The Bulletin of the Philadelphia Herpetological Society, Vol. 31
Gekkonid Lizards from Five Quarters Meet in Israel
Yehudah L. Werner
Israel harbors up to 12 species of Gekkoninae: Of 3 Mediterranean taxa, 2 Occur only in northern Israel; of 6 desert species, 4 are Saharan and inhabit SW Israel; 1 is local in the Levant; and 1 species is endemic to Mt. Hermon. Aspects of their biology are discussed. Additional species in Sinai and Jordan are mentioned.
What is the significance of the term "Five Quarters?" Gekkonid species actually converge on Israel from a greater number of geographical sources but the author was reluctant to overload the title of this paper with words. Israel is the Levant area, which forms a bridge between Europe, Asia and Africa. It has long been lauded as a meeting point for the floras and faunas of these continents or, in other words, of these different biogeographical regions (Haas & Werner, 1969; Fig. 1). Moreover, landscape and climate are extremely varied within Israel. These conditions greatly contribute to the richness of the local reptile fauna in addition to the relative richness which is normal for a small country (Smith, Chizar, & Smith, 1981). Whereas Israel makes do with only six or seven species of amphibians (European species which find their distribution limited by desert; Mendelsohn & Steinitz, 1945); it boasts some eighty species of reptiles (marine turtles excluded). To this figure the family Gekkonidae has contributed nine to twelve species, depending on who does the counting. All of these species belong to the subfamily Gekkoninae.
These species vary in their distributional range within Israel, in their abundance and ease of collection within that range, and in their hardiness in captivity. Hence, the extent of my experience with them is variable. During the past thirty-plus years, certain species have continuously inhabited the many cages in my "gecko room" at the Hebrew University of Jerusalem's Department of Zoology. Some individuals have survived up to twelve years and have often bred. These species have yielded much information, both from the field and laboratory work of students as well as from my own studies. At the other extreme are species which I have observed in the field only occasionally and which I have rarely exceeded a year's survival under our laboratory conditions. In the gecko room light and temperature are well regulated but humidity is not, as the Jerusalem climate is quite dry. For financial reasons, other insects only rarely augment the staple food of housefly maggots. This review is based on locality data (Werner, 1977) of specimens in the Zoological Museums of the Hebrew University of Jerusalem and Tel Aviv University and on the investigations of students, associates, and myself (quoted or still under way).
The Gekkoninae of Israel
One species, which occurs commonly throughout Israel, is the Turkish gecko, Hemidactylus turcicus (Fig. 2). Elsewhere it inhabits the Mediterranean countries (including those to the North, East, and South of Israel) and in recent decades has been spreading in the Western Hemisphere, including the southern U.S. (McCoy, 1970). In Israel it inhabits rocks, earth bank crevices, tree trunks and houses. Formally the populations of both the mesic (relatively humid) habitats in northern Israel and those in the southern deserts belong to the same subspecies, H. t. turcicus, but a current computerized investigation of the biometrical geographical variation of reptiles in the Near East (Kosswig, Laavee & Werner, 1976) has revealed morphological differences between northern and southern Israeli populations. Biological differences have also long been suspected: Eggs belonging to the Jerusalem population incubated at thirty degrees centigrade require 60 days to hatch, while those from the desert possess a longer incubation time. Frankenberg (1982) has recently studied the complex vocal behavior of H. turcicus.
On tree trunks in the northern part of the country (the southern limit is Hebron) is found the superbly camouflaged Bent-toed gecko, Cyrtodactylus (formerly Gymnodactylus) kotschyi orientalis (Werner, 1966; Fig. 3). This is the only Israeli gecko, which may clearly be considered an envoy from the north (Fig.4). Strangely, it is mostly restricted to an underdeveloped habitat of tree trunks in Israel. However, the many subspecies inhabiting islands and countries of southeastern Europe and southwestern Asia north of Israel, where tree trunks abound, live in rocks, ruins, etc., despite possessing the same tree-bark coloration. Again, vocal interaction between the sexes has been reported by Frankenberg (1978a).
Species Inhabiting Mt. Hermon, Jordan and Sinai:
An unusual member of the genus Cyrtodactylus, C. amictopholis (Fig. 5), was discovered on Mt. Hermon after this area became accessible to Israeli investigators. It appears to be endemic to this mountain at altitudes above fifteen hundred meters (Hoofien, 1967: Werner & Avital, 1980). This is a smaller gecko in which each clutch contains one egg rather than the usual two.
