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Anura:Microhylidae THE PHYLOGENY OF THE PAPUAN SUBFAIÏILY ASTEROPHRYINAE (ANURA: MICROHYLIDAE) by THO¡1IAS CHARLES BURTON, 8.4., B.Sc. (Hons)MeIb. Department of Zoology University of Adelaide A thesis submitted to the UniVersity of Adelaide for the degree of Doctor of PhilosoPhY JT'NE 1.9 8 3 To ChwLø SUMMARY THE PHYLOGENY OF THE PAPUAN SUBFAMILY ASTEROPHRYINAE (nruunR : MI cRoHyt-rrRE) The Asterophryinae is a subfamily of terrestrial and fossorial microhylid frogs restricted to the Papuan Sub- Region. It comprises 43 named species and subspecies in seven genera. A second microhylid subfamily, the Sphenophryninae, also occurs in the Papuan Sub-Region, and its relationship to the Asterophryinae is contentious- In this study I undertake a phylogenetic analysis of the Asterophryinae based on the results of an examination of the myology, osteology and external morphology of members of all of the genera, and also of members of the Sphenophryninae, other microhylid. subfamilies and the Ranoidea, which serve as out-groups at different levels of analysis. The Asterophryinae and Sphenophryninae form a monophyletic group (sensu Hennig, 19661 supported by two autapomorphies: (a) direct embryonic development within the egg capsule; and (b) fusion and enlargement of the palatine and prevomer. The monophyly of the Asterophryinae is supported by three autapomorphies: (a) posterior adherence of the tongue and its division into anterior and posterior sections; (b) fusion of elements of the mandible and displacement of the mentomeckelians from the anterior margin of the mandible; and (c) loss of a dorsal el-ement J-aa of the M. intermandíbuLaris. The monophyly of the Sphenophryninae is supported by only one character of dubious value: procoely of the vertebral column. The polarities of characters observed to vary among the asterophryines are assessed and a cladogram is constructed on the basis of shared and derived character states. This cladogram is converted to a Proposed classification in the format of the Annotated Linnean Hierarchy of Wiley lL979l. A number of taxonomic actions are shown to be necessary: (1) Resurrection of Mantophryne Boulenger 1897 to accommodate'three species of Phrynomantis with closer affinities to .4 sterophnys, HyLophorbus and' Phetohapsis than to other Phrynomantís species. (21 Transfer of Xenorhina doniae Lo Phrynomantís. (3) Erection of four tribes to accommodate monophyletic groups of genera: Asterophryini (Astenophrys, HALophot'bus, Møntophv,Ane and Phez'ohapsisl ¡ Barygenyini (BarygenA sl ¡ phrynomantini (Phngnomantisli and Xenorhinini lXenobatrachus anð. Xenorhína). (4) Redefinition of all genera in the light of newly described taxonomic characters. A number of morphological characters described for the first time in the course of this study appear to have potential for use in wider phylogenetic studies of the microhylid.s. Though a biogeographical synthesis is considered premature, a number of hypotheses are brought forward to iv explain aspects of the patterns of d.istribution of the Asterophryinae. v l I I 1 I I DECLARATION : l This thesis contains no material accepted I for the award of anY other degree or diploma in this or any other uníversity' To the best of my knowtedge this thesis contains no material previously published or written by another person, except when due reference is made in the text' vi ACKNOVJLEDGEMENTS Many individuals and institutions aided me at various stages during the completion of this project. My thanks are expressed to all of them. l4ike Tyler suggested and supervised this project and provided access to.specimens so that I could get it under way. Mike was an unfailing source of support and encouragement throughout the project. He read. closely and critically the drafts of the manuscript. Margaret Davies helped me greatly with my setting myself up for drawings, and. aided me by instruction and encouragement in that art and in the divers arts of osteology. Philippa Horton ted me back to the long-forgotten realms of histology, and helped me with proof-reading. These three bore much of the brunt of my tatest theories, mY successes (real and imagined) and failures, and for their support in times of triumph and despondency I am very thankful. other post- graduate students in the Department of Zoology, notably Jenny Gardner, Graeme Browning and Alice Wells lent their ears and gave useful advice and ideas when I needed them' Linda Trueb, in a five-minute conversation and. demonstration, showed me how to bring my chicken-scratchings to a publishable standard. Terry Schwaner lent me several books, gave some useful advice, and introduced me to the ideas of