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BONNER ZOOLOGISCHE MONOGRAPHIEN, Nr © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at Phylogenetic Systematics and Biogeography of the Carphodactylini (Reptilia: Gekkonidae) by AARON M. BAUER BONNER ZOOLOGISCHE MONOGRAPHIEN, Nr. 30 1990 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 Koenig herausgegeben und bringt Originalarbeiten, die für eine Unterbringung in den „Bonner zoologischen Beiträgen" zu lang sind und eine VeröffentHchung als Monographie rechtfertigen. Anfragen bezüglich der Vorlage von Manuskripten sind an die Schriftleitung zu richten; Bestellungen und Tauschangebote bitte an die Bibliothek des Instituts. This series of monographs, published by the Zoological Research Institute and Museum Alexander Koenig, has been estabhshed for original contributions too long for inclu- sion in „Bonner zoologische Beiträge". Correspondence concerning manuscripts for pubhcation should be addressed to the editor. Purchase orders and requests for exchange please address to the library of the institute. LTnstitut de Recherches Zoologiques et Museum Alexander Koenig a etabli cette serie de monographies pour pouvoir pubher des travaux zoologiques trop longs pour etre inclus dans les „Bonner zoologische Beiträge". Toute correspondance concernante des manuscrits pour cette serie doit etre adressee ä I'editeur. Commandes et demandes pour echanges adresser ä la bibliotheque de I'insti- tut, s. V. p. BONNER ZOOLOGISCHE MONOGRAPHIEN, Nr. 30, 1990 Preis: 36,— DM Schriftleitung/Editor: G. Rheinwald Zoologisches Forschungsinstitut und Museum Alexander Koenig Adenauerallee 150—164, D-5300 Bonn 1, Germany Druck: Rheinischer Landwirtschafts-Verlag G.m.b.H., 5300 Bonn 1 ISBN 3-925382-31-3 ISSN 0302-671X © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at Phylogenetic Systematics and Biogeography of the Carphodactylini (Reptilia: Gekkonidae) by Aaron M. Bauer BONNER ZOOLOGISCHE MONOGRAPHIEN, Nr.30 1990 Herausgeber: ZOOLOGISCHES FORSCHUNGSINSTITUT UND MUSEUM ALEXANDER KOENIG BONN © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at CIP-Titelaufnahme der Deutschen Bibliothek Bauer, Aaron M.: Phylogenetic systematics and biogeography of the Carphodactyhnie (ReptiUa: Gekkoniade) / by Aaron M. Bauer. Hrsg.: Zoolog. Forschungs- inst. u. Museum Alexander Koenig Bonn. — Bonn: Zoolog. Forschungs- inst. u. Museum Alexander Koenig, 1990 (Bonner zoologische Monographien; Nr. 30) ISBN 3-925382-31-3 NE: GT © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at CONTENTS Page Abstract 5 Introduction 5 Materials and Methods 13 Character Analysis 17 Cranial Osteology 18 Hyobranchial Apparatus 30 Axial Skeleton 32 Sternum, Pectoral Girdle and Forelimb 41 Pelvic Girdle and Hindlimb 45 Cloacal Bones 49 Coloration 51 Membrane Pigmentation 52 Eye 53 Ear 54 Endolymphatic System 55 External Digital Characters 56 Claws 58 Scalation Characters 59 Skin Folds and Webs 60 Preanal Organs 61 Cloacal Spurs 62 Tail 63 Reproductive Mode 65 Defensive Behavior 66 Results 67 Discussion 73 Biogeography of the Carphodactylini 77 Evaluation of the Phylogenetic Hypothesis 78 The Traditional View of Carphodactyline Evolution and Biogeography . 81 A Scenario for the Evolution of the Carphodactylini 83 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at Carphodactyline Species Accounts 88 Key to the Genera of Carphodactyline Geckos 89 Bavayia Roux 90 Carphodactylus Günther 97 Eiirydactylodes Wermuth 99 Hoplodactylus Fitzinger 103 Naultinus Gray 126 Nephrunis Günther 139 Phyllurus Goldfuss 155 Rhacodactylus Fitzinger 165 Acknowledgments 184 Literature Cited 186 Appendix A: Collection Acronyms 210 Appendix B: Species Examined 212 Appendix C: Character Matrix 218 I © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at 5 ABSTRACT The tribe Carphodactylini is composed of 49 species of geclcos of the subfamily Diplodactyhnae that are endemic to the regions of AustraHa, New Caledonia and New Zealand. The systematics of the group is approached through the methodology of phylogenetic systematics and a hypothesis of genealogical relationship is presented. The padless Australian genera Nephrurus (including Underwoodisaurus), Phyllurus, and Carphodactylus are the sister group of the New Caledonian and New Zealand car- phodactylines. The New Zealand genus Hoplodactylus is paraphyletic. The New Caledonian taxa {Bavayia, Eurydactylodes, and Rhacodactylus) form a monophyletic unit if the northern Australian genus Pseudothecadactylus is