A Contribution to the Knowledge of the Herpetofauna of Jordan. VI. the Jordanian Herpetofauna As a Zoogeographie Indicator

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©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at HERPETOZOA 9 (1/2): 71 - 81 Wien, 30. Juni 1996

A contribution to the knowledge of the herpetofauna of Jordan. VI. The Jordanian herpetofauna as a Zoogeographie indicator

Beitrag zur Kenntnis der Herpetofauna von Jordanien. VI. Die jordanische Herpetofauna als zoogeographischer Indikator

AHMAD M. DISI

KURZFASSUNG

Zoogeographisch gesehen erweist sich die jordanische Herpetofauna als heterogen, indem sie vier bio- geographischen Regionen entstammt (Orientalische, Paläarktische, Saharo-Sindische, Äthiopische Region) und vier unterschiedliche Ökozonen umfaßt. Dreiundneunzig Arten und Unterarten von Amphibien und Reptilien sind für Jordanien bekannt, die unterschiedliche Verbreitungsmuster zeigen. Die Mediterrane Ökozone beher- bergt die höchste Anzahl von Arten (35), gefolgt von der tropisch-sudanensischen Penetrationszone (16) und der Badyiah Ökozone (12). Jordanien ist nicht durch landschaftsmorphologische Grenzen von den umgebenden Ländern abgetrennt, sodaß keine diesbezüglichen Isolationsmechanismen wirksam sind, was zum Fehlen von Endemismen geführt haben dürfte. Immerhin gibt es im Ostmediterran als Gesamtheit drei Endemiegebiete: die Badyiah, die sich Jordanien mit Syrien, dem Irak und Saudi Arabien teilt, den Nordteil der Mediterranen Ökozone, der den Libanon, Israel, Syria und Jordanien einnimmt, sowie ein Gebiet, das den Südwesten Jordaniens, Israel and die Halbinsel Sinai umfaßt. Einige paläarktische Arten wie Coluber schmidti und C. ravergieri sind Relikte postglazialer Perioden und überlebten an ökologisch empfindlichen Refugialstandorten. Das Verbreitungsbild einiger Arten (Bufo viridis, Mauremys caspica rivulaia, Natrix tessellata, Coluber ventromaculatus, C. jugularis asianus, Walterinnesia aegyptia, and Vipera palaestinae) ist maßgeblich durch das Klima und anthropogene Einflüsse geprägt.

ABSTRACT

Zoogeographically, the Jordanian herpetofauna is heterogeneous, originating from four biogeographical regions (Oriental, Palearctic, Saharo-Sindian, and Afrotropical) and occupying four different ecozones. Ninety- three species and subspecies of amphibians and reptiles are recorded from Jordan, showing different distribution patterns. The Mediterranean ecozone harbours the highest number (35) of recorded species, followed by the Sudanian Tropical penetration zone (16), and the Badyiah ecozone (12), respectively. Jordan is not separated by natural boundaries from the surrounding countries, which prevents the operation of isolation mechanisms, and seems to result in the absence of endemism. There are, however, three areas of endemism within the Eastern Mediterranean region taken as one whole unit: the Badyiah shared by Syria, Iraq, Saudi Arabia and Jordan, the northern part of the Mediterranean ecozone shared by Lebanon, Israel, Syria and Jordan, and southwest Jordan, Israel and Sinai. Some Palearctic species such as Coluber schmidti and C. ravergieri are relicts of the postglacial period and continue to survive in refugal enclaves with delicate ecological patterns. The distribution of some species, such as Bufo viridis, Mauremys caspica rivulata, Natrix tessellata, Coluber ventromaculatus, C. jugularis asianus, Walterinnesia aegyptia, and Vipera palaestinae, has been greatly influenced by climatic as well as anthropogenic changes.

