodenheimeri Harrisonj Durban. Kus. Novit.,

S, Ft. 19, p. 261.

Holotype: ILIRH collection no. 1.2907, an adult male obtained at

Yotvata, Uadi Araba, I4O km N, of Eilat, Palestine on October 13, 195>9.

Common names: Bodeniieimer*s pipistrollo.

General distribution: Aden and Palestine.

The only known examples of this monotypic species from tho

Eastern Mediterranean Region vere obtained at tho typo locality

(Harrison, I960 and 196)4b).

Specimens examined: Only the type.

Identification: This is the smallest Middle Eastern pipistrelle,

FA 28-32 mm and GLS 10.5-12.0 mm. Color very pale buffy on the back, the belly whitish. First upper incisor bicuspid, first upper premolar displaced completely from tooth row.

Biology: Harrison (196ljb) believed this to be an endemic desert species. A femalo collected in April was found pregnant. Harrison

(op. cit.) noted this species at Yotvata, Palestine, flying at dusk low over the ground around tamarisk and eucalyptus trees. 128 Conus Nyctalus Bovrdich

1825. I?yc: lus Bowdich, Excursions in Madeira & Porto Santo,

p. 3 >.

Type specie : Nyctalus verrucosus Bowdich

Dental for/', la: 2-1-2-3/3-1-2-3 - 3U. t This j ‘.laoarctic genus reaches its southernmost limit of distribution along tho Eastern Mediterranean trhere it is rep- icsented by N. noctula lebanoticus, an endemic subspecies.

The genus has been considered congeneric with Pipistrellus by

Simpson (19b5>), but all European workers prefer to give this genus full generic status. However, Nyctalus would take priority over Pipistrellus if the two are congeneric,

Nyctalus noctula (Schreber)

177b. Vespertilio noctula Schreber, Saugeth,, I: pi, $2. text p. 166,

Type locality: France,

Common names: Common Noctula,

Distribution: Europe, southern USSR, Japan, China, Nepal,

Kashmir, Iran, Turkey, Lebanon and Palestine, A single specimen has been reported from Morocco (Laurent, 1937)# the only north

African record.

Only tho following subspecies has been recorded from tho

Eastern Mediterranean Region,

Nyctalus noctula lebanoticus Harrison

1962, Nyctalus noctula lebanoticus Harrison, Prdo,sZdol,Soci'Lend;,

139(2):337, pi. 1, 129

Fig. 6. Head of Nycatalus noctula lebanoticus,(HARR. 26.33^0),

collected at Natural Bridge, Faraya, Lebanon.

Holotype: BM collection no. 6l.lj06, an adult male obtained by R. E.

Lewis on July 29, i960, at Natural Bridge, Faraya, Lebanon. Three paratypes, two males and one female, collected at the same place and time.

General distribution: Lebanon and Palestine (vide Bodenheimer,

1958). 130

LetriLs and Harrison (1962) reported on tho first record of tho

species fr Lebanon J their material was later described in detail by Harr iso. (3.962). Bodenheimer (1958) cited a record by Festa

from Joric! e, Palestine, but sinco no spocimens are available

from this t >untry, he considered the report as very doubtful, * Specimens t.:cainined: Ij. Type series.

Identifications This is a medium size bat, FA $2-$6 mn and GIf> 18.U-19.7

mm. Ears ilioi’t, 3ii-19.mm_, and tragus widely expanded distally into

a club-shaped extremity (Fig.6).

Biology: The four specimens from Lebanon were obtained by shooting

into a fissure Tinder the natural Bridge (1600 m), 3 kn S.E, of

Faraya (7 km by road). Other crevices were inhabited by alpino

swifts or bats (Myotls blythi and Tadarida teniotis).

Genus Epteslcus Rafinesquo

1820. Eptosicus Rafinesque, Annals of Nature, p. 2.

Type species: Sptesicus mo3.anops Rafinesque » Vespeiijiio fuscus

Beauvois.

Dental foimula: 2-1-1-3/3-1-2-3 » 32

This genus is widely distributed in North America, Europe,

Africa, Asia and Australia. It is represented along tho Eastern

Mediterranean by two species, E, serotinus and E. bottae.

Following is a KEY to theso two species:

FA more than $2 mmj GLS more than 22 mm...... Epteslcus serotinus

FA less than 50 mm; GLS less than 13 mn,...... Epteslcus bottao 131

Epteslcus serotinus (Schreber)

177h« Ves- :xvbilio serotinus Schreber, Saugeth,, I:pi, ^3, text

p. 3->7.

15?£l. Epti leus serotinus Schreber; Ellerman and Morrison-Scott,

Choc list, p. 1$7.

Typo locall ■/: France,

Common name : Serotino Bat,

Distributio. : Europe, southern USSR, Korea, China, Mongolia, Kashmir, northern Iniia, Afghanistan, Iran, Turkey, Lebanon, Palestine,

Morocco, Tunisia and Libya,

Only tho nominate form is pertinent to this study,

Sptesicus serotinus serotinus (Schrebcr)

Synonymy under species.

Typo material: Not examined.

General distribution: Europe, Turkey, Lebanon and Palestine,

Allen (1915) and Lewis and Harrison (l?62) noted this species at Chtaura and Beirut in Lebanon, Bodenheimer (1935 and 1958) recorded it from Palestine (Jerusalem and Tel Aviv) and Lebanon, while

Harrison (I961|b) repoi'ted on specimens from Tel Aviv and Uadi Amud in Palestine,

Soecimens examined: 1, LEBANON: Beirut (1 Hi),

Identification: This bat is appreciably larger than E, bottae,

FA 52-56 mm and GLS 22-23 mm. Two incisors and four cheekteeth be­ hind the canine,- Color dark bro-.n on tho back slightly paler on tho belly.

Biology: Pregnant females, collected in late April and early

May (Harrison, 1961(b), had a single embryo each. Nothing is 132 known of t] ’ liabitat of this species in this region,

Eptosicus bottae (Peters)

1869. Vest .cus bottao Peters, Kbor. Pi’euss, Alcad, V,Tiss,, p, L1O6.

Typo localJ '.y: Yenen, Arabia,

Com;,ion narae : Botta^ Serotino Bat,

Distributio Yemen, Iraq, Egypt, Palestine, Iran, Afghanistan,

Mongolia, southwestern USSR, Rumania and Switzerland,

, .E, b, Innasi is tho only subspecies pertinent to this study,

Eptesicus bottae innesi (Lataste)

1887, Vesperugo (Vesporus) innesi Lataste, Ann. Mus, Stor, Mat,

Genova, It:62^,

1951, Eptesicus isabellims innesi Lataste; Ellerman and

Morrison-Gcott, Checklist, p, 1^6,

1961i, Eotesicus bottae innesi Lataste; Harrison, Mammals of

Arabia, l:l){0,

Lectotype : Ei collection no, 19.7»7«35>20, an adult femalo collected at Cairo, Egypt,

General distribution: Egypt and Palestine,

Harrison (l963c)rocorded 'this species for tho first time in this region from Yotvata, Palestine,

Specimens examined: 1, PALEST DIE: Yotvata (1 HARR),

Identification: Appreciably smaller than E. serotinus, tho only other serotino recorded from this region, FA LjO-ljl mm and GIS l6-17mm.

Biology: Harrison obtained tho only specimen from Palestine 133

"at duak f.l -ing abovo an area of irrigated farr.iland with linos of tamarisk and eucalyptus trees, surrounded by sandy desert".

Genus Obonycteris Peters

1359. Otoi'.cteris Peters, liber. Preuss. AJcad. V/iss., p, 223.

Tjrpo specie-Otonycteris homprichii Peters.

Dental fora Haj 1-1-1-3/3-1-2-3 ** 30.

This monotypic genus is restricted to North Africa and south­ west Asia.

Otonycteris hemprichi Peters

1859. Otonycteris hemprichii Peters, Kber. Preuss. Alcad. VJiss,,

p.223.

Type locality: None given, but as the type was collected by Kernprich and Ehrenberg one can assume that Egypt is the typo locality.

Common names: Henprich's Long-eared Bat.

Distribution: Algeria, Tunisia, Libya, Egypt, Sinai, Palestine,

Jordan, Syria, Iraq, Saudi Arabia, Iran,Afghanistan, Russian

Turkestan and Kashmir.

Four subspecies are recognized to date. The nominate sub­ species from North Africa, 0. h, cinerea in central Asia and 0. h. petersi and 0, h. jin from the Arabian Peninsula. Tho relationship between tho last two subspecies is. not clear, as 0. h. petersi is

known only from two specimens. However, so far it appears that

0. h. jin, restricted to tho central and western parts of tho

Arabian Peninsula, represents large sized populations, while

C. h. petersi represents small sized eastern populations (Cheesman

and Hinton, 192Uj Harrison, 1961(b).

Only tho former subspecies is pertinent to this study. 13U

Otonycteris hemprichi jin Cheesman and Hinton

192b. Qtoryetorio jin Cheesman and Hinton, Ann. Kag. Kaldlist.,lU(9) :5U9.

19^1. Oton, eteris hemprichi jin Cheesman and Hinton; Ellerman and.

Korr." Jon-Scott, Checklist, p, 180.

Holotype! Bli ;ollection no. 2U.0.2.2., an adult male coLlccted at

Hufuf (200 lb), Hasa, Saudi Arabia, on December 10, 1923, by

Major R. E. Cheesman,

General distribution: Palestine, Sinai, Jordan and Saudi Arabia.

s Bodenheimer (19I?3) referred a specimen collected by E, Schmitz

from Wadi el Hukallih, Palestine, to 0, h. hemprichi. Harrison

(l9chb) recorded specimens from Shen Ramon; Palestine. Atallah

(1966 and 1967a) reported on three spocimens collected at Azraq

cd Druz, a new record for Jordan.

Specimens examined: 9. JORDAN: Azraq ed Druz (3 SA). SYRIA:

Qariatoino (1 BZM). PALESTINE: near Jerusalem ( 1 FMNH). Also

four paratypes from Hufuf, Saudi Arabia at tho BM.

Identification: Ears very long exceeding 3^ mn in length re­

sembling Plecotus sp. The ears in the latter species, however,

are joined across tho forehead, free in 0. hemprichi. FA 61-69

mm CIS 23.£-2f>.£ mm.

Biology: I found tlireo pregnant females, each with two embryos,

in an old deserted hut near the Azraq Oasis, Jordan, on May 2, 1966.

The hut is located on the basalt slope at the edge of tho hammada.

As X entered four bats flew and hung in a close by date palm tree.

The next day I obtained three specimens from the interstices

between the stones, of which the hut walls were constructed.

Zahavi (vide Harrison, 196!jb) found taro specimens in a similar 13!? habitat at hen Ramon in the Nccev. Theso collection sites indicate tho high ar dity and temperature tolerance of this species. Judging from tho si; i of embryos examined, I estimate the time of birth at early Jr,. -•,

Genus Plecotus E, Geoffroy

I8l8, Picci vUS Geoffroy, Description de l1Egypt, 2:112,

Type species: Vesperbilio auritus Linnaeus,

Dental formula: 2-1-2-3/3-1-3-3 « 36

. Ellerman and Morrison-Scott (19^1) maintain that P, auritus

(Linn,, 1753) is the only representative of this genus in the Old

World, Lanza (i960) demonstrated that this species actually includes tvro sibling species, P. ward! Thomas, 1911 (considered a subspecies of P, auritus by Ellerman and Morrison-Scott) and

P. auritus, Lanza (op, cit,) distinguished P, wardi from tho latter by its larger skull, more inflated tympanic bullae and the shape and size of the baculum. Recent authors concur irith

Lanza's rearrangement but show that P, austriacus (Fischer, 1829) has priority over P. irardi,

Stebbings (196?) presents excellent criteria for positive identification of P, auritus and P, austriacus,

Plecotus austriacus (Fisher)

1829, Vespertilio auritus austriacus Fisher, Synops, Hamm.,

p. 117.

Typo locality: Vienna, Austria,

Common names: Grey Long-eared Bat, Long-eared Bat, 136

Distributic : Ths exact distribution of this species in relation to

P. auritus s not clearly understood. Doth species range throughout

Europe, sou hern USSR, Japan, China, Kashmir, Afghanistan, Iran,

Turkey, Syj. a, Palestine, Sinai and North Africa southward to Sudan and Ethiop5 ,

Harris- n (1963q) referred three specimens collected in Iran to

P, vardi ( P. austriacus) on the basis of cranial measurements and the shr..;;e and size of the baculum which is shown to satisfy

Lanza’s criteria for this species (Harrison, 0£. cit. Fig, 1 a and b). In this paper Harrison indicated that the status of P, christiei

Gray, from North Africa and the Eastern Mediterranean Region, cannot be assessed without examining the baculum. However, Harrison (ipoljb) later referred the form chrisieri to P, austriacus on the basis of measurements alone,, I have examined the baculum in two specimens collected at Mogharet Khureitun (Cave of Adullam) Palestine, and found the baculum to agree in every respect with Lanza’s and Harrison's figures for P* austriacus. Thus there is no doubt that christiei is a pale subspecies of P, austriacus rather than P. auritus,

The following is the only subspecies pertinent to this stud^

Plecotus austriacus christiei Gray

1838, Plecotus christii Gray, Mag, Zool, Bot,, 2:h95*

1931* Plecotus auritus christiei Gray; Ellerman and Morrison-

Scott, Checklist, p. 131

196b, Plecotus austriacus christiei Gray; Harrison, Mammals of

Arabia, 1:178. 137 Holotype: ’ i-I collection no, 66a, obtained in North Africa by

D, Christie, Skin only in a poor condition.

General distribution: Egypt, Cyrenaica, Tunisia, Algeria,

Morocco, Stv'an, Ethiopia, Sinai, Palestine and Syria,

Tristr^n (l866a and 1881:) reported P, auritus as "common in all the hill-country of Palestine, especially in caves and tombs about Bethlehem and Jerusalem, and by the sea of Galileo",

Bodenheimer (1920, 1935 and 1953) maintained that this species is common in Palestine and referred these populations to tho form christiei, Aharoni (1930) recorded specimens from tho Dead

Sea Basin, -while Harrison (19614b) listed tho following localities in Palestine: 10 km N. Eilat, Eilat, Cave of Adullam, Timna, and Nahal Hever (Judean Desert), Trouessart and Kollmann (1923) obtained a specimen from Djeroud, Syria,

Specimens examined* U. PALESTINE: Cave of Adullam, 10 km

S, E, Jerusalem (3 SAj 1 BM),

Identification: Ears large (35-39 mm) connected across the forehead, FA 334»1 mm and GLS 16,5-17,5 mm.

Biology: This bat inhabits caves in desert and steppe areas, singly or in small numbers. At Mogharet Khureitun (Cave of Adullam) I found theso bats hanging solitarily on the ceiling of the cave, 20 m high, along with Rhinolophus blasii, Harrison (196ljb) recorded early pregnancy in a female collected in March at Nahal Amram,

10 km N, Eilat, Negev, Stebbings (1966a and 1966b) for a description of the habits and ecology of this species in England)^ 133 Genus Hiniopterus Bonaparte

1837* Mini- ■-’torus Bonaparte, Fauna Ital., I: fasc, 20.

Typo specie j Vespertilio ursinii Bonaparte « Vespertilio schroiborsi Kuhl,

Dental fore la: 2-l-2-3/3-l~3-3a 36. # This v .despread Old World genus is represented in the Palaearctic

Region by t' .0 f ollovdng species.

Hiniopterus schroiborsi (Kuhl)

1819. Vesperbilio schreibersii Kuhl, Am. Wetterau, Ges. Naturk,

U, 2:18^.

Typo locality: Kulmbaser Cave, mountains of Southern Bannat, Hungary.

Coixicn names: Schreibcrs* Long-winged Bat, Schreibers1 Bat, Long­ winged Bat,

Distribution: Europe, Asia, Malaysia, Australia.:., Madagascar and most of Africa,

Eastern Mediterranean populations represent tho following subspeciess

Hiniopterus schroiborsi pallidus Thomas, 1907

1907. Hiniopterus schreibersi pallidus Thomas, Ann. Mag, HalHtV20(7);197,

1956. Hiniopterus schroiborsi pulchor Harrison, J. Mammal.,

37(2):26l. From Ser'Amadia, Kurdistan. (Harrison, 19614b),

Holotype: BM collection no. 7.7.114.7., on adult female obtained on

March 25, 1907, on the southern shore of tho Caspian Sea, Iran,

General distribution: Iran, Turkey, Iraq, Syria, Lebanon,

Palestine and Jordan, 13?

Tristi a (l83li) and Aharoni (1930) recorded this species from caves overlooking tho Jordan Valley. Bodenheimer (1920, 1935 and.

1953) repoi .od finding long-winged bats common throughout Palestine and Jordan, Ucttstein (1913) noted it from Aleppo, Syria, Loris and

Harrison 0)62) from 2 km E. of Amchite, Lebanon, and Harrison (1956a and 1959a) from Jerash, Jordan, Harrison (1961}b) reported on specimens from Tel el Kurdoni, near Akko and the Herzlia Cave in

Palestine. Atallah (1969b) recorded specimens from Anjar and

Mogharet Saleh, Lebanon and Dor (19l(7) from owl pellets in

Palestine.

Specimens examined: 108 LEBANON: Mogharet Saleh, 2 km E, Amchite

(UO SAj 6$ AUBj 2 BM)j Anjar (l SA).

Identification: A medium size bat, FA UU-U8 mm and GLS lU.5-16.0 mm. Lower jaw with six cheekteeth behind the canine. Second phalanx of third finger about three tines as long as tho first phalanx.

Biology: At Mogharet Saleh I found this bat in large colonies roosting with Rhinolophus forrunequinum, R. ouryale, Hyotis myotls, M, blythi and Rousettus aegyptlacus. These colonies do not migrate but hibernate at this locality. At a cave near

Anjar a single long-cringed bat was found roosting with 225 M. myotls. Harrison (1956a) reported that their flight is rather si-rift and continues late at night. Nothing is known about tho reproductive biology of this species in the Middle East. Family HDLOSSIDAE Gill, 18?2

This mily is widely distributed ranging over most of tho world be tv- n latitudes k5>°il and U5°S. It is represented in tho

Eastern He' ..terrancan Region by a single species, Tadarida teniotis,

Genus Tadarida Rafinesquo lOllj, Tad. ./Ida Rafinesque, Precis. Scm., p.

Type species: Cephalotes teniotis Rafinesquo.

Dental fon.sula: 1-1-2-3/2 or 3-1-2-3 « 30 or 32.

Tho follovring species is tho only Palaearctic representative of this genus found in tho area of study.

Tadarida teniotis (Rafinesque) l8lU. Ceohalotes teniotis Rafinesque. Precis. Son., p. 12.

Type locality: Sicily, Italy.

Common names: European Free-tailed Bat.

Distribution: Southern Europe, southwestern USSR, Korea, Japan,

Taiwan, China, Afghanistan, Iran, Iraq, Turkey, Lebanon, Palestine,

Jordan, Egypt, Morocco and Algeria.

This species has six lower incisors instead of four. Following is the only subspecies pertinent to this study:

Tadarida teniotis ruepoelli (Temminck)

Py50?53 rupelii (sic) Temminck, Mon. Mamm., I?22U, pi. 13,

1951. Tadorida teniotis rnnpelll Temminckj Ellerman and Morrison-

Scott, Checklist, p. 13fi.

Tupe: Not examined, from Egypt,

General distribution: Iraq, Lebanon, Jordan, Palestine, Egypt,

Morocco and Algeria, Trist\ \ (183U) recorded Nyctlnomus cestonii Savi (»^* tenlotia) an caves along the Jordan Valley. Bodenheimer (193I> and 1938) - ported on specimens from Yagur, Herzlia and Jerusalem,

Schmicdokn ht (1906) from Jericho, and Harrison (19614b) from

Jerusalem, -i if fa and Beit Guvrin in Palestine. Lewis and i Harrison (5 .'62) repoi'ted on few specimens from Lebanon taken at

Natural BrJ ige, near Faraya, while Atallah (1967a) found several colonies at Faidhat cdh Dhahikiya, the north end of Uadi Sirhan,

Jordan,

Specimens examined: Hi;. JORDAN: Faidhat edh Dhahikiya (h SA).

LEBANON: Natural Bridge, near Faraya (10 AUB).

Identification: Tail projects conspicuously beyond the border of the interfemoral membrane, Federate size bat, FA 3U-6U mn and GLS 23-23 mm.

Biology: Lewis and Harrison (1962) found this species at an altitude exceeding 1,600 m occupying small fissures under the

Natural Bridge, near Faraya, Lebanon. I found these bats common along tho north end of Uadi Sirhan in extreme desert conditions. They roosted in largo coloni.es in narrow crevices in tho chalk cliffs surrounding the VJadi, They were easily located by their squeaks and guano accumulated at the bases of theso cliffs,

Lewis and Harrison (loc. cit.) placed parturition in this species at mid-June judging from tho size of embryos in two pregnant females collected May 31, 1961 at tho locality men­ tioned abovo. Each femalo held a single embryo. Ordor MGOMOHPHA

The orler includes two recent families: Leporidae (rabbits

and hares) and Ochotonidae (pikas). The monotypic, holarctic

family Ochotonidae is widely distributed in Palaearctic Asia east

of the Ural Mountains and the Caspian Sea, Ochotona rufescens

(Gray) which inhabits the mountainous regions of Iran represent

the westernmost record of the family. It is interesting, however,

that this species does not extend farther west into Turkey, Iraq,

and the Fastcm Mediterranean Region, especially as the mountainous habitats in these countries are not much different from those in

Iran, The family Leporidae achieves an almost world trade dis­

tribution especially as a result of human introductions (ex,

Australia) .

Family LEPORIDAE Gray, 1821

Rabbits and Hares

Two genera represent this family in the Palaearctic Region,

The monotypic genus Oryctolagus represented by 0, cuniculus (Linn,),

the European rabbit an introduced species, and the genus Lepus, with many species of hares native to all continents except South America

and Australia,

ill 2 1)43

Genus Lepus Linnaeus, 1758

1758. Lee'. Linnaeus, Syst. Nat., 10th ed., 1:57.

Typo speci< : Lepus timldus Linnaeus,

Three species, L. eurcpaeus, L. capensis and L, arabicus, have been rep or! .d fi’om the Eastern Neditorranean Region (Ellerraan and

Morrison-S'.ott, 1951). Gureev (196)4) restricted the range of L, capensis t< Africa, while Fetter (1959 and 1961a), Yon-Tov (196?) and Angerm;.iin ({^ers. coiiin.) believed that L. capensis and L. europaeus r.ro conspecific. The third species L. arabicus, a desert fora, has been given full specific status on the basis of its highly inflated bullae and smaller dimensions (See Yom-Tov, op. cit. for details).

However, Nr, D. L. Harrison (pers, coran.) informs mo that he finds a clino in characters used to separate L, arabicus from the northern and eastern populations of the L. capensis-europaeus group (See also discussion under L. c, arabicus). If this is valid,then all three species are conspecific. In this paper both arabicus and suropaeus are considered junior synonyms of L, capensis.

Lepus capensis Linnaeus, 3.758

1758, Lepus capensis Linnaeus, Syst, Nat., 10th ed,, 1:58.

Type locality: Cape of Good Hope, Africa,

Common names: Cape Hare, Tolai Haro, Arabian Hare (L. c, arabicus),

Syrian Hare (L. c. syriacus), Arnab or Arnab Barri (/a’abie),

Distribution: Africa, Europe and temperate Asia.

Two subspecies are recognized in the geographical area covered. L. c, syrL?AU3 a brush and foi'csfc land inhabitant restid-cted to the r'editei'j axiean Region and L. e. arabicus a desert adapted form,

Lenus capensis s^rriacus Ehrenberg, 1033

1833. lopuM syriacus Elirenberg, Sjanb. Phys. Haam,, 2 sig. u.

1951. Lepu-i europaeus syriacus Ehrenberg, 1833i Ellenaan and

Morrison-Scott, Checklist, p. U35«

Typo: Hateri ■ .1:»Froih "Mt'i ? Lebanon; •

General distribution: Turkey, Lebanon, Syria, Jordan and Palestine.

Tristram (l866 and 19^6) recorded this hare as common in the Mediterranean Region of Palestine. Lewis,

Lewis and Atallah (1967)reportod this subspecies from Lebanon and the Heins plain in Syria. Yom-Tov's (196?) specimens from his localities, numbered 1 through 33 and through 6l refer to this form.

Specimens examined: 19. LEBANON: Hear Chmistar (12 AUB), Also seven specimens from Turkey (Bitlis, Horsing and Urfa) at the

FMNH.

Identification: Large hares, GLS OU-101 ram and HP ll£-l6!> mm.

Percent bulla longth/occipitonasal length (as defined by Yom-Tov, loc. cit.) is less than lU.O#. Guard hairs on the back are double banded (brown-black-brown-black tip) or completely black, while the underfur is single banded with grey baso and a black tip. This gives the animal an overall light grey color. Biology: V'tis haro occurs on tho Coastal Plain and the- Upland

P.egions nor'li of Beersheba to the west and Jerash to the east*

Lei;is, Leiri-J and Atallah (196?) found these hares uncommon in

Lebanon diu; to excessive hunting pressure and intensive defor­ estation as a result of expanding cultivation.

