A Guide to Reptiles and Amphibians of Egypt

A Guide

to the

Reptiles and Amphibians

of Egypt

A Guide

to the

Reptiles and Amphibians

of Egypt

Sherif Baha El Din

The American University in Cairo Press

Cairo — New York

Copyright © 2006 by

The American University in Cairo Press

113 Sharia Kasr el Aini, Cairo, Egypt

420 Fifth Avenue, New York, NY 10018

www.aucpress.com

All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the publisher.

Photographs and illustrations by the author unless otherwise indicated.

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Contents

Acknowledgments

Foreword

Introduction

The Egyptian Setting

Evolution of Egypt’s Landscape and Its Herpetofauna

The Contemporary Egyptian Herpetofauna

Biogeography

Species Richness and Distribution

Herpetofaunal Habitats of Egypt

Desert Habitats

The Western Mediterranean Coastal Desert

Wadis and Mountains

Gravel Plains

Sand Dunes

Oases

Wetland Habitats

Littoral Habitats

Marine Habitats

Arable and Urban Landscape

Conservation

Methodology

Notes on the Text

Classification and Nomenclature

Species Included

Species Maps

Abbreviations

Amphibia

Order ANURA (Frogs and Toads)

Bufonidae

Genus Bufo

Bufo dodsoni, Dodson’s Toad

Bufo kassasii, Nile Valley Toad

Bufo regularis, Egyptian Toad

Bufo viridis, Green Toad

Ranidae

Genus Ptychadena

Ptychadena mascareniensis, Mascarene Ridged Frog

Ptychadena schillukorum, Sudan Ridged Frog

Genus Rana

Rana bedriagae, Levant Green Frog

Rana saharica, Saharan Green Frog

Hylidae

Genus Hyla

Hyla savignyi, Savigny’s Tree Frog

Reptilia

Order SQUAMATA: Suborder SAURIA (Lizards)