KEY WORDS Jordan: zoogeography, herpetofauna, relict and endemic species

INTRODUCTION

The herpetofauna of Jordan consists and distribution, providing limited infor- of 93 species and subspecies belonging to mation pertaining to the animals' ecology 23 (5 amphibian and 18 reptilian) families. (HART 1891; BARBOUR 1914; SCHMIDT, Most previous studies on the herpetofauna 1930; PARKER, 1935; HAAS, 1951, 1952; of Jordan dealt mainly with systematics WERNER 1971, 1988; Disi 1983, 1985, ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at

72 A. M. DlSI

1987, 1991, 1993; DlSI & al. 1988; AL- ranean region has witnessed intensive geo- ORAN & AMR, 1995). : . logical events that are reflected in paleo- Distribution, composition and abun- biological effects (TCHERNOV 1988). The dance of the biota in the Mediterranean biotic configuration of the region was region are. the product of long episodes of modified by extensive plate tectonics changes resulting from evolution, adapt- which split Jordan and Arabia from ation and migration (ROBINSON 1972). Palestine and Sinai, and the latter from These processes were greatly shaped and Africa. Furthermore, climatic changes oc- affected by several factors: curred resulting from the Pleistocene Jordan is a. crossroad of four zoogeo- glaciation (ROBINSON 19y2). The anthro- graphical realms (Afrotropical, ' Saharo- pogenic factors are man's introduction of Sindian, Oriental, Palearctic). The south of exotic species on the one hand and Jordan is a part of the Levantine Land- destruction on the other. All above factors bridge. Moreover, Jordan comprises a Have contributed to the heterogeneity of great variety of plant communities, and the herpetofauna of Jordan (Disi 1987). geomorphological as well as climatic con- .-.-•;,, In the present study, the zoogeo- ditions which allowed the formation of graphy of Jordan is discussed, based on the four ecozones in this restricted area, (AL- distributional patterns of the herpetofauna. EISAWI Ì985). Hence* Jordan appears to be Endemic and relict species, and the major one, of thè richest and most heterogenous environmental changes affecting the abund- natural areas in the temperate region of thé ance and distribution of reptiles and am- world. In ; addition, the Eastern Méditer^ / phibians are enumerated and highlighted.

MATERIALS

The present work is based on reptile M., Germany; Staatliches Museum für and; amphibian specimens collected be- Tierkunde, Dresden, Germany; Zoologi- tween May 1977 and July 1995. The bulk sche Staatssammlung, München, Germany; of this collection is deposited at the Jordan Zoologisches Forschungsinstitut und Mu- University Museum, Department of Bio- seum Alexander Koenig, Bonn, Germany, logical Sciences, Amman. Additional spe- Zoologisches Museum der Alexander- cimens have been examined from the fol- Humboldt-Universität, Berlin, Germany; lowing museums: Jordan Natural History Museum d' Histoire Naturelle,. Genève, Mûsîèum,' Yarmouk University, Irbid, Jor- Switzerland; Natural History Museum, Ba- dan;, British Musuem of Natural ^ History, sel, Switzerland; Naturhistorisches Muse- Londonv England, Natur-Museum und For- um Wien, Austria. schungsinstitut Senckenberg, Frankfurt a.

ZOOGEOGRAPHY OF AMPHIBIANS AND RBPTILÊS OF JORDAN. ••-/• >•-•:•;' ' • • • A REVIEW -, -•• ••-•.•••..> •. (figure 1, table 1).; • • .