Syrian hares are nocturnal, talcing cover in the daytime under vegetation or in between rock .ledges* Nothing is known of tho re­ productive biology of this species in the Middle East*

Lepus capensis arabicus Ehrenberg, 1833

1833. Lepus arabicus Ehrenberg, Syob. Phys. Mamm., 2: sig. r.

1833. Lepus sinaiticus Ehrenberg, Symb. Phys. Mamm., 2: sig, t,

near Mt. Sinai. NEW SYNONYMY.

1867. Locus judeae Gray, Ann. N. H., 20:222. Palestine. NEW SYNONYMY*

TypeMaterial:''Frbm'Qunfidha'(19-Ni hL E)‘, Saudi Arabia,

General distribution: Sinai, southern Palestine, Jordan, Saudi Arabia,

Iraq and Kuwait,

Tristram (l866aand 188U) restricted the distribution of this form to the Bead Sea Basin and erroneously recorded L. aegyptiacus and

L. isabollinus, two African representatives of L. capensis, from southern Palestine. Bodenhoimer (1920, 1935 and 1953) repeated

Tristram's records, Aharoni (1930) and Hart (1891) found this hare abundant in southern Palestine.

Atallah (l9o7a and c) recorded specimens from the Azraq and El-Jafr Basins in Jordan. There appears to be no record of this form in Iraq. I have, however, seen two road kills on the Hj6 desert road southeast of Rutba that could be referable to this subspecies. Yon Toy's (196?) localities irr.ibcred 33 through £3 and

59 through C2 refer to the distribution of this form in southern

Palestine,

Specimens e: amined: 19. JORDAN: Qasr Amra (5 SA)j Jabal Aseikhem

(1 SA)j near Qasr ^mra (2 SA). Also four specimens identified as L. a, ar.iolcus from 5 mi S. E. Hodeida, Yemen, two L. c, sinaiticus fjrom St. Catherinet-1'onastory, Sinai, and one L. c, aegyptiacus from Qena, Isna, Egjgjt at the FHNH, and four L. arabicus from Badanah, Saudi Arabia af AUB.

Systematic remarks: Yom-Tov (196?) distinguished two populations in Palestine, a largo northern mountain form and a small southern desert form, and two species L. capensis and T», arabicus. Ho referred both populations to L, capensis indicating that the northern form has been previously referred to L, ouropaous syriacus, but he does not discuss the subspecific status of the southern form.

He reported L. arabicus from a single specimen taken at Eilat with l6t8% bulla length/occipitonasal length (B.L,P./O.N.). Ho differentiated between L, arabicus and L, capensis on tho basis of tho bullar ratio according to the proceeding formula (arabicus greater than 16% and capensis less than 16%), His figures (og. cit,,

Table 3), however, clearly show a perfect cline from individuals with largo bullae in the south to small bullae in tho north. It thus appears to bo clear that Yom-Tov’s L, arabicus is an individual with a largo bulla representing the southern population of L. canensis. This southern form has also been designated as L, c. sinaiticus, described from Mount Sinai (Tristram, 186^ Bodenhoimer, 1920, 1935 nd 1953j Aharoni, 1930)

Moasu'i 'i'lcnba and color descriptions of L. £. sinaiticus natch in c ;ry respect those from Jordan, previously identified as a» a‘. blcus (Atallah, 1967a). X have examined specimens from north' n Saudi Arabia, Jordan and Sinai and feel compelled to refer al .1 of these specimens to L, capensis arabicus, tho earliest available name.

Other hares from southern Palestine have been referred to

L. .jndeao Gray, L, aegyptius Do snares t and L. isabellinus

Cretzschmar (Tristram, l866aand I831j; Hart, 1891 j and Bodenheimer,

1920, 1935 and 1953). These are discussed separately belows

1, Locus .judoae Gray: This form is known from a single specimen

collected by Tristram in Judea. Tristram (1881:) indicated

that this form is very close to the Persian L. craspedotis

Blanford considered a subspecies of L, arabicus by EUerman

and Morrison-Scott (1951). As Yom-Tov (196?) found specimens

from the Judean Desert identical to the southern population

here named L. c. arabicus, L. judae Gray will be considered

a junior synonym,

2, Lepus aegyptius Dosmarest: Petter (1963 and 196?) treated this

name as a nomen dubium. EUerman and Morriscn-Scott (1951)

reduced aegyptius to a subspecific level under L. capensis

and restricted its distribution to Egypt and the Sudan, indicat­

ing that Bodenheimer has recorded it from Palestine, Bodenheimer

(1920, 1935 and 1953) based his record on Tristram's (188U)

erroneous identification. 3. Lopus --'ibollinus Cretr^hinar: This fora is considered a

subspcclos of L, caponsis restricted to the northern Sudan

(Potter, 1967)• Records from Palestine must be erroneous.

Identifies Mon: Small hares, GLS 73-8J4 wm and HF 88-121 mm.

Percent bu3 a length/occipitonasal length more than

Color buffy bro’-m.

Biology: I found this hare common throughout the Eastern

Plateau. It is abundant where cover is available as in vradi systems and near an oasis. They are strictly nocturnal in habits, but a few were seen during the day as they dashed cut of their hiding places when approached. Their hiding places consist of a small depression tho size of thoir body, dug along the base of a shrub or a clump of grass.

Young individuals and lactating females were collected early in May, 1966, at the Azraq Basin. Asdcll (1961t) reported that in South Africa L. capensis is polyestrous and breeds in late winter or early spring producing 3 or U litters a year. I.ro such dataabe available from the Middle East, Order RODENTIA

Thirty five species of rodents, representing seven families,

are reports 1 in this paper from the Eastern Mediterranean Region.

Three additional species, Citellus citellus xanthoprymnus Bennett,

Glis glis (Linn.) and Apodemus flavicollis (Melchior), uhich have been for seme time associated with the Eastern Mediterranean Region are discussed and in all three cases reports of these species are believed to be erroneous.

Following is a KEY to Eastern Mediterranean rodent families:

1. Large animals, GLG over 100 mm and HB over £00 mm; pelage sparse,

very coarse, with much of the dorsal surface covered with quills,

...... -...... HYSTRICIDAE

Gmail animals, GIB less than $5 mm and HB loss than 1|00 mm; pelage

short and dense (spines are present in some mouse-sized species

(Acomys), but not modified into long quills)...... *...... ,.2

2. HB over 200 mm; tail fully haired, thick and bushy; infraorbital

Canal narrow not transmitting any part of masseter medialis muscle

...... SCIURIDAB

HB less than 200 mm; tail naked or sparsely haired; infraorbital

canal enlarged transmitting masseter medialis muscle...... 3

3. HF elongated, over 5h mm in length; tail at least 1-1/2 times HB;

infraorbital foramen greatly enlarged, insido width 3-U mm,

height 6-8 mm...... DIFODIDAE i$o

HF not tlongated, never exceeding U2 mm in length; tail less

than 1-1/2 times Iffi; infraorbital foramen moderately large,

inside ’ddth 1-2 mm, height 2-U mm...... U

It. Cheekteeth It/It; black stripe through eye...... GLIRIDAE

Cheektevth 3/3J no black stripe through eye...... 5

5. Tail much shorter than hind foot, if seen externally; eyes

minute, hidden by fur; external ear (pinna) absent; fossorial.

...... SPALACIME

Tail longer than hind foot; eyes well developed, easily visible;

external ear (pinna) present; terrestrial, occasionally arboreal ...... 6

6. Molar teeth with tubercles arranged in three longitudinal

series.«..»...... MURIDAE

Molar teeth flat crowned or with tubercles arranged in two

longitudinal series...... CRICETIDAE

Arabs refers to all rodents other than squirrels, mole-rats,

jerboas and porcupines either as mice (Far or Far El-Khala)

or rats (Jurd or Jardoun).

Family SCIURIDAE Gray, 1821

Squirrels

This family is vridely distributed throughout the Old World except for the Austr.cLlian Region and most of the desert parts of

North Africa and Arabia. It is represented in the Eastern Mediter­ ranean Region by a single species, Sciurus anomalus, an element of the Irano-Tauranian fauna.

Tristram (183U) erroneously recorded ground squirrels, Citellus citellus, as "Exceedingly abundant on the sandy and stony plains 1S1 of the uplands of Noab and Gilead, burro^dLng generally in the neighborhood of ruins.” This species, however, is not known south of the Anatolian Plateau in Turkey, and it is more than probable that Tristram's records refer to jirds or sand rats.

Genus Sciurus Linnaeus

17!?8. Sciurus Linnaeus, Syst, Nat., 10th ed., 1:63.

Typo Species: Sciurus vulgaris Linnaeus.

This genus is represented by two species in the Palaearctic

Region, S. vulgaris in Europe and northern Asia and S. anomalus in southwest Asia.

Sciurus anomalus Schreber

17^3. Sciurus anomalus Schreber, Saugeth., 14:781.

Type locality: Sabeka, 2$ km southwest of Kutais, Georgia, Caucasus.

Common names: Persian Squirrel, Sinjab (Arabic).

Distribution: Caucasus, Turkey, Iran, Iraq (Harrison, 19?6a), Syria,

Lebanon, Palestine and Jordan.

Th

■”1951:1477)/ the nominate form from Turkey and the Caucasus, S. a. pallescens from the Zagros fountains in Iran and Iraqi Kurdistan v and S. a. syriacus from the Eastern Mediterranean Region.

Sciurus anomalus syriacus Ehrenberg

1828. Sciurus syriacus Ehrenberg, Symb. Phys., I, pi, 8.

1951. Sciurus anomalus syriacus Ehrenberg) EUerman and Morris on-

Scott, Checklist, p. hll»

Tyre material: Not examined, from Lebanon.

General distribution: Lebanon, Syria, Palestine and Jordan. 152

Tristr .1 (1866a and 188U) and Bodenhoimer (1920 and 1935)

recorded th 3 squirrel from northern Palestine, Lebanon and Mt,

Kermon and .Damascus in Syria, Bodenheimer (1950) contended that

the species became extinct south of the Taurus Mountains, Turkey,

Levis, Levri; and Atallah (196?) however, reported its occurrence at 1 Faraya, Kfaraina, Becharre, Laklouk, Ain Zhalta, Barouk Cedars and

Hadeth Cedars in Lebanon,

Specimens examined: 3« JOPJDAN: Dibbine Ranger St,, northwest of

Jerash (1 SA), LEBANON: Natural Bridge, near Faraya (1 AUB);

Kfarzina (1 AUB),

Identification: This is tho only mammal in this region easily

identified by its bushy tail. Color of back Chickadee Gray, belly Amber Yellow.

Biology: This squirrel is relatively abundant in the Ajlun

Mountains, Mt. Hermon and the Lebanon Mountains, but restricted to cedar, oak or pine forests, Lewis, Lewis and Atallah (196?) maintain that the species can be found active any time of the day feeding on acorns and pine or cedar seeds. They construct their nests in trees or in rock crevices. Young are born in April and

Hay,

Family CRICETIDAE Rochebrune, 1883

Hamsters, Voles, Gerbils

This family is represented in the Eastern Mediterranean Region by three subfamilies including seventeen species. Following is a

KEY to subfamilies: 1!?3

1. Tail 3 ,33 than 10% HBj tail sparsely haired, lacking a terminal

pencil upper incisors not grooved...,.,...... 2

Tail 1 nger than 10% HBj tail fully haired with a prominent

terair .1 pencil; upper incisor grooved (plain in Psammomys)

...... Gerbillinae

2. Cheek ouches present} cheekteeth low crowned and rooted......

...... Cricetinae

Cheek ouches absent} cheekteeth high crowned and rootless

...... Micro ie

Subfamily Cricetinae Murray, 1866

Hamsters

This subfamily includes about 60 genera of New World rats and mice, true hamsters and zokors. It is represented in the

Eastern Mediterranean by two species representing two genera of true hamsters. Following is a NET to these genera and species:

Tail short and inconspicuous, usually less than 20 mm in length}

GLS over 30 mm} color of back golden-yellow......

...... Mosocricetus auritus

Tail over 20 mm in length} GLS less than 28 mj color of back

grey...... Cricetulus migratorius

Genus Cricetulus Milno-Edwards

1867. Cricetulus Milno-Edwards, Ann. Sci. Nat., 7:376.

Type species: Cricetulus griseus Milne-Edwards.

Seven species are recognized in this Palaearctic genus. All are primarily eastern Asiatic in their distribution except C. migratorius which extends westwards to Greece and the Eastern

Mediterranean Region 15U

Cricetulus migratorius (Pallas)

1773. Mus raigratorius Pallas, Reise, 2:703.

Type local ';y: Lower River Ural, Western Siberia.

Common nam a: Migratory Hamster, Gray Hamster.

Distributi n: Southern USSR from Ukraine to Novosibirsk Province,

Chinese Turkestan, Kashmir, Afghanistan, W. Pakistan, Iran,

Turkey, Gr oce, Bulgaria, Syria, Lebanon, Palestine and Jordan.

Aharoni (3.932) recognized two subspecies along tho Eastern

Mediterrancanj C. m. vemula Thomas in Turkey and northern Syria and C. m. cinerascens (Wagner) in Palestine, eastern and southern

Syria, However, Neuhauser (1936) restricted tho dark colored form vernula to the southeastern coast of the Black Sea south- wnrd to Erzerum in Turkey, and referred all specimens from south­ ern and eastern Turkey to tho latter subspecies, I have examined

15 specimens (SA and USNM) from Adana, Talas, Ulu Dag and Cehenem

Dere in western Turkey and can find no differences between these populations and those from Syria and Lebanon (see Appendix I).

Cricetulus migratorius cinerascens (Wagner)

I8I18. Hypodaeus cinerascens Wagner, Arch,fur Nat,, I:l8U.

191? 1. Cricetulus migratorius cinerascens Wagner; Ellerman and

Morrison-Scott, Checklist, p, 622.

Type material: Not examined, from Syria.

General distribution: W. Pakistan, Afghanistan, Iran, southern and southwestern Turkey, Syria, Lebanon, Palestine and Jordan.

Aharoni (1932) regarded the species as rare in Syria and

Palestine, recording specimens from Hama and Karyatein in Syria

Ras Baalbek and Sidcn in Lebanon and Jerusalem, Palestine. Bodenheimer (193!> snd 19^3) believed the species to bo common throughout the Mediterranean Region. Osborn (1965) and Lehmann

(1966a) ro orded specimens from Aleppo, Syria; and Lewis, Lewis and Atall- 1 (196?) from tho Eekaa Valley and Laklouk in Lebanon as well as Aleppo.

Other records.-Tristram (l80lj) and Bodenheimer (1920),

Specimens examined: 1:2 LEBANON: Becharre Cedars (3 SA)j

American University of Beirut Farm, Bekaa Valley (6 AUB; 13 SA);

Laklouk (1 AUB), Sidon (1 BZM); Baalbek (1 BZM). SYRIA: near

Aleppo (I4 AUB; 3 BZM); Hama(3 BZM); Karyatein (l BZM); PALESTINE:

Jerusalem (2 BZM). Also 15 specimen from Turkey, exact localities listed in text.

Identification: GLS 23.5-27.5 mm, but usually less than 26 mm.

Color of back usually Charcoal Grey with an inconspicuous dark median stripe, belly greyish white. Young specimens are much greyer on the back while mature adults are more of a Grey Stone color.

Biology: Specimens were collected during the winter and spring

(October through May) in Lebanon at altitudes from 800 n to

2,900 m. They vrore trapped along grain fields or on rocky waste grounds, associated with large colonies of Heriones tristramj at low altitudes and Hicrotus guentheri on gentle mountain slopes•

They are strictly nocturnal and rather sluggish in their movements.

One pregnant female was collected in late March and many immatures were taken in April at Aleppo, Syria. Of thirteen specimens collected December 1, 1961: in the Bekaa Valley only 156.

ono immati. o was found. Adult females ’..’ere not pregnant while adult male j had enlarged testes measuring approximately 10X7 mm.

Burrows distinct, with entrances descending almost perpendicular to the suiCaco* Examination of cheek pouches indicates a pre­ ference fc.* grass and wheat seeds.

Genus T'esocricetus Kehring

IO98. Menocid.cetus Kehring, Zool. Ana., 21:li9li..

Type species: Cricetus nigricans Brandt a Kesocricetus nigriculus

Nehring

This monotypic genus is confined to a very small area of south­ eastern Europe and southwestern Asia.

Mosocricetus auratus (Waterhouse)

1339. Cricetus auratus Waterhouse, Proc. Zool. Soc. London, p. 57,

1951. Mosocricetus auratus Waterhouse; Ellerman and Morrison-

Scott, Checklist, p. 630,

Type locality: Aleppo, Syria,

Coiranon names: Golden Hamster, Syrian Hamster.

Distribution: Syria, Turkey, Iran, Caucasus, Bulgaria and Rumania,

Aharoni (1932) restricted the nominate form to the environs of

Aleppo, Syria, and recorded M. auratus brandti, described from the

Transcaucasus, from northern Palestine and Lebanon. Her record for Lebanon is based on tho following statement of Tristram

(lOBh:12) where under his discussion of Cricetus nigricans Brandt

(■* M. a. nigriculus Nehring) he maintained: "This Caucasian species was found by Dr, Roth near Lebanon". Also though she maintained that this fora reaches its southernmost distribution 1$7 at Mctullrh in northern Palestine, she had no specimens to sub­ stantiate her findings. To my knowledge no specimens of M, auratus have been taken south of Aleppo, Syria,

Kesocricetus auratus auratus (Waterhouse)

Synonymy u; dor species,

Typo materjalt Not examined.

General distributions Northern Syria and the Anatolian Plateau,

Turkey. Aharoni (1932) and Osborn (1965) reported on specimens collected ;,t Aleppo, Syria.

Other iocords,- Tristram (1866a and 18314), Bodenheimer

(1920, 1935 and 1958).

Specimens examined: 5. SYRIA: Aleppo (2 BZM; 2 AUB; 1 USNM).

Identification: Color of back golden yellow, belly Woodash with a golden yellow throat stripe. Tail short, subequal to HF. CBL

33-37 mm.

Biology: Much has been written on the biology of this species in captivity, but very little is known of its habits and habitat in tho wild. Aharoni (1932) and Neuhauser (1936) found Syrian hamsters living in grain fields in tho Steppe Region. Aharoni

(op. cit.) also maintained that they live singly occupying deep burrows with several entrances and many storage compartments.

Asdell (196U) reviewed all that is known of the reproductive biology of the species.

Subfamily Microtinae Killer, 1906

Voles

This holarctic subfamily is represented along the Eastern

Mediterranean by two genera and four species. Both genera, Arvicola 158

and Micro1 "3, arc widely distributed throughout Europe and temperate

Asia read ‘.ng their southernmost point of distribution in Asia,

in Palest? .e. Follovring is a KEY to these genera:

Tail long'-a* than l/2 HBj HF over 30 mm; GLS over 35 nun..,..*......

...... ».Arvicola « Tail less than 1/2 HB; HF less than 2h mmj GLS less than 3k mm...

...... Hicrotus

Genus Arvicola Lacepcdo

1799. Arvicola Lacepede, Tab, des Mamm., p. 10.

Type species: Hus amphibius Linnaeus

Ellerman (19hl) restricted this genus to the Palaearctic

Region, recently however Zinmermann (1955) and Hooper and Hart

(1962) referred Hicrotus richardsoni (do Kay), of northwestern

United States, and southwestern Canada, to this genus.

Hinton (1926) in his revision of the subfamily Microtinae recognized ten Palaearctic species, as belonging to the genus

Arvicola. Ellerman and Morrison-Scott (1951) reduced all

Palaearctic species to a subsoccific status under A. terrestris.

Arvicola terrestris (Linnaeus)

1758. Hus terrestris Linnaeus, Syst. Nat., 10th ed,, I:6l.

Type locality: Upsala, Sweden,

Common names: Water Vole.

Distribution: Europe except for Ireland and many of the smaller islands, USSR Westwards from the Lena River, Persia, Turkey,

Syria and Palestine. 1S9

Trist, (l88h) listed the trater vole as a member of the

Palestinian fauna, but Aharoni (1932) raised some doubt as to tho existe. ce of this species in Palestine and maintained that the specie reaches its southernmost point of distribution at Lako

Antioch, u- uthern Turkey. Bodenheimer (1935:101) contended that the report of the water vole from Palestine was based on incorrect identification. However, he later (1950:185-6) listed this species from Palestine based on some skeletal remains found in pellets of tho barn owl reported by Dor (19U7)» Osborn (1962: Fig. 1) repeated these records,

0. Haas (pers, comm.) informed me that this species is common in the Lake Hule Valley in northern Palestine, however, I have no specimens at hand from this locality and to my knowledge there are no examples of this species from the geographical area covered in this study in any European or American museum.

Genus Hicrotus Schrank

1798. Hicrotus Schrank, Fauna Bioca, 1:72.

Type species: Hicrotus terrestris Schranck » Hus arvalls Pallas

Following is a KEI to the three species recognized from the

Eastern Mediterranean Region:

1. Tail approaching l/2 HBj ear length 15 mm or more5 sole of

hind feet with six tubercles...... M. nivalis

Tail 1/5 to l/U HB; ear length Hi mm or less; sole of hind

foot with five tubercles...... 2

2. Tympanic bullae length 10 mm or moro; lateral inflation of

mastoid portion of bullae conspicuous when viewed from abovo

M. socialis 160

Tympanic bullae length 10 ran or less; mastoid portion of

bullae poorly developed, not easily seen from above,...,

...... M, guentheri

Ognev (19^0) and Lay (196?) suggested that Hicrotus guentheri is only a subspecies of Hicrotus socialis, however, Kowalski (1950)>

Osborn (1952), Gromov ot al (1963), Atallah (1965), and Lehmann

(1966a) among others regarded the two species as distinctly separate,

Hicrotus nivalis (Martins) l8!i2. Arvicola nivalis Martins, Rev, Zool., p, 331,

Type locality: Faulhom, Bernese Oberland, Switzerland,

Common names: Snow Vole,

Distribution: The species is restricted to tho following mountain ranges in southern Europo and southwest Asia: Pyrenees, Alps,

Carapathians, Balkans, Caucasus, Elburz, Taurus, Lebanons and

Anti-Lebanons.

Aharoni (1932:212) believed the holotypo of Hypudaeus syriacus

Brants, 1927 to be nearly identical to Hicrotus nivalis (Martins,

I81i2) and reduced syriacus to a subspecies of M. nivalis, M, syriacus, however, predates M, nivalis by fifteen years and thus would take priority over nivalis. Bate (I9h5:l5l) maintained that it is by no means certain that syriacus was based on a form of M, nivalis, and Ellerman and Morrison-Scott (1951:708) listed

5* ?yrlaGU3 a3 a nomen dubium while Kowalski (1953:270) asked for tho abolishment of this forgotten name. Bate, Elleman and

Morrison-Scott and Kowalski based their evaluation of H. syriacus 161 solely on ■•haroni's remarks. I have examined tho holotype of H. syriacus ( 3ZH no. I3l6 from Syria collected by Hemprich and

Ehrenberg), of which only the left upper molar row was located, and found it to resemble M. socialis - M« guentheri group and is deiinitely not a form of H. nivalis.

Ahaivai (loc. cit.) recognised three subspecies along the

Eastern Mediterranean; syriacus on Mb. Lebanon, hermonls on Mt.

Hermon and pontius on the Ansariyah Mountains. X have examined specimens from all throe mountains ranges and refer all Eastern

Mediterranean populations to tho following subspecies: (M. n.

•pontius is an acceptable subspecies, but is restricted to the

Pontine Mountains, Turkey).

Hicrotus nivalis hormonis Miller

1908. Hicrotus hermonis Miller, Ann. Mag. Mat. Hist., 1(8):103,

1932. Chionomys nivalis syriacus Brants; Aharoni, Z. Saugetierk.,

7(2):212. NEW SYNONYMY.

19^1 • Hicrotus nivalis hermonis Miller; Ellerman and Morrison-

Scott, Checklist, p. 693.

Holotype: BM collection no. 6U.8.17.31#» an adult male preserved in alcohol collected on Mt. Hermon by Rev. H. B. Tristram.

General distribution: Lebanon and Syria.

Tristram (l80it) recorded this form from Mt. Hermon, Aharoni

(1932) from Kafrun, Syria; Bate (19U5) from Mt. Lebanon, and

Loris, Lewis and Atallah (196?) from the Natural Bridge, near

Faraya and Laklouk in Lebanon. 162

Other records.- Miller (1908), Bodenheimer (1920, 193^ and

1958), EUerman (ISW), Koualaki (1958) and Atallah (1965).

Specimens examined: 55. LEBANON: 5 km S. \J. laklouk (1 SA);

Refuge of Akl, laklouk (1 REL); Cedars, N. E. Becharre (0 SA);

Western slopes of Mb. Hemon, 3 km E. Rachaya (1 SA); Jabal

Safha, elc/. 1350 m, 2 kr.i N. W. Kfar Qouq (6 SA); 1 km W.

Mreijat (2 SA); 2 km N.W. Dalir El-Eaidar (2 SA); Jabal El-ICnisse,

elev, 19C0 m, (7 SA); S. slopes of Jabal Sanine, olev, liiOO m,

(9 SA); Natural Bridge, Faraya (6 SA; 6 AUB), SYRIA: Kafrun (1

BZM). Also l\ specimens from Taberga, Caucasus (HZM) and the holotype of M. nivalis pontius (BM).

Identification: Tail about l/2 HB and ear length exceeding 15 mm.

General appearance grey, olive grey in other microtines in the area.

Biology: Snow voles are only on rare occasions found on mountain

ranges below 1,300 m. In Lebanon they were found on mountain

ranges below 1,300 m. In Lebanon they were found from mountain peaks exceeding 2,700 m descending the slopes to 1,300 m, except

for a single specimen collected at 1,150 m near Rachaya. Burrow

entrances are always concealed under rocks and boulders, the

characteristic habitat of this species.