A third member of the genus, C. scaber, narrowly skirts Israel. It occurs in Jordan to the east and in the western ports of Sinai to the west (Werner, 1973: 1982). Other geckos which approach but do not enter Israel are in Jordan and include Asaccus elisae (formerly Phyllodactylus elisae), Pristurus flavipunctatus quweirensis, and Stenodactylus grandiceps (both described by Haas and Werner, 1971). In western Sinai are found Hemidactylus flaviviridis and Tarentola mauritanica (Werner, 1973, 1982). Among the Sinai species, Cyrtodactylus scaber and Hemidactylus flaviviridus must have reached the port towns on the Gulf of Suez through shipping long the Red Sea.
Several other gecko species enter Israel from the southeast or the southwest. Unlike Hemidactylus and Cyrtodactylus these are almost entirely non-climbing inhabitants of desert grounds. Blandford's ground gecko, Bunopus (formerly Alsophylax) blanfordi (Fig. 6) and the Persian ground gecko, Ceramodactylus (or Stenodactylus) doriae, occur in Israel only in the Arava Valley (from the Dead Sea to the Gulf of Elat; Fig. 4). This is the western limit of their respective ranges in southwestern Asia. Of the two, Cermodactylus is the ground-dweller, showing in extreme measure the supposedly related adaptations of locomotion: a belly held off the ground by long limbs, an almost undetachable tail, and particularly large eyes (Werner, 1965b, 1968, 1969: Werner & Broza, 1969).
Of the Saharan geckos entering the southern deserts of Israel from the west, Petrie's gecko, Stenodactylus petrii closely resembles Ceramodactylus morphologically and in being strictly arenicolous. The two species are never sympatric. The less arenicolous elegant gecko, Stenodactylus sthenodactylus (Fig. 7) with similar but less extreme adaptations, occurs throughout Israel's southern deserts, coexisting with both of the above taxa. Its range extends to the north of the desert, along the light soils of the coastal plain in the west and the warm Jordan Valley to the east (Fig. 4). Ceramodactylus and the two Stenodactylus species are the main forms observed at night on the desert sands. The pigmy Saharan, Tropiocolotes steudneri (Fig. 8) of Sphaerodactylus size, lives in a variety of habitats throughout the extreme desert. This species also lays a single egg per clutch. One of the persisting riddles of Israeli herpetology is whether the reputedly more clearly cross-banded T. nattereri is actually T. steudneri or a sibling species.
The Fan-toed Geckos:
All of the above description patterns are more or less readily explainable and are paralleled by other animals and plants. The Fan-toed geckos belonging to the genus Ptyodactylus (Fig. 9) present a more complex picture. These are the largest and most conspicuous geckos in Israel, occupying rocks, cliffs, stone walls, and crevices in hard earth banks (Werner, 1965a). The genus Ptyodactylus is widespread throughout the northern half of Africa (as far south as Cameroon), east to Iran, Pakistan, and Arabia, and north to northern Syria. They are easily recognizable as belonging to the genus and are characterized by bifid expanded fans at the tips of all toes. This is the only Israeli genus besides Hemidactylus possessing expanded toes. Taxonomy within the genus, although not completely neglected (Loveridge, 1947), is in need of revision.
Three forms of Ptyodactylus are readily distinguishable within Israel. Usually considered subspecies, they have all been regarded as synonyms by Wermuth (1965). However, after personally studying many aspects of these forms for 30 years (as has Frankenberg for the past 12 years), it appears that the only condition they lack for being accepted as full-fledged species is a formal presentation of their case (which is in preparation). P. hasselquistii puiseuxi occurs in the north (Galilee; Fig. 10). It is diurnal (Frankenberg, 1978,1979b) with relatively small eyes (Werner, 1969) and sexual dichromatism. The males are beautifully blackish with white dots and the females gray with lighter and darker dots, although both sexes vary their darkness according to the temperature. Most of the rest of the country is inhabited by P. h. guttatus which is more or less crepuscular, basking especially in the cooler hours or seasons (Frankenberg, 1978b, 1979b). In obvious relation to its being less diurnal are the significantly larger eyes compared to P. h. puiseuxi (Werner, 1969; Fig. 10). These two species of Ptyodactylus, along with Cyrtodactylus kotschyi, are the only geckos in Israel which regularly bask in direct sunlight. The other species tend to be nocturnal or crepuscular, the exact timing of main activity shifting with the seasons (Frankenberg, 1978b). P. h. hasselquistii inhabits warm canyons near Elat and around southern Sinai and is strictly nocturnal (Frankenberg, 1978b, 1979b). The eyes are not significantly larger then the P. h. guttatus (Werner, 1969). The sexes differ only slightly and are colored with orange cross bands. In general habits this is a much more slender, long-snouted, and long-tailed animal than the two other forms.