Mediterranean Ecozone • • true concerning rainfall, and vegetation types and density. The highest rainfall is The Mediterranean ecozone extends found in the Ajlun mountains (600 mm from Urn Qias in the north through the annually), an ecozone characterized by Ajlun mountains, the hill regions of Amon terra rosa and/or rendzina soil. These soil and Moab to the Edom mountains region types are the richest in the area, suited for in the south, including the upper Jordan cultivation and dense well-grown forests. Valley. Many creeks and wadies drain this The vegetation here is essentiality formed hilly region from east to west and lead to by the forests: pine, evergreen oak, the Jordan River/the Dead Sea and Wadi deciduous oak and Juniperus. Araba. The southern mountains are higher A total of 35 species of amphibians than thè northern ones, while the reverse is and reptiles were found to inhabit this ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at Herpetofauna of Jordan VI. The Jordanian herpetofauna as a Zoogeographie indicator 73 ecozpne (2 amphibians, 2 tortoises, 13 liz- 'Wadi Araba and Southern Desert ards and 18 snakes); 32 of these species originate from the Palearctic fauna, and 3 Wadi Araba and the Southern Desert are of Arabian and Saharo-Sindian origin of Jordan are also known as Sudanian- (table 1). Although the Mediterranean eco- Tropical penetration zone. This ecozone zone represents less than 20% of thé total extends from Al-Karama in the north area of the country, it harbours 37.6 % of through the Dead Sea depression, Wadi the known amphibian and reptile species. Araba (Jordan Rift Valley) as far as The abundance of vegetation, forested southern Jordan. It is bordered by Wadi mountains, wadi systems along with vari- Rum and the Al-Disi region. This ecozone ous types of microhabitats permitted sev- is characterized by the lowest annual rain- eral species to seek refuge in this ecozone. fall (50 mm), and highest mean annual Plentiful insects, rodents, and vegetation maximum air temperature (Wadi Araba, are available all over the year as a source 30.8 °C) in the country (National Atlas of of food. Jordan, Part I, 1986). The soil of the Southern Desert is primarily of granite Badyiah stone, aeolian sand, and sand dunes, while that of Wadi Araba is mainly alluvial sand The Eastern Desert 'Badyiah', is part and gravel carried by flashing floods from of the Syrian Desert and forms the centre the surrounding uplands. The. wadies end- of the Saharo-Syrian-Sindian Desert. The ing in Wadi Araba build up wide alluvial Arabic term 'Badyiah' is more descriptive fans (ATALLAH 1977). Qas (Playa) are than 'desert', because the area is capable of found in both southern Jordan and Wadi supporting vegetation and animal life. Araba, and formed where a single basin Nevertheless, the limiting factor here is the receives surplus rain water and silt drained availability of water. This ecozone extends from the surrounding regions (wadies) as a through Syria, Iraq, Jordan, and northern result of erosion. These Qas are deprived Saudi Arabia. The Badyiah soil is primar- of vegetation and fauna. The vegetation of ily formed of limestone with flints scat- this ecozone is related to tropical elements tered over the surface, or basalt pebbles such as Accada sp., Calotropis procera and boulders that resulted from volcanic persica and Haloxylon persicum (AL-EI- outcrops centered around Jabal Druz (Al- SAWi 1985). . Arab) (ATALLAH 1977). This ecozone The Jordan Rift Valley is part of the forms the largest surface area of Jordan. Syrian-African Rift. It is regarded as a Annual rainfall is around 100 mm and passageway between Eurasian and African most, of the vegetation cover is concen- faunal elements, and a gateway for north- trated in wadiesXAL-ElSAWi 1985), ward and southward dispersal (TcHERNOV Twelve species of reptiles (9 lizards & YOM-TOV 1988). The southern end of and 3 snakes) are restricted to this ecozone this ecozone is part of the Levantine Land- (table 1). Ten are of Arabian and two are bridge between the Mediterranean area and of Saharo-Sindian faunal origin. Notwith- the Arabian desert and represents a 'Bio- standing the vastness of the Badyiah com- geographic Filter' (POR 1987). The width pared with the other ecozones, the number of this filter has been varying in accord- of species present here comprises only ance with the fluctuating geological and 13% of the total herpetofaunal species of ecological changes. POR (1987) also indi- Jordan. This ecozone is known for its cated that the Levantine Landbridge can be scarce and scattered vegetation as well as regarded as the most prominent focal point its flat open surface with a reduced number in the distributional history of the modem of microhabitats. Reptiles are usually con- terrestrial biota of the globe, because it fined to narrow strips where vegetation connects the major old world continental prevails along wadi beds or depressions landmasses. The Levantine Landbridge is that receive more water in the form of the amalgamation place of biota of the floods than do other surrounding areas. Palearctic, Afrotropical, Saharo-Arabian, For no obvious reason, the False Horned and Oriental zoogeographical realms, re- Viper, Pseudocerastes persicus fieldi is sulting in high species diversity, compared to other temperate regions of the same size restricted to the Badyiah ecozone. ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at 74 A. M. Disi