They are strictly nocturnal. Young are bom in May and

early June. Where they occur,snow voles are not abundant and unlike other Eastern Mediterranean microtines they are not

conspicuously colonial.

Hicrotus socialis (Pallas)

1773. Hus socialis Pallas, Reise Russ. Reichs, 2:705.

1827. Hypudaeus syriacus Brants, Hot. Gesl. d.-Huizen, p. 92.

Syria. NEW SYNONYMY (see discussion under M. nivalis). 163

Type local L-y’: Grassy regions of desert by the Ural River*

Common nam-ss Social Vole.

Distributi .ns Southwestern USSR, Turkey, Iran and Syria.

Bate (19)-i5) recorded M. socialis from Laklouk and Becharre in Lebanon based on examination of owl pellets collected at i those locnllties. Bodenheimer (1958) maintained that H. socialis paradoxus (Ognev and Heptnor, 1928) inhabited the higher altitudes of Mb. Lebanon and the Anti-Lebanon Mountains, Kowalski (1958) collected a few specimens from Mt, Sanine and Jabal Kammouha in

Lebanon.

Atallah (1965) and Levris, Lewis and Atallah (196?) showed that there is a considerable variation in cranial and external measurements of'Hicrotus guentheri taken at different altitudes.

High altitude populations were found to bo small,resembling M, socialis in measurements but cranially exhibiting all character­ istics of M. guentheri. This may suggest that previous records of socialis from Lebanon have been due to erroneous identification.

•Dr. D. L. Harrison compared some of Kowalski's material from

Lebanon referred to M. socialis and found it identical to the high altitude populations of M. guentheri (pers, comm.). Bate's material was not examined but intensive trapping at tho same localities yielded only M. guentheri. I have trapped for micro­ tines throughout Lebanon and collected over 500 specimens of M, guentheri, but not a single M. socialis. It thus appeal’s that

M. socialis is not a member of the Lebanese fauna. I have examined six specimens, two adults and four young, of K. socialis collected by REL at Aleppo, Syria. This appears to be the only record of this species along the Eastern Mediter­ ranean. My sample is inadequate for subspecific determination.

Hicrotus guentheri (Danford & Alston)

1880. Arvlcola guentheri Danford and Alston, Proc, Zool. Soc.

Lond,, p, 62.

1917. Hicrotus phllistinus Thomas, Ann. Mag. Hat. Hist., (8)19:li£0.

Elcron, Palestine. (Bodenheimer, 19h9).

1932. Hicrotus guentheri Danford & Alston; Ellerman and Morrison-

Scott, Checklist, p. 696.

Type locality: Marash, Turkey.

Common names: Gunther’s Vole, Mediterranean Vole, Levant Vole.

Distribution: Greece, Turkey, Syria, Lebanon and Palestine.

All Eastern Mediterranean populations have been referred to the nominate form (Ellerman and Morrison-Scott, 1951; Atallah,

1965>; Lewis, Lewis and Atallah, 1967).

Hicrotus guentheri guentheri (Danford & Alston)

Synonymy under species.

Syntypes: Two, EM collection no. 80.b.9.21., an adult male, and no. 8O.I4.9.22., a young, collected at Marash, Turkey in February,

1879.

General distribution: Turkey, Syria, Lebanon and Palestine.

This is a common species throughout the Mediteri’anean Region.

It has been reported from Palestine (Tristram, 188U; Thomas, 1917;

Bodenheimer, 1920, 1935 19li9, 1953 and 1959; I. Aharoni, 1930;

B. Aharoni, 1932) and Lebanon (Allen, 1915; Bate, 19li5; Kowalski, 16$

19$3j AtaO .a.h, 196^; Lehmann, 1966; Lmds, LeirLa and Atallah, 1967)*

Specimens examined: 203. LEBANON: American University of Beirut.

Farm, Eck' \ (36 AUB); 2 km N.N.W. Dahr El-Eaidar (l6 SA; 16 AUB);

6 km E. F; ..’aya (10 AUBj 16 SA); Cedars, E. of Becharre (l AUBj 3 SA)j

3 km E, of Rachaya (2 SA); Jabal El-Knisse, elev, 1900 m (1 SA); 2 km N.W: of An jar (20 SA); S. slopes of Jabal Sanine, dlev. 1,700 m

(2 SA) and elev. %000 m (9 SA); Refuge of Akl, Laklouk (32 AUB);

Hasbaya (1 FKNH). PALESTINE: U kmN.W. of Beit Fajjar (6 SA);

Ramallah (2 BZM); Jaffa (1 BZM); Ekron, south of Jaffa (TYPE of M, phj.listlnus at BM; 3 BZM); Jesreel (6 BZM); Mishmar Haemeg (2 SA);

Also 18 specimens from Azaze, north of Aleppo, Syria at BZM.

Identification: Mastoid portion of auditory bulla poorly developed and not visible in dorsal view. Bullae less than 10 mm in length, over 10 mm in M. socialis. Tail longer than IIF approximately l/Ij to l/$ HB length. Hairs on back with grey bases and yellow­

ish brown tips, usually with a reddish tinge if taken at altitudes over 1,600 m. Hairs on ventral surface yellowish or with white tips and grey bases. The transition between tho dorsal and ventral coloration along the flanks is either distinct (ht high1 elevations) or gradual (at low altitudes). •• Biology: Gunther's voles are the most common microtine rodents in the Mediterranean Region, occurring from tho coastal plain to above 2000 motex’s in tho mountains. They tend to be associated with grassy areas and agricultural fields but also occur with almost equal frequency in uncultivated mountain valleys- in close 166

association with Berboris erotica L« and other mountain shrubs.

Their burrow systems are extensive, with many entrances and

interlacing tunnels. These tunnels are five to eight centimeters beneath the surface of the soil. Burrow systems of family groups

in any ono colony overlap with each other so thoroughly that it

is virtually impossible to separate one system from another, presumably individuals use each others tunnels extensively.

Similar finds were reported by Ondrias (1965) in Greece,

Field observation showed that this species tends to be active throughout the day as well as tho night with peaks of activity occurring in the morning and evening. The majority of the surface activity is associated with short, rapid forays away from tho entrance of the burrow to obtain food. This food, consisting of succulent vegetation, when available, is then taken into tho burrow to bo eaten. Although an abundance

of succulent vegetation seems necessaiy for peak population, colonies at higher altitudes (where water may bo lacking during the late summer months) seem to be able to subsist to a large extent on dry grass and other dry vegetable material. At low altitudes largo populations may be found around rivers and other water sources.

This species lias occasionally been referred to locally as the "Plague Vole", a name applied to it by virtue of its high fecundity. Collections in Lebanon indicate that reproduction takes place throughout tho year at lower altitudes, with only brief intorruptions during tho cold season at higher altitudes. 16?

Dissections of tventy-seven pregnant females show an average of

8 embryos p r individual but as many as 17 have been reported

(Bodenheimc •, 19l|9) in some specimens. Tlais latter nuiaber is excep­ tional and ith but eight mammae on the adult female, many young would probably perish,

t The ne..its are usually located at the termimis of a lateral branch of the burrow system at a depth which appears to bo deter­ mined by the drainage of the soil. In wot areas, the nests may bo situated within five to eight centimeters below the surface, while in drier soils, they may occur at depths to one meter. The nest is constructed of finely shredded grass with a lining of similar but more finely shredded material,

A large population of these voles may become of economic importance through their voracious feeding habits. Unless some effort is made to control them,they can rapidly denude large areas of such cultivated crops as alfalfa (Aharoni, 1932}

Podenheimer, 1914? and 193*9). This has certainly been the case in such locations as the A.U.B. Farm in .the Bekaa Valley,Lebanon, where a combined program of irrigation and poisoning has failed to eliminate the population from the alfalfa fields.

From field observations, there is little doubt that this species is heavily preyed upon by hawks, owls, snakes and carniv­ orous mammals and they probably constitute the bulk of these predatordb diets.

Subfamily Gerbillinao Gray, 182^

Gerbils and Jirds Along the Eastern Mediterranean members of this subfamily are typically inhabitants of the Desert and Steppe Regions. 163

Their gan~ -al coloration is essentially light brown resembling the color xf the soil on which they live. Thirteen species rep­ resenting 'our genera are reported from this region. Following is a KEY t) the four genera pertinent to this study:

1. Upper Incisors with a longitudinal groove; tail more than

803 of HB...... 2

Upper .incisors smooth; tail less than 803 of

...... Psammomys

2. Large forms, GLS usually over 30 mm; adult cheekteeth ex­

cessively hypsodont and flat crowned, showing no signs of

the original tubercles...... 3

Small forms, GlS less than 28 mm; adult cheekteeth not be­

coming extremely hypsodont, but with clear traces of the

two original tubercles...... Gerbillus

3. Soles of hind feet naked; tail bushy; tympanic bullae

highly inflated with suprameatal triangle inconspicuous

(Fig. 7DK...... Sckeetamys

Soles of hind feet at least partly haired; tail thinly

haired with a terminal pencil; tympanic bullae if inflated

then suprameatal triangle is also highly inflated.,...,,,.

...... Heriones

A most interesting study on zoogeographical and ecological relationship of gerbils and jirds inhabiting the Saharo-SIndian

Desert Region is presented by Fetter (l?6lb), while those in

Palestine have been reviewed by Zahavi and Wahrwan (1957) •, 169

Fig. 7. Fostero-lateral view of skull. A. Meriones libycus B. Heriones crassus C. Meriones tristrami D. Sekeetamys

calurus 170

Genus Gerbillus Demsmarest l80)j, Gerbillus Desm-arest, Nouv. Diet. H. N., p. 2lj.

Type species: Gerbillus aegyptius Desmarest « Dipus gerbillus

Oliver.

Six species are reported from the Eastern Mediterranean

Region. Follov.-ing is a KEY to these species,

1. Soles of hind feet naked...... 2

Soles of hind feet fully haired...... ,U

2. Small, CDL less than 20 mm; HF less than 21 mm; tail less

than 100 mm in length...... 0. henleyi

Large, CDL over 21 mm; HF over 22 mm; tail over 100 mm in

length...... 3

3. Bullae inflated, mastoid region easily visible from above;

zygomatic arches almost touching auditory meatus...... G. nanus

Bullae not inflated, mastoid region not visible from above;

zygomatic arches widely separated from auditory meatus......

...... 0. dasyurus

I4. Small, GLS less than 23 mm; HB 70-100 mm, never over 100 mm,5

Large, GLS over 30 mm; HB 95-130 mm..,...... ,G. pyrairridusn

5. Dorsal pelage uniform; 2N “ h3 chromosomes...... G. .gerbillus

Dorsal palago greyish in meddle; 2N = I4O chromosomes.

...... G. allenbyi

Gerbillus dasyurus (Wagner)

I8b2. Meriones dasyurus Wagner, Arch, Nat. 8,1:20.

1951. Gerbillus dasyurus Wagner; Ellerman and Morrison-Scott,

Checklist, p. 633. 171

Type loca’ity: Sinai.

Common nar .3: VJagner’s Gerbil.

Distribution: Sinai, Palestine, Jordan, Saudi Arabia, Iraq, and

Syria.

Havri/on (l9!?6d) described Gerbillus dasyurus mesopotamiae

from Iraq. Hatt (19!?9) referred two specimens from Lake

Habbaniya and Basra in Iraq to this subspecies. Lay (1967:173)

examined Hatt's specimens and referred them to G. nanusj

also Harrison (pers. comm.) has changed his views and now regards mesopotamiae as a-subspecies of 0, nanus.

Lehmann (1966a) described G. dasyurus nalmyrae, a light

colored form from Palmyra, Syria and G. dasyurus leosollicitus,

a dark colored form from Dier-el-Hajar, 2!? km S, E. of Damascus,

Syria, I have examined both holotypes (Aid), paratypes and many

series of specimens taken at different localities in the Eastern

Mediterranean Region and found tremendous variation in the coloration

of this species at different localities or when living on different

soil types. Specimens collected on the dark basaltic rocks are

slightly melanisticj those collected on sand are very light brown

colored while those collected on the rocky steppes are sort of an

intergrade. Specimens from southern Jordan and the Syrian Desert

(excluding the basalt desert) are identical to the nominate fom described from Sinai., Those from the environs of Jerusalem, Pales­ tine, are darker, but occasionally light color forms are found.

Others from the Eastern Uplands in northern Jordan are still darker than the Festern Uplands specimens and,as mentioned above,those on the basalt rocks are melanistic. 172

G. d, olmyrae io identical to the nominate subspecies in every r< ;pect including color, G. d, leosollicitus, though dark r than the nominate form, vrill be considered synonymous ith it because the sole use of color as a criterion f r subopecific differentiation in this species is not sour especially as pelage color seems to depend entirely on local conditions, in particular coil types,

All Eastern Mediterranean populations are here referred to the nominate form,

Gerbillus dasyurus dasyurus (Wagner)

Original citation under species,

1966. Gerbillus (Dipodillus) dasyurus palmyrao Lehmann,

Zool, Feitr., 12(2):288. Palmyra, Syria. NEW SYNONYMY,

1966, Gerbillus (Dfoodillus) dasyurus leosollicitus Lelimann,

Zool, Beitr., 12(2):288. Deir-el-Hajar, Syria. NEW SYNONYMY.

Type material: Not examined.

General distribution: Sinai, Palestine, northern Saudi Arabia,

Jordan, western Iraq and Syria.

Mehring (I90]p) described Dipodillus dasyuroides from the mountains of Moab, Jordan. Allen (1915) recorded specimens from

Petra, Wadi El-Hasa, Ain Musa and Bir. ed-Doleh in Jordan.

Aharoni (1932) examined Nehring's material and two additional specimens from Ghor es-Safieh, Palestine and found them to con­ form to the type and description of Wagner’s gerbil from Sinai.

Ellerman (I9I48) recorded the species from the Dead Sea Basin,

Zahavi and Vahrman (195?) from Has en Nakura, Eilon, Haifa,

Wadi Ara, Jerusalem and several localities in the Negev and Wadi Areba in Palestine, Lehmann (1966a) from Aleppo, 173

Naharia, ?.'> Ion S. E. Danascus and Palmyra in Syria and Atallah

(1967a and c) from El-Jafr and tho Asraq Basin in Jordan.

Lewis, Lewis and Atallah (1967) reported on the possibility of the occurrence of the species in southern Lebanon but could not obtain any specimens.

Other records.- Aharoni (1930), Fetter (19^7) and Bodenheimer

(1958).

Specimens examined: lli7. PALESTINE: Bethlehem (2£ SA); Beit

Sahur (£ SA)j N. end of Dead Sea (11 SAj U AUB; 1 BZM)j

Jericho (3 BZM)$ Ibn Ubcid (1 SA)j k hm N.W. Beit Fajjar (2 SA).

JORDAN: 6 km S. Jerash (1 SA)j El-Jafr (3 SA)j Azraq ed Druz

(3 SA); 3-Jl km N. VJ. Azraq-Shishan (30 SA); Petra (19 AUB); 5 km S.Rasen Negeb (1 SA); 3U km N. Aqaba (8 SA); 5-10 km E.

Wadi Musa (3 SA); Moab (Holotype and one paratype of Dipodillus dasyuroides Nehring, BZM). SYRIA: Palmyra (Holotype and three paratypes of Q. d. palmyras Lehmann, AKM; 10 AUB); Aleppo

(5 AKM); 13.5 km S. E. Aleppo (2 AUB); 25 km S.E. Damascus

(Holotype and three paratypes of 0. d, leosollicitus Lehmann,

AKM).

Identification: Hairs on belly white and hairs on the back have long grey bases with a subterminal brovm band and black tip.

The black tip varies in length in different populations living on different soil types. Tail with a well developed terminal pencil. GLS less than 20 mm. Tympanic bullae not inflated, not projecting beyond the level of the occiput. Soles of hind feet usually less than 25 mm; soles naked. 17>4

Biology: .gner's gerbil ia the only member of the genus that penetrates tho Mediterranean Region, but even here it is restrict ed to the >uthem and wanner sections. They are common through­ out the St ppo and Desert Regions where they were collected in a variety -.f habitats including wadi systems, silt dunes, lime-- stone hamn .da, basalt desert, cultivated land and rock slides.

They have been noted to utilize burrows of either Psammomys

obeous or ''erionos libycus in the desert (Atallah, 1967a), but in the steppes and along cultivated land their burrows are usually constructed under rocks and behind stone walls.

Young, usually five, are born between late March and early

June in the desert and from late April to July in the steppes.

One specimen collected near Bethlehem, Palestine, however, was

found lactating in November,

Gerbillus nanus Blanford

1875, Gerbillus nanus Blanford, Ann, Mag, Nat, Hist,, l6(h):312

Type locality:' Gedrosia, W. of Gwadar, V/, Pakistan.

Common names: Baluchistan Gerbil,

Distribution: The Saharo-SinAian Desert Belt southwards to

Somalia in eastern Africa,

Only the following subspecies is recognized in tho

geographical area covered,

Gerbillus nanus arabium (Thomas)

1913. Dipodillus arabium Thomas, Ann. Mag. Nat. Hist., 2(9):6l,

1951, Gerbillus nanus arabium Thomas; Ellerman and Morrison-

Scott, Checklist, p. 633. 1#

Holotype: Jt Collection no, 10,3.12.1., an adult male collected

at Tebuk, "nudi Arabia on 3rd January 190? by D. Carruthera.

General Di bribution: Arabia and deserts of Palestine.

Allen (1915) and Bodenheimer (1935) referred specimens

from Aqaba j Jordan to Dioodillus quadrimaculatus (O, n'anus),

Ellerman (!.9b3) recorded specimens from 60 mi south of Beersheba, fetter (1957) from b'adi Araba and Zahavi and Wahrman (1957)

from near ocdom, Hatseva, Mahal Hemda, Yotvata and Mahal Titnna

in southern Palestine.

Other records.- Aharoni (1932), Bodenheimer (1958)» Potter

(1961b) and Harrison (1968c).

Specimens examined: None from the geographical area covered.

Two from Qaisumah, northeastern Saudi Arabia (SA),

Identification: Hairs on back with grey bases and Walnut

Taffy tip, belly white. Tail with a well developed terminal

pencil. GLS greater than 28 mm. Tympanic bullae inflated,

projecting appreciably beyond the occiput. Soles of hind feet

naked, usually over 25 mm in length.

Biology: Along the Eastern Mediterranean this gerbil is

restricted to tho Wadi Araba, where Zahavi and Wahrman (1957)

recorded dense populations on salt flats with burrows con­

structed beneath Nitraria retusa and similar shrubs. Lewis,

Lewis and Harrison (1965) found this gerbil living commensally with two species of jirds, Morionea crassus and M. libycus.

Lay (1967) recorded pregnant and lactating females in late

November to early January in Iran. No such dataare available

from Palestine 176

Gorblllus henleyi (do Winton)

1903. Dip'-dillus henleyi do V.'inton, Nov. Zool., 10:2814.

Typo locality: Zaghig, Wadi Natroun, Egypt.

Oommon narces: Pygray Gci'bil,

Distribution: Algeria, Egypt, Sinai, Palestine and Jordan.

Only twelve specimens have been taken from tho area of study, these wore provisionally assigned to the greyish sandy subspecies 0. h. mariae (Atallah and Harrison, 1967).

Gerbillus henleyi mariae (Bonhote)

1909. Dipodillus mariae Bonhote, Proc. Zool. Soc. Lond., p. 792.

1951. Gerbillus henleyi mariae Bonhote; Ellerman and Monison-

Scott, Checklist, p. 63I4.

Holotype: BM collection no. 9.7.1.14b• obtained on March 2£, 1903# at the Mokattam Hills, near Cairo, Egypt.

General distribution: Egypt, Sinai, Palestine and Jordan.

Harrison (I963f) recorded a single specimen about 10 km

E. Dimona, Negev. Atallah (196?a) recorded another specimen from

Paidhat edh Dhahikiya, south of the Asraq Basin at the northern border of Wadi Sirhan in Jordan. Ten additional specimens were collected at El-Jafr, Jordan (Atallah, 1967c). All these collec­ tions vero studied in some detail by Atallah and Harrison (1967).

Soecimens examined: 12. PALESTINE: 10 km E. Dimona (l HARR).

JORDAN: Faidhat edh Dhahikiya (1 SA); El-Jafr (10 SA).

Identification: This is the smallest Eastern Mediterranean gerbil,. GLS less than 22.£ mm. Soles of hind feet naked, less than 20 mm. Color identical to G. n. arabium. Terminal pencil 177 thin and t: irsely haired. Tympanic bullae inflated, projecting beyond tho level of the occiput.

Biology: tallah (1967c) obtained some of his specimens from dykes surx unding barley fields on an agricultural desert re­ search fat 1 at El-Jafr, Jordan. Burrow entrances were no larger tha 1 those occupied by lizards, Acanthodactylus spp., being not tore than 1-2 cm in diameter. Others were trapped at burrow entrances of Heriones crassus (also Harrison, 1963f),

M. libycn-' or Psammomys obesus«

Immatures were collected in May and June. One female collected in June held six embryos, another held four early embryos and was lactating at the time, and a third was lactating.

Gerbillus gerbillus (Olivier)

1800, Dipus gerbillus Oliver, Bull. Soc, Philom. Paris, 2:121.

Type locality: Giza Province, Egypt.

Common names: Lesser Egyptian Gerbil.

Distribution: Morocco eastwards across the Sahara t mi,

Palestine, and Jordan.

Pettor, Seydian and Mostaschfi (1957:113) end Potter (1968: ll) include Iraq and eastern Iran in the range of this species based on specimens of G, cheosmani Thomas, which they consider a synonym of G. gerbillus. This synonymy, however, has not yet been established.

Thomas (1919c) described Gerbillus bonhote! from northern

Sinai, considering it similar to G. andersoni except for a slightly 178

larger bulla. Ellerman (19U8:303) did nob find this character

consistent and placed bonhotei as a synonym of Q. gerbillus

andersoni 'To Winton, as did Elio man-’ To rris o n-S c o 11 (195l:63U)«

Atallah and Harrison (1967:317) recorded six specimens from

Jordan: "slightly larger than the nominate race...Tho auditory

bullae are inflated ventrally as in Gerbillus cheosmani, but

still cannot be confused with latter, where the bullae are

inflated both ventrally and posteriorly (mastoid region), pro­

jecting considerably beyond the occipital condyle. This des­

cription coincides with that of Gerbillus gerbillus andersoni

as in Sotzer (1958 V*. Due to lack of comparative material

at the time, however, Atallah and Harrison (op. cit} did not

assign their specimens to any particular subspecies. Fetter

(1968:12) listed andersoni as a synonym of G. pyramidun and

noted (p. 10) that bonhotei could bo a synonym of Gerbillus

allenbyi. The distribution of G. gerbillus in relation to G.

allenbyi in Palestine has been studied in detail by Zahavi and

Wahrman (1957)« These studies conclude that G. allenbyi is res­

tricted to the coastal dune area while G. gerbillus extends from north Africa into Sinai and the Negev to Wadi Araba. The two

species, slightly different morphologically, were found to differ

in their diploid chromosome number as vrell as in the sex deter­ mination mechanism, G. gerbillus with 2Na )-i3, XYT chromosomes in tho male and G, allenbyi with 2N4;0, XY chromosomes in tho male. 17?

I hiv ) examined the holotypes of Gerbillus gerbillus, G, andersoni; G, bonhotei and G, cheesmani (EM and BZM) and tend to refer specimens from Wadi Araba and the adjacent highlands to the fom bonhr .ei regarded as a subspecies of G. gerbillus.

Gerbillus gerbillus bonhotei Thomas

1919. GoiM-llus bonhotei Thomas, Ann. Mag. Nat. Hist., 3(9):£60.

Kolotypej EM Collection no. 19.5.7.5., an adult female collected at Khabra ,ibu Guzour, S. E. of El-Arish, northern Sinai, on

Dec. 1918 by Cap. S. S, Flower.

General distribution: Sinai and Wadi, Araba in Palestine and Jordan.

Zahavi and Wahrman (195?) recorded specimens from the

Sekhor dunes, south of Beersheba, £ km S, Halutsa, 1 km S,

Bir Mashabim, 3 km N. W. Nitsana and 1 km E. Yotvata in southern Palestine and Atallah and Harrison (1967) from a largo wadi, 77 km N, Aqaba on the Aqaba-Maan desert highway in Jordan.

Other records.- Bodenheimer (1935 end 1958).

Specimens examined: 26 JORDAN: 77 km N. of Aqaba on tho

Aqaba-Mann desert highway, 22 km N. of Al~Quweira (5 SA)j i

S. Ras en Negeb (1 SA). Also 20 specimens of G. g. gerbillus from Egypt at FMNH.

Identification: Color of back Inca Gold, belly white. Tail with a thin and inconspicuous terminal pencil. GLS 25-28 mm. Tympanic bullae slightly inflated and not projecting beyond the occipital condyle. Hind feet usually over 30 mm in lengthj soles covered with hair 10O

Biology: hia form is more or less restricted to shifting or stable sat' 1 formations throughout Wadi Araba and north-western

NegeV (Zal' xl and Wahrman, 195?) and the Eastern Uplands bordering -radi Araba (Atallah and Harrison, 196?).