Gekkonidae

Genus Cyrtopodion

Cyrtopodion scabrum, Keeled Rock Gecko

Genus Hemidactylus

Hemidactylus flaviviridis, Yellow-bellied Gecko

Hemidactylus foudaii, Elba Gecko

Hemidactylus mindiae, Mount Sinai Gecko

Hemidactylus robustus, Red Sea Gecko

Hemidactylus sinaitus, Speckled Gecko

Hemidactylus turcicus, Turkish Gecko

Genus Pristurus

Pristurus flavipunctatus, Semaphore Gecko

Genus Ptyodactylus

Ptyodactylus guttatus, Spotted Fan-toed Gecko

Ptyodactylus hasselquistii, Egyptian Fan-toed Gecko

Ptyodactylus ragazzii, Sahelian Fan-toed Gecko

Ptyodactylus siphonorhina, Saharan Fan-toed Gecko

Genus Stenodactylus

Stenodactylus mauritanicus, Northern Elegant Gecko

Stenodactylus petrii. Sand Gecko

Stenodactylus sthenodactylus, Elegant Gecko

Genus Tarentola

Tarentola annularis, Egyptian Gecko

Tarentola mauritanica, Moorish Gecko

Tarentola mindiae, Qattara Gecko

Genus Tropiocolotes

Tropiocolotes bisharicus, Bishari Pigmy Gecko

Tropiocolotes natterei, Natterer’s Pigmy Gecko

Tropiocolotes nubicus, Nubian Pigmy Gecko

Tropiocolotes steudneri, Steudner’s Pigmy Gecko

Tropiocolotes tripolitanus, Tripoli Pigmy Gecko

Agamidae

Genus Agama

Agama spinosa, Spiny Agama

Genus Laudakia

Laudakia stellio, Starred Agama

Genus Pseudotrapelus

Pseudotrapelus sinaitus, Sinai Agama

Genus Trapelus

Trapelus mutabilis, Changeable Agama

Trapelus pallidus, Pallid Agama

Trapelus savignii, Savigny’s Agama

Genus Uromastyx

Uromastyx aegyptia, Egyptian Dabb Lizard

Uromastyx ocellata, Ocellated Dabb Lizard

Uromastyx ornata, Ornate Dabb Lizard

Chamaeleonidae

Genus Chamaeleo

Chamaeleo africanus, African Chamaeleon

Chamaeleo chamaeleon, Common Chamaeleon

Lacertidae

Genus Acanthodactylus

Acanthodactylus boskianus, Bosc’s Lizard

Acanthodactylus longipes, Long-footed Lizard

Acanthodactylus pardalis, Egyptian Leopard Lizard

Acanthodactylus scutellatus, Nidua Lizard

Genus Latastia

Latastia longicaudata, Long-tailed Lizard

Genus Mesalina

Mesalina bahaeldini, Mount Sinai Lizard

Mesalina brevirostris, Short-snouted Lizard

Mesalina guttulata, Small-spotted Lizard

Mesalina martini, Red Sea Lizard

Mesalina olivieri, Olivier’s Lizard

Mesalina pasteuri, Saharan Lizard

Mesalina rubropunctata, Red Spotted Lizard

Genus Ophisops

Ophisops elbaensis, Mount Elba Snake-eyed Lizard

Ophisops elegans, Elegant Snake-eyed Lizard

Ophisops occidentalis, Western Snake-eyed Lizard

Genus Philochortus

Philochortus zolii, Grass-loving Lizard

Genus Pseuderemias

Pseuderemias mucronata, Anseba Lizard

Varanidae

Genus Varanus

Varanus griseus, Desert Monitor

Varanus niloticus, Nile Monitor

Scincidae

Genus Ablepharus

Ablepharus rueppellii, Rueppell’s Snake-eyed Skink

Genus Chalcides

Chalcides ocellatus, Ocellated Skink

Chalcides cf. humilis, Saharan Ocellated Skink

Genus Eumeces

Eumeces schneideri, Golden Skink

Genus Scincus

Scincus scincus, Sandfish

Genus Sphenops

Sphenops sepsoides, Audouin’s Sand-skink

Genus Trachylepis

Trachylepis quinquetaeniata, Bean Skink

Trachylepis vittata, Brideled Skink

Order SQUAMATA: Suborder SERPENTES (Snakes)

Typhlopidae

Genus Ramphotyphlops

Ramphotyphlops braminus, Flower-pot Blind Snake

Genus Typhlops

Typhlops vermicularis, Greek Blind Snake

Leptotyphlopidae

Genus Leptotyphlops

Leptotyphlops cairi, Cairo Worm Snake

Leptotyphlops macrorhynchus, Beaked Worm Snake

Leptotyphlops nursii, Anderson’s Worm Snake

Boidae

Genus Eryx

Eryx colubrinus, Theban Sand Boa

Eryx jaculus, Javelin Sand Boa

Colubridae

Genus Dasypeltis

Dasypeltis scabra, Egg-eating Snake

Genus Dolichophis

Dolichophis jugularis, large Whip Snake

Genus Eirenis

Eirenis coronella, Crowned Peace Snake

Genus Hemorrhois

Hemorrhois algirus, Algerian Whip Snake

Hemorrohis nummifer, Coin-marked Snake

Genus Lycophidion

Lycophidion capense. Wolf Snake

Genus Lytorhynchus

Lytorhynchus diadema, Diademed Sand Snake

Genus Macroprotodon

Macroprotodon cucullatus. Hooded Snake

Genus Malpolon

Malpolon moilensis, Moila Snake

Malpolon monspessulanus, Montpellier Snake

Genus Natrix

Natrix tessellata, Diced Water Snake

Genus Platyceps

Platyceps florulentus, Flowered Racer

Platyceps rogersi, Spotted Racer

Platyceps saharicus, Saharan Cliff Racer

Platyceps sinai, Sinai Banded Snake

Genus Psammophis

Psammophis aegyptius, Saharan Sand Snake

Psammophis punctulatus, Speckled Sand Snake

Psammophis schokari, Schokari Sand Snake

Psammophis sibilans, African Beauty Snake

Genus Rhynchocalamus

Rhynchocalamus melanocephalus, Palestine Black-headed Snake

Genus Spalerosophis

Spalerosophis diadema. Diadem Snake

Genus Telescopus

Telescopus dhara, Egyptian Cat Snake

Telescopus hoogstraali, Hoogstraal’s Cat Snake

Elapidae

Genus Naja

Naja haje, Egyptian Cobra

Naja nubiae, Nubian Spitting Cobra

Genus Walterinnesia

Walterinnesia aegyptia, Black Desert Cobra

Atractaspididae

Genus Atractaspis

Atractaspis engaddensis, Palestine Burrowing Asp

Viperidae

Genus Cerastes

Cerastes cerastes, Horned Viper

Cerastes vipera, Sand Viper

Genus Echis

Echis coloratus, Burton’s Carpet Viper

Echis pyramidum, Carpet Viper

Genus Pseudocerastes

Pseudocerastes persicus, False Horned Viper

Order CROCODYLIA (Crocodiles)