in the world. TcHERNOV (1988) stated that GASPERETTI (1988) accepted the Saharo- confrontation of the Palearctic and Afro- Syrian-Sindian region as a biogeographical tropical communities did not cause any realm of its own, which has a very large extinction or any massive biotic turn- distinctive fauna separated from the Pale- over in the southern Levantine meeting arctic and Afrotropical realms. JOGER area. ARNOLD (1987) mentioned that chan- (1987) suggested that the Irano-Turnian ging of the topography and expansion of province (Iran) should be classified as a the desert area southward resulted in sepa- subregion of the Saharo-Syrian-Sindian re- ration of North Africa and Arabia. Some alm, and this is followed in the present species crossed this barrier (JPtyodactylus study. TCHERNOV (1988) indicated that the hasselquistii, Acanthodactylus boskianus, only opportunity for a large-scale success- Spalerosophis diadema, Mesalina olivieri, ful colonization by the Saharan psammo- Varanus griseus, Psammophis schokari, phile taxa might have taken place in the Malpolon moilensis, Lytorhynchus diade- last (Wurm) glacial. ma). Moreover, ARNOLD (1987) stated Sixteen reptile species inhabit Wadi that Wadi Araba might have acted as a Araba and southern Jordan (eleven species barrier to desert forms in the moist pluvial of lizards and five of snakes). One species periods. This barrier resulted from topo- is of Palearctic origin, while seven, six, graphic and other abiotic factors. Some and two are of Saharo-Sindian, Arabian, North African species failed to pass be- and Afrotropical origin, respectively. This yond Wadi Araba (Stenodactylus stheno- type of multiple reptilian origin accords dactylus, Sphenops sepoides, Psammophis with the explanation of the geomorpho- aegyptia, Uromastyx ornata), while some logical changes and mass movements of the Arabian forms did not cross Wadi Araba in biota (KOSSWIG 1955; ARNOLD 1987; JO- the opposite direction (Stenodactylus gran- GER 1987; POR 1987; TCHERNOV 1988). diceps, Acanthodactylus grandis, A. robu- Three species of snakes (Telescopus dhara, stus, A. schmidti, Mesalina brevirostris). Atractaspis microlepidota engaddensis, Some other species (Bunopus tuberculatus, Echis colorât us), inhabiting this biotope Acanthodactylus opheodurus, Atractaspis succeeded in reaching the Mediterranean microlepidota engaddensis, Coluber ele- ecozone. gantissimus, Eirenis coronella, Pseudocer- astes persicus fieldi) extended beyond Wa- Irano-Turanian ecozone di Araba into southern Israel and Sinai but did not cross the Suez Canal (ARNOLD Phytogeographically, the Irano-Tur- 1987). anian ecozone is a strip of variable width surrounding all the Mediterranean ecozone DE JONG (1976) indicated that the except its north. Mean annual rainfall is Sahara is 'a life zone', a biome and not a less than 150 mm. The Irano-Turanian re- transitional zone between the warm-tem- gion is not distinguised zoogeographically perate part of the Afrotropical and the from adjacent bioclimatic ecozones, just Palearctic realms. He also indicated that forming a transitional zone between the the Saharan belt acts as a barrier prevent- Mediterranean and the surrounding eco- ing faunal exchange between the two above-mentioned biogeographic zones. In zones. SAINT GIRONS (1982) pointed out that - from a zoogeographical point of his discussion of the zoogeography of the view - the Irano-Turanian ecozone in Israel Middle East, KOSSWIG (1955) raised the is of disputed validity. problematic issue of defining natural boundaries of the region, and considered it This ecozone does not have its own a subjective judgement. TCHERNOV (1988) entity since it does not possess a specific stated that the isolation of the Sahara as a fauna as do other ecozones in Jordan. The unique biogeographic province started dominant type of soil here is loess and cal- during the Pliocene, and that there is no careous. general agreement about the boundary Common species between the Palearctic and the Saharo- Arabian-Sindian realm. Twenty-seven species inhabit more ARNOLD (1987), JOGER (1987), and than one ecozone (twenty species occupy ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at Herpetofauna of Jordan VI. The Jordanian herpetofauna as a Zoogeographie indicator 75

, Y

Mediterranian ! • \;:^..^ Irano-Turanian

z.ro\• ... •.•••••.•'••;•••.•','• •. • • • •' >• [|—f^The Badyiah P771 Wadi Araba & Southern Jordan ^•^ (Sudanian Penetration) lOOKm

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Fig. 1: Main ecozones of Jordan. Abb. 1: Die Hauptökozonen Jordaniens. ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at 76 A.M. DIS!