The different daily activity times typical of the three Ptyodactylus (Fig. 10) may actually stem from a similarity in temperature requirements. From preliminary work (Werner & Goldblatt, 1978) it is suspected that they all have similar preferred body temperatures of approximately thirty-two degrees Centigrade. In cooler Galilee this is attained by diurnal basking (P. h. puiseuxi ), in the warmest locations by nocturnal activity (P. h. hasselquistii) and in the intervening areas by intermediate behavior (P. h. quttatus). By now more than a direst adaptation to temperature has evolved, for the animals also retain their characteristic daily cycles under standardized uniform conditions (Frankenberg, 1978b, 1979b).
The three Israeli as well as other Ptyodctylus species share many characteristics. They (males especially) are loudly vocal, and the females adhere the eggs to a rock substrate, the only Israeli geckos to do so (Werner 1965a). The three forms differ not only in body proportions, details of scutellation, coloration, extent of sexual dichromatism, and timing of daily activity, but also in additional behavior and/or physiological traits. In relation to eye size and activity time, they differ in the response of the pupil to varying light intensity (Frankenberg, 1979a) and in visual acuity at varying light intensities (Frankenberg, 1981). The males of the three species emit distinctly different multiple-click calls (Frankenberg, 1974; Werner, Frankenberg & Adar, 1978). As yet unstudied is the observation that individuals of P. h. hasselquistii are much more nervous and aggressive than the others.
In summary, the geckos of Israel instructively demonstrate the heterogeneous composition of the local herpetofauna: northern, Mediterranean forms include Cyrtodactylus kotschyi and Ptyodactylus h. puiseuxi, whereas Hemidactylus turcicus is circum-Mediterranean. Arabian forms are Bunopus blanfordii and Ceramodactylus doriae. The two Stenodactylus and the one or two Tropiocolotes are Saharan; Ptyodactylus h. hasselquistii is tropical-African (Sudanese or Ethiopian). P. h. guttatus, ranging from Egypt to Jordan, appears to be a local product. Loveridge (1947) implied it might be a hybrid between the other two forms. A true endemic species is Cyrtodactylus amictopholis.
I am indebted to the countless individuals who donated specimens; to successive animal caretakers, lastly Daniel Guata and the late Shemuel Levy; to successive personal assistants, lastly Nehama Rivlin; to successive curatorial assistants, lastly Iris Jacobi; to successive departmental photographers, lastly A. Niv; to Prof. H. Mendelsshon and Dr. M. Goren of Tel Aviv University, for free use of the collection under their care; to my home University, for its support through thirty years (the recent forced cessation of this support now threatens the live collection); and to the Society for the Study of Amphibians and Reptiles as well as Tnuva Dairy (Jerusalem) for their recent support of the live collection.
Frankenberg, E. 1974. Vocalization of males of three geographical forms of Ptyodactylus from Israel (Reptilia: Sauria: Gekkoninae). J. Herpetol. 8: 59-70
Frankenberg, E. 1978a. Calls of male and female tree geckos, Cyrtodactylus kotschyi. Isr. J. Zool. 27L 53-56
Frankenberg, E. 1979b. Interspecific and seasonal variation of daily activity times in gekkonid lizards (Reptilia, Lacertillia). J. Herpetol. 12: 505-519.
Frankenberg, E. 1979b. Influence of light and temperature on daily activity patterns of three Israeli forms of Ptyodactylus (Reptilia: Gekkoninae). J. Zool. Lond. 189:21-30.
Frankenberg, E. 1981. Optomotor responses of three congeneric gekkonid lizards having different daily activity times. J. Zool. Lond. 193: 147-156.
Frankenberg, E. 1982. Vocal behavior of the Mediterranean house gecko, Hemidactylus turcicus. Copeia. 1982: 770-775.
Frankenberg, E. and Y. L. Werner. 1979. Lunar cycle affecting daily activity rhythm in a gekkonid lizard, Ptyodactylus Isr. J. Zool. 28: 224-228.
Haas, G. and Y.L. Werner. 1969. Lizards and snakes form southwestern Asia, collected by Henry Field. Bull. Mus. Comp. Zool. 138: 327-406.