Wiley' Rev. Br. John Marks, headmaster of Rostrevor college, gave me time off to complete this project with generous conditions, and some of my colleagues, particularly Mary Meredith and Peter Taylor, bore ad'ded' burdens as a result of vaa my absence. Mary often acted as a sounding-board for my ideas. Department of science and Industry supported my field- trip to Papua New Guinea and the Department of Biology, UPNG, supported my visa application. In PNG I was very fortunate to be acconìmodated, helped in the field and in other ways, and befriended by Dennis and Krista Black, Roy and Margaret Mackay, Rupert Stocks and AII-en Allison' Specimens were provided by the following institutions: American Museum of Natural History (R.G. Zweifel); Australian Museum (H.G. Cogger); Museum of comparative Zoology (J. Rosad.o) ; Rijksmuseum van_Na_tuurlijke Historie, Leiden $6.6fla Avslral,an l4useom (il'Ty/e") ; (M. Hoogmoed) r/ University of- eÑG (r,. HiII, D. B1ack). Keith IvlcDonald provÍded líve specimens from Queensland' Ross McDonald. took radiographs of specimens deemed too precious to dissect. The Australian Museum supported my work with a student grant, and I was supported also by a research grant from the UniversitY of Ad'elaide- This project could not have begun without the pioneering work of Dr. R.G. Zweifel, who placed, the taxonomy of the Asterophryinae on a f irm basis. He \^las prompt and generous in the provision of specimens, transparencies, advice and encouragement. That he went out of his way to aid me in a reworking of his major study is a mark of his humility and scientific honestY. It was my wife, cherie, who suggested that I take time off work to finish this project. cherie bore the diminution viii of our income, mY strange time-table, my bizarre preoccupations, and her increased parental burden with loving tolerance. She was always ready to discuss my work and its meaning and her opinions were valued highly. Jonathan, our son, who regards crePuscularity and an interest in frogs normative for adult males, rarely caused distraction and anxiety sufficient to disrupt my work hours, and he gave me much joy between times. ax TABLE OF CONTENTS Page VOLUIT{E 1 SUMMARY .Irr DECLARATION vi ACKNOWLEDGE}'IENTS vii CHAPTER 1- - INTRODUCTION 1 1.1 The Microhylids of the Papuan Zoogeogtaphic Subregion 1 7.2 HistorY 2 1. 3 The lr{yology and osteology of the New Guinea MicrohYlids Ll_ L.4 Aims of this Stud'Y 15 CHAPTER 2 - MATERIALS AND METHODS t7 2.L Materials Available T7 2.2 Materials Examined' 18 2.3 Ivlethods: Morphology 19 2.4 Methods: Field StudY 22 2.5 , Method.s: PhYIogenY 22 2.6 BiogeograPhY 30 2.6.1 Origin of the New Guinea microhylids 30 2.6.2 Biogeography within New Guinea 35 CHAPTER 3 - RESULTS: THE MUSCULATURE OF PHRYNOMANTTS STICTOGASTER 38 3. 1 lvluscles of the Head 3B 3.2 Muscles of the. Trunk 45 3.3 lr{uscles of the Pectoral Girdle and Forelimb 48 3.4 l"luscles of the Pelvic Girdle and Hindlinb 63 x Page CHAPTER 4 - RESULTS: VARIATION IN MUSCLE CHARACTERS 82 4.L AsteroPhrYinae 82 4.1.1 Muscles of the head 84 4.L.2 l"luscles of the trunk 95 4.I.3 Muscles of the pectoral girdle and forelimb 98 4.1.4 Muscles of the petvic gird'le and hindlímb 100 4.2 SphenoPhrYninae 106 4.2.1 Muscles of the head 107 4.2.2 Muscles of the trunk 113 4.2.3 lvluscles of the pectoral girdle and' forelimb l_ 14 4.2.4 Muscles of the pelvic girdle and' hindlimb 116 4.3 Other MicrohYlids 119 4. 3. 1 l4uscles of the head 119 4.3.2 Muscles of the trunk L23 4.3.3 Muscles of the pectoral girdle and forelimb t24 4.3.4 Muscles of the pelvic girdle and híndlimb L25 VOLUME 2 CHAPTER 5 RESULTS: THE OSTEOLOGY OF PHRYNOMANTTS ST TCTOGASTER L2B 5.1 Cranium L2B 5.2 The Mandible 1,32 5.3 Hyoid and LarYnx 133 5.4 Pectoral Girdle 133 xl_ Page 5.5 Forelimb and Manus l_34 5.6 Vertebral Column 135 5.7 Pelvic Girdle 1_36 5.8 Hindlimb and, Pes 137 CHAPTER 6 - RESULTS: VARIATION IN SKELETAT, CHARÀCTERS 138 6.1, AsteroPhrYinae 138 6.1.1 Cranium 138 6.1.2 Mandible 145 6.1.3 Hyoid and larYnx I46 6.L.4 Pectoral girdle L46 6.1.5 Forelimb and manus r46 6.1.6 Vertebral column r47 6.1.7 Pelvic girdle L47 6. 1.8 Hind.limb and Pes 148 6.2 Sphenophryninae ]-49 6.2.1 Cranium r49 6.2.2 Mandible 153 6.2.3 Hyoid and. IarYnx t_5 3 6.2.4 Pectoral girdle 153 6.2.5 Forelimb and manus r_54 6.2.6 Vertebral column L54 6.2.7 Pelvic girdle 155 6.2.8 Hindlimb and Pes 155 6.3 Other Microhylids 155 6.3.1 Cranium 156 6.3.2 Mandible r_59 6.3.3 Hyoid and larYnx 159 6.3.4 Pectoral girdle 159 6. 3.5 Forelimb and manus 160 6.3.6 Vertebral column 160 6.3.7 Pelvic girdle 161 6.3.8 Hindlimb and, pes 161 CHAPTER 7 - RESULTS: SUPERFICIAI MORPHOLOGY L62 xLr Page CHAPTER 8 .
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