two ecozones). This may be due to (1) ad- topes of Jordan. Three are anuran amphib- aptability of the species to colonize dif- ians and their presence is affected by the ferent habitats with different biotic and availability of water bodies (table 1). The abiotic factors (Disi 1987); (2) the borders other four, {Hemidactylus turdcus, Chal- between the four ecozones in Jordan are cides ocellatus, Mabuya vinata, Psamm- not well defined, thereby allowing inter- ophis schokari), attain a wide but discon- mingling of faunal elements among ad- tinuous distribution in Jordan, their pres- jacent ecozones. KOSSWIG (1955) indicated ence being favoured by moist terrain; they that there are difficulties in distinguishing are often found in and around farms, or natural biogeographic barriers within the near human settlements. Similar observ- Levantine region. Some species, extend ations were made by ARNOLD (1987) in their range through the Irano-Turanian Arabia, and he suggested that this scattered ecozone (Malpolon monspessulanus insig- type of distribution resulted from man's nitus), or, through wadi systems pene- introduction. According to JOGER (1987) trating into the Mediterranean ecozone Hemidactylus turcicus,. and H. flaviviridis (Echis coloratus, Atractaspis microlepi- are widely distributed and are easily intro- dota engaddensis). (3) Climatic conditions duced by man, SCHATTI & GASPERETTI are not stable (AL-ElSAWl 1985), and (4) (1994) indicated that Psammophis schokari anthropogenic factors may also play a role is particularly abundant in some areas and in this respect. Some species such as Co- is possibly the most frequently encountered luber ventromaculatus, Walterinnesia ae-> snake in Arabia where it occupies different gyptia and Bufo viridis are described as habitats. WERNER (1987) stated that C. agricultural followers (Disi 1987, 1993). ocellatus and P. schokari have been found Seven species are found in all bio- throughout Israel. /> ,

RELICT SPECIES

The distribution of amphibians and Al-Arab (Al-Druz) in Syria, and in Leba- reptiles in the Middle East compared to non as well (BISCHÖFE; pers. comm.). that in Jordan suggests that one amphibian Further taxonomic work is needed to clar- (Pelobates syriacus) and three reptile spe- ify the status of L. laevis and L. kulzeri. cies (Lacerta kulzeri, Coluber ravergieri, The populations of Coluber ravergieri and C. schmidti) are relict species. P. syriacus C. schmidti are relicts from postglacial was collected from two localities in north- periods and occupy a habitat which is ern Jordan, which are isolated from the ecologically similar to that of previous nearest populations in Syria. This species (postglacial) conditions. Both species are continued to survive in a very delicate in complete isolation in the remote moun- ecological habitat which necessitates im- tainous areas of the southern Jabal Al- mediate protection. There are debates on Arab, separated by a considerable distance the classification of L. kulzeri, but the from the known ' area of distribution population of this lacertid lizard from Pet- (ScHÄTTi & AGASIAN 1985; DISI 1993). ra is clearly different from those in Jabal

ENDEMISM

KOSSWIG (1955) stated that there are ing countries by natural barriers, which difficulties in distinguishing natural bio- prevents the operation of isolation mechan- geographic barriers within the Levant. He isms and seems to result in the absence of also indicated that the Levantine region is endemism. There are, however, three areas a transitional zone between the Palearctic of endemism in the Eastern Mediterranean and the Sahara-Arabian desert belt, with region taken as a unit: (1) An area with complex mosiac patterns of distribution. Micrelaps muelleri, Typhlops simoni, and Jordan is not separated from- the surround- Chalcides guentheri which is shared by ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at Herpetofauna of Jordan VI. Jordanian herpetofauna as Zoogeographie indicator 77 northern Jordan, Israel, Lebanon, and the illustrated by the presence of Coluber ele- western part of Syria. A fourth endemic gantissimus; (3). the Syrian Desert exem- species, Cyrtopodion amiciopholis, has. plified by the presence of Stenodactylus been reported only, from Mt. Hermon grandiceps, Acanthodactylus robust us, À. (WERNER 1988; SIVAN & WERNER 1992); , tristrùmi,'Tràpélus' persicus fieldi, and Lau- (2) Southern Jordan bordering th^e north- :daìdastellio'picea. ;' west of Saudi Arabia and southern Israel,