Hoofien, J.H. 1967. Contributions to the herpetofauna of Mount Hermon, No. 1. Cyrtodactylus amictopholis n. sp. (Sauria, Gekkonidae). Isr. J. Zool, 16: 205-210.
Loveridge, A. 1947. Revision of the African lizards of the family Gekkonidae. Bull. Mus. Comp. Zool. 98:1-469, pls. 1-7.
McCoy, C.J. 1970. Hemidactylus turcicus (Linnaeus) Mediterranean gecko. Catalog of American Amphibians and Reptiles, Vol. 87: 1-2.
Mendelssohn, H. and H. Steinitz. 1945. Contributions to the ecological zoogeography of the amphibians in Palestine. Rev. Fac. Sci. Univ. Istanbul., Ser. B, 9: 289-298.
Smith, H.M., Chizar, D. and R.B. Smith,. 1983. Comparison of regional taxonomic densities. Bull. Phila, Herp. Soc., 29: 9-13.
Wermuth, H. 1965. Liste der rezenten Amphibien und Reptilien: Gekkonidae, Pygopodidae, Xantusiidae. Das Tierreich. Lfg. 80, de Gruyter, Berlin.
Werner, Y.L. 1965a. Ueber die israelischen Geckos der Gattung Ptyodactylus und ihre Biologie. Salamandra. 1: 15-25.
Werner, Y.L. 1965b, The comparative caudal osteology of some gekkonid lizards from Israel. Isr. J. Zool. 14: 286-301.
Werner, Y.L. 1966. Cytodactylus Kotschyi orientalis in Israel. Lacerta. 24: 94-96.
Werner, Y.L. 1968. Regeneration frequencies in geckos of tow ecological types (Reptilia: Gekkonidae). Vie et Milieu, Ser. C19: 199-221.
Werner, Y.L. 1971. Lizards and snakes from Transjordan, recently acquired by the British Museum (Natural History). Bull, Br. Mus. Nat. Hist. 21: 213-256, 6 pls.
Werner, Y.L. 1973. The reptiles of the Sinai Peninsula. Introductory notes for students in the course "Introduction to the Fauna (Fishes, Amphibians and Retiles)." Dept. of Zoology, The Hebrew University of Jerusalem, Jerusalem, Israel. IV + 47pp. 1 pl. (Hebrew text, English abstract and key to Eremias; maps with scientific names).
Werner, Y.L. 1977. Manual mapping of locality records - an efficient method. J. Bioqeoqr. 4: 51-53.
Werner, Y.L. 1982. Herpetofauna survey of the Sinai Peninsula (1967-1977), with emphasis on the Saharan sand community, in Herpetological Communities: a Symposium of the Society for the Study of Amphibians and Reptiles and the Herpetologist's League, August 1977, ed. N. Scott. U.S. Fish and Wildlife Service, Wildlife Research Report 13: 153-161.
Werner, Y.L. and E. Avital. 1980. The herpetofauna of Mt. Hermon and its altitudinal distribution. Isr. J. Zool. 29: 192-193.
Werner, Y.L. and M. Broza. 1969. Hypothetical function of elevated locomotory postures in geckos (Reptilia: Gekkonidae). Isr. J. Zool. 18: 349-355.
Werner, Y.L., Frankenberg, E. and O. Adar. 1978. Further observations on the distinctive vocal repertoire of Ptyodactylus hasselquistii cf. hesselquistii (Reptilia: Gekkonidae). Isr. J. Zool. 27 176-188.
Werner, Y.L. and A. Goldblatt. 1978. Body temperature in a basking gekkonid lizard, Pytodactylus hasselquistii (Reptilia: Lacertilia: Gekkonidae). J. Herpetol. 12:408-411.
Werner, Y.L., Kosswig, C. and D. Lavee. 1976. Computerized mapping of animal distribution and of morphological variation. Isr. J. Zool. 25: 201-202.
Photos and maps linked in this document: Fig.1, Fig.2, Fig. 3, Fig. 4, Fig. 5, Fig. 6, Fig. 7, Fig. 8, Fig. 9, Fig. 10.
Published in the Bulletin of the Philadelphia Herpetological Society [ISSN 0553-9587] - Volume 31, (dated 1983-84, printed 9/87) © 1987-1997 Philadelphia Herpetological Society. All rights reserved. May be reproduced for individual educational use. All other uses require author or society permission in advance. Links to this page welcome.
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