Crocodylidae

Genus Crocodylus

Crocodylus niloticus, Nile Crocodile

Order TESTUDINES (Turtles and Tortoises)

Testudinidae

Genus Testudo

Testudo kleinmanni, Egyptian Tortoise

Cheloniidae

Genus Caretta

Caretta caretta, Loggerhead Turtle

Genus Chelonia

Chelonia mydas, Green Turtle

Genus Eretmochelys

Eretmochelys imbricata, Hawksbill Turtle

Genus Lepidochelys

Lepidochelys olivacea. Olive Ridley Turtle

Dermochelyidae

Genus Dermochelys

Dermochelys coriacea, Leatherback Turtle

Trionychidae

Genus Trionyx

Trionyx triunguis, Nile Soft-shelled Turtle

Emydidae

Genus Trachemys

Trachemys scripta, Red-eared Terrapin

Gazetteer of Egyptian Localities

Glossary

Bibliography

Index of Scientific Names

Index of English Names

Acknowledgments

Many friends and colleagues have helped me throughout this project in various ways. I would like to thank them all: Moustafa Fouda, Waheed Salama, Omar Attum, Ahmed Riad, Tom Cole, Richard Hoath, Mohamed Kassas, Alan Resetar, Hala Barakat, Esther Wenman, John Grainger, John and Patsy Gasperetti, Gabriel Mikhail, Francis Gilbert, Barry Clarck, John Gerhart, David and Heddi Merrie, Ibrahim Helmy, Ahmed Disi, Loutfy Boulos, Mohamed Ibrahim, Kaori Odani, Jeff and Linda Millington, John and Susan Warberton, Jose Rosado, Harold Voris, Steve Goodman, Nick Arnold, Mohamed Gad, John Poynton, Pierre-Andreà Crochet, George Zug, Patrick Osborne, Zuhair Amr, Jens Vindum, Van Wallach, Ivan Ineich, Rainer Günther, Ulrich Joger, Alan Leviton, Osama Ghazali, Saleh Mahmoud, Wolfgang Böhme, Yehuda Werner, Jiři Moravec, Herman in den Bosch, Fekri Hassan, Colin McCarthy, Hymen Marx, Lyle and Gloria Rosenzweig, Alison Clarck, Max Kasparek, Stephen Spawls, Wolfgang Wüster, Roberto Sindaco, Samy Zalat, Gian Lorenzo, Dave Ferguson, Michael Pearson, Mohamed Sharif Khan, Tim Wacher, Ali Hamdy, John McEachern, Philippe Geniez, Moustafa Saleh, Samir Ghabbour, Ahmed Gamal, Hans Schneider, Jim Buskirk, Paul Williams, Wael Abed, Amr Sharawi, Jacopo Sinibaldi, Neil Hewison, Mohamed El Saghir and all the rangers in the protected areas of Egypt.

I am especially indebted to my wife and family who have been patient and unrelentingly supportive.

Special thanks go to Omar Attum, Roberto Sendaco, Gabriel Mikhail, Mindy Baha El Din and Daniel Müller who provided invaluable photographic materials.

Foreword

The knowledge of our biodiversity resources is still far from complete. Understanding and accounting for our biodiversity is essential if we are to develop meaningful and effective strategies and actions for their future conservation. The present treatise is an excellent example of biodiversity survey, which updates and consolidates our knowledge about an important part of the Egyptian fauna, the herpetofauna. This fauna deserves special attention particularly the amphibia, turtles and tortoises, which are among the most threatened groups of biota worldwide.