;' t IMPACT OF ENVIRONMENTAL CHANGES ON LOCAL RANGE EXPANSION AND SPECIES ABUNDANCE Man-induced environmental changes, for housing and, agricultural purposes through agricultural projects in the eastern which led to decreasing biodiversity and Badyiah, southern Jordan and Wadi Araba, extermination of breeding sites; of Pelo- resulted in the expansion of the known bates syfiäcus. Moreover, extensive use of range of distribution for certain species insecticides, herbicides, and many other (Bufo viridis, Coluber ventromaculatus, chemicals as fertilizers or for sterilization Walterinnesia aegypiia), called agricultural of soil in agricultural projects, permits followers (Disi 1987). Meanwhile polluted transfer of these chemicals through' food water bodies in the Jordan Valley increased chains; several trophic levels may be ;af- the population density and distribution area fected by this. In addititon, high cöncen- of Mauremys caspita riyulàtàì On the trations rnay b^ accumulated in the bodies other hand, for other species, ecological of organisms occupying higher trophic changes have resulted in a rapid decline of levels.. Ini the Jordan Valley, I saw dead natural habitats due to overgrazing, urban rodents out of their burrows on several oc- expansiòn, unorganized construction plans, casions, and most probably chemicals have mismanagementent of the highlands, and been the, cause,of death., PÀZ (.1987) re- fire. All these factors led to deforestation, ported that the number of breeding pairs of soil erosion and desertification, resulting Grey. Heron (Ardea cinerea)' was; -high in in a patchy fragile ecocomplex, and affect-—, 1953 ancl ,1954, but'the; population was ing the abundance of lacertid lizards in the exterminatedin 19,04, apparently as a re- Edom Mountains. JONES (1981) indicated suit of the birds feeding on poisoned ro- that grazing induced, structural changes in dents. T^e amount of herbicides, insecti- vegetation which reduced lizard abundance , cides and, fertilizers imported and used in and diversity. And WERNER (1988) stated 1995 was several/times higher "than that that decrease of woodland areas in Israel used in the sixties and- seventies- of the was followed by decreasing abundance ,of , centu|y; Jordan' bias increased its' use of Laceriq\ media israelica. Another example, t ',•fertilizers height-fold since ,1978.; (HAT- of habitat destruction in Jordan is land use OUGH-BORAN & Disi 1995).. .•.,:W..-.-v.i':.\

•••/- -,- -J- v*:*i; ,:''•Ì:Z'-'.:.': L,..>••>'•'.'•.'•/ •••r/i '.x ;-.::Vi '•.•;Viy»i".>\r«'i

<} + CONCLUSIONS.^:' • *->?-^'s •>«• ; '•*»* •^-^.•..-

In. spite of extensive work on the spe©iesi'.,o.f«iüievgen^ herpetofauna of Jordan in the last decade ^/iara'complex^ 'Malpolònymòilensis and (DISI, Ï983, 1985, 1987, 1991, 1993; AMR Mä0SfcnîJlß^siila^y0ij0tust as & al., J994; AL-ORAN & AMR, 1995), th^Wlt^'nie^^^^^^^M^/aJWa, zoogeographical overview is still limited. and the Rana cf. ridibunda complex; .^ This isiïhe first detailed study dealing with {^} T/l7-'- ^Kiterpretatiön'öf-fine patterns of the herpetogeography of Jordan,, and, al- thevîiodl^Ë^tëra^^ though ^certain aspects may still need fur- ^-|iòrir',hasj s'beenr^Êy^pî^r^y^riASêquate ther elabortation, the following conclu- herjsetological/ inventories^fronr some sur- sions niay be drawn: ^ rûndiä^.j^^tn^^lh'^^^^g^tmg that lf:[ Several taxonomic^problems are moTt^fi^lßf'^^ a still unsolved as exemplified by. certain^ mòre^"aTOurateianâ,4ëtoifâ understanding ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at