Near J.-Quveira they were found in a large wadi, sparsely vegetated nd covered with blown sand. Burrow entrances, 3 to

1: cm in di imeter, were usually located beneath shrubs. At this locality Koriones crassus was found to share the same habitat, while at higher elevations (near Rasen Negeb) they were found along with rock dwellers such as Gerbillus dasyurus and Heriones tristrami. Nothing is known of the habits and reproductive biology of this species.

Gerbillus allenbyi Thomas

1918. Gerbillus allenbyi Thoms, Ann. Mag. Nat. Hist., 2(9 ):m6.

Holotype: BM collection no. lh.9.29.5, a yoxing adult male collected at Rohobot, near Jaffa, Palestine by I, Aharoni.

Common names: Allenbyi Gerbil.

General distribution: Central coastal plain of Palestine

(Orni and Efrat, 1966:33)*

Aharoni (1932) recorded specimens from tho type locality and Jaffa. Zahavi and Wahrman (1997) obtained specimens from

Ma’agan Mikhael, Qeisariya, Hadasin, S. VJ. Herzliya, Eenei

Beraq, Holon, Rishon le Zion, Palmahim, Nitsanlm and near

Ashqelon.

Other records.-Thomas. (1918), Bodenheimer (1939 and

1990), Ellerman (19U8) and Potter (1996a and 1997). 131

Specimens .gained: U. PALESTINE: Rohoboth (1 BZM)j Jaffa

1 (BZTl)j T L Aviv (1 BZM); no exact locality, labelled Ras

Fashkha by ustake (See Aharoni, 1932) (l BZM)«

Idcntific;. ..on: Color of back gray, belly white. GLS less than 28.£ .i. Tympanic ballao not projecting beyond tho occipital I’ondylo. Hind feet usually less than 28 tm in length; scles hairy. See Potter (1956iond 1957) for more detailed comparison with other Palestinian gerbils.

Biology: Zahavi and Wahrman (1957:35U) reported this endemic species as "veiy common along tho coastal plain in all kinds of sandy biotopes: shifting dunes, stable sands, sandy soils and even sand-stone hills'*. Nothing is known of the reproductive biology of this species,

Gerbillus pyramldum I. Geoffroy

1825. Gerbillus pyramldum I. Ceoffroy, Diet, Class, II. N.,

7:321.

'I

Typo locality: Giza Province, Egypt,

Common names: Greater Egyptiari Gerbil.

Distribution: Morocco acres3 tho Sahara to Egypt, Sinai and sonthern Palestine.

Thomas (1919c) described Gerbillus fluwori from northern

Sinai. Aharoni (1932) reduced this form to subspecific status under G. pyramldum and Ellerman (I9ij8) treated it as a synonym of 0, pyramldum tarabuli Thomas known from Libya and Egypt. 182

Zahavi and i-'ahrr.mn (19^7) and Wahrman and Zahavi (1958)

examined cl.-omosome configurations in Palestinian and North

A.frican poj ilations of 0, nyramidum and defined three allopatric

sibling gn, os vrithin tho pyramidum complex, x-rilthout assigning

natr.os, as f -llows:

1. C<.:->stan Plain Group: 2N “52, This form has been

recorded fiom tho Judean coast, south of River Yargun only,

however, it is believed that it extends farther south to Gaza,

It is abundmt on coastal sand dunes,

2. Negev Group: 2N “ 66. This form is abundant in

northern Negev (near Beersheba, Nitsana and Tureibe) and Sinai,

These wore found to inhabit loose sand dunes as well as compact

sand deposits.

3. Algeria Group: 2N “ 1|0, The distribution of this form

in North Africa is still unestablished. This form will eventually

retain tho original name, the proceeding two forms need to bo

named.

Identification: Color of back light brown, belly white,

GLS larger than all Eastern Mediterranean gerbils, usually over

31 mm,in length. Hind foot usually over 32 mm in length; soles hairy.

Biology: Both groups in Palestine aro restricted to either

shifting sand dunes or stable sand formations (Zahavi and

Wahrman, 1957:36h). The gestation period is 25 days (Petter,

1961b). 133

Genua Sekeetamys EUeman

15h7. Soke.-tenys Ellerman, Proc. Zool. Soc. Lond., 117:271.

Tyne specie. : Gerbillus calurus Thomas.

Ellen.’ a (l^Ij?) erected Sekeetamys as a new subgenus of the genus Ikriones cand referred Gerbillus caluims to it.

VTassif (19!?ljb) reviewed tho systematic history of G. calurus ard maintained that this gerbil belongs to tho subgonus

Dipodillus of the genus Gerbillus. Petter (1956b) elevated

Sokootamys to a full generic level on morphological grounds.

Additional evidence vas later supplied from cytological studies

(Wahrman and Zahavi, 1955J Zahavi and Vahrman, 1957).

Sekeetamys calurus (Thomas)

1892. Gerbillus calurus Thomas, Ann. Hag. Nat. Hist., 9(6):76.

1951. Heriones calurus Thomasj Ellerman and Morrison-Scott,

Checklist, p, 638.

1956. Sekeetamys calurus Thomas; Pettor, Mammalia j 20(ii):Ii20.

Distribution: Sinai, eastern Egypt, southorn Palestine and Jordan.

Only tho nominato form is pertinent to this study,

Sokcebamys calurus calurus (Thomas)

Synonymy under species,

Holotype: An adult female in BM collection, preserved in alcohol. Not examined.

General distribution: Sinai, southern Palestine (Negev and

Judean Desorb) and southorn Jordan (Eastern Uplands).

Zahavi and Wahrman (1957) recorded specimens from Metsada, vicinity of Rosh Zahar, Nahal Boqeq, Makhtesh Ramon, Har Aref,

Sedc Abraham, Nahal Timna, Bir Ora and Nahal Sholomo in tho Central Hi" and Southern Hills of the Negev and tho Judean

Desert, Pa' ratine. Atallah and Harrison (196?) obtained a single spO' Vren fror.i the Eastern Uplands in southern Jordan.

Other 'ecords.- Bodenheimer (19^3).

Specimens examined: 18 JORDAN: 3h Ion N. of Aqaba (l SA).

Also 17 specimens from the vicinity of St. Catherine's

Monastery, Sinai (USNM and FMNH).

Genus Meriones Illiger l8ll. Meriones Illiger, Prodx*. Syste. Hamm., p. 82.

Since tho publication of Chaworth-Kusters and Elleman's

(I9h7) revision of the genus Heriones many systematic changes were proposed and substantiated (Petter, 1956a; Matthey, 1957J

Zahavi and Vahrman, 1957} and others), most of these are summarized in Nadler and Lay (1967:286), who recognized fourteen species of which only five are reported from tho Eastern Mediterranean Region.

Following is a KEY to these five species;

1. Tympanic bullae highly inflated extending beyond the occipital

condyle; suprameAtal triangle idLth the posterior processes

separate (Fig. 7B)...... M. crassus

" Tympanic bullae smaller, not extending beyond the occipital

condyle; suprameatal triangle with the posterior processes

joined (Fig. 7A and C)...... 2

2. Zygomatic arches almost touching the auditory meatus; supra­

meatal triangle large and bullae inflated (Fig. 7A); large,

GLS usually over 38 Km*,.,...... 3

Zygomatic arches widely separated from the auditory meatus; 18$

supr?rit .tal triangle small and inconspicuous, bullae moderately

inflate i (Fig. 7C)j small, GlS usually loss than 30 mm...... It

3. Hind Cj ,v3 darkj tail reddish, with tip black tufted,...,......

...... M. libycus

Hind claws palej tail cream colored, with black tuft extending

at leafb l/2 the length of the dorsal surface of tail......

...... M. sacrament!

It. Soles of hind feet completely haired; tail usually equal to

HB,...... *...... H. vinogradovl

Soles of hind feet naked at heel; tail longer than HB......

...... M. tristrami

Heriones tristrami Thomas

1892. Heriones tristrami Thomas, Ann. Hag. Nat. Hist., 9(6):ll|8.

1903. Heriones blackleri Thomas, Ann. Mag. Nat. Hist,, 12(7):l09.

Smyrna, Turkey. (Zahavi and Wahrman, 191>7).

Type locality: Dead Sea, Palestine.

Common names: Tristram's Jird.

Distribution: Northern Sinai, Palestine, Jordan, Syria, northern

Iraq, northwestern Iran, Turkey and Caucasus.

Aharoni (1932) recognized three subspecies in this region: bodenheimer! Aharoni from the Ansariyah Mountains, a small grey colored form, kariatinl Aharoni from Karyatein, Syria, a large light colored form with an inflated bulla-,' and the nominato form from Palestine and Jordan, .a medium size, sand colored form. Petter

(l96lb:l|8) recognized two subspecies: M. t, tristrami from Syria southwards to Sinai and M. t. blackleri Thomas penetrating slightly into northern Syria but more or less restricted to Turkey and 186

Kill'd is tan. Petter (op. cit.) does not, however, discuss the position o.; bodenheimori or kariatinl.

H. tr trami blackleri, with its white tipped tail is readily di inguished from other subspecies of tristrami in this geographic; area. This form has been reported from several localities In southern Turkey (Lehmann, 1966a) and undoubtedly it must pen :trate into Syria. However, as yet there has' been no specimen i to substantiate this statement and this fom is not included in this paper.

M. tristrami bodenheimeri, known only from two specimens, appears to bo only a color variant of tho nominate fom. Spec­ imens from Jordan, Syria, Palestine and Lebanon, however, show a striking variation in coloration. As an example, the following are color descriptions of three specimens taken on different soil types and localities, the first two localities are not more than 5 km apart from each other, in Jordan.

Locality Color of back*

Basalt Desert, Azraq ed Druz Pecan Brovm Limestone rolling hills, near Tawny Airman

Oasis, Azraq-Shishan Grey 13

*Color on the flanks usually lighter, belly white.

Aharoni (loc. cit.) also separated bodenheimeri from the nominato race on the basis of greatest skull lengthj 36.5 - 37*7 wm in the nominate fom and 33.0 - 35.0 mm in bodenheimeri. Lewis, Lewis and

Atallah (1967: Table VII), however, show that the greatest length of skull of specimens taken at five localities in Lebanon and Syria 187 varies botvoen 32.3 - 39.U rom. M. tristrami bodenlieimeri vrill thus bo treated is a junior synonym of M. t, tristrami.

M. tri''trami kariatinl, however, is considered valid. Thus, two subspecies are recognized in the Eastoi’n Mediterranean, as follows:

Heriones tristrami-tristrami Thomas

Synonymy under species, also tho following:

1932. Meriones tamarlcinus bodenheimeri Aharoni, Z. Saugetierk., 7.197

. Kafrun, Syria. NEW STNONYMY.

Holotype: BM Collection no. 61i.8.17.35., an adult male in alcohol collected near the Dead Sea, Palestine, by Caron H. B. Tristram.

General distribution: Northern Sinai, 1’alestine, Jordan, Syria and Lebanon.

Thomas (1892b) reported this form from the Dead Sea and Mt.

Carmel in Palestine, Allen (1915) from Shobok and Tafila in

Jordan, and Bir ed Doleh in Syria. Aharoni (1932) from Jaffa,

Beersheba and Jesreel Valley in Palestine and Kafrun, Syria;

Misonne (1957) from near Tel el Abiad, Syria; Zahavi and Wahrman

(1957) from thirty-six localities in northern Palestine; Lehmann

(1966a) from Amman, Jordan; Atallah (1967a) from Azraq ed Druz and Azraq-Shishan; and Lewis, Lewis and Atallah (1967) from

Beirut, Dahr el Baidar, Ain Zhalta and AUB Farm, Bekaa, in

Lebanon, and Aleppo, Syria.

Other records.- The two specimens on which Thomas (1892b) based his description vrorc identified by Tristram (l88Ii) as

Gerbillus taoniurus and Psammomys tamarlcinus« Nehring (1901), 103

Bodenheimoi (1920, 1935 and 1950), Bate (I9l45)> Ellerman (19h0) and Petter (1957 and 196lb).

Specimena c amined: 153 LEBANON: AUB Farm, Bekaa Valley

(35 AUB)j B,irut (6 AUB)j Dahr el Baidar (1 AUB; 1 SA)j Ain

Zhalta (1 iv’3); 3 km E. of Rochaya (l SA). PALESTINE: Bethlehem

(12 SA); BoLt-Sahur (3 SA); Dier Ibn Ubaid (1 SA); Ekron

(2 BZM); Rohoboth (5 BZM); Tabgha, Tiberias (2 BZM); Jerzeel

Valley (2 BZM). SYRIA: Aleppo (l BZM; 5 AUB); Kafrun (2 BZM).

JORDAN: £-10 km S. of Ras en Negeb (lh SA); 2 km N. Azraq-Shishan

(1 SA); Azraq-Shishan village (l SA); Azraq ed Druz (1 SA);

Schniller Boarding School near Amman (2? SA). Also 3it specimens from Turkey (Tosya, Inevi, Eregli, Tarsus and Talas) at BZM and

FMNH.

Identification: Soles of hind feet naked at heels, HF 31-30 mm.

Tympanic bullae not inflated; suprameatal triangle small and inconspicuous. Claws whitish. Terminal tuft on tail very short.

GLS usually less than 30 mm.

Biology: Tristram's jird is common throughout the Steppe Region at altitudes below 1,300 m in tho Mediterranean Region. Burrows aro usually constructed on tho side of hillocks and mounds of earth rather than on level ground. Each burrow system has at least two entrances and a nest, about 20 an in diameter, located at the terminus of one of the passages. Nests excavated in north­ ern Iraq are figured in Petter (1961b:Fig. 38). 189 Irrcmtv ;os were collected in June, July and August* Two

pregnant females collected in July had 8 and 6 embryos respectively.

Gestation pnriod 2h days. The baculum has been illustrated by

Argyropulo (1939).

This primarily nocturnal species feeds on the surface but

occasionally food is dragged to their burrows. They have, however, never been observed to build large stores of food in their burrow system as other species of jirds do (Lewis, Lewis and Atallah, 1967).

Food is essentially grain and other seeds as well as green and dry

stems and leaves,

Bodenheimer (1935, 19h9 and 1959) maintained that M. tristrami showed synchronized fluctuations with vole outbreaks. Initial observations in Lebanon (American University of Beirut Farm, Bekaa), however, do not support Bodenheimeri observations. At peak populations, Gunther's voles were found to extend their range from cultivated fields to adjacent hills forcing a reduction in H, tristrami populations inhabiting these hills,

Meriones tristrami kariatinl Aharoni

1932, Meriones tamarlcinus kariatinl Aharoni, Z, Saugetierk,, 7:197.

1951. Meriones blackleri kariatinl Aharoni, 1932J Ellerman and

Morrison-Scott, Checklist, p. 6ljl,

Holotype: BZM collection no. l}2309, an adult female collected at

Karyatein, Syria by I, Aharoni on 7-9 February, 1930.

General distribution: Known only from Karyatein and Palmyra, Syria.

Lewis, Lewis and Atallah (1967) recorded this subspecies from

Palmyra, Syria. 190

Other records,- Zahavi and Wahrman (1957073).

Specimens xamined: 12.SYRIA: Palmyra (12 AUB),

Identified' ‘.on: Resembles the nominato form, except for their

light sanu.; color and slightly more inflated bullae. Terminal

tuft on ta l less conspicuous than the nominato form.

Biology: ihis form is restricted to extreme arid conditions

in the northern parts of the Syrian Desert. Specimens from

Palmyra were collected on loose sand accumulated at the bases

of shrubs and dato palms.

Meriones vinogradovi Heptner

1931. Meriones vinogradovi Heptnor, Zool. Anz., 9b:122.-

Type locality: Azerbaijan, Iran

Common names: Vinogradov’s Jird,

Distribution: Northwestern Iran, Transcaucasus, northeastern Turkey

and northern Syria (vide Misonne, 1957 and 1959)•

M. vinogradovi is only provisionally listed here as a member of

the Eastern Mediterranean fauna. As no specimens belonging to this

species from the geographical area covered were foundj additional collecting in northern Syria is warranted for confirmation of

Misonne’s (loc.cit.) records.

Meriones libycus Lichtenstein

1823, Meriones libycus Lichtenstein, Verz. Doubl, Mus. Berlin, p, 5*

Typo locality: Near Alexandria, Egypt,

Common names: Libyan Jird.

Distribution: Algeria, Libya, Egypt, Palestine, Jordan, Syria,

Iraq, Saudi Arabia, Iran, Afghanistan, Russian and Chinese

Turkestan. Its existence in southorn Palestine and Sinai is not 191 r yet establi ;ad,

.Aharon (1932) recorded Iferionos libycus aacramonti Thomas and cry' .ourus legori Aharoni from many localities south of

Jaffa, Pal-, vino, ELlerman and Morrison-Scott (19^1:6U6) referred tho form 1c ari to M, libycus« However, Zahavi and Wahrman (1957:

362) exaraii.i d some ofAharini's material and placed legeri in synonymy in h M. saeramenti, which was given specific status on the basis c its characteristic chromosome number of U6j hh in M, libycus anii 60 in M, crassus, tho two species with which it has been associated.

Only the following subspecies is recognized along the Eastern

Mediterranean* Meriones libycus syrius Thomas

1919. Meriones syrius Thomas, Ann, Mag, Nat, Hist,, 3:268,

1951• Meriones libycus syrius Thomasj Ellerman and Morrison-

Scott, Checklist, p, 6)45.

Holotype: BM collection no. 5,7*2.2,, an adult male from Karyatein,

Syria, collected by D. Carruthers on March 3, 1905*

Gonex’al distribution: Syria, Jordan, Iraq, Saudi Arabia, and possibly Palestine,

Thomas (1919b) recorded syrius from 320 km E. of the

Bead Sea, northern Saudi Arabia and Aharoni (1932) from Dier el

Zor and Karyatein in Syria. Zahavi and Wahrman (1957:362) referred specimens from Zahr ol Rubin (corrected to Nahal Rubin),

Palestine, previously assigned to this subspeeies (Aharoni, 1932}

Chaworth-Musters and Ellerman, 19b7} Ellerman, 19b8) to M, saeramenti,

and maintained that the species is not a member of tho Palestinian fauna, Atallah (1967a and c), however, found M. 1. 3yrlus abundant 192

throughout ./ns Azraq Basin and El-Jafr in Eastern Jordan.

Other ,’ecords .-Misonne (195?) and Fetter (1957).

Soecinens caminod: 56 JORDAN: 5 km N. Al-Hasa (1 SA)j

12 km N. A’ Ilasa (1 SA)j km E.N.E. Qasr Ariira (7 SA)j 24: km

S. Shaumar Agric. Res, St. (11 SA)j Azraq-Shishan (l SA)j

El-Jafr (iJi SA)j U5 to N. Maan (7 SA). SYRIA: Dior el Zor

(1 BZM)j Ka./atoin (5 BZM).

IdentifiGa;Ion: Soles of hind feet naked at heels, HE 37~^0 ran.

Tympanic h.Ilae inflated ventrally, mastoid region not projecting

beyond the occiput, supramcatal triangle large (Fig. ?A).

Clara dark. Tail reddish irilth a rail developed terminal tuft.

GLS usually over 39 mm.

Biology: Libyan jirds are common throughout the Syrian Desert

and the more southern parts of the Eastern Uplands. They are

locally abundant forming largo colonies in uadis or along high-

uaysuhora water runoff allows a greater amount of vegetation than

surrounding land. The same habitat is usually sha d by at

least two of the following: Keriones crassus, Gow. llus dasyttnis,

G, henleyl, G, nanus and Psammomys obesus. Gerbillus spp, often

construct their bmrora as an offshoot from the burrows of these

jirds, both utilising the same entrances,

Libyan jirds construct elaborate burrow systems with numerous

entrances.and storage compartments. These burrows are usually located

beneath shrubs. The entrances are narked on the outside by a pile

of excavated soil and dry plants. Activity has been recorded at 193 every hour t1 the day or night, with peaks in the early noming and late ai ‘.vrnoon hours. They are easily identified in the field by th ir reddish tails which are normally carried above the body vh a they run,

Lewis, Lewis and Harrison (1965) noted that Donkey Melon, i Citrullus o vl.ocynthls (L.), is the preferred food of this species in northern Saudi Arabia, Observations in eastern Jordan support this stater'.:nit. Donkey Melon,- , however, can only be found in late winter and early spring and for the rest of the year these jirds feed on stems, leaves and seeds of almost every species of desert shrub within reach,

Atallah (1967c) observed that the breeding season ends between late May and mid-Juno,

Merlones crassus Sundevall

181(2, Meriones crassus Sundevall, K. sv. Vetensk, Akad.

Kandl,, p. 233♦

Type locality: Ain Musa, Sinai,

Common names: Sundevall's Jird,

Distribution: The Saharo-SIndian Desert Belt from the Sahara to northern India,

Only the nominate form is pertinent to this study.

Meriones crassus crassus Sundevall

Synonymy under species.

Type material: Mot examined.

General distribution: Northeastern Egypt, Sinai, Palestino, northern Saudi Arabia, southern Syria and southeastern Iraq.

The species is listed as a member of the Palestinian fauna by Allen (1915),I. Aharonl (1930), B. Aharoni (1932), Eodenheimer 19U

(193^ and 1950), ;md Potter (1957). Zahavi and Wahman (1957) recorded C[. .'ciciena from aovoral localities throughout tha Negev, south of a Ine between Kamshit-Revivim. Atallah (1967a and c) reported it .from the Azraq Basin and EL-Jafr in eastern Jordan.

Specimens i .mined: 26 JORDAN: 22 km N. Al-Quweira (1 SA)j

Azraq-Shish m (1 SA)j 2 km N. W. Azraq-Shishan (2 SA); 3 km

E.N.E. Qasr Arara (U SA); 2 km S. Shaumari Agric. Res. St.(5 SA);

Ii5 km W. El Jafr (2 SA); El-Jafr (1 SA). Also 10 specimens from the vleinity of St. Catherine’s Monastery, Sinai, at the FMNH.

Identification: Soles of hind feet naked at heel, HF usually 3Z~

36 mm. Tympanic bullae greatly inflated projecting considerably beyond the occiput, suprameatal triangle very large (Fig. ?B).

Claws pinkish. Tail shorter than HB with a very thin terminal black or brown tuft, GLS usually less than 39 mm.

Biology: Sundevall13 jirds occupy the same habitat as Meriones libycus as well as the bare hammada and sandy areas. On the hammada their burrows could be seen from long distance, are their, mounds of red sand provide a sharp contrast wiJ; a black flint of the hammada. '

This species is strictly nocturnal and never forms largo colonies as Meriones libycus or Tsammomys obesus. Burrow systems are elaborate, usually with many storage compartments and three or four entrances located at the base of a shrub. Tunnels are usually 3-U cm in diameter. Lewis, Lewis and Harrison (1965:73) describe the nest of this species as follows: "Usually it is situated somewhere near the point of maximum depth of the 19$ system and <.s either in a blind chamber pff the main tunnel or at the conflm .co of tvro or more tunnels. The nests, unlike those of most of he other species, are frequently composed of rags and paper, oft', i lacking vegetable material completely." Diagrams of burrow t.,/stems of this species in Iran and Egypt are shown in i Fetter (19$J and 1961b).

Irnmatr.ros were collected in April.

Meriones sacramenti Thomas

1922. Meriones sacramenti Thomas, Ann, Mag. Mat. Hist., 10(9)

1932. Meriones erythrourus legcrl Aharoni, Z, Saugetierk., 75202*

Wadi el Abiad, southwest of Beersheba, Palestino. (Zahavi

and Wahrman, 1957:362)

Holotypa: BM collection no. 22,10.14.1., an adult male collected from 10 mi south of Beersheba, Palestine, on July 17, 1922 by

P. A, Buxton.

Common namest Palestine Jird,

Aharoni (1932) and EUerman (19^1) reduced Meriones sacramenti to subspecies of M. libycus. Chaworth-Mustors and EUerman (1914?) transferred sacramonti to the M, crassus group. Fetter (1957) and

Zahavi and Wahrman (195?) independently retained M. sacramenti as a full species on morphological and cytological grounds.

General distribution: Known only from the coastal plain south of the

River Yargon and the northern Negev in Palestine,

Aharoni (1932) recorded specimens from Wadi el Abiad and Jaffa, while Zahavi and Wahrman (1957) recorded it from Holon, Rishon le Zion, Palmahim, Nir Gallim, 1 km south of Bir mashabim and

Nahal Lavan ( « Wadi el Abiad) in Palestine* 196

Other ocordoThomaa (l9?.2b), Bcdenheimor (1935 and 1958),

Chavrorth-Mu .itera and Ellorman (I9h7), Elleman (I9h8) and Patter

(1957 and 3 6lb). Misonna (1957) reported M. sacranrenti from northern Sj .-ia« This record, however, has been shown to be erroneous ( 'ahavi and Wahman, 1957:362) inasmuch as sacramenti v is endemic to southern Talcstino.

Specimens examined: 9 PALESTINE: Wadi el Abiad, southwest of Beersheba (8 BZM); Rishcn le Zion (1 SA).

Identification: Soles of hind feed naked at heel, HF 35-Ul ran.

Tympanic bullao as in M. libycus. Claws pinkish. Tail shorter than HB with a black tuft extending from its tip to at least

1/2 length of tail. GLS usually over hO mm.

Biology: This endemic species is confined to the coastal dunes and semi-desert areas of southern Palestine. Nothing is known of the habits and reproductive biology of this species.

Genus Psammomys Cretzschmar

1828. Psammomys Crotsschmar, Ruppell Atlas, p, £6.

Type species: Psammomys obesus Cretzschmar.

Tvro species are currently recognised in this genus (Ranck, 1968), but only the following is pertinent to this study.