The introductory chapter provides a synoptic outline of the scene of Egypt: its diverse deserts, oases, Nile Valley and Delta, wetlands, and coastal lands. This sets an ecological background of the survey that follows. The main bulk of this book provides an authoritative systematic survey including keys for identification. For every species the text provides detailed diagnostic features with notes on habitat and ecology, conservation status, geographic range, distribution maps in Egypt, and an excellent selection of photographs.

This is a most welcome contribution to the natural history of the Egyptian biodiversity, which provides students, researchers, conservation technicians, and the layman with a valuable and up to date reference.

This outstanding publication deserves well-earned acclaim. The author, Dr. Sherif Baha El Din, and the publisher, The American University in Cairo Press, are to be commended for this valuable contribution to the Egyptian natural history studies.

Mohamed Kassas

Emeritus Professor of Plant Ecology

University of Cairo

Former President of the World Conservation Union (IUCN)

Introduction

The Egyptian fauna has been documented and investigated for a long time, probably longer than any other fauna in the world, dating back to pharaonic times. The herpetofauna received special attention from the ancient Egyptians, who depicted many reptile species in their inscriptions, such as Cobras Naja sp., Horned Viper Cerastes cerastes, Nile Crocodiles Crocodylus niloticus, and Dab Lizards Uromastyx sp. Throughout the eighteenth and nineteenth centuries Egypt was a popular destination for naturalists and explorers, becoming an important source of material used in describing many taxa after the establishment of the Linnaean nomenclature system. In more recent times, Egyptian herpetofauna has been the subject of four comprehensive reviews. Anderson’s (1898) classic work was the earliest comprehensive treatment of the Egyptian herpetofauna. Although it is now over one hundred years old, it remains the best researched and documented account of the country’s reptiles and amphibians. Many of Anderson’s observations on very difficult and complex groups such as Ptyodactylus have proven to be accurate, and reflect his keen understanding and careful scrutiny of these animals. Flower (1933) presented the first herpetofaunal list for Egypt within its modern-day boundaries (Anderson’s Egypt was more generously proportioned and included parts of today’s Sudan). He updated the taxonomy and nomenclature of previously reported taxa, adding several species that were new records for the country. Marx (1968) was mainly concerned with an extensive collection of reptiles and amphibians acquired in Egypt by the United States Naval Medical Research Unit number 3 (NAMRU 3) between the 1940s and 1970s, largely deposited in the Field Museum of Natural History (Chicago) and the National Museum of Natural History (Washington, D.C.). These collections were the first to include extensive material from Egyptian localities outside the Nile Valley and Delta, where much of the previous field-work had been focused. This work presented the first detailed distributional data on the herpetofauna of Egypt and reported several taxa new to the country. The latest account by Saleh (1997) largely summarizes previous knowledge, depending heavily on distributional data presented by Marx (1968), but without much re-evaluation or verification of the records at hand. One of this publication’s significant contributions is the provision of fairly good photographic representations of many Egyptian taxa. There are also numerous shorter publications that deal with single species, smaller regional faunas, or review the taxonomy of certain groups represented in the country.

After a preliminary evaluation of the contemporary knowledge of local herpetofauna, it was immediately obvious, that despite the fairly extensive coverage Egypt has received over the years, there were many gaps in both taxonomic understanding and geographical coverage. Without a fairly accurate understanding of the basic taxonomic units at hand and their geographical distribution, any subsequent ecological or biogeographical interpretations are likely to be flawed. Some of the existing gaps in knowledge were important elements that would render an ecological interpretation of herpetofaunal communities highly inaccurate. The case of the sister species Acanthodactylus scutellatus and A. longipes (the latter only recently recognized in Egypt [Baha El Din 1994a, 1996]) is illustrative in this respect. Without recognizing that what was thought to be one taxon is in fact composed of two species, the understanding of sand dune lizard communities in Egypt would be considerably different from what we know about them now.

This volume summarizes the more important observations of the author over the past twenty years of field experience in Egypt and the available relevant literature in an attempt to update current knowledge of this important component of the Egyptian fauna. This work is by no means the final word on Egypt’s herpetofauna. There are still many puzzles to be resolved and many ongoing changes in the taxonomy of many groups, thanks to the wider application of molecular techniques.