78 A.M. Disi

Table 1 : Distribution of the herpetofauna in the main ecozones in Jordan. The indication of the zoogeographical affinity is based on data from ARNOLD (1987), JOGER (1987), WERNER (1987, 1988), LAMARCHE & CLEMENT (1988), GASPERETTI (1988), and SCHÂTTI & GASPERETTI (1994). Ecozones: M - Mediterranean, B - Badyiah, W & S - Wadi Araba and southern Jordan. Zoogeographical affinity: A - Arabian, Af - Afrotropical, P - Palearctic, SS - Saharo-Sindian. (+) - Species penetrates into the periphery of the indicated ecozone. Tab. 1: Verteilung der Herpetofauna auf die Hauptökozonen Jordaniens. Die Angabe der zoogeo- graphischen Affinität basiert auf Daten aus ARNOLD (1987), JOGER (1987), WERNER (1987, 1988), LAMARCHE & CLEMENT (1988), GASPERETTI (1988), und SCHÄTTI & GASPERETTI (1994). Ökozonen: M - Mediterran, B - Badyiah, W & S - Wadi Araba und südliches Jordanien. Zoogeographisches Element: A - Arabisch, Af - Äthiopisch, P - Paläarktisch, SS - Saharo-Sindisch. (+) - Die Form dringt in die Peripherie der Ökozone ein. Taxon Ecozone / Ökozone Zoogeograph. M B W&S affinity / Affinität Amphibia Urodela Salamandridae Triturus vittatus (JENYtiS, 1835) + P Amphibia Anura Pelobatidae Pehbates syriacus BOETTGER, 1889 + P Bufonidae Bufo viridis LAURENTI, 1768 + + + P Hylidae Hyla savignyi (AVDOUW, 1827) + + + P Ranidae Rana cf. ridibunda PALLAS, 1771 + + + P Reptilia Testudines Cheloniidae Chelonia mydas (LINNAEUS, 1758) Gulf of Aqaba Eretmochelys imbacata (LINNAEUS, 1766) Gulf of Aqaba Dermochelyidae Dermochelys coriacea (LINNAEUS, 1766) Gulf of Aqaba Emydidae Mauremys caspica rivulata (VALENCIENNES, 1833) + P Testudinidae Testudo graeca terrestris FORSKAL, 1775 + P Reptilia Squamata Sauria Gekkonidae Bunopus tuberculatus BLANFORD, 1874 + A Cyrtopodion kotschyi orientate STEPANEK, 1937 + P Cyrtopodion scaber (HEYDEti, 1827) + A Hemidactylus turcicus turcicus (LINNAEUS, 1758) + + + P Pristurus rupestris BLANFORD, 1874 + SS Ptyodactylus guttatus HEYDEH, 1827 + + A Ptyodactylus hasselquistii ÇDOHNDORFF, 1798) + + SS Ptyodactylus puiseuxi BOUTAN, 1893 + + A Stenodactylus doriae (BLANFORD, 1872) + A Stenodactylus grandiceps HAAS, 1952 + A Stenodactylus sthenodactylus (LICHTENSTEIN, 1823) + SS Tropiocolotes steudneri (PETERS, 1869)(?7". nattereri STEIND., 1901) + Af Chamaeleonidae Chamaeleo chamaeleon recticrista BOETTGER, 1880 + + Af Agamidae Laudakia stellio brachydactyla (HAAS, 1951) + P Laudakia stellio picea (PARKER, 1935) + A Laudakia stellio stellio (LINNAEUS, 1758) + P Phrynocephalus arabicas ANDERSON, 1894 + A Pseudotrapelus sinaitus HEYDEN, 1827 + + A Trapelus pallidus haasi (WERNER, 1971) (+) + + A Trapelus persicus fieldi (HAAS & WERNER, 1969) + A Uromastyx aegyptia microlepis BLANFORD, 1874 + + SS Lacertidae Acanthodactylus boskianus (DAUDIN, 1802) (+) + + SS Acanthodactylus grandis BOULENGER, 1909 + A Acanthodactylus opheodurus ARNOLD, 1980 (+) + + A Acanthodactylus pardalis (LlCHTENSTHN, 1823) + SS Acanthodactylus robustus WERNER, 1929 + A Acanthodactylus schmidti HAAS, 1957 + A Acanthodactylus tristrami (GÜNTHER, 1864) + A Lacerta kulzeri MÜLLER & WETTSTEIN, 1932 + P Lacerta laevis GRAY. 1838 + P_ ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at Herpetofauna of Jordan VI. The Jordanian herpetofauna as a Zoogeographie indicator 79

