© Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at A REVISION OF THE LIZARDS OF THE GENUS ACANTHODACTYLUS (SAURIA: LACERTIDAE) by ALFREDO SALVADOR BONNER ZOOLOGISCHE MONOGRAPHIEN, Nr. 16 1982 Herausgeber: ZOOLOGISCHES FORSCHUNGSINSTITUT UND MUSEUM ALEXANDER KOENIG BONN © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at BONNER ZOOLOGISCHE MONOGRAPHIEN Die Serie wird vom Zoologischen Forschungsinstitut und Museum Alexander Koe- nig herausgegeben und bringt Originalarbeiten, die für eine Unterbringung in den Bonner Zoologischen Beiträgen" zu lang sind und eine Veröffentlichung als Mono- graphie rechtfertigen. Anfragen bezüglich der Vorlage von Manuskripten und Bestellungen sind an die Schriftleitung zu richten. This series of monographs, published by the Zoological Research Institute and Museum Alexander Koenig, has been established for original contributions too long for inclusion in ,, Bonner Zoologische Beiträge". Correspondence concerning manuscripts for publication and purchase orders should be addressed to the editor. L' Institut de Recherches Zoologiques et Museum Alexander Koenig a etabli cette serie de monographies pour pouvoir publier des travaux zoologiques trop longs pour etre inclus dans les ,, Bonner Zoologische Beiträge". Toute correspondance concernant des manuscrits pour cette serie ou des com- mandes doivent etre adressees ä Pediteur. BONNER ZOOLOGISCHE MONOGRAPHIEN, Nr. 16, 1982 Preis 30,- DM Schriftleitung/Editor: H. Ulrich Zoologisches Forschungsinstitut und Museum Alexander Koenig Adenauerallee 150-164, 5300 Bonn, Germany Druck: Ferd. L. Doepgen, 5010 Bergheim © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at A REVISION OF THE LIZARDS OF THE GENUS ACANTHODACTYLUS (SAURIA: LACERTIDAE) by ALFREDO SALVADOR BONNER ZOOLOGISCHE MONOGRAPHIEN, Nr. 16 1982 Herausgeber: ZOOLOGISCHES FORSCHUNGSINSTITUT UND MUSEUM ALEXANDER KOENIG BONN © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at CONTENTS 1. Introduction 5 2. Material 5 3. Methods 6 4. Acknowledgements 7 5. Genus Acanthodactylus Fitzinger 8 6. Key to the species 8 7. Species groups 10 micropholis group 10 boskianus group 10 yemenicus group 11 tristrami group 12 grandis group 12 erythrums group 12 pardalis group 13 scutellatus group 13 cantoris group 14 8. Historical resume 15 9. The species of Acanthodactylus 19 A. micropholis 19 A. boskianus 23 A. schreiben 38 A. masirae 42 A. opheodurus 45 A. felicis 50 A. yemenicus n. sp 53 A. erythrurus 57 A. savignyi 66 A. Wanci 69 A. boueti 73 A. guineensis 77 A. pardalis 80 A. bedriagai 85 A. busacki n. sp 88 A. maculatus 92 A. spinicauda 97 A. tristrami 101 A. robustus 106 A. grandis 110 A. scutellatus 113 A. aureus 122 A. dumerili 126 A. longipes 132 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at A. gongrorhynchatus 136 A. haasi 139 A. arabicus 143 A. schmidti. 146 A. blanfordi 151 A. cantoris 155 10. Summary 159 11. Zusammenfassung 160 12. Literature cited 161 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at 5 1. INTRODUCTION Acanthodactylus is recognized by persons within the field as one of the most difficult genera of the family Lacertidae. The present state of its systematics could be described as chaotic. Seeing that these lizards are the most conspic- uous and abundant in the desert and arid regions of northern Africa and south- western Asia, it seems unusual that there are so few publications dedicated to the genus. Several recent studies marked by mistaken identification of species have only confused the issue more. For example, in 1936 Angel reported specimens of A. pardalis in Syria when actually they were A. grandis. In 1957 Haas assigned to A. fraseri specimens which Leviton & Anderson (1967) described later as A. gongrorhynchatus. In 1961, Haas discussed a specimen of A. scutellatus which was actually A. grandis. Busack (1975) studied what he thought to be A. parda- lis but which actually turned out to be A. scutellatus. As far as this present work is concerned, I here propose a rearrangement which corrects much of the past and present confusion but yet at the same time does not overlook the possible need for future revisions. 