I hope this volume will help stimulate further field-oriented research in the Egyptian academic and amateur natural history world, and bring a better appreciation of the important ecological role this fauna plays and the urgent conservation action it needs.

The Egyptian Setting

Egypt occupies the northeastern corner of the African continent, with a surface area of just over one million square kilometers (1,019,600 km), or about 3% of the total area of Africa (Zahran and Willis 1992). The country is situated within the largest and driest desert region on the globe, with one of the world’s largest rivers running through it. Egypt can be divided into four major terrestrial physiographic regions: the Nile Valley, Western Desert, Eastern Desert, and Sinai (Kassas 1993); and two marine regions: the Mediterranean and Red Sea. Perhaps the most outstanding feature of Egypt’s landscape is the Nile River, the principal and most important source of fresh water in the country. The Nile splits Egypt into two almost equal, but quite different, parts. Egypt east of the Nile has much variation, including the country’s highest mountain peaks; west of the Nile the landscape is generally featureless, largely made up of vast expanses of flat open desert dotted with scattered oases. Egypt also has long coastlines on the Red Sea and the Mediterranean, two seas with very different marine ecosystems.

Egypt enjoys a wide variety of topographic features, ranging from the rugged mountains of South Sinai and the Eastern Desert (up to 2,641 m) to the Qattara Depression (134 m below mean sea level), to flat featureless gravel plains and complex hilly country, contributing greatly to the diversity of Egypt’s habitat types.

Situated between latitudes 32°N and 22°N, Egypt falls mostly in the temperate zone, since less than quarter of its land area lies south of the Tropic of Cancer (Said 1990). Egypt largely belongs to the hyper-arid region of the Sahara (Ayyad and Ghabbour 1986), characterized by hot, almost rainless climate (temperatures can range between -4°C and 53°C). The Sahara generally has low relative humidity and is the most extensive region in the world with an average of over ten hours of sunshine per day throughout the year (Smith 1984). Egypt can be divided into three general climatic zones: the Mediterranean coastal belt with 70-200 mm annual rainfall and a mean temperature range between 9.5°C-29.7°C; the Middle Egypt belt, south to 29°N, with 1-35 mm annual rainfall and a mean temperature range not much higher than that for the Mediterranean zone; and an Upper Egypt belt with an annual precipitation of less than 3 mm, and mean temperature of between 9.5°C-41.8°C (Goodman and Meininger 1989). Climate is largely correlated with latitude and proximity to both the Mediterranean and the Red Sea (Ayyad and Ghabbour 1986). Altitudinal effects are only observed at a few high mountains in South Sinai, the mountains overlooking the Red Sea, and Gebel Uweinat (Ayyad and Ghabbour 1986).

Most precipitation is received between November and April over much of Egypt, concentrated in December and January. The frequency and amount of rainfall generally increases northward: rainfall occurs every year in the Mediterranean coastal desert, while in the interior of the Western Desert rainfall is highly infrequent and unpredictable, perhaps falling once every decade. Some Red Sea mountains receive orographic precipitation on their seaward (eastern) side, most notably Gebel Elba.

Evolution of Egypt’s Landscape and Its Herpetofauna

The contemporary Egyptian landscape is the outcome of millions of years of geological evolution. However, the events that took place since the early Miocene (25 MYBP) have had the greatest impact in shaping the present-day structural framework of Egypt (Said 1990). It is in this period that Egypt reached roughly its current shape and size, the Nile River was born, and the Arabian Plate drifted away from Africa to open up the Red Sea.

The Red Sea, the principal migration barrier between Arabia and Africa, is relatively young (Joger 1987). Early Miocene rifting produced the Gulf of Suez and widened the Red Sea (Said 1990). By the Middle Miocene (some 12 MYBP), the Red Sea was connected with the Gulf of Suez, both of which were flooded with Mediterranean water (Said 1990). After severing its connection with the Mediterranean in the Pliocene (some 7-5 MYBP), the entire Red Sea was inundated for the first time by the waters of the Indian Ocean, through the Strait of Bab El Mandab (Said 1990).