regarded as a subgenus of Rhacodactylus. Systematic accounts and summaries of all biological hterature relating to each species are presented. The tectonic history of the southwest Pacific region is in harmony with the hypothesis of carphodactyline relationships. The primary division of the tribe into Austrahan and Tasmantis Uneages was probably brought about by the opening of the Tasman Sea about 80 mybp. By the Oligocene marine ingressions isolated New Caledonia from nor- thern New Zealand, thus splitting Hoplodactylus. Carphodactyline geckos highlight the antiquity, endemism and biogeographic significance of the herpetofauna of the southwest Pacific, and New Caledonia in particular. INTRODUCTION Although a great deal of systematic work has dealt with the place of gekkonids among squamates and with higher order relationships among geckos, little has been produced with respect to inter- and intrageneric relationships within the tribe Carphodactylini. The carphodactylines as currently construed include the Australian genera Nephrurus, Phyllurus, Carphodactylus and Pseudothecadactylus, the New Zealand genera Naultinus and Hoplodactylus, and the genera Bavayia, Eurydactylodes and Rhacodac- tylus from New Caledonia and the Loyalty Islands. In addition, Underwoodisaurus (Australia) and Heteropholis (New Zealand) are also occasionally recognized as distinct taxa at a generic level. The present work is an attempt to summarize that which is known about carphodactyline biology and to erect a hypothesis of relationship upon which subsequent evolutionary morphological studies may be based. (In the pages that follow, in order to avoid confusion, nomenclatural changes implemented as a result of this study are used consistently from the start — e.g., Nephrurus milii instead of Phyllurus milii or Underwoodisaurus milii). The earliest reviewers of lizard taxonomy were perplexed by the taxonomic affinities of the first described carphodactyline, Phyllurus platurus (White 1790). Schneider (1797), Daudin (1802), La Cepede (1804), and Merrem (1820) all placed the species in an agamid genus, either Agama or Stellio. Gray (1825) was uncertain as to its affinities, but suggested hnks with either the agamids or gekkonids. Bory de Saint-Vincent (1825) © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at 6 I described a second species, P. milii (= Nephrurus milii), recognized its similarity to P. | platurus, and assigned both to the Gekkonidae. i Fitzinger (1843) distributed the four species known to him into three genera in two i families of the Ordo Ascalabotae. The family Stenodactyh, essentially containing all i geckos without dilated scansorial subdigital pads, included Gymnodactylus miliusii and ' Gonyodactylus platurus, the latter genus being distinguished by a prominent kink in the digits. The remaining taxa, Hoplodactylus duvaucelii and Hoplodactylus (Rhacodactylus) leachianus, were included among the Platydactyli, a mixture of all of the padded geckos without divided or lobed scansors. This classification system, Hke most of those which followed for over one hundred years, made no attempt at j phylogenetic analysis but rather pigeon-holed species primarily on the basis of external j digital characters. i Gray (1845) keyed Phyllurus and Naultinus to the same first subdivision of geckos while separating Platydactylus (including Rhacodactylus and Hoplodactylus duvaucelii). ' Girard (1858) relied on artificial divisions to yield information about natural groups | ("the genus Naultinus is a Stenodactylian: hence widely distinct from Hoplodactylus, which ... is a Platydactylian"). Boulenger (1885a) accorded no special recognition to the group now regarded as the j Carphodactylini. His division in the "Catalogue of Lizards" was based on the order ' in which genera fell out in the family keys. Nevertheless, the structure of the keys, while largely artificial, may imply some phylogenetic information (Russell 1976). Boulenger's primary divisions were based upon pedal structure and the carphodactyUnes known at I the time were distributed among five different major groups of geckos. Notably, divi- sion VII hsts the genera Naultinus, Hoplodactylus and Rhacodactylus in sequence. These are immediately preceded by Lepidodactylus which at the time included the j species later transferred to Bavayia. Although clustered together because of digital i similarities, the particular order of these genera may reflect Boulenger's
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