Psammomys obesus Cretzschmar

1823, Psammomys obesus Cretzschmar, Ruppell Atlas, p, 58, pi. 22.

Type locality: Near Alexandria, Egypt.

Common names: Fat Sand Rat, Sand Rat.

Distribution: Throughout the Sahara eastwards to Sinai, Palestine,

Jordan and Saudi Arabia, 197

Thorn.it; (1902) described Psiranomys terraosanctae from four specimens a .preciably larger than P. obesus from the Dead Sea Basin.

Aharoni (19 2:193) reduced tcrraesanctae to a subspecific level under P. oi jus, recognizing tvo subspecies in southern Palestine, separated b/ 015.

Boderihjimer (1958:18^) regarded both subspecies as synonymous.

I have compared specimens from Jordan, Syria and Palestine with others from Sinai, the Eastern Desert Governorate and the Western

Desert Gove onorate in Egypt and found them identical in every respect (Appendix I), Greatest length of skull varies tremendously within populations, and it appears that Thomas (1902) based his des­ cription on very old individuals that were compared with young adults of the nominate form. Thomas (og. cit.) and Aharoni (1932) regarded terraesanctae as the largest Psammomys, while Setzer (1963s

5$) maintained that "it is the smallest of the subspecies of sand rats occurring in Egypt". P, o. terraesanctae will be treated hero as a junior synonym of the nominate form as suggested by Bodenheimor

(op. cit.).

Psammomys obesus obesus Cretzschmar

Original citation under species.

1902. Psammomys terrae-sanctae Thomas, Ann. Mag. Nat. Hist., 9(7):363.

Dead Sea, Palestine, (Eodenheimer, 19!?8).

Type' material: Not examined.

General distribution: Algeria, Tunis, Libya, Egypt, Sinai, Saudi

Arabia, Palestine, Jordan and Syria.

Allen (1915) recorded a single specimen from Ain Abu Heran, north of Aqaba, Jordan. Eodenheimer (1920, 1935 and 1958) reported 198 it as conmon noar the Dead Sea, southern Judea, the Negev and the

Syrian Desert vrhile Aharoni (1932) recorded it from Ain Gedi and

Sedom near Mie Dead Sea, Palestine and Karyatein, Syria. Zahavi and Wahrman (1957) collected specimens at Wadi Qilt, Kallia, Ain

Gedi, Sedom. Tureibe Plain, Beerot Oded, Nahal Mor, Nahal Nafha and Yotvata in Palestine and Atallah (1967a and c) from the Azraq

Basin and El-Jafr in Jordan,

Other records,- Tristram (1866a and I88I1), Elleniian (19^8) and better (1957 and 196lb).

Specimens examined: 63. JORDAN: 5-10 km N, Jiza (2 SA)j 62 km

N. Maan (1 SA); 26 km N. Maan (1 SA); Maan (1 SA); 10 km N. Ai-

Has a (1 SA); 2-b km S. Shaiunari Agric, Res. St. (7 SA); El-Jafr

(h SA); h$ km N. Maan (2 SA). PALESTINE: Near Qumran, N, end of Dead Sea (3 SA; 12 AUB); Ain Gedi (l BZM). SYRIA: Karyatein

(7 BZM), Also 21 specimens from Sinai (El-Arish) and Egypt

(Morsa Matruh, El-Alamein, Buheira, Port Said and Wadi El-P.okham) at KTNH.

Identification: Upper incisors plain (grooved in all other representatives of the subfamily in this region). Tail short, usually less than 75# of HB. HF usually 37-hl mm, claws black.

GLS over I4O ran in adults. Soles of hind feet naked at heel.

Biology: Fat sand rats are more or less strictly diurnal.

Sizable colonies are common in wadi systems, alluvial fans and sandy areas. The following quotation from my field notes regarding the habitat of this species may be worthy of mention: 199

"December 2'} V)6$: - Psamnioinya is very common all along tho highway bet-, een Amman and Maan and to a lesser extent between

Maan and Ao ba (in Jordan). Populations are concentrated in large colon :s in sandy areas,however, atypically a large population .'.s. found on a rocl

0. henleyi i nd occasionally M. crassus.

Members of this species can bo seen at any time of the day sitting at the entrance of their burrows sun-bathing or feeding.

Other habits noted are the sole use of one hand in grasping food, tho wiping of food before eating and tho combing of the fur with their fore feet (Zahavi and Wahrman, 1957:363).

Burrow systems arc more or less circular with entrances in all directions of the compass and many connecting tunnels and storage areas (See Fetter, 1952 for details). Immaturos were collected in May and June, ’while a pregnant female collected in mid January held six embryos.

Family SPALACIDAE Cray, 1821

Mole Rats

This monotypic family is restricted to southeastern Europe, southwestern USSR, Turkey, Iraq, Eastern Mediterranean Region,

Egypt and Libya. 200

Genus Spalax Guldenstaedb

1770. 5pal x Guldenstaedb, IIov. Com, Acad, Imp, Sci, Petrop,, ill, I:hlO

Type specie : Spalax mlcronh thalmus Guldenstaedt

Of th . three species recognized by Slleman and Morrison-

Scott (195i / only tho following single species has been reported from the K tom Mediterranean Region,

Spalax ehrenbergi Nehring

1898, Spal. x ehrenbergi Nehring, S, B. Ges, Nat, Fr. Berlin, p.

178, pi. 2.

Type locali ty: Jaffa, Palestine.,

Common nair.es: Palestino Mole Rat,

Systematic remarks: Recent examination of chromosome numbers by

Revo (1969) and Wahrman, Goitein and Kevo (1969) separated four allo- patric populations from chromosomal and mating data in Palestine and adjacent lands as follows:

1, 2N“52 from the Upper Galilee Hills,

2, 21,^4 from tho Gulan Heights and Mt, Hermon.

3, 2N=53 from central Palestine, a triangle between Tiberias, Haifa .and Jaffa,

b, 2!M=60 from Samaria, Judea and northern Negev,

If those foui* chromosomal populations were to bo regarded as sibling species as suggested by Nevo (on, cit.:b8b), then s« ehrenbergi will bo applied to the central Palestinian population while the remaining three populations need to be named. However, all four populations, must be treated as a single taxonomic unit until more complete data are reported by tho foregoing investigators or additional 201

studies arc made. On morphological grounds North African populations

(Egypt and .ibya) have been referred to S. ehrenbergi aegyptiacus

Nehring, uh i.le all Eastern Ilediterranean populations were referred to the nominal 5 form (EUerman and Korrison-Scott, 1951)«

Gener.al dio .x’ibution: Syria, Lebanon, Palestine, Jordan and Iraq

(vide Harrison, 1956a).

Records of this species are numei'ous. The majority of these records, however, are based on sightings of the characteristic mounds rattier than actual collection of specimens, Tristram (1866a and 1881}), Eodenheimer (1920, 1935 and 1958) and Aharoni (1930) report the species as common throughout cultivated land in Palestine.

Bate (19U5) recorded skull fragments from owl pellets collected on

Kt. Lebanon and Dor (I9h7) from Palestine. EUerman (I9h8) recorded specimens from several localities in Lebanon, Syria, Jordan and

Palestine. See also Nevo (1961) for distribution and habitat proferance.

Other records.- Pocock (1917),Misonne (1957), Mountfort (1965),

Lehmann (1966a) and Lewis, Lewis and Atallah (1967).

.Specimens examined: 30. LEBANON: Beirut (3 USNM; 1 SAj 1 AUB)j

Saraain el Faouqa (3 SA); Nabi Chite (l SA); Laklouk (1 REL);

AUB farm, Bekaa (2 AUB). PALESTINE: Bethlehem (1 SA); Jerusalem

(Ij BZM); Tabgha, near Tiberias (3 BZM); Smuas (l BZM); Jaffa

(7 BZM); Safyeh (1 BZM)* JORDAN: 60 mi east of Dead Sea (1 AUB).

Identification: Body elongate, taU and external ear (pinna) absent. GLS usually less than hi mm. Fossorial. 202

Biology: I is fossorial species is comon throughout tho

Kediterranc a and Steppe Regions, It is limited in its distri­ bution to I' a south and east by the extreme deserts: tho

Judean Dcsc. t, the Negev and tho Syrian Desert. Nevo (I96l) observed th .b the southern limit of distribution of mole rats in Palestin; coincides with the 100 mm isohyot, which he believed wr ; responsible for the extreme scarcity of bulb and com plants, on which mole rats are solely dependent for food.

This also c ,pears to be the case in southern Jordan.

Palestine mole rats were observed at altitudes from 300 m below sea level to about 2,000 m above sea level.

Tho habits and reproductive biology of this species have been studied by Nevo (1961 and 1969) in Palestine and Lewis,

Lewis and Atallah (196?) in Lebanon. Parturition occurs from early February at low altitudes to early April at higher alti­ tudes, An average of 3-U young are reported by Nevo (1961) and

2-3 by Lewis, Lewis and Atallah (op. cit.). Prior to parturi-- tion elaborate breeding mounds are built. These are elevated some 19-20 cm above the ground with the nest centered about

5-10 cm below the surface and surrounded by many storage compartments connected by many passage ways. Copulation takes place in the breeding mound and the gestation period has been estimated as 23 days (Novo, 1961:139).

The extent of their surface activity is not known. Nevo

(1961) maintained that they can bo frequently seen on the surface 203 day or nip.li during tho months of March and May (possibly young adults soar*, ling for thoir fuburo territory) or during autumn months (adults searching for a mate). Skeletal remains recovered from owl pc bets are further proof of their nocturnal surface activity (D .be, 19U5 and Dor, 19h7). Other predators are domestic

i dogs and wild carnivores which destroy the breeding mounds seeking the young. These predators are probably attracted to tho nests by the squeaki. g of the young.

Adults maintained in a cage display an aggressive behavior to any moving object introduced into their cage. They were found to relish fruits, bread, onion and cheese. When two adults are placed together they fight bitterly until one dies, no canni­ balistic behavior was observed. Burrowing behavior and control measures have been described by Reed (1950) and Watson (1961) respectively.

Family MURIDAE Gray, 1821

Old World Rats and Mice

This family is represented in the Eastern Mediterranean by nine species representing five genera. Following is a KEY to these genera:

1. Dorsal pelage spiny...... Acomys

Dorsal pelage normal...... 2

2, Small forms, IIP and CDS loss than 30 mmj maxillary tooth-row

less than 6 mm...... 3

Large forms, HF and CIS over 30 mmj maxillary tooth-row over

6 mm...... U 3 . !'>. a Ghorb| less than 35 mmj HF less than 20 mmj with tii 30 roots j small, half the length and width of

...... MU3

Er s long, more than l£ mmj HF over 20 mmj with four

or five rootsj more than half the length and width of i 15 ...... Apodemus

h. T vj.1 short, less than l/2 TLj ears short, less than 20 mmj

ft osorial...... Mesokia

Tail appreciably longer than l/2 TLj ears long, over 20 mm

terrestrial and arboreal...... Rattus

Genus Apodemus Kaup

182?. Aoodemus Kaup, Skizz Europ. Thienvel, I: l5h.

Type species: Hus agrarius Pallas.

Throe species belonging to this genus have been reported from this regionj these are: Apodemus mystacinus, A. sylvaticua and

£• flavicollla (Aharoni, 1932). Thero is no doubt of the occurrence of the first two species, but all records of A. flavicollis o questioned. Allen (I9l£) was first to report this species g with A. mystacinus and A. sylvatlcus from tho lower slopes <

Mt. Hermon. I have examined most of his collection but find only representatives of the last two species. Aharoni (1932:183) described A. flavicollis pohlei from Kafrun, Syria. Weuhauser

(1936), Eodenheimer (19!?8) and Lehmann (1965) considered pohlei a junior synonym or a valid subspecies of A. mystacinus. 20£

Bodenhninisr (19^3) listed A. flavicollis as a member of - the Palestinian fauna based only on fragmented skulls in owl pellets collected at

Mt. Carmel by Dor (I9h7). However, since reliable determination of

A.flavIcoH's requires intact skulls and better skins, it is unlikely that this species occurs in tho Eastern Mediterranean Region. The southernmost population of A. flavicollis has been reported from

Villayet Rina in northeastern Turkey, almost 1,000 km from Palestine.

Following is a KEY to the two eastern Mediterranean speciest

HB less than 100 mmj HF less than 2h mmj GLS less than 2k mm

...... A. sylvaticua

HB over 100 mmj HF over 25 mmj (115 more than 25 mm in adults

...... A. mystacinus

Apodemus mystacinus (Danford & Alston)

1377. Hus mystacinus Danford and Alston, Proc. Zool. Soc. bond.,

p. 279.

1951. Apodemus mystacinus Danford and Alston j EUerman and Morrison-

Scott, Checklist, p. 56U.

Type locality: Zcbil, Bulghar Dagh, Turkey.

Common names: Broad-toothed Field Mouse.

Distribution: Greece, Yugoslavia, Bulgaria, Turkey, Syria,

Lebanon, Palestino and Jordan,

Lehmann (1965) referred specimens from northern Syria and southern Turkey to the form A. mystacinus pohlei Aharoni, previ­ ously regarded as a junior synonym of the nominate form

(Weuhauser, 1936 and Eodenheimer, 1953). Lewis, Lewis and Atallah

(1967) referred thoir specimens from Lebanon and Syria to the 206

nominate f( ;.i« Great amount of variation in cranial and external neasuremen' > can be found among Eastern Mediterransan populations

(loc. cit. ublo II) and awaiting more detailed study I provision­

ally liefer 11 these populations to the nominate form*

xtemus mystacinus mystacinus (Danford & Alston)

Synonymy u- ’or species also the following:

1932. Amo, -mas (Sylvaemus) flavicollis pohlei Aharoni, Z,

Saut .tierk*, 7:183. Kafrun, Syria. (Neuhauser, 1936).

Type material: not examined.'

General distribution: Cilician Taurus Mountains, Turkey, Syria,

Lebanon, Palestino and Jordan.

Allen (1915) obtained fourteen specimens from the base of

Mt. Hermon. Neuhauser (1936) and Bodonheimer (1958) considered

Aharoni's (1932) A. flavicollis pohlei from Kafrun, Syria, to bo a mystacinus and Bodenheimer (op. cit.) recorded it from Jerusalem,

Palestino. EUerman (19U8) referred specimens from Baalbeck and

Shcmlan in Lebanon to A, f. pohlei, while Lehmann (1965) referred specimens from Syria and Palestino to A. mys tacinus pohloi.

Lewis, Lewis and Atallah (196?) reported tho nominate form from

Mahr el Kelb, Ainab, Ain Zhalta, Natural Bridge near Faraya and

Laklouk in Lebanon and Kassab, Syria.

Other records.- Bodenheimer (1935) Wahrman and Ritte (1963) and Osborn (1965).

Snecimens examined:132 .LEBANON•Natural Bridge, noar Faraya (16

AUBj 13 SA)j 2 km N. of Bater (U SA); Reis el Assi, Source Orontes

(1 SA); Afka (2 SA); El-Rvmeileh, 7 km W. of Laklouk (16 SA);

Cedars, noar Bochorro (1 SA); 1 km S. of Lake Karaoun (1 SA); 207

Jabal Safin, 2 km N. W, Kfar Qouq (10 SA); Laklouk (8 AUB); Rachaya

(2 FMNH); / -.nab (6 AUB); Ain Zhalta (6 AUB); ICnhr el Kelb (8 SA);

1 km W. of Treijat (6SA). PALRSTINB: li Ion H. Vf. Beit Fa j jar

(1)4 SA). f .RIA: Kaa.aab (17 AUB). JORDAN: Dibbine (l SA).

Identification: Color of adult back Drab, darker along tho mid-dorsal surface, belly dull vhite. Young uniformly Cub colored on back. Individual hairs on belly with grey bases.

GLS usually over 28 mm, HF over 23 mm. Tail slightly longer than HB,

Biology: The broad-toothed field mouse is confined to the forest belt of tho Mediterranean Region. It is found at all altitudes from 100 n (Mahr el Kelb, Lebanon) to 2,$00 m, whero a single specimen was obtained at Becharro Cedars, Lebanon.

It is strictly nocturnal, with peaks of activity following sunset and at dawn. It burrows under rocks and feeds primarily on pine seeds, acorns and carob pods. Two pregnant females were collected in September, each with $ embryos. Lactating females were collected in June and August, while juveniles were obtained in April, May, Juno, July, September and October. These data suggest a continuous breeding season from early April through late October, with probably two litters per season. Three litters per season have been reported in Yugoslavia (Miric, 1966).

The species shares the same habitat with Crocidura russula,

Mlcrotus nivalis, Eliomys melanurus, Apodemus sylvatlcus and occasionally Mas muscuius and Rattus rattus, especially at low 208 elevations ?ar villages.

Anodcnus sylvaticns (Linnaeus)

17^8. Mus ylvaticus Linnaeus, Syst. Kat., 10th ed,, 1:62.

Type locality: Upsala, Sv/eden.

Comnon name i: Common Field Mouse.

Distribution: Europe excluding northern Scandinavia and

European USSR, Kazakhstan, Iran, Iraq, Turkey, Atlas Mountains in north Africa and tho Eastern Mediterranean (vide Zimmermann,

1962).

Many questions have been raised regarding the validity of

Aoodemus sylvaticus and A. flavicollis and numerous workers have transferred subspecies from one species to the other.

Both species have been recorded from Turkey: A. flavicollis saturatus Neuhauser from northeastern Turkey and A. sylvaticus tauricus (Barrct-Hamilton) ranging over most of Turkey (Neuhau­ ser, 1936). EUerman (l9i|3) did not deal with the form saturatus but accepted Nauhauser's views regarding A. sylvaticus tauricus while EUerman and Morrison-Scott (19^1) accept both forms as used by Nauhauser. Lelmann (1966a:302) believed all A, sylvaticus and A, flavicollis in Turkey were hybrids and designated them as a ’'semi-species'’, Aoodemus sylvaticus (tauricus) saturatus

Neuhauser. The rules of nomenclature, do not allow for such a system, nor do I- agree that the Turkish populations cannot bo separated. 209

Tho fc a tauricus has been given full specific status by some author:? (see Larina(195>8)for details). I have examined many specin' as of Apodemus sylvatlcus and A. flavicollis and compared specimens from Turkey previously referred to tauricus with both species. Both sylvaticus and flavicollis appear to represent distinct species and tauricus definitely belongs to tho sylvaticus group. Eastern Mediterranean populations, hero referred to A. sylvaticus tauricus, are slightly smaller and are lighter in coloration than Turkish populations. A detailed study may warrant recognition of a separate subspecies in tho

Eastern Mediterranean.

Apodemus sylvaticus tauricus (Barret-Hamilton)

1900. Mus sylvaticus tauricus Barret-Hamilton, Proc. Zool. Soc.

Lond,, p. 1|12.

19f>l« Apodemus sylvaticus tauricus Barret-Hamilton; Ellorman

and Morrison-Scott, Checklist, p. £71*

Holotypo: BM Collection no. 77.8.13.9.# collected at Zebil,

Taurus Mountains, Turkey on January 5# 1876 by C. 0. Danford.

General distribution: Turkey, Syria, Lebanon and Palestine,

Tristram (1866a and I88I1) reported on a single specimen collected "in the plains" in Palestine, however, both Aharoni

(1932) and Nauhauser (1936) considered this record erroneous.

Allen (191^) obtained specimens from Shiba, Rachaya and Ain

Kovsha in Lebanon, Bodenheimer (I93f> and 1958) listed tho species as a member of the Palestinian fauna, but he did 210 nob aay if I.o had actually obtained specimens. Lehmann (196£) recorded sj. cimens from Kastel Maaf, Syria, which he identified as A, sylv:. i.cus iconicus Heptner, Lewis, Lewis and Atallah

(1967) coll eted eleven specimens at Laklouk and Natural Bridge near Faraya in Lebanon.

Other ocords.- Wahrman and Ritto (1963).

Specimens o.camined; 21? LFBUTON: Natural Bridge, near Faraya.

(6 SA; 7 AliJ); Cedars, near Becharro (3 SA); Jabal El Knisse, olev. 1,900 m (1 SA); Jabal Sanine, near Ouadi ed Delb, elev,

1,^00 n (h SA); S, Slope of Jabal Sanino, elev. I,lj00 n (2 SA);

Laklouk (2 AUB).

Identification: Color of back Beech, belly white. Individual hairs on belly white with grey bases and occasionally a yellow streak, 10X2 mm, is present on the breast. GLS less than 26 mm,

HF less than 23 mm. Ear short, less than 18 mm.

Biology: Tho common field mouse is restricted to elevations above 1,200 m in tho Mediterranean Region. It inhabits rocky mountain slopes and cedar forests with the more comnon Apodemus mystacinus and Mlcrotus spp. Burrows are usually constructed near or beneath shrubs and rocks, but occasionally they were trapped among Mlcrotus guentheri colonies in the valleys.

Juveniles were obtained in July, while two pregnant females collected in July and October had four embryos each. This suggests continuous breeding during the spx-ing and summer.

They feed on leaves and steins of various mountain shrubs and cedar cones, when available. 211

Genus Rattus Fischer

1303. Rattvs Fischer, Nat. Nus. Nat. Raids, 2:128.

Type specie. : Mus dccuirianus Pallas » ITus norvegicus Ecrkcnliout.

Follow: ug is a KEY to the two species recognized in tho

Eastern Med .berranesn Region:

Tail shorter than HBj temporal ridges on brain case

approximately parallel...... R. norvegicus

Tail longer than HBj temporal ridges on brain case widely

curved...... R. rattus

Rattus rattus (Linnaeus)

17£8. Hus rattus Linnaeus, Syst. Nat., 10th ed., I:6l.

Type locality: Sweden.

Common names: Houso Rat, Black Rat.

Distribution: A commensal form distributed throughout the world, introduced by man.

Aharoni (1932) recognized three subspecies in this area of study, as follows:

(1) R. r. alexandrinus (Desmarest) with brown dorsum not sharply separated from the grey or yellowish white venter. An inhabitant of dry regions chiefly as human commensals,

(2) R. r. frugivorus (Rafinesque) with greyish brown dorsum sharp­ ly separated from white or yellowish white venter. An inhabitant of moister habitats feeding primarily on fish and snails.

(3) R. r. flaviventids (Brants) with bright reddish brown dorsum not sharply separated from tho dirty, yellowish venter. An inhabitant of desert areas. Also a smaller form, GLS not exceeding ti3.£ mm in 212 adults (mor. than h7.5 mm in the remaining two subspecies).

Ellerm' n and Morrison-Scott (19ijl 5^81-2) considered that aloxandrinu- and frugivorus were merely color forms but treated them as subrpecies, while Bodenheimer (1958:182) placed flavlventrl-'i in synonymy with R. r. frugivorus. Specimens from

Palestine arid Lebanon were found to satisfy the color requirements of either alexandrinus or frugivorus or were intermediate (Lewis,

Lewis and Atallah, 196?), but in all cases GLS less than h2.0 mm suggested flaviventrls. I have subsequently examined a few specimens from northern Saudi Arabia (Dahran) with tho typical yellow belly of flaviventris which seems quite distinct from any Lebanese or Palestinian specimens.

Our understanding of Rattus rattus populations is unsatisfactory for the Eastern Mediterranean and future studies may indicate the presence of a single subspecies in the whole area. I presently believe that Aharoni*s (loo, cit.) flaviventris is probably frugivorus and that, at best, two color forms can be distinguished in this area, as follows:

Color of back gray brown, flanks buff gray, belly white or yellowish; color transition along the flanks with a distinct line of demarcation; in fields rarely in houses,..,..,.,,,..

...... '.'frugivorus " ' color form

Color of back olive brovm, flanks drab, belly yellowish gray (siroccot); color transition along the flanks indistinct; in houses rarely in fields.,,.,...... '.'alexandrinus'* color form 213

"alexandrinus” color form

1819. Mus lexandrinus Desmarest, Nouv Dist. H. N., 29:li7.

19^1. Rat^ .3 rattus alexandrinus Desmarest; EUerman and

Mon '.son-3cott. Checklist, p. £81.

General di;: tribubion: A commensal form distributed throughout the * world.

Record j of this color form from the Eastern Mediterranean are scarce, Aharoni (1932) reported it to be present from Hama,

Syria to southern Turkey, but had only a single specimen collected at Hama while Bodenheimer (193!? and 19!?3) considered it tho commonest rat in Palestino.

Other records.- Bodenheimer (1920) and Lewis, Lewis and

Atallah (196?).

Specimens examined: 9. PALESTINE: Beit-Sahur (9 SA).

Identification: Belly greyish yellow, dorsum olive brovm.

GLS 37-142 mm. IIP short, usually less than 36 mm.

Biology: Rats of thellalexandrinusl, color form are typically found near houses in the Mediterranean Region. They are active at all hours of the day or night. Their burrows are usually concealed under stones or along houses. They breed all year round.

"frugivorus11 color form lOlii. Huscuius frugivorus Rafinesque, Precis des Decouv. et

Travaus Simiol., p. 13.

1951. Rattus rattus frugivorus Rafinesque; EUerman and Morrison-

Scott, Checklist, p. 582. General dit 'oribution: Throughout Eurasia and North Africa.

Aharoni. (1932) recorded specimens from Hadera and Jericho in Pales tii-j, however, she didn't give any data regarding theso specimens 5i tho appendix. She recorded R, r. flaviventris from several localities in Palestine and Lebanon, but Neuhauser (1936) found her specimens to represent this color form.