2. MATERIAL This review is based on the examination of more than 6.000 lizards belonging to the genus Acanthodactylus. The majority of them are preserved in alcohol while some are skeletons. I was also able to examine radiographs of more than 420 specimens. Below is a key for the abbreviations of the collections exam- ined. AMNH American Museum of Natural History, New York. ANSP Academy of Natural Sciences, Philadelphia. BM British Museum of Natural History, London. BMI Bagdad Museum Iraq, Bagdad. CAS California Academy of Sciences, San Francisco. CM Carnegie Museum of Natural History, Pittsburgh. EBD Estaciön Biolögica de Donana, Sevilla. FMNH Field Museum of Natural History, Chicago. KU Kansas University Museum of Natural History, Lawrence. LACM Los Angeles County Museum of Natural History, Los Angeles. MCZ Museum of Comparative Zoology, Cambridge. MNHNP Museum National d'Histoire Naturelle, Paris. MVZ Museum of Vertebrate Zoology, Berkeley. NHMW Naturhistorisches Museum, Wien. TCWC Texas Cooperative Wildlife Collection, Texas. © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at 6 UCM University of Colorado Museum of Natural History, Boulder. UF University of Florida, Florida State Museum, Gainesville. UMMZ University of Michigan Museum of Zoology, Ann Arbor. USNM United States National Museum, Washington. TAU Tel Aviv University, Tel Aviv. ZFMK Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn. A catalogue number for each specimen will appear next to the abbreviation of the collection. In those instances where more than one specimen is included under the same catalogue number, the number of the specimens involved can be found in parentheses. 3. METHODS My examination of specimens was aimed at studying the following charac- ters: Size; relative proportions of head, limbs, digits, and tail; number of supraoculars. Presence of lanceolate cavity between the frontanasal and the middle of the frontal. Number of rows of granules betweeen the supraoculars and the superciliaries. Number of superciliaries. Size of the subocular keel. Number of supralabials anterior to the subocular. Subocular contact or lack of contact with the upper lip. Pectination of the eyelids. Size of the scales on the lower eyelid. Number of supratemporals, with or without keel. Size and shape of the temporal scales and the presence or lack of keel. Presence or absence of tympanic scale. Relative size of ear opening and the presence or lack of pectination on the anterior border. Size, shape and number of dorsals and the presence and size of their keels. Size and shape of the late- rals. Size, shape, and row placement of ventrals. Size, shape, and number of scales on the anterior upper surface of the tail and the presence or lack of keels. Number of rows of scales on the fingers. Number of lamellae and carinae on the fourth toe and its lateral pectination. Number of femoral pores on each side of the body and their arrangement or not in one continuous row across the body. Variation of color pattern and coloration in young, subadults, and adults. The osteological characters which were studied are the number of presacral vertebrae, number of cervical ribs, the number of sternal ribs, the number of non-autotomic vertebrae of the tail, and the number of vertebrae with double processes on the tail. It was necessary to choose a region which has been well investigated and from which we have abundant material available from several different locali- ties. Northern Egypt fulfilled these conditions since enormous collections have been gathered there by R.E. Kuntz, W. H. Wells and G.M. Malakatis on © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at 7 behalf of the United States National Museum (Washington, D.C.) and by H. Hoogstraal on behalf of the Field Museum of Natural History (Chicago). For a given locality the variation of characters, as well as the different combinations of character states, was studied for the entire sample. In this manner, three dif- ferent taxa corresponding to A. boskianus, A. scutellatus, and A. pardalis were obtained. This same operation was followed for all the remaining localities in Egypt, the findings being consistent with the former results. As our area of investiga- tion grew towards northwest Africa and towards southwest Asia, new taxa were added to the list. Special attention was given to the variation of character states and their different combinations in allopatry as well as in sympatry, revealing that for some cases nothing more was involved other than populations of the same species. In addition to the already troublesome transcription of Arabic names of lo- calities into English, other localities situated in the middle of the desert had no names at all and could only be referred to by the longitude