Table 1: Continued. Tab. 1: Fortsetzung. Taxon Ecozone/Ökozone Zoogeograph, M B W&S affinity / Affinität Lacerta media israelica PETERS, 1964 P Mesalina brevirostris microlepis (ANGEL, 1936) A Mesalina guttulata guttulata (LICHTENSTEIN, 1823) SS Mesalina olivieri schmidti (HAAS, 1951) SS Ophisops elegans blanfordi SCHMIDT, 1939 P Ophisops elegans ehrenbergi WlEGMANN, 1835 P Ophisops elegans elegans MENETRIES, 1832 P Scincidae Ablepharus kitaibelii (BlBRON & BORY, 1833) P Chalcides guentheri BOULENGER, 1823 A Chalcides ocellatus (FORSKAL, 1775) SS Eumeces schneiderii pavimentata (G.-ST.HlLAIRE, 1827) SS Eumeces schneiderii schneiderii (DAUDIN, 1802) SS Mabuya vinata (OuviER, 1804) ?SS(A) Ophiomorus latastii BOULENGER, 1887 P Scincus scincus meccensis WlEGMANN, 1837 A Sphenops sepsoides (AUDOUIN, 1827) SS Anguidae Ophisaurus apodus PALLAS, 1772 Varanidae Varanus griseus (DAUDIN, 1802) ss Reptilia Squamata Ophidia Leptotyphlopidae Leptotyphlops macrorhynchus (JAN, 1860) ss Typhlopidae Typhlops simoni (BOETTGER, 1879) p Typhlops vermicularis MERREM, 1820 p Boidae Eryxjaculus (LINNAEUS, 1758) ss Colubridae Coluber elegantissima (GÜNTHER, 1878) A Coluber jugularis asianus (BOETTGER, 1880) P Coluber nummifer REUSS, 1834 P Coluber ravergieri MENETRIES, 1832 P Coluber rhodorachis (JAN, 1865) ? Coluber rogersi (ANDERSON, 1893) P Coluber rubriceps (VENZMER, 1919) P Coluber schmidti NlKOLSKY, 1909 P Coluber ventromaculatus GRAY, 1834 A Eirenis coronella (SCHLEGEL, 1837) SS Eirenis decemlineatus (DUMERIL & BlBRON, 1854) P Eirenis lineomaculatus SCHMIDT, 1939 P Eirenis rothi JAN, 1863 P Lytorhynchus diadema (DUMERIL & BlBRON, 1854) SS Lytorhynchus kennedyi SCHMIDT, 1939 ss Malpolon moilensis (REUSS, 1834) ss Malpolon monspessulanus insignitus (GEOFFROY, 1827) p Micrelaps muelleri BOETTOER, 1880 p Natrix tessellata (LAURENTI, 1768) p Psammophis schokari (FORSKAL, 1775) ss Rhynchocalamus melanocephalus (JAN, 1862) p Spalerosophis diadema cliffordi (SCHLEGEL, 1837) ss Telescopus dhara (FORSKAL, 1775) ss Telescopus fallax syriacus (BOETTGER, 1880) p Telescopus nigriceps (AHL, 1924) Atractaspididae A Atractaspis microlepidota engaddensis HAAS, 1950 ( Elapidae Af Walterinnesia aegyptia LATASTE, 1887 ( Viperidae Cerastes cerastes (LINNAEUS, 1758) SS Echis coloratus GÜNTHER, 1878 ( SS Pseudocerastes persicus fieldi SCHMIDT, 1930 A Vipera palaestinae WERNER, 1938 ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at 80 A.M. DlSI

of the true distribution of various species. problems to be faced. 3. There are debates concerning 5. Some authors hâve considered the zoogeographical stem of some species the generic level, while others have util- e. g., Chamaeleo chamaeleon, Pseudotra- ized the specific level for their Zoogeo- pelus sinaitus and others, which may hin- graphie, conclusions. However,_ the species der any comprehensive conclusion being • ; is an objective and real unit which exists in drawn. .;.'(:-> ! nature, and is the basic one*in taxonomy 4. Complexity of geological and and evolution, while other (higher) taxo- climatic effects in the Eastern Mediterra- - , nomic units are arbitrary. So, the species is nean areas and absence of fossil records of the only unit that should be utilized in zoo- the present reptile and amphibian groups geographical studies. :;.:v..' (ARNOLD 1987; TCHERNOV 1988) are other

ACKNOWLEDGMENTS 'I'Would like to thank Dr. Z. AMR and Dr. W. staff of the museums indicated in this article for their BÖHME (Bonn) for reading the manuscript and for valuable help/ Finally,''I'Kami grateful,"tó all who making instructive comments on the text. I would helped in collecting the specimens used in this study, also like to emphasize my great indebtedness to the ' ' ' ••'- '• • ' ' • , .

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DATE OF SUBMISSION: February 27th, 1996 Corresponding editor: Heinz Grillitsch

AUTHOR: Prof. Dr. AHMAD M. DlSI, Department of Biological Sciences, Faculty of Science. The University of Jordan, 11942, Amman, Jordan.