During the late Miocene, some 6-7 MYBP, the sea retreated from much of Egypt’s land as a consequence of the Messinian crisis, during which the Mediterranean Sea dried out (Said 1993). This was caused by the blockage of the Strait of Gibraltar, preventing the Atlantic waters from entering the Mediterranean. Over several thousand years the Mediterranean waters evaporated, leaving it a hollow salt-encrusted basin, its bottom several thousand meters below the current sea level (Said 1993).This dramatic event had an enormous role in shaping the modern landscape of Egypt, which experienced intensive erosion, leading to the deepening of local drainage systems, excavation of the oases depressions, and eventually the establishment of the Nile River’s first course (Said 1990). This early Nile eroded a deep and winding canyon, which reached some 4,000 meters below current sea level at its outlet somewhere below the contemporary Delta (Said 1993). After reestablishment of the connection with the Atlantic and refilling of the Mediterranean some 5.4 MYBP, the Eonile canyon was transformed into a long marine body, which extended as far south as Aswan (Said 1993). Over the next two million years this long arm of the Mediterranean became increasingly estuarine and gradually filled with fluvial sediments, reaching north to the Delta. The function of the Nile as a local barrier was interrupted in the early Pleistocene when Egypt was completely desiccated and the river stopped flowing intermittently over a period of a million years (Said 1993). Eventually the river resumed its flow vigorously some 800,000 BP, making its first connection with the Ethiopian Highlands (Said 1993); thus the function of the Nile as a conduit of African fauna and flora was only established relatively recently. The size, course, and vigor of the river varied greatly through the Quaternary, until it established its current form.

Several other smaller rivers dissected the land of Egypt in various periods of the Miocene. One of these flowed northward from the Gilf El Kebir Plateau toward the Mediterranean, draining large areas of the Western Desert. The remains of this river system now lie concealed below the sands of the Great Sand Sea (Said 1990, El-Baz et al. 2000). Other rivers drained the Red Sea mountains, most importantly Wadi Qena.

The great Western Desert depressions probably initially formed parts of an ancient hydrographic network, which drained the extensive limestone plateau. Consequentially a combination of erosional processes worked together to enlarge and excavate these depressions, which represent the only relief over much of the Western Desert plateau today.

The origin of the sands of the huge dune fields and sand sheets of the Western Desert has been the topic of much debate. Most recently El-Baz et al. (2000) proposed that the sands originated as a result of combined fluvial-aeolian erosional action. During past wetter climates, water action eroded the vast sandstone country in the interior of the Western Desert in areas like the Gilf El Kebir and beyond. Ancient watercourses discovered below the sand-covered surface flowed to the north, carrying the sand with it. Subsequent arid periods combined with northerly winds blew the sand southward again. This cycle continued over millennia, building up the volume of available sand.

The Quaternary period probably had the greatest tangible effect on the contemporary habitat and species diversity of Egypt, as it has done in other temperate regions of the northern hemisphere. Cyclical climatic changes characterized the Quaternary, altering between cold (glacial) phases, where sea level was low, and warm (pluvial) phases, where sea level was high. At least seven pluvials are recognized in Egypt, generally corresponding with worldwide episodes of higher sea levels and a warmer climate (Said 1990). Knowledge of changes in sea levels over geological time has long been considered essential to the understanding of the distribution of both terrestrial and aquatic organisms (Voris 2000). The eustatic sea level changes of the Pleistocene, which reached a maximum lowering of 120 meters below current sea level (Joger 1987), are known to have greatly affected land-mass configurations throughout the world. At the local Egyptian level the Pleistocene sea level fluctuations probably did not have a substantial impact on the land-sea configuration or on the movement of fauna and flora. Most significant is that probably much of the Gulf of Suez became exposed and most Red Sea islands were connected to the mainland. At the regional scale, Africa and Arabia might have been intermittently connected during the maximal Pleistocene lowering, which would have established a land bridge across the Strait of Bab El Mandab, allowing the exchange of biota between the two sides.

The advent of the Pleistocene epoch brought to Egypt a pattern of aridity that set the tone for the current climatic conditions that prevail in the country today (Said 1990). The severity of the aridity of the early Pleistocene was so great that the Nile stopped flowing into Egypt (Said 1993). Although Egypt enjoyed several pluvial phases, the effects of such warm rainy periods appears to have been restricted to the southern part of the country, where the SudanoSahelian savannah belt migrated northward into southern Egypt (Said 1990). During glacial phases the northern part of the country enjoyed higher precipitation and came under the influence of the Mediterranean pre-Sahara steppe (Said 1990).