Other records.- Bodenheimer (193£ and 19^3) and Lewis, Lewis and Atallah (196?).

Specimens examined: 7 LEBANON: International Airport, Beirut

(li SA)j American University of Beirut, Campus (2 SA)j Nahr el

Kelb (1 SA).

Identification: Sams as "alexandrinus", but sharply bicolored with a white or yellowish belly. HF usually over 3£ mm.

Biology: These rats were found abundant along a stagnant stream running west of Beirut International Airport, Several burrows were located beneath mimosa trees (Acacia cyanophylla) common in this area, especially on sand dunes near the stream.

A few rats were seen climbing these trees in broad daylight

(3 p.m.) and later several nests were located in tho trees.

The nests wore approximately 2,$ m above ground made of densely packed mimosa leaves and twigs. A nost collected in

April was found to contain six young. A mother frightened at the beginning ran away but soon returned to defend her young.

At this locality Meriones tristrami and Mus muscuius wore common.

The specimen from Nahr el Kelb was collected from a stone wall along a side stream, running through citrus groves. 21$

Rattus norvcKlcus (Borkenhout)

1796. Hus lorvoglcus Borkonliout, Outlines N. H. Gt. Britain

& I eland, I:$.

19$1« P.ab: ,:s norvegicus Borkenhout; Elleman and Hox-rison-

Sco^ t, Checklist, p. $89.

Type locali ty: England.

Ccmr,ion nan s: Nox^ay Rat, Broivn Rat.

Only the noninato form is recorded from the Eastern

Mediterranean Region.

Rattus norvoaicus norvegicus (Berkenhout)

Synonymy under species.

Type material: Not examined.

General distribution: World-wide through introduction by man, a native of Palaoarctic Asia.

Tristram (l866a and 188U) recorded Hus decumanus (“R. norvegicus) as comraon throughout Palestino, while Aharoni (1932) maintained that she did not find this species any place in Palestine.

Bodenheimer (193$ and 19$3) reported that this form had established itself at Haifa, Jaffa and Tel Aviv, three major ports, Lehmann

(1966a) and Lewis, Lewis and Atallah (196?) obtained specimens from

Beirut, Lebanon.

Specimens examined: 3. LEBANON: Beirut (1 SA; 2 AUB).

Identification: Tail shorter than HB. HF usually more than liO mm.

Color of back greyish brown, belly greyish white with a distinct lino of demarcation along the flanks between tho two surfaces.

Temooral ridges on skull run parallel to each other, widely curved in Rattus rattus. 216

Biology: I own rats aro atnmdant to ;ill port cities along tho

Mediterranr a. They can be seen at night and occasionally during the day no; • sewer outlets and garbage piles. They have been able to ponetrat deeper inland (Lewis, Lewis and Atallah, 196?) and will probab ;/ become more abundant in the future.

Genus Hus Linnaeus

1753, Hus ‘‘.tanaeus, Syst, Nat,, 10th ed,, 1:^9,

Typo specie.: Hus muscuius Linnaeus,

Hus muscuius Linnaeus

17f>8. Hus mus cuius Linnaeus, Syst, Nat., 10th ed,, 1:62,

Type locality: Upsala, Sweden.

Common names: House House, Far el Beit (Arabic),

Distribution: World-wide through introduction by man.

Two color forms are recognizable in the Eastern Mediterranean

Region, theso are: (l) a dark grey form with a greyish white belly and an indistinct lino of demarcation between tho two surfaces, and

(2) a bicolored form with white or yellowish belly and olive green back. Both forms aro usually found together and equally abundant around human habitations, while the bicolored form predominates

(over 95%) in the feral state especially in the Steppe and Desert

Regions, The bicolored for^-VThich was,described,from Syria by

Brants (1827:125) as Hus praetextus , was considered a subspecies of H. musculus (Ellorman and Morrison-Scott, 1951:6o6). Tho dark colored commensal form has been referred variously to several subspecies by aharoni (1932), Bodenheimor (1935 and 1958), and 21?

Lewis, Lewi 5 and Atallah (196?), but will bo treated here as only a color vav'.ent of H. muscuius praetextus.

The effects of probable repeated introductions of Mus muscuius, over thousaids of years, are not presently understood. However, the survival advantages of a lighter, bicolored form in feral conditions, under the present conditions of aridity, are more obvious.

Mus muscuius praetextus Brants

1827. Kus praetextus Brants, Gesl. der Kuizen, p. 12J>,

195>1. Mus mus cuius praetextus Brants j Ellerman and Morrison-

Scott, Checklist, p, 606.

Holy type: BZM collection no. I63U, collected by Hemprich and

Ehrenberg from Sakhara, Syria. Skull broken.

General distribution: Not yet established, probably most of the Middle East.

Allen (1915) and Aharoni (1932) recorded specimens from several localities in Palestine, Syria, Lebanon and Jordan. Tristram

(1866a and l831j), Bodenheimer (193£ and 1953) and Ellerman (19^0) from

Palestine, Lehmann (1966a)frcm Lebanon, Syria and Jordan, ahd Lewis,

Lewis, and Atallah (1967) from Lebanon.

Specimens examined: 70 PALESTINE: Bethlehem (13 SA)j Beit

Sahur (10 SA)j near Qumran, N. end of Dead Sea (6 SA)j h km

N. W, Beit Fajjar (2 SA). JORDAN: Axraq ed Druz (7 SA)j

A.zraq-Shishan (2 SA)j El-Jafr (1 SA)j 10 km N.E. Amman (1 SA).

LEBANON: ABB Farm, Eckaa (20 SA); near Mar Abda (h SA)j Ras el Assi (2 5A)j 1 1cm, S. of Lake Qaraoun (2 SA). 218

Idontifica! .on: CIS usually loss than 22.5 run, ears short less than 13 run. Tail usually subequal, but sometimes equal to HP* Color variable, the two color forms were described v .der species discussion.

Biology: The house mouse is vridely distributed throughout the Eastern Mediterranean Region. It has been collected at a variety of habitats, but it is usually found in moist habitats as along water courses, cultivated fields and oasCs. Specimens were taken at burrow entrances of Meriones tristrami and

Microtus guentheri in Lebanon. Atallah (1967c) recorded four specimens sharing the habitat with Gerbillus henleyi,

0. dasyurus and Meriones libycus at El-Jafr, Jordan.

Genus Acomys E, Geoffroy

1838. Acomys Geoffroy, Ann. Sci. Nat. Paris, Zool., 10:126,

Typo species: Mus cahirinus Desmarest.

This genus is primarily confined to the African continent and southwest Asia. Following is a KEY to the three species recognized in the Eastern Mediterranean Region.

1. Ventral surfaces of feet black; tail short, averaging less

than 90 mm; spines covering the entire dorsal surface; ears

covered with short hairs and spines...... ,..,.,...... 2

Ventral surfaces of feet yellowish-brown; tail long, averag­

ing more than 95 mm; spines not extending as far anteriorly

as the head; ears naked,,A. dlmidiatus 219

2. Golden- allow over-all coloration; baculiti terminating in

a bony -rifid with threo distinct ossicles.,,A, russatus

Black o or-all coloration; baculum terminating in a car-

tilagin-:i3 trifid...... A, leTvisi

Acomy dimldiatus (Cretzsclimar)

1B26, Mus imidiatus Cretzschmar, Ruppoll Atlas, p. 37» taf,

13, tig. a,

19$1» Acaiw's cahirinus dlmidiatus Cretzschmar; ELlerraan and

Monlson-vGcott, Checklist, p, 6lJj.

1937. Acomys dlmidiatus Cretzschmar; Setzer, J, Egypt, Publ,

Hlth. Assoc,, 3b(3)J93.

Type locality: Sinai,

Common names: Cairo Spiny Mouse, Sinai Spiny Mouse,

Distribution: North Africa, southwest Asia and India,

Only the nominate form is pertinent to the studied area.

Acomys dlmidiatus dlmidiatus (Cretzschma**)

Synonymy under species.

Type material: Not examined.

General distribution: Sinai, Palestine, Jordan, southern Lebanon,

Syria, southern Turkey (Lehmann, 1966b), southwestern Iran and

Saudi Arabia.

Tristram (1866a and lOCb) observed the species in the Dead

Sea Basin and the hills of Moab, Allen (1915) studied specimens obtained at Aqaba, Petra, Tafila and Wadi Kerak in Jordan.

Aharoni (1932) recorded it from Tabgha on Lake Tiberias, Jerusalem, 220 Ghor es Sat .ah and Hisma in Palestino and Moab, Jordon,

Ellemun (1 Ii3) studied EM collection and I'ecorded specimens from Wadi Z rqa, Jordan and Jerusalem, Palestine, Potter (195U) recorded it from Bir Hindis, Wadi Masai and Jerusalem in Palestine,

Atallah (l$v 6b) recorded it as common near cultivated fields and houses n semiarid areas, and Levris, Le-vrls and Atallah

(1967) repo..'ted it from Sour In southern Lebanon, I am not aware of on/ records from Syria, but it is conceivable tint it may.; occur there especially as it has been taken in southern Turkey

(Lehmann, 1966b)

Other records,- Wahman and Zahavi (19i>3)»

Specimens examined: 8Ii PALESTINE: Bethlehem (29 SA)j Beit-

Sahur (18 SA)j Deir Mar Saba (1 SA)j Ain Faschkha (9 SA)j U km

H. Beit Fajjar (2 SA)j Jerusalem (2 BZM); Hisma (5 BZM); Tabgha,

Lake Tiberias (h BZ?-^ Ain Gedi (U BZM); Jericho (l BZM). JORDAN:

3J4 len N. Aqaba (1 SA), Also 8 specimens from St. Catherine’a

Monastery, Sinai .

Identification: Soles of hind feet not black. Tail length usually exceeds 95 Km in adults. Color of back Toast, belly white (grey in soma commensal individuals) with a distinct line of demarcation between the two surfaces. Spines on back more or less restricted to the posterior parts of the body. 221

Systematic omarks t Specimens from the Eastern Mediterranean

Region are .-lightly different from those from Sinai. Ear and tail 1-. igth average longer (see Appendix I) and the pelage is lighter in the Sinai specimens. A more intensive study of these animals may warrant separation of the Eastern Mediterranean and

Sinai popu? vtions into more than one subspecies.

Biology: Cairo spiny mice are common throughout the Steppe

Region of southern Palestine, less common in Jordan and

Syria. They occupy a variety of habitats varying from rock slides under extreme desert conditions to cultivated fields and houses in villages and occasionally in cities. My anal­ ysis of stomachs as well as field and laboratory observation indicates that these mice consume a large amount of animal matter along with the normal diet of green leaves and stems.

In semiarid areas an abundance of snail shells usually marks the nearby burrow entrance which is . usually concealed beneath rocks or stone walls, I have observed cannibalistic behavior in the laboratoiyand stomach contents of a lactating female collected at Bethlehem in February 196£ contained fur of a young (possibly its own).

Spiny mice breed all year round. Littor size varies between 3 and Ij.,

Acomys russatus (Wagner) lOhO. Mus russatus Wagner, Abh Bayer Akad. Wiss., 3:195, pi. 3,

fig. 2.

1951. Acorn?/- russatus Wagner. Ellerman and Morrison-Scott,

Checklist, p, 616.-

Type locality: Sinai. 222

Common naif • Golden Spiny Mouse.

Distributi' : Eastern Egypt, Sinai, Palestine, Jordan and Arabia.

Only 1 to nominate form has been recorded from the geographical area covert

Acomys russatus russatus (Wagner),

Synonymy ui lar species.

Typo mteri.'.l: Not examined.

General distribution: Eastern Egypt, Palestine, Jordan and vrestern Sar.di Arabia.

Tristram (1866a and l88ib) found golden spiny mice common in

Sinai, but only near the southern end of tho Dead Sea in Palestine.

Nehring (1901b) obtained a specimen at Moab, Jordan and another from Ain Gedi, Palestine. Aharoni (1932) maintained that the range of the species extends from Egypt to Jerusalem, Palestine. Her records, however, do not list any specimens from the environs of

Jerusalem, nor any other locality in Palestine.

Other records.- Bodenheimer (1920, 1935 and 1958) and

Atallah (1967a and 1969a)

Specimens examined: 31* JORDAN: Moab (1 BZM)* PALESTINE: Ain

Faschkha (h SA)j Chor es Safieh (1 BZM), Also 25 from St.

Catherine's Monestery,' Sinai '(FMNH).

Identification: Color of back golden yellow, bally white.

Soles of hind feat black. Dorsal surface coinpletoly covered with spines. Tail short, usually less than 80 mm in length. 223 Systematic omarks: The populations from the Judean Cesort are

appreciably smaller and lighter in coloration than the nominate

form descr.i ed from Sinai. I have just submitted a paper for

publication describing a new subspecies from tho northern end

of tho Deaf! Sea (Atallah, 1969a)

Biology: 'i ic golden spiny mouse inhabits steep rock slides in

semiarid a;eaS. It is diurnal in habits with peaks

of activity occurring in the morning and late afternoon.

They share the same habitat with Acomys dlmidiatus, but the latter

species is nocturnal in habits.

Areas occupied by these mice are for the most part devoid

of green vegetation at least ten months of the year. Seeds,

leaves and stems are usually stored under rocks serving as

feeding stations,also a largo amount of animal matter is

consumed.

Acomys leva si Atallah

1967. Acomy lewlsi Atallah, J. Mammal,, 1j0(2):298.

Holytypo: FMNH collection no. 99i|21, an adult male collected

at 3 km N. W. Azraq-Shishan, Jordan on April 2lt, 1966 by

S.I. Atallah.

Common names: Black Syrian Desert Spiny Mouse, Lewis’ Spiny Mouse.

General distribution: Known only from tho typo locality, probably widely distributed throughout the basalt desert in southern

Syria and northeastern Jordan. 22h

Hcmsloy and George (1966) recorded two specimens from the type local! y of le^-rlsi identified as Acomys cahirinus, a north

African spe ;ies, but Atallah (1967b) maintained that they are referable .o lewisi,

Speciraens examined: 11, JORDAN: 3 km N. W. Azraq-Shishan

(6 SA)} Azriq ed Drua (3 SA)} Ij krti N» W, Azraq-Shishan (2 SA).

Identification: Color uniformly blackish grey. Soles of hind feet black* Tail short, usually less than 80 mm in length.

The baculm lacks tho partially or completely ossified terminal trifid of Acomy russatus and A. dlmidiatus (Atallah,

1967b:Fig. 1).

Biology: lewis' spiny mouse abounds along the edge of the basalt desert where it adjoins the limestone hammada, as well as in gardens around human dwellings. It shai'es the same habitat with Bliomys rr.elanurus, Gerbillus dasyurus and Mas mus cuius.

Genus Nesokia Gray l8h2. Nesokia Gray, Ann. Mag. Nat. Hist., 10 (l):26!j«

Typo species: Arvicola indica Gray.

This monotypic genus is primarily a central Asiatic form, intermittantly distributed in moist habitats of the Middle East,

Nesokia indica (Gray)

1830. Arvicola indica Gray, Illustr. Ind. Zool., I: pl« xi.

Type locality: India,

Common names: Short-tailed Bandicoot Rat, Short-tailed Mole Rat, Distributi.! j From Russian and Chinese Turkestan eastward to Egypt*

Tho Pr '.stinian population of this species, N. indica bacheri, is separate from N, indica suilla confined to Egypt, N, indica buxtonl fr<. southern Iraq and northern Arabia, and N. indica nyosorus fa. n northern Iraq and northeastern Syria (vide Misonne, •1 ' 1 1 "" r ^ « 19^7) by e-. inses of desert.

N, in< oa myosorus Wagner (l8h£) was described from a single specimen c< .looted by tho botanist Kotschy labelled "Syria".

Wagner nam-d it Koriones myosorus, but latei' Nehring (1901b) referred it to Nesokia. Ellerman (19^8^808) though unacquainted with this form suggested that it may bo tho earlier available name for II. i, buxtoni Thomas, but Ellen;’an and Morrison-Scott

(I9!?l:6l9) listed both subspecies as separate. Misonno's

(1957) specimens from Tell Abiad along the Syrian-Turkish border are tho only examp3.es of myosorus from Syria other than the holotypo. This subspecies will not bo dealt with in this papor, as it has not yet been recorded from the area of study.

Only tho following subspecies is recognised along the Eastern

Mediterranean,

Nesokia indica bacheri Nehring

1897. Nesokia bacheri Nehring, Zool. An?.. No. 5U7i503*

1951. Nesokia indica bacheri Nehring) Ellerman and Morrison-

Scott, Checklist, p. 620. 226

Typo mater.' &: Wot examined, from Ghor es Safieh, Palestine.

General di: tribution: Palestine and Jordan.

Wohrii j (1899) obtained seven specimens from Moab, Jordan.

Aharoni (1S;32) referred Tristram’s (10820 record of a Nesokia from the Dead Sea region to this form, she also studied some of

Nehring’s material•

Other records. Nehring (1901b), Bodenheimer (1920, 193$ and

1958) and Ellerman (19240).

Specimens examined: 11, PALESTINE: Ghor es Safieh (3 BZM).

JORDAN: Moab (8 BZM).

Identification: Tail usually shorter than l/2 HB, Color of back reddish brown, belly yellowish grey with a white breast-band between the forelegs. GLS more than I4O mm, ZB more than 2$ mm.

Fossorial.

Biology: Bandicoot rata aro restricted to warm and moist habitats. Thus throughout the Middle East one finds populations of this species only at certain localitieswhero their stringent habitat requirements aro fulfilled. Nothing.is known of tho habits and reproductive biology of the Eastern Mediterranean populations. I failed to locate a population in tho lower

Jordan valley, even after extensive search and trapping.

Family GLIRIDAE Thomas, 1897

Dormice

Three species belonging to this family have been recorded from the Eastern Mediterranean: Eliomys molanuruo, Dryomys nitedula and 227

Olin glia. Recent records of the first two species are abundant while the 1 'ird species recorded by Tristram (18o6a and 188U) has not be; i taken or observed for almost a century. Tristram

(l866a:89) vaintained "The great Dormouse is very abundant in the oasis of thi Jordan valley, especially about Jericho, where it has its nest in every dom tree. Through some oversight we did not bring a specimen homo} but it has been mentioned by several writers, from

Russell dov.iwards. At Jericho it was very lively in winter when disturbed". This record was repeated by Aharoni (1930:337) and

Bodenheimer (1935>:S)£). Recently Dr. Haas at the Hebrew University located a passage in Seetzen (l85U:Vol. 2, p» 311) of which Tristram's record appears to be the exact translation. U. J. Seetzen was not a zoologist and he may have been describing a fat sand rat (Psammomys), or the garden dormouse (Eliomys nelanurus). Following.^ is a KEY„ to the two species currently recognized in this region:

1. C3L over 28 mm} tail shortly haired at base, tufted at tip.

.Eliomys malanurus

2, CBL less than 2£ mm} tail uniformly bushy. .Dryomys nitedula

Genus Eliomys Wagner

181(0. FT ion vs Wagner, Abh. Bayer Akad. Wiss., 3:176.

Type species: Eliomys nolanurus Wagner.

Two species are recognised in this genus: E. melanurus along tho Eastern Mediterranean from Asiatic Turkey to Sinai and E, quercinus in southern Europe and North Africa. Bodenheimer

(19^8:181) suggested reducing melanurus to a subspecies of E. quercinus as did Neithanmer (1959)• This does not, however, appear to be the case, as both species are distinct} melanurus has a 228

highly ini bed bullae and lacks the white banner on tail of quercinus.

Eliomys melanurus Wagner

18140. Ell ,;/w (Myoxus) melanurus Wagner, Abh. Bayer Akad.

Wls ,, p. 1?6, pi. 3, fig. 1.

Type mter >1: Not examined, from Sinai.

Common nam j: Southwest Asian Garden Dormouse.

General dl bribution: Sinai, northwestern Arabia, Palestine,

Jordan, Sy,la, Lebanon and southern Turkey.

Tristram (1866a and l88li) recorded the species as common in the Jordan Valley and the Dead Sea Basin, but he obtained only

Wo specimens from Moab. Allen (1911)) reported it from Rashaya and Ain Hersha in Lebanon, Bodenheimer (1920, 1935 and 1958) from northern Palestine and southern Jordan, Ellerman (19U8) from Karyatein, Syria, Hemseley and George (1966) and Atallah

(1967a) from the Asraq Basin, eastern Jordan and Lewis, Lewis and Atallah (1967) from Laklouk and Natural Bridge near

Faraya in Lebanon,

Other records.- Missono (1957).

Specimens examined: 19 SYRIA: Karyatein (1 BM). LEBANON:

Jabal Kammoah, near source (2 SA); Cedars, near Becharre (l SA);

Jabal Safha, 2 km N. W. of Kfar Qouq (6 SA); 1 km W. Mreijat

(1 SA); Jabal Knisse (5 SA); Laklouk (1 REL; 1 SA); Natural Bridge near Faraya (1 REL),

Identification: Color of back smoke brown in young and subadults.

Grey 31 in adults. Belly greyish white, eyes with a black stripe 22? continuing alow the ear. Tail with a terminal black tuft extending l/2 tail If -.gth, proximal l/2 grey. GL3 usually more than 30 mm.

Arboreal,

Biology: Tiria species is vridely disti'ibuted throughout the

Eastern Mec)itorranean from the desert to mountain tops (2,£3>Om at

Cedars, near Pecharro). Populations, however, are highly local­ ized and restricted to rocky area, especially on mountain slopes.

They share tho same habitat with Microtus nivalis and Apodomus spp, in the mountains and Gerbillus dasyurus and Acomys sp. in tho desert.

Stomachs examined contained more animal than vegetable matter. Among tho multitude of arthropods recovered, one stomach was found to contain the remains of a gekko,

Ptychodactylus hasselqnisti (Donndroff).

The garden dormouse is strictly nocturnal, and judging from trapping results it appeared to be active only the earlier part of the night. .Immature3 were collected in July.

Genus Dryomys Thomas

1906, Dryomys Thomas, Proc. Zool. Soc. Lend., 2:3i|8.

Type species: Mus nitedula Pallas.

A monotypic genus.

Dryomys nitedula (Pallas)

1779. Mus nitedula Pallas, Nov. Spoc. Quad, Glir, Ord., p.88.

Type locality: Region of Lower Volga, Russia,

Common names: Forest Dormouse.

Distribution: From Switzerland and Germany eastward to Russia, and Chinese Turkestan. 230

I have .^nly nxaminod two specimens belonging to this species from the gc* ;rat>hical area covered. These specijtens are here provisional'/ assigned to D. n. phrygius solely on geographical grounds.

Dryomys nitedula nhrygius Thomas

1907. Dryoii/s nitedula phrygius Thomas, Ann, Mag. Nat. Hist.,

20(7 ):li07.

Type material: Not examined; from Murad Dagh, Ushak Province,

Turkey.

General distribution: Turkey and northern Palestine.

Tristram (1866a and 1881;) listed this species as a member of the Palestinian fauna. Bodenheimer (19!?8) reported on a single specimen from Upper Galilee in the collections of Tel Aviv

Museum. Mevo and Amir (1961 and 1961:) studied in some detail the distribution and biology of this species in Palestine.

Specimens examined: £. PALESTINE: Saar (1 FMNH); Upper

Galilee (1 FMNH). Also three specimens from Ulu Dag, Turkey

(USNM).

Identification: A small dormouse with TL less than 200 mm and GLS less than 2£ mm, Bmmish grey general appearance with a bushy tail. Arboreal.

Biology: All that is available on tho biology of this species has been published in Nevo and Amir (1961:). Garden dormice were found to be restricted to the evergreen maquis of Quoreus calliminos-Pistacia pales blna. It is primarily nocturnal. 231 arboreal an omnivorousi. The male and the female share in build­ ing the nos } but the male leaves before the young are bom.

Each female has two to three litters per year, with an average of three young, The breeding season io interrupted by winter months during which dormice occasionally go through short periods of light hibernation.

Family DIPODIDAE Waterhouse, I81j2

Jerboas

Members of this Palaearctic subfamily are confined to desert and steppe regions. Hind feet are elongated and tail very long, adapted for bipedal locomotion, .

Two genera and species are recognized in the Eastern Mediterranean

Region. Following is a KEY to these two species;

1. Bullae feebly inflated, mastoids not appearing in dorsal aspect

of skull; cheekteeth li/3; five-toed; upper incisors smooth..

...... Allactaga euphratica

2. Bullae greatly inflated, mastoids appearing prominently in

dorsal aspect of skull; cheekteeth 3/3; three-toed; upper

incisors grooved...... Jaculus .jaculus

The two Arabic names jarboua and yarboua are applied to all members of this family.

Genus Jaculus Erxleben

1777• Jaculus Erxleben, Syst, Regn. Anim., p. Uoln

Type species: Jaculus orientalis Erxleben.

This genus is restricted to the Saharo-Sindian Desert Belt, with only the following species recorded from the Eastern 232

Mediterrajiean Region.

Jaculus jaculus (Linnaeus)

17!jB. Hus jaculus Linnaeus, Syst. Hat,, 10th cd., 1:63.

Type locality: Giza pyramids, EgjT>t.

Common names: Lesser Egyptian Jerboa, Three-toed Jerboa.

Distribution: North Africa and southwest Asia.

