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The Comparative Anatomy and Systematics of Mesozoxc Sphenodontidans

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The Comparative Anatomy and Systematics of Mesozoxc Sphenodontidans THE COMPARATIVE ANATOMY AND SYSTEMATICS OF MESOZOXC SPHENODONTIDANS XIAO-CHUN WU BIOLOGY DEPAR~~ENT McGILL UNIVERSITY, :I{ONTREAL QUEPEC, CAN!\DA A thesis submitted to the Faculty of Graduate Studies and Research in partial fulfilment of the requirements for the degree of Doctor of Philosophy. @ XIAO-CHUN WU, 1991. ( PRBPACB ( This thesis is centred around an anatcmical study of the craniai osteoloqy of the Mesozoic sphenodontidans, Asiacephalosaurus wangi, Raro;ugalosaurus mcqilli, Dianosaurus petilus and HomoeosaurU3 maximiliani. The first two are newly discovered and have not been studied previously. The third, previously described only in a preliminary fashion, was misinterpreted as a protorosaur. Althouqh many specimens of H. maximiliani have been described (Cocude-Michel, 1963; Faber, 1981) since the species was first named by Meyer (1845), the newly exposed dorsal part of the skull of a new specimen has provided many new cranial features. Consequently, it was necessary to examine, draw and describe aIl of these sphenodontidans. The descriptions are aIl original and present a virtually complete picture of the cranial osteoloqy of these qenera for the first time. In addition, the different patterns of the middle ear apparatus of the Sphenodontida and the functional problems of the temporal reqion in the Lepidosauria have been considered. Although use is made of the literature to obtain backqround data, the conclusions drawn are oriqinal unless explicdtly stated otherwise. Finally, the detailed descriptions of the four qenera have permitted a reassessment of the phyloqeny of the Sphenodontlda. After revlewinq the llterature, original conclusions ara qiven. ( i ACKNOWLEDGEMENTS This study owns its existence to the supervision of Dr. Robert L. Carroll who critically read and edited several drafts of this manuscript, and l am greatly indebted for his interest and encouragement. l would l ike to express my appreciation to the other members of my dissertation committee--Drs. Joan Marsdon, Robert E. Lemon, and David M. Green--for their time, effort and patience, and especially to Dr. D. M. Green for his comments and help in obtaining a preserved specimen of the living Sphenodon punctatus. l am very grateful to Dr. Robert Holmes for his proofreading the manuscript and improving it considerably. l am aiso grateful to professor Ai-lin Sun (Institute of Vertebrate Paleontology and Palaeoanthropology, Academia Sinica, Beijing, China) and Dr. Sankar Chatterjee (The Museum of Texas Tech University, Lubbock, Texas) for their suggestions and encouragements. Furthermore, i t is a pleasure to acknowledge Drs. R. carroll, Hans-Dieter Sues (Smithsonian Institution, Washington, D. C.), R. Holmes, D. M. Green, Denis Walsh, Stephen Godfrey (Tyrrell Museum, Drumheller, Alberta), pierre -Yves Gagnier and Mrs. Michael Debraga, Kebang Lin, and Edward Hitchcock for stimulating discussion concerning origin and evolution of the impedance matching middle ear, the form and function of the temporal region of the Lepidosauria, and interrelationships within the Sphenodontida. l have particularly benefitted from Mr. K. Lin's and Dr. ii ( D. M. Green' s knowledge of Mix parsimonious Algorithm in PHYLIP Package and other computer programs. l thank Dr. Pierre-Yves Gagnier for his help translating the French version of the abstracto My thanks are also due to Mrs. Pamela Gaskill for her initial guidance in draft~ng the figures and encouragement, Dr. David Berman (carnegie Museum, Pittsburgh) for permit to study the material of Homoeosaurus maximiliani, Dr. C. H. Dauqherty (the School of Biological Sciences, victoria University, Wellington, New Zealand) for providing a preserved specimen of the living Sphenodon punctatus (from Stephen Island), Mr. F. L. Li (The Kunming Branch of Chinese Academy of Scjences, Kunming, Yunnan, China) for providing preserved specimens of Japalura laviceps (from Sichuan province), Mrs. Messrs R. Lamarche and G. L'heureux for photographie work, Mrs. Z. J. Wang, H. Wang (The Culture Museum of Lufeng County, Yunnan, China) and F. yi (Dawa Elementary School, Lufeng, China) for their help during rny field work. In addition, l have received considerable assistance from the staff, especially from Ingrid Birker and Marie La Ricca, at the Redpath Museum. Finally, l would like to thank my wife, Wen-ying Lu, for her encouragement and support throughout the preparation of this thesis. This research was made possible by financial support from Natural Sciences and Engineering Research Council of Canada grants to Dr. R. L. Carroll. ( iii 'l'ABLI OP COK'l'BII'1'8 PREFACE ••••••••• · ................................... Page i ACKNOWLEDGEMENTS · ................................... ii LIST OF ILLUSTRATIONS ................................ vii LIST OF TABLES ....................................... xiii ABSTRACT •••• .. .. .. .. .. .. .. xiv INTRODUCTION ......................................... 1. MATERIALS AND METHODS .............................. 6 Asiacephalosaurus wangi · ....................... 11 Systematic Paleontoloqy · ....................... 11 Description · ................................... 12 a. General Features of the Skull 13 b. Dermal Bones of the Skull Roof 18 c. Palatoquadrate . .. .. .. ..... 27 d. Oermal Bones of the Palate .............. 29 e. Braincase · ............................. 33 f. Mandible ................................ 40 ÇJ. Stapes •.. · ............................. 43 h. Dentition · ............................. 43 i. Postcranial Skeleton .................... 45 Rarojugalosaurus ~cgilli ............................. 47 Systematic Paleontoloqy · ....................... 47 Description ..................................... 48 a. General Features of the Skull . ......... 49 b. Dermal Bones of the Skull Roof .......... 53 c. palatoquadrate . ........................ 59 iv d. Dermal Bones of the Palate ............... page 59 ( e. Braincase · .............................. 64 f. Mandible . · ............................. 64 q. Dentition · ............................... 66 Dianosaurus petilus Younq, 1982 .••••••••••••••••••••• 67 History of study .••..••...••••.•.••....••••...•• 67 Systematic Paleontoloqy ......................... 68 Redescr ipt iorl ......................... 69 a. General Features of the Skull •. ......... 69 b. Dermal Bones of the Skull Roof ••••.•.•.• 73 c. Palatoquadrate •.••••••.•• . ............. 77 d. Dermal Bones of the Palate • 1 •••••••••••• 77 e. Braincase · ............................. 81 f. Mandible • · ............................. 82 g. Dentition · ........................... 82 SPECIMEN kEFERRED TO Homoeosaurus maximiliani Meyer, 1845 ................................................. 84 History backqround of the Specimen .............. 85 Description ...••.••... ........................ 86 a. Dermal Bones of the Skull Roof ........... 86 b. Palatoquadrate ••••.••..•• ............... 99 c. Dermal Bones of the Palate .............. 99 d. Braincase · ............................. 100 e. Mandible . · ............................. 101 f . Dentition · ............................. 101 g. Postcranial Skeleton .................... 103 ( Reconstruction of the Skull • .................... 107 v Assessment of the Principal Differences Between C.M. 6438 and Previously Studied Specimens ••... page 109 COMPARATIVE MORPHOLOGY OF THE MIDDLE EAR REGION OF THE FOSSIL GENERA AND ITS IMPLICATIONS FOR THE AUDITORY EVOLUTION OF THE SPHENODONTIDA ............. 114 FUNCTIONAL ANALYSIS OF THE TEMPORAL REGION ••..••.... 134 Cranial Muscles Innervated by the Trigeminal Nerve ....•••...........•.............•......... 135 a. Adductor Mandibulae .•.......•..•.••.... 136 b. Constrictor Internus Dorsalis ...... .... 155 Anatomical Differences of Cranial Muscles Between Sphenodon and Lizards . ............. .... 156 Relations Between the Adductor Jaw M\!scles and Associated Bony Structures ...•......•....•..... 164 Adductor Jaw Muscles in Primitive Sphenodontidans 174 Reduction of the M. Pterygoideus Atypicus 177 Retention of the Fixed Quadrate in the Sphenodontida ... 184 INTERRELATIONSHIPS WITHIN THE SPHENODONTIDA ...••.... 189 CONCLUSIONS •.......•.••....••...••..•............... 208 REFERENCES •••.•••..••••••••.•••••••••.••..••••••...• 210 ABBREVIATIONS .. • . • • . • • • . • • . • . • . • . • . 223 vi LISTS or ILLUSTRATIONS Figure page 1 Geological map of Lufeng Basin .................. 7 2 Sphenodontidan-bearing beds in Lower Lufeng Formation ................................... 9 3 Asiacephalosaurus wangi, dorsal skull restoration 14 4 Asiacephalosaurus wangi, 1ateral skull restoration .. .. 15 5 Asiacephalosaurus wangi, ventral skull restoration . .. 16 6 Asiacephalosaurus wangi, occipital restoration 17 7 Asiacephalosaurus wangi, lateral view of skull 19 8 Asiacephalosaurus wangi, lateral view of partial skull ................................ 21 9 Asiacephalosaurus wangi, dorsal view of skull 23 10 Asiacephalosaurus wangi, anatomy of cephalic surface of quadrate •.•...•••..•••.••...•...• • . .. 26 11 Asiacephalosaurus wangi, occipital view of sku1l 28 12 Asiacephalosaurus wangi, ventral view of skull .. JO 13 Asiacephalosaurus wanqi, ventral view of partial skull ................. I!' •••••••••••••• 31 14 Asiacephalosaurus wangi, lateral view of braincase ..... .. 37 15 Asiacephalosaurus wangi, medial view of 1eft mandible ............................... .. 39 16 Asiacephalosaurus wangi, maxillary dentition ••.. 44 vii Figure page 17 Rarojugalosaurus mcqilli, dorsal skull restorat ion .. .. 50 18 Rarojugalosaurus
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