Poorly developed drainage, lack of playa deposits, and the preservation of old salt deposits all indicate that northern-central Egypt (e.g., in the vicinity of the Qattara Depression) has been arid for longer periods than other parts of the country (Said 1990). This could be the result of the region being too far north for the southern influence of pluvial periods and too far south for northern climatic effects. It is possible that these long-arid parts would have formed refugia for arid-adapted fauna and flora, during extensive pluvial episodes. Across the Sahara, such refugia probably gave rise to new taxonomic units during wet periods when arid-adapted taxa probably became isolated in pockets of desert. A similar process took place in refugia formed around the Mediterranean basin during glacial phases (Blondel and Aronson 1999).

Pluvial phases transformed much of the Egyptian deserts into an open grassy savannah with Acacia trees and scattered shallow wetlands, a habitat not unlike that of the Sahelian region today. Large African fauna, such as giraffe, elephant, rhinoceros, and ostrich have been documented in various parts of the Egyptian Sahara (Gautier 1993). Fossil remains of one amphibian and twelve reptiles, mostly of Afrotropical affinity, were collected from middle Palaeolithic sediments at Bir Tarfawi in the southern Western Desert (Mlynarski 1993, Szyndlar 1993). The following taxa have been identified: a frog of the family Ranidae, lizards of the family Lacertidae and Scincidae, Agama sp., Eryx sp. (probably E. muelleri), three snakes of the family Colubridae, Causus sp. viper and Crocodylus sp. (probably C. niloticus) (Szyndlar 1993). Three Chelonians were also identified: Plusios cf. adansoni, Geochelone cf. pardalis, and Cyclanorbis cf. senegalensis (Mlynarski 1993). The presence of amphibians, crocodile, and the Soft-shelled Turtle C. senegalensis all indicate the presence of extensive wetlands in this part of the Sahara, which were hydrographically connected with other water bodies in Africa.

The distribution of various species fluctuated with the shrinkage and expansion of their respective habitats. Eventually, with the onset of the current hyper-arid conditions some 4,900 YBP (Butzer 1976), almost all Afro tropical species retreated further south, as climatic conditions became too xeric for them in the Sahara. Only relict populations of a few species, such as Philochortus zolii, Tarentola annularis, and Echis pyramidum still persist in isolated Saharan oases. Some Afrotropical species found in the Nile Valley today probably represent relicts that were left behind when the surrounding terrain became a desert. Similarly, there are several northern species that extended southward into Egypt during glacial maxima and retreated during the interglacials, often leaving behind relict populations. The best known are examples from the flora, such as funiperus phoenicea in North Sinai.

The Contemporary Egyptian Herpetofauna

The contemporary Egyptian herpetofauna encompasses at least 118 species, composed of nine species of amphibians, sixty-one lizards, thirty-nine snakes, one crocodile, seven species of turtles and a tortoise. In total the current work adds almost 20% to the previously reported fauna, including some twenty species not reported in the last published herpetofaunal review of the country (Saleh 1997).

Contributions to the herpetological knowledge of Egypt leading up to this volume include six new species: Bufo kassasii, Hemidactylus foudaii, H. mindiae, Tarentola mindiae, Tropiocolotes bisharicus, and T. nubicus. An additional fourteen species are reported from Egypt for the first time (in this and previous publications by the author): Acanthodactylus longipes, Hemorrhois algirus, Hemidactylus robustus, H. sinaitus, Leptotyphlops nursii, Mesalina martini, M. pasteuri, Ophisops occidentalis, Psammophis punctulatus, Ptychadena schillukorum, Ptyodactylus ragazzii, Rana saharica, Ramphotyphlops braminus, and Stenodactylus mauritanicus. The status of several taxa in Egypt was confirmed, cither positively (e.g., Tropiocolotes nattereri) or negatively (e.g., Uromastyx acanthinura). Three taxa recognized previously at subspecific level arc elevated to the specific level: Stenodactylus mauritanicus (previously S. s. mauritanicus), Ptyodactylus siphonorhina (previously P. guttatus var. siphonorhina), and Ptyodactylus ragazzii (previously Ptyodactylus hasselquistii ragazzii). Chalcides populations inhabiting southern Egypt, and extending across the central Sahara and the Sahel, are recognized as a distinct species, for which the name Chalcides cf. humilis is proposed. Many nomenclatural adjustments are also made according to the most up-to-date taxonomic understanding available.