Two subspecies are recorded from tho Eastern Mediterranean

Region: J. jaculus vocator from the Syrian Desert and J.

jaculus schluoteri from southern Palestine. The two subspecies

are separated geographically by the Rift Valley and the

adjacent uplands. Bodenheimer (1958) reported the nominate

subspecies from the Negev region, however, there appears to

be no evidence for its presence there. It is thus omitted

from this study,

Jaculus jaculus vocator Thomas

1921. Jaculus loftusi vocator Thomas, Ann. Hag. Nat. Hist., 8(9):)4lil.

1951• Jaculus jaculus vocator ThomasJ Ellerman and Morrison-Scott,

Checklist, p, 539•

Holotype: EM collection no. 0.5»22.3*> a young adult male taken near Muscat, southern Arabia.

General distribution: Arabia, southwestern Iraq, Jordan and Syria.

Thomas (1922) described J. Jaculus syrius from Karyatein,

Syria, this form is considered a junior synonym of J. jncuius vocator (Elleman, 19U8 and Ellerman and Morrison-Scott, 1951).

Atallah (196?a and c) found this form common in eastern Jordan.

Other records.- Tristram (1881: as Dipus hirtioes) md

Ellerman (19)48), 233

Specimens <..camined: $$ JORDAN: El-Jafr (22 SA); 2-3 km

N. E. of A. aq-Shishan SA)j Azraq-Shishan (h SA)j $ km

W. Azraq-Si ..shan (l SA)j near Qasr Amra (3 SA). SYRIA: Palmyra

(!? AlIB)j D' r es Zor (3 BZM); Karyatein (Holotype of J, jaculus syrius, BM). Also Vy specimens from northern Saudi Arabia (AUB).

Identification: Color of back Drab, belly white. HP usually less than 6l mm. Banner on tail with a white tip and a brown or black subterminal band. GLS less than 3U mm.

Biology: Threertocd jerboas are restricted to the Syrian Desert plateau. They are strictly nocturnal, becoming active ohly after dark. A good part of their activity, however, is usually restricted to the first three or four hours of darkness.

The burrow system usually has two entrances, both of which are kept closed during the day by a thin layer of packed soil as a protection against predators and the desert heat. Tho burrow is usually between 5>0 to 100 cm in depth irith the nest located at the deepest point. From the main burrow descending toward the nest, many side burrows run up toward the surface and end blindly about 3-U cm from the surface. Such burrows are used for escape, and at many times while digging or driving near tho entrances, frightened jerboas were found to break through one of these blind burrows onto tho surface and run to a close burrow. 23h

A fen-' e collected in April held four embryos; young vero obtained in Juno, In the laboratory they are reported to breed once every .hree months, with a litter of three or four young and a gestation period of 2£ days or less (Lewis, Lewis and

Harrison, 1^65:67)• An adult male collected in northern Saudi

Arabia in 3 y'63 is still living in tho laboratory. Courtship and copulation behavior has been described by Lewis, Lewis and Harrison (op, cit.).

Bedouins seek jerboas for food. Other important predators aro the fox and wild cat. Jaculus jaculus schlueteri (Nehring)

1901. Pious s chill tori Nehring, S. B. Ges, Nat. fr. Berlin, p. 163.

195>1. Jaculus jaculus schlliterl Nehring; Ellerman and Morrison-

Scott, Checklist, p. 539. Type material: Not examined, from Palestine.

General distribution: Southern Palestine.

Bodenheimer (1935 and 1953) restricted this jerboa to the sand dunes along the coastal plain in southern Palestine, and maintained that it is replaced to tho south by the nominate sub­ species. These findings, however, aro conjectural as he did not obtain any specimens from the Negev to compare with those from the coastal plain.

Other records.- Ellerman (19h8)# 23S

Specimens omined: 1. PALESTINli: Jaffa (1 BM).

Identifica! .on: This fom appears to be slightly larger than

J. j. vocal .1% The only specimen examined had a qLS of 3ii.O mm.

HF. measur. ..cuts are reported to range between 66-70 mm, usually le: ; than 6l in vocator. 1 1 ■ r ■“ -■ ■ ■ •

Biology: .jthing io known of the habits of this form.

Collection localities indicate its preference to rapidly shifting sand dunes.

Genus Allactaga Cuvier

1036. Allactaga Cuvier, Proc. Zool. Soc. Lond., p. ihl.

Type species: Mus .jaculus Pallas « Dipus sibiricus major Kerr.

The systematic status of southwest Asian populations of

Allactaga spp. have been recently revised by Atallah and

Harrison (1963). These workers reduced A. willimsi Thomas to a subspecific status under A. euphratica Thomas. Four sub­ species were recognized, of which only the nominate form is recorded in the geographical area covered.

Allactaga euphratica Thomas

1881. Alactaga euphratica Thomas, Ann. Mag. Nat. Hist., 8(£):l£.

1897. Allactaga williamsi Thomas, Ann. Mag. Hat. Hist., 20(6):309.

Van,eastern Turkey. (Atallah and Harrison, 1968)

Typo locality: Iraq.

Common names: Euphrates Jerboa, Five-toed Jerboa,

Distribution: Iraq, Jordan, Syria, Turkey, Kuwait, northern

Saudi Arabia, Iran, Transcaucasia and Afghanistan. 236

Only i. 2 nominate form is recorded from the Eastern

Mediterran i Region.

Allactaga euphratica euphratica Thomas

Synonymy tu or species.

Syntypes: 'no, Erl collection no. ^0.10.21.10., an adult male, and no. $0. 0.21.6., an adult female. Both specimens were obtained by the Euphrat j Expedition, 1835-7. They bear no exact date or locality.

General distribution: Iraq, except for tho southeastern comer (east of the River Euphrates south of Baghdad) and Kurdistan, Kuwait, north­ ern Saudi Arabia, Jordan, Syria and southeastern Turkey to Urfa and

Mardin,

Tlissone (1957) obtained specimens from many localities in northeastern Syria in the vicinity of Tell Abiad. Atallah (1967a and c) recorded it from tho Azraq Basin and north of Maan in eastern Jordan, .and Atallah and Harrison (1968) from Palmyra, Syria.

Specimens examined: JORDAN: 5-10 km N.E. of Qasr Amra (1 SA); U km

E.H.E. of Qasr Amra (l SA); 30 km E.Mafraq (1 SA); km N. Maan (l SA);

Aritman (l BM). SYRIA: Karyatein (3 BM; 1 AUB); near Palmyra (2 HARR;

1 SA; 1 AUp); Syrian Desert (2 BM)»

Identification: A medium sized five-toed jerboa; CIS 30-3U mm and HF

5l~6l mm. Color of back Tawny, belly white. Banner on tail has threo definite bands, a short proximal white band, an elongated dark brown band, and a white tip. Ears long, usually over 30 mm in length. 237

Biology: Fj./e-toed jerboas share tho same habitat with Jaculus jaculus, but are far less cornnon than the latter. They are strictly nocturnal becoming active only during the late hours of the night.

They are usually encountered while feeding in low depressions of the hammada, where annual grasses grow.

The burrow system is essentially as described for Allactaga bobrinskli (Gabilaev, 1963:790). The nest is located at the deep­ est point of the main burrow. One or two side escape burrows aro usually constructed but they end blindly a short distance below the surface. A single burrow entrance is maintained, but is kept plugged during the day. A specimen dug out of its burrow around noon on

Kay bth, 1966, near Hafraq was found in a torpid state and took five minutes to become active again.

Two females collected in April held nine and six embryos each, while another was lactating.

Family HYSTRICIDAE Burnett, 1830

Porcupines

This Old World family is represented in the Eastern Mediterranean by one species, Hystrix indica.

Genus Hystrix Linnaeus

17^3. Hystrix Linnaeus, Syst. Nat., 10th ed., 1:96.

Typo species: Hystrix cristata Linnaeus

Early records of porcupines from the Eastern Mediterranean were referred to the African species, H. cristata (Tristram, 1866a,

1368 and I88I1), Ellerman (19h8:766) referred all southwestern 233

Asiatic po apines to H. leucura Icucura, but Elleman and Morriscn-

Scott (1931 519) considered this fom synonymous with H. indica indica»

All recent ocords are referred to the latter species (Bodenheimer,

19£3j Lewif-'j Levris and Atallah, 1967).

Hystrix indica Kerr .

1792. Hyat ix cristata var. indica Kerx’, Anim. Kingd,, p. 213. Based

on C.’.ollie^ Buff on, 1781, 7: pi. 206.

19^1. Hysi-rix indica indica Kerrj Elleman and Morrison-Gcott,

Checklist, p. 3>19.

Type locality: India.

Common names: Indian Crested Porcupine,

Distribution: Southwest Asia eastward to Russian Turkestan and India,

Only two subspecies are recognized by Elleman and Morrison-

Scott (1951:^20); H. indica hirsutirostris Brandt in Transcaucasia and the nominate subspecies throughout the rest of the species range,

Hystrix indica indica Kerr

Synonymy under species.

Type material: Hot examined.

General distribution: Southwest Asia, Afghanistan, India, Ceylon and Nepal,

Tristram (1866a, 1863 and 1881:), Aharoni (1930) and

Bodenheimer (1920, 1933 and 1938) list the porcupine as a member of the Palestinian fauna. Elleman (19lj8) referred tho three forms,

H. hirsutirostris aharoni!, H. hirsutirostris schmidtzi and H. mesopotsmlca, described by Muller to H. leucurua (» H. indica), 239

Lewia, Lew/ and Atallah (196?) reported on several specimens collected in Lebanon,

Specimens omined: 1^. PALESTIMR: Emmaus (Holotype of H. hirsut­ irostris al renii Muller, BZM); Ain Dschacier, N. W. Dead Sea (Holo­ type of H. ili'sutirostris schmidtzi Muller, BZM); Rehoboth (2 BZM);

Jabal Abdu'i Aziz (2 BZM); Ain Fawar, Kadi Kelt (1 BZM); Sllwan

(1 BZM); V’ Li Svenit (1 BZM). LEBANON: Kartaba (1 SA; 2 AUB);

Bekaa Vail. / (l AUB); Nobi Chito (2 AUB).

Identification: Dorsal surface and tail covered vith quills, black and white banded. This is the largest rodent in Asia with GLS more than 120 mm, cheekteeth Ij/lt, hypsodont and rootless.

Biology: Tho Indian crested porcupine is uncommon, but widely distributed throughout the Mediterranean Region and to a lesser ex­ tent in the Steppe Region, It is usually found during tho day occupying small caves or burrows but becoming active at night.

Porcupines are killed by villagers because of crop destruction.

They aro valued as food. Order CAPJJIVORA

Sevent :n opecies of carnivores arc here reported from tlio

Eastern Hedjterranean Region (Table 1,). These however, do not include the following species which aro believed to be extinct in this Region today: a. Ursus a- ctos syriacus Hemprich and Ehrenberg, 1820: The SYRIAN.

BROVHi BEAR has been reported from the mountainous regions north

of Palestine, especially tho Westei*n Uplands, Carruthers (190?

and personal diary) described the habits and habitats of this

species on Kt. Hormon and tho Anti-Lebanon Mountains, A mount­

ed specimen at AUB was obtained by Carruthers north of Zabadani,

Syria in 1905# Schmitz (1912, vide Bodenheimer, 1958) described

the hilling of a bear on Mt, Hermon, tho last authentic record

of the species, Talbot (I960) maintained that ’’Bears are still

•ooportod from the slopes of Mt, Hermon,” This record appears

to bo erroneous, I have asked many people living on the west-

era slopes of Mt, Hermon about sV^'ihg boars, but only a few old

men reported seeing some at the beginning of the century. b, Aclnonyx jubabus (Schrober, 1776): The CHEETAH is undoubtly

extinct in this region. Tristram (1066a and 108U) maintained

that it .was scarce, but more common in tho Ajlun Mountains.

Harrison (1968b) reported that "this is one of the most urgent 2hl

censor .bion problems, perhaps already too late. An inhabitant

of open desert and stoppo, it was last definitely reported in

195>0 i( if, Saudi Arabia (llorrison-Scott, 1951) and may yet exist

in tho emote desert tracts •whore tho frontiers of Jordan, Iraq

and Sai Arabia meet,"

Folio-. Ing is a K3I to species, genera and families of carni­ vores proem utly occurring along the Eastern Mediterranean:

1, Hind feeb with four digits...... *.9

Hind feet with five digits.,.,..•♦*•*.••.•2

2. Total number of teeth 28 or 30; claws retractable; upper camas-

sial tho dominant cheektooth, with only one small functionless

molar behind it; muzzle short; five digits on forefeet, (Family

Felidae)...... 6

Total number of teeth I{2; claws nob retractable; upper carnas-

sial with two well developed molars behind it; muzzle long; four

digits on forefoot, (Family CanidaeJ,,,...,.....,,,,..,.,...,.,3

3. Size largo, GLS usually over lii5 mm; pupils round; postorbital

process thickened, convex dorsally; tail less than 1/2 HB.

(Genus Canla)...... U

Size small, GLS visually less then lUiO mm; pupils elongate; post-

orbital process thin, concave dorsally; tail at least l/2 HB.

(Genus Vulpes)...... 5 2)\2 1(« Very l.- ;o, GLS aiid HF over 10O mj cingulun on outer edgo of If*’

inoonSj ouous...... Genie lupus

Not as arge, GLS and HF less than 17$ mraj cingulum on outer edge of li'- v .11 developed...... Canis aureus

$, Back of oars black or dark brovm; medium size, GIS usually more

than D. > rran, ...... Vulpes vu3.pes

Back of oars came color as head and body; smaller, GLS less than

110 rim ...... Vulpes rucppelli

6. Largo, GLS over 16$ mm; pelage distinctly marked with hollow

rosettes.Panthera pardus

Smail, GLS less than 130 mm; pelage indistinctly marked with bars

and spots. (Genus Felis)...... 7

7. Ears distinctly tufted at tips with long black hair; back of ears

black; upper jaw with three cheekteeth behind the canine......

...... Felis caracal

Ears indistinctly tufted, tufts never over 1$ mm in length; back

of ears not black; upper jaw with four cheekteeth behind the

canine...... 8

8. Small, GLS less than ICO mm; tail longer than 1/2 HB,,

...... Felis silvestris

Large, GLS over 110 mm; tail shorter than 1/2 HB......

...... Fells chaus

9. Snjjital crest well developed, forming a keel like ridge; upper

camassial the dominant cheektooth with one small molar behind

it. (Family Hyaenidae)...... Hyaena hyaena 2h3

Sa$$ital ,’GSt poorly developed; upper carnasaial poorly develop­

ed with no or two molars behind it, the first almost equal or.

larger 11 size than the carnassial...... ,10

10,.One upper molar; T less than 2/3 HB, (Family Kustelidae),,,,,,,12

Two upper nolnrs; T more than 2/3 HB, (Far,lily Viverridae),,,,,, 11

11, Pelage spotted, tail ringed; no bony tube to auditory orfice;

jugal and postorbital processes widely separated,,,,,,.,,,,.,,,,

...... Genetta genetta

Pelage not spotted, tail not ringed; with bony tube to auditory

orfice; jugal and postorbital processes almost encircle tho orbit,

...... Herpestes ichneumon

12, Large forms, CIS over 7!? inn and HB over UCO mm; total number of

teeth 33, 36 or 32...... lit

Small forms, GLS less than 60 mm and HB less than 350 mm; total

number of teeth 3U...«...... 13

13, GLS over lr5 mm; pelage marbled bro-.m and yellow; pterygoid pro­

cess connected with bullae,,.,...... Vormela peregusna

GLS less than hO mm; pelage uniformly brown on the back, belly

white; pterygoid process short, not reaching bullae...,...... ,.,

...... Mustela nivalis iJi, One lower molar; dorsal surface of body and head white, tho rest

of tho body jot black...... Mellivora capensis

Tiro lower molars; color more or less uniform, not as above.,.,..

...... 15

15. Foot webbed, adapted for swimming; premolars h/3» total number

of teeth 36.*...... Lutra lutra

Feet not webbed, plantigrade; prenolars UAi> total number of

teeth 33...... 16 GIS les:.: than 90 innj tail longer or as long as l/2 HB

i-.artes foina

GLS ovoi* 109 mm; tail short, less than l/3 HB .Moles moles Order HZRAGOIDJSA

This ..?riean order ia rapresontcd by only a single living

family (Pr. '^aviidae), three genera and eleven species of hyraxes

(Anderson >' Jones, 1967:363). Of these only Procavia capensis

has ponetr; ::ed into southwestern Asia, where two subspecies have

been recorded} P. c. syriaca (Schreber) from Sinai, Palestine and

Lebanon and P. c. jnyakari Thomas from Saudi Arabia and Yemen.

P. c. syriaca can be easily distinguished from all other mammals in tho area by its vestigial tail, black fleshy pads on

soles of feet, nails rather than claws on toes, and tusk-like

incisors.

216 Order ARTIODACTILA.

Five £> jcics belonging to two fojvriJLies of even-toed ungulates are reported .Cron tho Eastern Mediterranean Region (Table l). Three others. Damn u«mat Caereolus canreolus and Oryx leucoryx, though ccmnon in biblical times and persisting until the first decade of this century have since become extinct. Following is a short sumary of previous records of these three species:

Family Corvidae a, Dana darn mesopotarnica (Brooke, 1875): The FALLOW DEER still

survives in the Zagros Mountains, Iran, However, its former

range included Turkey, Syria, Iraq, Palestine, Jordan and

Lebanon (Hasselquist, 175?J Tristram, 1866b and iQOLtj Bodenhoi-

mer, 1920 and 1958; Harrison, 1963a), Tho following quotation

from lUCH's Red Data Book (1966) summarizes its current status:

uSo rare that until a few years ago this doer was believed to

be extinct. Some 30 of them were discovered in 1957 and 1958

,., in Iran, the rare survivors seemingly owe their safety

to the density of tho forest screens ,,,u b. Capreplus~capreolus coxi Checsman & Hinton, 1923: The ROE DEER

may still persist in northern Syria as it Is still reported as

common 'in southern and eastern Turkey and Kurdistan, However,

no specimens wore taken or seen in the geographical area studied

21:6 2U7

since tl second decade of this century. Previous to this,

records ror.i this area are abundant (Tristram, 1866a, 18?6

and 18S!i 5 Carruthers, 1^09J Podenheimer, 1920j Schnitz,

vide Anc ,, 19^6).

Family Bovidao

0• Oryx IcH'-oryx (Pallas, 1777): The MIABIM OliyX once common through­

out the Arabian Peninsula is now extinct except for scattered herds

in the southeast (Carruthers, 1935J Bodenhcimer, 1958j Talbot, I960;

Mountforb, 1965j Harrison, 1960a and b),

Follo-tring is a KEY to the five species presently occurring'Jji this region:

1. Upper incisors presentj upper and lower canines forming long

tusksj no horns, (Family Suidae...... ,5us scrofa

Upper incisors absent; lower canines incisiforn, upper canines

absent; males and usually females with horns, (Family Bovidae)

...... 2

2. Adult males bearded; adult male horns more than 360 mm long,

curved backwards like a. scimitar; contrasting white and black

markings on legs...... Capra ibex

Adult males without beards; adult male horns less than 320 mm

long, straight or with a slight sigmoid curve; legs uniformly

cream or white colored. (Genus Gagella)...... ,,.3

3. Males with a goitrc-liko throat swelling during breeding sea­

son; horns of adult males close together at the bases (less than

13 mm), widely divergent at tips,.,......

flazolla subgutterosa 2h8 Hales m goiterodj horns of adult males wider apart at their

bases (; eater than 13 inm), less iddelj'- divergent at their

tips...... h

It. Horns oT adult males long (265-310 mm), horn of adult females

also lo' ; (175“2ltO mm); nasal bones overlap with posterior

« eztrenu.J os of the premaxillae for a long distance......

...... Gaaolla dorcas

Horns of adult males short (150-269 mm), horns of adult females

short (05-135 mm)j nasal bones scarcely overlap with posterior

extremities of premaxillae...... Gagella gazella DOMESTICATED AIID IiJTROL'UOED K^HLIIS

Thirtr ;U cpocies of doriasticated and introduced mammals arc presently found along the Eastern Mediterranean Region, These are listed 1 ;low in a systematic order and discussed:

Order Lagomorpha

1, Oiyctolagus cuniculus (Linn.): ETOOPEAN RABBITS are not raised

commercially any place in tho Middle East, They are, however,

commonly raised by villagers and farmers for their own consump­

tion, They are. valued only as a food item, as little value is

placed on tho pelts. Stock populations are brought from Europe.

Order Rodentia

2, Myocastor coypus (Molina): NUTRIAS are abundantly farmed in

Israel for their pelt. A Syrian trapper offered me a specimen

in the summer of 1963 captured along the Jordan River south

of Lake Hule, most probably on escapee from one of the big

ranches,

Strict measures must bo taken to keep this species frai establishing wild populations, which may cause devastating havoc to land and agriculture, as lias been the case in tho United States

(Laycock, 1966),

Order Carnivora

3, Canis faailiaids (Linn.): Most villagers, shepherds and bedouins

keep DOGS for protecting their property and herding livestock.

2U9 2|?0

Tho d.o(j: ire aggressive and uould attach said bite anyone

approach' ng a bedouin canip or a fara house unless restrained

by theii 'iiasters. They are usually kept on leashes during tho

day tirnc and released at night in order to chase away jackals,

foxes, .a’cupinos and mongooses which feed on various vegetables

grown fee human consumption as well as chickens and other farm

animals,

U, Fells c^i’^us (Linn.): CAT3 aixj very common in every village and

city in the Kiddle East. Some are kept as pets, but the majority

are wild living off garbage and small vertebrates. Hatt (1959)

and Atallah (l9

wild cats, Fells silvestris, any place whore the two species

meet.

Order Perissodactyla

5. Equus aslnus (Linn.): DONKEYS are maintained by villagers and

bedouins for riding and as boasts of burden. They are occasion­

ally used in ploughing, especially by poor farmers. Their feces

are collected and dried to bo used for fire or as a fertilizer.

6. Equus caballus (Linn*): Numerous HORSES of different breeds are

maintained by Arabs for riding, pulling carts and as beasts of

burden. They are still used by a section of the Jordanian Army

known as the "Fursan”.

Order Artiodactyla

7. Sus scrofa (Linn.): Though the Middle East is considered to bo

the principal center of pork avoidance in the Old World, one can

still find DOMESTIC PIGS raised throughout this region. They are 251

espccicO.. f common aromd Jerusalem and in Lebanon, two places

whore C: Lstian Arabs predominate over Moslems and Jews*

0. Comelus romsdarins(Linn*): SINGLE-HUMPED C/JIELS have been

completf.. / domesticated. They ran go from North .Africa east­

wards tl ou/jh Arabia to India* Largo numbers are kept by the

bedouins for riding and as beasts of burden. I have seen

herds of almost one hundred individuals feeding in wadi sys­

tems or un pastures near oases throughout tho Desert Region.

Tho Desert Police in Jordan have for many years relied solely on camels for transportation in and out of tho desert, but more recently Land Rovers seem to bo slowly replacing camels.

9. Bubalus bubalis (Linn.): The distribution of the INDIAN or

MATER BUFFALO is restricted by the inability of these animals

to survive without adequate supply of water. They are common­

ly kept by farmers living along water courses or oases. They

are extensively used in faming; also buffalo’s milk is

considered much superior to cow’s milk.

10. and 11. Bos taurua Linn, and Bos indlcus Linn,: DOMESTIC

CATTLE of many breeds are kept abundantly for their milk,

meat and skin. Beef, however, is much cheaper than other

kinds of meaty

12. Capra hirers (Linn.): Large herds of GOATS, 200 and over, can

be seen any place from mountain tops to extreme desert. The

species is considered one of the principal factors contributing

to the present degraded state of plant cover throughout tho

Eastern Mediterranean Region. The intensive grazing pressure 252

of this ;pecies has resulted in complete deforestation vhich

is usual ly follovrcd by soil ei'osion ultimately leading to

bare uiid or a Mediterranean scrub formation.

13. Ovls av:' os (Linn.): SHEEP are somewhat more selective in their

feeding than goats and apparently do not cause as drastic

changes in habitat. They normally feed on grasses and broad­

leaved sihrubs. This latter includes young tree seedlings,

necessary for forest regeneration.

Tho Jordanian Government has recently increased the value of sheep and reduced the value of goats in an attempt to encourage shepherds to roly solely on sheep. Also slaughter houses are asked by the Government to slaughter a greater number of goats than sheep,

Mr. Tsher Qalyubi (pors. comm.) informs me that the Government hopes to reduce the number of goats to less than one third in a ten year period. ZOOGEOGRAPHIC,\L SUM?4ARY

Tho ni ity-four species recorded from tho Eastern Mediterranean

Region can ho treated under five headings as follows: (** indicates

an endemic epeciesj -”<• indicates an endemic subspecies)

I. Element,-j of the PALAEARCTIC REGION:

The seventy-three species representing this region aro hero divided into three components corresponding to the three phyto-

gcographical regions recognized in southwestern Asia (see section

on "General Ecology and Phytogeography" for details).

A. The Mediterranean Region,- Thenammalian fauna character­ istic of this region resembles that of tho Euro-5iborlan Region.

Tho majority of the species appear to have penetrated tho Eastern

Mediterranean Region during pluvial periods of the Pliestocano, and many have since evolved into distinct subspecies.

Following is a list of species and subspecies characteristic of this region:

Erineceus europaens concolor

Crocidura leucodon judaica

Crocidura lasia

Crocidura russula monacha

* Grocidui’a suaveolous portal!