With 118 species, the herpetofauna of Egypt is an important part of the country’s biological diversity, comparable with the similar number of terrestrial mammalian species and about 150 species of breeding birds. Reptiles are particularly prominent, widespread, and well adapted to life in the arid desert environments that make up almost 90% of Egypt’s territory. In large parts of Egypt’s deserts the only vertebrates occupying them are reptiles. This is especially true in the extremely arid parts of the Western Desert.

Biogeography

Biogeographically, the Egyptian herpetofauna is fairly heterogeneous. This can be attributed to the fact that Egypt lies at the juncture of four major biogeographical regions: Saharo-Sindian, Irano-Turanian, Mediterranean (these three generally considered as subregions of the Palaearctic), and Afrotropical. The majority of the Egyptian herpetofauna is of Palaearctic affinity. The Saharo-Sindian component making up the majority of these species is well represented in Egypt’s vast deserts, while Mediterranean and Irano-Turanian elements occupy fairly small areas along the Mediterranean coast and in the Sinai highlands respectively. There is also a fairly strong Afrotropical influence extending from sub-Saharan Africa along the Nile River and the Red Sea coast and mountains (or in some cases representing relict populations). Marx (1968) found that the largest proportion of the Egyptian herpetofauna (about 68%) is shared with southwest Asia, while 40% is shared with the rest of North Africa and 37% with northern East Africa.

Egypt has no major terrestrial centers of endemicity either for flora or fauna. Bufo kassasii is the only species of the herpetofauna apparently endemic to Egypt, but there are several restricted-range species (species largely restricted to Egypt, but also extending into neighboring countries), found largely in or near the country’s hot spots of terrestrial biodiversity.

At the larger regional scale, Egypt lies at a meeting point of Earth’s largest land blocks, making it of special importance as a land bridge for the movement of terrestrial biota (Bowen and Jux 1987). At the local level the geography of the Egyptian landscape has many physical and ecological barriers, and conduits for biota, which make for a complex biogeographic setting.

Egypt is bounded from the north and east by the Mediterranean and the Red Sea respectively, with only a limited land connection with Arabia through Sinai. However, the Dead Sea Rift, which runs from the head of the Gulf of Aqaba to Lebanon’s highlands, represents a real barrier to the movement and exchange of many biotic elements between Arabia and the Sahara (Joger 1987). From the south and west Egypt is connected with the rest of the African continent, without any apparent physical barriers, but the extreme aridity of the Sahara prevents almost all sub-Saharan biota from reaching the country. The seemingly simple, monotonous Saharan landscape is in fact a patchwork of various substrate types, representing a complex pattern of barriers and conduits for various species of the fauna and flora. For example, the extensive dunes in the Western Desert, while perhaps a barrier to some species characteristic of hard substrates, represent corridors for psammophilous species’ movement.

The Nile River is also a primary biogeographical barrier for several terrestrial desert taxa, while also acting as a corridor along which many Afrotropical species (primarily species associated with wetland and mesic habitats) have spread northward to reach the Mediterranean. Similarly, the Red Sea has acted as a corridor for some Afrotropical terrestrial elements, though to a lesser extent.

Species Richness and Distribution

The non-marine herpetofauna of Egypt (113 species) is moderately rich with respect to its land area (1,019,600 km). In comparison, some ninety species have been recorded in Israel and Palestine, with an area of only 28,000 km (Werner 1988) and about 98 species have been recorded in Morocco (including the Western Sahara), with an area of 710,850 km (Bons and Geniez 1996). These differences in relative species richness can be attributed largely to habitat diversity and productivity, but biogeographical factors probably play an important role in determining species richness at the regional level. Israel and Palestine, being at a meeting point of four biogeographical regions, Mediterranean, Saharo-Arabian, Irano-Turanian, and Sudanian (Werner 1987), have a particularly rich herpetofauna in a relatively small area.

Herpetofaunal species richness plotted on a half-degree