Rhinolophus ferraunequlmm ferrumoqu.-imta

Rhinolophus hipposidercs minimus 2$h

* Rhino] :Aus eui'yale .judaicus

«■ Myoti .nyotis rnacrocephalicua

Kyotir omarpinatus ernarpinatus

Myoti' catjaccinii burcschi

Myoti:. nattorori hovcli

Pipis rollua ninistrellus pipistrellus'

Piois’-rollus kuhlii ikhvanius

Pipistrollus savii caucasicus

Nyctalua noctula Isbanoticus

Eptesicus serotinus serotinus

Plocotus austriacus christiei

Arvicola terrestria

■a Microtus nivalis hermonis

Microtus guenthari guentheri

Apodemus nr/stacinus rtr/stacinus

Apoderrrus sylvattojs tauricus

Ellonys melanurus

Canis lupus pallipes

Vulpes vulpes palaestina

Hustela nivalis

Males males

» Felis sylvestris tristrami

Sus scrofa libycus 255

B. T.: Irano~Tauranian Region.- This region though distinct in terms oO its flora, its fauna is quite poor and many of tho species aiv feared ^d.th either the Mediterranean or the Saharo-

Sindian Re; ons. The seventeen species characteristic of this region can .5 assembled in four groups, according to theii' present di;. lihution, as follours:

(1) . Maiim'irestricted to tha Xrano-Tauranian Regions

Hemicchinus auritus calligoni

Mesocricetus auratus auratus

Microtus sccialis

Meriones vir-e-gradovi

(2) . Mammals penetrating into the Saharo-Sindian Region:

Hemicchinus auritus aegyptius

Gazella subgis.tterosa marica

(3) . Mammals penetrating into- tho Mediterranean Region:

Hyotis blythl omari

Rntesicus bottao innesl

* Sciurus anomalus syriacus

Cricetulus migratorius cinerascens

Sealax ehrenbergi

Dryotnys nitedula phrygius

Canis aureus syriacus

* Vormela peregusna syriaca

-* Martes foina sjcriaca 256 di). Mam- j penetrating into the Mediterranean Region as veil as tho

Saharo-Sind an Region:

Tadarida teniotis rueppelli

Meriones tristrami tristrami

* Hyaena hyaena syriaca

G, The Saharo-Sindian Region: The mammalian fauna characteristic of this region includes species adapted for life in the desert.

Following is a list of these species.

Paraechinus aethiopicus pectoralis

Paraechinus aothiopicus ludlowi

Rhinopoma hardulckei cystops

Rhinopoma microphyllum microphyllum

Asellia trldens tridens

»» Pipistrellus bodenheimeri

Obcnyctcris homprichi .jin

* Gcrbillus dasyurus dasyurus( satisfactorily

penetrated the preceding- two regions)

Corbillus nanus arabium

Gerbillus henleyl mariao

Gerbillus gerbillus bonhotei

Gerbillus allcnbyi

Gerbillus pyramldim

•><■ Sokcetamys calurus calurus

* Meriones tristrami karietini

» Meriones.libycus syrius

Mevionos erassus crassus 257

3H* I'nrionoa sacramenti

Psaimomys obesus obesua

Acomya russettus russatus

## Aconya leirlsl

Jaculua jaculua vocator

* Jaculus jaculus schluoterl

Allactaga euphratxca euphratica

* Vulpes vuluca arabica

Vulpea lateppolll sabaea

Gazolla gasella

Gazella dorcas

II. Elements of the ETHIOPIAN REGION:

Nine species represent this region in the area of study, these are:

Rousettus aegyptiacus aegyptiacus

Taphozoua nudiventria nudlvcntria

Nycteris thebaica thobaica

Rhinolophus clivosus clivosus

Ac cry s diirddiatus dirnidiatus

Mellivora capensis

Genetta genetta t e rrac a anc tao

Horpostis ichneumon ichne\imon

Procavia canensls syriaca III. Elcr, .03 of tho ORIENTAL REGIONS

Only •! o species characteristic of this region have saticfacto'i ly penetrated southx;estem Asia into the Eastern

Kediterror, .n Region, these are:

* Nesokia indica bacherl

Hystrix indica indica

Tiro S', iitional species, Felis caracal and F, chaus, listed in the foilcuing section may belong here.

IV. Pluriregional speciesi

A few species can bo termed pluriregional as they aro shared by two or all three zoogeographical regions listed above, these are

Suncus etruscns etruscus

Rhinolophus blasii blnsii

Miniopterus schroibersi pallidus

-x- Lcnus canons is syriacus

* Lepus capensis arabicus

Lutra lutra seistanlca

Fo^3 caracal schmltzi

* chaus furax

Panthera pardus

-x- Caora ibex nubiana

V. Commensal species:

Rattus rattus (alexandrinus and fru^ivorus color

forms)

Rattus nor/egicus norvegicus

Hus muscuius praetextus BIBLIOGRAPHY

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28o Appendix I. List of measurcnents. Kean (x) and extremes (Kin - Kax) are given for every measurement, when more than three specimens vere examined. Abbreviations of measurements are defined in text (pp. 59-pl). S.D. - Standard Deviation.

SPECIES Ho. TL T KF E CBL A3 33 10 M

INSECTIVORA

Erinaceus euronaeus- 13 •v* 251 25 h7 28 5h.9 32.9 22.8 13.8 143.3 Kin 223 21 hh 23 52.5 30.8 21.5 13.1 bO.O Kax 315 32 53 32 59.3 36.0 25.0 15.8 W.O

Hemiechinus auritus 5 X 188 21 31 36 143.5 25.3 19.1 11.1 33.9 calligoni Min 171 11 29 3h lt2.5 22.5 13.2 10.6 31.5 Max 205 30 3U 39 au.6 27.2 20.1 11.8 35.5 aenyptius 2 182 25 30 33 38.2 21.9 17.0 10.1 29.5- 222 20 3h. h3 145.5 26.2 19.1 10.7 35.0

M Paraechinus aethionicus 6 mr 223 2h 3h 50 50.5 29.6 21.5 ll}.6 33.0 pectoralis Kin 197 20 31 hh h9.h 29.0 20.14 12.9 37.2 Max 255 30 36 55 52.6 30.U 2li.2 15.6 29.2 ludlowi 2 255 32 37 he 51.7 30.0 21.1 12.0 38.2 2'u9 20 35 h3 147.3 28.0 22.1 12 .li 36.8 SPECIES No. TL T T/TL /., HF E GLS BB IC K % Crocidura leucodon Eastern Mediterranean h X 102 3U 33 12 9 20.h 8.9 a.2 12.8 Min 100 31 31 12 8 20.0 8.2 3.5 12.3 Kax 105 38 36 12 10, 21.2 9.7 a.6 13.2

Turkey 11 2 113 37 32 lii _ 20.0 9.0 a.2 12.1 Kin 105 29 27 lii mm 19.1 8.7 h.O ii.a Max 12U hi 37 Hi .5 20.9 9.6 h.h 13.1 S.D. 6.h7 5.53 2.91 0,1k - 0.60 0.28 0.10 o.as

Crocidura lasia 3 122 hi 3U 13 22.0 9.6 a.5 13.7 121 ilO 33 13 10 22.1 a.7 ia.2 13U b2 31 13 12 22.7 10.2 5.a i5.o

Crocidura russula Eastern Mediterranean 52 X 115 li5 39 13 9 19.0 9.0 a.i 12.2 Kin 95 37 35 11 6 17.9 7.9 3.3 11.2 Max 137 53 h2 15 11 20.1 9.1 a.a 13.8 S.D. 7.56 3.56 5.17 0.98 0.95 O.hU 0.31 o.ia 0.52 T$L° T» SPECIES No. TL T ' v.' KF GLS- BB IC M

Crocidura russula Turkey 63 120 h5 33 13 8 19.2 8.8 lt.1 12.0 Min 100 37 3U 11 6 18.0 B,h 3.7 io.5 Max 2li$ 59 li3 15 10 20.7 9.3 h.3 13.3 S.D. 8.82 5.01 2.25 2.37 1.83 0.63 0.2h o.Ht 0.53

Crocidura suaveoleus 1 92 35 38 12.5 9 17.0 7.5 3.7 11.3

A Suncus etruscus 3 79 27 3li O 6 13.1 6.1 2.8 6.3 77 26 3U 6.5 5 13.2 6.0 2.8 7.9 63 26 33 8 5.5 12.6 5.6 3.0 8.1

No. TL T KF FA E GLS BB ZB M

CHIROPTEP.A Rousettus acsyptiacus 27 X 151 15 23 92 23 1:2.9 16.9 25.6 33 .Ii Min 13U 10 20 87 21 ii0,9 16.2 23.6 32.0 T'ax 168 25 26 99 25 U5.1 17.8 27.6 35.2

Rhinooosia microphylluu 1 hB Hi 65 19 20.2 3.2 11.6 Hi.2 YU?

SPECIES Ko. TL T HF FA E GLS BB ZB M

RhinoDona hardwicksi 23 X 125 61 13 56 20 17.8 7.3 10.2 11.9 Min no 52 12 51 17 17.1 6.9 9.8 11.6 Max 1145 70 13 60 22 18.5 7.7 10.6 12.6

Ta-ohozons niidiventris 3 125 36 20 75 23 am - 32 19 76 23 29.0 11.8 16.2 21.3 «• “ 13 75 22 — mm — 21.1

Nycteris thebaica 1 - 52 9 li6 28 18.9 8.2 11.0 12.5

Rhinolo'Dhns I'erruneauinvn 50 X io5 37 12 58 25 23.7 9.0 12.3 16.3 Kin 92 32 n 53 23 22.8 8.5 11.9 15.3 Max 118 U3 n 62 27 2a .6 9.7 12.7 17.1

Rhinoloohns clivosns (from Harrison,196hb:83) 2 33 8 hi 20 20.2 8.1 9.9 13.5 81 23 10 hi 18 19.2 8.1 10.0 13.1 SPECIES No. TL T HF FA E GLS BB ZB M

Ehinolonhus X 6h 26 7 36 16 15.1: 6.2 6.9 9.h hi-Doosidero's Min ^6 23 6 35 H: i5.o 6.2 6.8 9.1 Max 69 29 8.5 33 IS 15.9 6.3 7.0 9.3

Hhinolo-chn" enryale 30 X 7h 27 9 16 22 18.8 8.2 9.3 11.7 Min 63 23 8 15 20 17.8 7.6 8.2 11.2 TO f? Max 8u 31 11 1:8 23 19.9 9.0 10.3 • t

Rhinolophus blasii 16 X 73 27 9 1:6 18 19.3 8.2 8.8 11.9 Min 65 2h 8 1:5 17 19.0 7.8 8.6 11.2 Max 66 31 11 1:7 19 19.6 9.1 9.3 12.2

A.sellia tridens 79 21 8 50 19 19.0 7.3 10.5 12.7 78 21 10 52 21 18.9 7.0 10.2 13.5 8U 22 55 51 21 13.6 7.0 10.3 12.3

mm Kyotis rcyotis 30 X 136 55 16 66 28 25.8 10.2 15.6 19.8 Min 128 h? 13 53 27 2k.9 9.6 i5.o 29.1 Max 116 62 17 71 29 26.8 10.6 16.1 20.2 SPECIES No, TL T HF FA E GLS B3 Z3 K

Kyotis blvthi 30 fc 126 57 13 60 25 22.3 9.5 13.9 17.0 Kin 115 51 12 55 23 21.6 19.2 13 .b 16.2 Kax 139 65 16 63 28 22.8 9.9 15.b 17.6 -

Kyotis emar^inatus 2b X 8b bo 9 bo 15 16.0 7.1 9.5 11.8 __ 85 36 8 37 lb 15.7 7.0 9.1 11 .b Kax 92 bb 11 b3 17 16.5 7.3 9.9 12.2

Kyotis caoaccinii IS X 86 38 12 bl 13 15.3 7.5 3.8 10.7 Kin 80 3b 11 bO 12 lb.9 7.1 8.1 10.2 Kax 91 bo 13 b3 is 15.8 8.0 9.b 11.7

Kyotis nattereri lb X 87 bl 9 39 18 15.8 7.6 9.7 11.6 Kin 8b 37 8 38 16 15.3 7.3 9.3 11.1 j/tcuv 9b b5 10 bO 19 16.2 7.9 10,0 12.1

Pioistrellv.s ■cioistrellus 2 70 30 5 31 12 12 .1 5.9 7.1 8 76 35 6 30 10. 11.6 6.1 7.2 7.6

i 2B7

SPECIES No. TL T HF FA E GLS B3 Z3 M

^inistrellis kuhlii 33 x 31 36 6 3k 12 13.3 6.6 7.7 9.7 Hin 6? 30 5 31 10 12.7 6.3 7.0 9.1 Max 90 k3 7 37 Ik 13.? 7.7 8.6 10.2

Finistrellus savii 3 87 32 6 33 10 •* 82 32 6 32 11 13.1 6.2 — 9.3 83 30 7 32 10 13.7 6.3 •» 9.6

Pinistrellus 1 7k 3k 6 32 12 12.0 5.8 7.2 .3.2 fcodenheineri

Nyctalus noctula U x 131 3k 11 3k 16 19.1 9.9 13.3 15.0 Min 128 $0 11 32 Ik 18.k 9.8 13.0 Ik,2 Max 135 31 12 56 19 19.7 10.1- 13.6 I5.k

Sptesicus serotinus 1 119 kl 10 56 21 22.0 9.6 13.6 16.0

^ rk,,%r* 1 100 ko 9 k2 17 16.6 7.2 10.k 12.0 SPECIES No TL T HF FA E GLS BB Z8 K

Otonycteris henrprlchi $ 131 Sh 13 65 kl 2k .k 10.3 Ik.8 17.2 Min 116 51 11 61 33 23.6 9.9 Ik.3 16.8 Max Ik 3 57 Ik 69 k2 25.k 10.9 15.3 18.1

Plecotus austriacus Ii 92 k6 8 kl 38 17.1 8.1 8.5 11.0 Min 81 kl 7 39 35 16.7 7.6 8.1 10.k Max 97 k8 9 k2 39 17.2 8.k 8.7 11.6 yiniooterus 60 X 113 57 11 k6 12 15.3 7.9 3.k 11.1 schreibersi Hin 10k 51 10 kk 11 Ik.8 7.7 S.l 10.7 Max 128 61 12 k8 13 15.3 8.1 3.7 11.6

Tadarida teniotis 10 X 138 50 11 62 32 2k.1 11.5 13.8 17.0 Min 133 U6 11 59 31 23.2 11.1 13 .k 16.6 Max 1$0 53 13 6k 3k 2k.9 11.3 Ik.2 17.5 SPECIES No. TL T H? E GLS CSL Z3 DM LM M

IAGGMORPHA Lenus c^-osnnis srrriacus 10 X 579 77 133 112 92.7 81.9 a2.6 15.6 16.0 71.9 Min b9o 72 122 100 87.8 77.2 IiO.O 13.8 Ik.8 68.3 Max 620 89 lh5 126 93.2 83.5 kk.9 17.1 17.5 76.3

arabicus 8 X b57 6h lOh 103 77.3 68.2 37.1 13.2 13.6 59.3 Min 393 5h 83 92 65.2 57.0 32.9 12.7 13.2 k3.9 Max U93 75 111 112 83.8 7k.3 39.5 13.8 Ik.2 65.9

RODEJITIA Sciurus ancr.altis 2 3b8 lUo 60 29 Ii7.5 k5.8 26.2 9.k 9.k 31.1 353 iho 56 31 hi.3 k5.5 30.0 9.h 9.3 33.3

Cristnlus nipratoriua Syria ic itebaaon 20 X 116 22 16.£ 17 2k .k 2k.0 13-k 3.3 3.9 16.3 Min 10k 19 16 15 21.8 20.7 11.k 3.3 3.k Ik.2 Max 128 36 17 19 27.0 27.0 Ik.7 3.k k .2 13.5 S.D. 7.36 k.59 0.50 1.16 l.k9 1.58 0.37 0.17 0.20 1.1k SPECIES No. TL T HF E GLS CBL ZB UM m M

Cricetnlus nigratorius Turkey lb 3 1214 23 17 18 2U.7 2ii.2 13 .li 3.3 3.3 15.8 Min 112 22 15 17 23.1 22.2 12.7 3.7 3.7 lh .6 Max lltO 35 19 20 26.2 25.9 lb.3 h.O h.l 17.2 S.D. 3.77 3.39 1.09 1.00 1.20 1.33 0.59 0.10 0.10 0.89

Kesocricetus auratus It X 133 Hi 19 21 3li.8 3ii.8 19.2 5.7 5.8 2h.9 Min 128 13 19 21 33 .li 33.li 18.6 5.6 5.7 22.6 Max 137 15 19 22 36.3 36.3 19.3 5.7 5.8 26.6

Mlcrotus nivalis $0 X 173 57 21 16 23.3 27.6 15.6 6.6 6.3 19.6 Min 2b9 b.9 20 lii 27.1 26.0 H;.8 6.1 5.6 13.3 Max 198 70 22 18 29.9 29.6 17.0 7.1 7.1 21.3

mm Mlcrotus guentheri 120 X 137 25 19 12 27.1 26.7 15.5 6.3 6.3 19.0 Min lilt 20 16 10 2lt.3 23.9 13.3 5.5 5.6 16.9 Max 165 36 21 lii 29.7 29.7 17.8 7.0 7.3 21.6 SPECIES No. TL T HF E GLS CBL ZB UK IK M

Gerbillus dasyunis 100 X 20b 111 2b 13 2b .7 23.2 12.5 3.6 3.6 15.1 Kin 169 91 22 12 21.9 20.3 11.b 3.b 3.3 13 .b Kax 232 129 25 13 26.3 25.0 15.0 b.O b.l 16.2

Gerbillus henlcj'i 11 X lb9 83 19.5 9 21.2 18.9 12.0 2.6 2.6 12.1 Kin 132 73 18 6 20.1 17.9 11.3 2.3 2.h 11.0 Kax 172 92 20 10 22.2 19.7 12.9 2.8 2.7 12.8

Gerbillus gerbillus 6 X 22b 127 30 16 26.8 25.5 15.7 b.O b.O 16.9 Kin 205 112 29 13 25.1 23.9 lb .5 3.8 3.9 15.8 Kax 230 132 31 16 37.8 26.7 16.2 b.2 b.l 17 .b

Gerbillus allenbyi 3 183 110 22 13 26.1 2b.2 3.3 3.6 16.5 21b 122 27 15 2b.2 22.5 lb.7 3.7 3«9 15,0 210 120 26 16 25.9 2b.3 15.1 3.7 36 16.3

Sekeetarr.'o calums 18 X 267 lb6 33 22 33.5 31 .b 18.2 5.1 5.0 20.1 Min 253 131 32 19 32.0 29.7 17 .b 5.0 b.S 13.8 Kax 283 162 33 23 3b .b 32.6 18.9 5.3 5.3 21,2 29%

SPSCISS No. TL T HF E CIS CBL ZB UN I2-' M

Keriones tristrani 100 :< 269 132 314 20 36.3 35.2 19.7 5.3 5.3 22.3 Kin 22U 115 31 13 32.8 31.8 17.7 h.6 a.5 2o.a Kax 312 160 37 22 39.h 38.5 22.8 5.3 5.8 25.0 od

Xerionss libycus hO S 318 150 39 20 39.3 37.14 22.3 5.7 5.9 2)4. H

Kin 270 130 37 18 36.1 3U.1 20.h 5.3 5.3 22. ir\ Kax 351 171 a2 23 h2.1 ho.0 2k.2 6.5 6.L 26.

Ncriones crassns 13 x 2a? 113 32 16 35.8 32.3 19.7 5.7 5.7 21.4 Kin 232 105 31 15 3a.5 30.8 19.0 5.2 5.2 20.6 Max 275 137 3a 19 37.8 3a .a 20.7 6.3 6.7 23.2

Keriones sacrar.onti 2 27a 128 ao 20 39.9 33.a 23.5 6.0 6.1 25.8 290 135 ai 21 ao.5 33.9 22.9 6.2 6.0 25.3

Psamiorryn obesuo h$ x 286 12a 33 16 li3.8 142.5 26.1 6.8 6.0 28.9 Jordan and Palestine Kin 252 105 36 ia 39.5 38.7 23.9 6.3 6.3 214.6 Max 3ii5 150 ai 17 U8.2 l46.U 28.U 7.2 7.5 33.2

Sinai and Egypt 12 2sa 121 37 15 ai.2 39.6 2a.9 6 ° 6.9 27.7 1 .0.11 263 106 35 13 38.a 36.7 23.3 6.6 6.7 25;3 Kax 319 13a 39 17 a3.o a2.2 26.5 7.2 7.2 30.0 SPECIES No. TL T HF E CIS CBL Z3 BH LM K

Scalax ehrariberpri 23 mr 180 23 37.8 37.8 23.0 6.9 7.0 32.5 Kin 15b — 21 - 33.3 33.3 25 .b 5.7 6.1 27.0 Hax 200 -• 26 — bS.8 b8.3 33*8 8.7 3.0 37.1

•w Arjoderrus rrrstacinus 100 Nryv 237 12b 25 20 29.3 28.8 lb.7 b.6 b.6 13.7 i-ian 19b 100 23 16' 27.8 25.3 13.5 b.3 b.< 17.1 Kax 27b 151 27 22 32.3 31.5 15.3 5.0 5.0 20.2

A-oodernus sylvaticus 25 X 133 97 22 17 23.8 22.9 12 .b 3.7 3.7 15.7 Non 1U8 75 20 16 21.5 20.6 11.2 3.5 3.b 13.6 Kax 205 n5 23 18 25.7 25.0 13.b 3.3 b.O 17.2

Rat't'is rattus 19 3C 387 202 36 2b b0.8 bO.3 19.9 6.6 6.6 26.3 Kin 360 183 33 22 33.1 37.5 19.2 6.3 6.3 25.7 Max b22- 212 b2 26 b2.0 bl.3 20.6 7.1 6.5 27.7

Rattns norvoricuo 3 332 167 bO 20 bb.2 bb.2 6.5 7.0 30.6 3b 2 150 bo 21 b6.6 b6.6 27.5 7.b 7.3 30.5 b22 202 bb 21 51.2 b6 26.1 8.2 8.2 3b.b SPECIES No. TL T H? E C-LS CBL ZB UM LK X

Xus rras cuius 70 X Ihh 66 17 13 20.U 19.7 10.7 3.3 3.2 13.3 Kin no U7 16 11 19.2 18.h 9.6 3.2 3.0 12.2 Max 18U 85 19 Ik 22.U 21.3 11.6 3.5 3.U 1U.2

Ac err/'s dimidiatus 70 200 9h 20 19 29.9 23.3 1U.7 li.6 ii.U 19.2 Kin 192 8h 18 17 28.1 26.2 13.9 U.l U.l 17.6 Max 227 111 21 21 32.6 31.3 15.7 i:.9 U.7 20.3 J

Acorrys russatus X — u 151 67 19.5 19 27.2 25.8 13.8 h»9 U.7 17.7

Judean Desert Kin H 1U3 67 19 18 26.8 26.6 13.5 ii.9 U.7 17.1

Max 165 67 20 20 27.8 27.U Hu 3 5.0 U.8 18,1H

Sinai «rw mm HuO lu3 03 11 182 73 20 21 27.5 U.6 17.3

Kin - O 168 56 19 19 26,6 13.3 U.6 u.u 17.0

Max 139 81 21 22 29.1 mm lii.6 5.0 5.0 19.1 H S.D. 5.63 7.2U 0.57 0.U3 0.76 mm o.ko 0.10 0.13 0.58

Ac errors lcr:d.si 22. X 182 67 19 18 28.2 26.7 Hi .3 5.2 5.1 18.6 Kin 177 61 19 17 27.8 25.9 13.9 5.o U.9 18.3 Max 192 7k 20 19 28.6 27.3 lit.6 5.U 5.2 13.9 SPECIES No. TL KF GLS CBL Z3 XTA LM M

Nssokia indica 11 r* 23U 13U 39 17 U6;a 16.2 29.6 9.h 9.2 36.1 Min 195 133 38 17 Ul.7 Ul.7 27.5 8.8 8.7 33.5 Max 263 135 Uo 17 U3.2 U8.2 31.1 10.7 9.9 33.5

mm Elionys nolanurus IS X 218 99 26 27 31.0 30.2 19.1 5.2 5.2 19.9 Kin 195 81 25 2U 29.2 27.5 17 .U h,8 5.0 17.7 Max 2U0 112 27 30 33.0 32.2 20.6 5.U 5.U 21.5 O

Dryonys nitodula 2 195 105 21 13 2U.U 23.6 15.0 3.8 U.o 15. CO 183 100 20 12 2U.2 23.8 15.0 3.6 3.8 15. -

Jacnlus .jacalus vocator ibO X 293 178 60 23 33.0 29.0 21.7 h.7 h.8 20.U Kin 261 159 56 21 31.3 27.9 19.5 li.U U.U 19.U Max 326 197 62 26 3U.0 29.8 22.7 5.0 5.1 21.7 nllactaya ouphratica 15 •v 299 176 55 35 32.0 29.5 22.3 6.U 6.5 22.6 Kin 26? 158 51 30 30.2 27.8 21.0 6.0 6.2 20.7 Max 320 199 61 U2 33.5 31.7 23.2 6.8 6.9 23.6 SPECIES No, TL T HF S GLS CBL ZB UM LM M

Hystrrx 'indica 17 X 761; 167 96 35> 1U2.2 133.8 76.6 30.8 32.6 107.9 Min 660 105 80 30 128.0 121.9 72.2 2h,2 29.8 93.2

Max 790 2l;0 105 h6 li;8.7 119.9 60.9 32.8 39.3 119.9vn»