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CO 2 ^^5' ^v >: I I '^NOIini TUTION *^N0liniIlSNI_NVIN0SHlllMS*^S3 yV8 a l^L I B R AR I ES*^SMITHSONlAN INSTITUTION ^ CO ^ z \ ^'^ ^ 5 c'J 5 ^ QL 66fi iiversity of California Publications L25U4 1987X ZOOLOGY Rept. Volume 118 Phylogenetic Systematics of Iguanine Lizards A Comparative Osteological Study by Kevin de Queiroz PHYLOGENETIC SYSTEMATICS OF IGUANINE LIZARDS A COMPARATIVE OSTEOLOGICAL STUDY KEPT. Phylogenetic Systematics of Iguanine Lizards/ A Comparative Osteological Study by Kevin de Queiroz A Contribution from the Museum of Vertebrate Zoology of the University of California at Berkeley yti»%K«^*^ UNIVERSITY OF CALIFORNIA PRESS Berkeley • Los Angeles • London UNIVERSITY OF CALIFORNIA PUBLICATIONS IN ZOOLOGY Editorial Board: Peter B. Moyle, James L. Patton, Donald C. Potts, David S. Woodruff Volume 118 Issue Date: December 1987 UNIVERSITY OF CALIFORNIA PRESS BERKELEY AND LOS ANGELES, CALIFORNIA UNIVERSITY OF CALIFORNIA PRESS, LTD. LONDON, ENGLAND ISBN 0-520-09730-0 LIBRARY OF CONGRESS CATALOG CARD NUMBER: 87-24594 © 1987 BY THE REGENTS OF THE UNIVERSITY OF CALIFORNIA PRINTED IN THE UNITED STATES OF AMERICA Library of Congress Cataloging-in-Publication Data De Queiroz, Kevin. Phylogenetic systematics of iguanine lizards: a comparative osteological study / by Kevin de Queiroz. cm. — p. (University of California publications in zoology: v. 118) Bibliography: p. ISBN 0-520-09730-0 (alk. paper) 1. Iguanidae—Classification. 2. Iguanidae—Evolution. 3. — Iguanidae ^Anatomy. 4. Anatomy, Comparative. 5. Reptiles— Qassification. 6. — — Reptiles Evolution. 7. Reptiles Anatomy. I. Title. II. Series. QL666.L25D4 1987 597.95—dc 19 87-24594 CIP Contents Li^f of Illustrations, vii List of Tables, x Acknowledgments, xi Abstract, xii INTRODUCTION 1 Historical Review, 1 Goals of This Study, 10 MATERIALS AND METHODS 1 3 Specimens, 13 Phylogenetic Analysis, 13 Basic Taxa, 14 The Problem of Variation, 14 Construction of Branching Diagrams, 16 IGUANINE MONOPHYLY 18 COMPARATIVE SKELETAL MORPHOLOGY 2 1 Skull Roof, 21 Palate, 39 Braincase, 44 Mandible, 49 Miscellaneous Head Skeleton, 59 Axial Skeleton, 69 Pectoral Girdle and Sternal Elements, 81 Pelvic Gridle, 86 Limbs, 89 Osteoderms, 89 NONSKELETAL MORPHOLOGY 92 Arterial Circulation, 92 Colic Anatomy, 93 External Morphology, 94 vi Contents SYSTEMATIC CHARACTERS 100 Skeletal Characters, 100 Nonskeletal Characters, 104 CHARACTER POLARITIES AND THE PHYLOGENETIC INFORMATION CONTENT OF CHARACTERS 1 06 ANALYSIS OF PHYLOGENETIC RELATIONSHIPS 1 17 PreHminary Analysis, 1 17 Lower Level Analysis, 122 PHYLOGENETIC CONCLUSIONS 130 Preferred Hypothesis of Relationships, 130 Character Evolution within Iguaninae, 130 COMPARISONS WITH PREVIOUS HYPOTHESES 132 DIAGNOSES OF MONOPHYLETIC GROUPS OF IGUANINES 135 Iguaninae Bell 1825, 135 Dipsosaurus Hallowell 1854, 141 Brachylophus Wagler 1830, 143 Iguanini Bell 1825, 145 Ctenosaura Wxtgmonn 1828, 146 Sauromalus T)\xvi\€n\ 1856, 157 Amblyrhynchina, new taxon, 160 Amblyrhynchus Bell 1825, 163 Conolophus Fitzinger 1843, 165 IguaninaBell 1825, 167 Iguana Laurenti 1768, 168 Odwra Harlan 1824,170 Appendix I: Specimens Examined, 175 Appendix II: Polarity Determination Under Uncertain Outgroup Relationships, 179 Appendix III: Polarity Determination for Lower Level Analysis, 185 Appendix IV: Polarity Reevaluation for Lower Level Analysis, 187 Literature Cited, 191 List of Illustrations FIGURES of North American after Mittleman 1 . "The phylogeny and relationships iguanid genera," (1942), 6 2. "Grouping and possible phylogeny of the genera of iguanids occurring in the United States," after H. M. Smith (1946), 7 3. "Phylogenetic relationships of the Madagascar Iguanidae and the genera of iguanine Hzards," after Avery and Tanner (1971), 9 4. Etheridge's phylogeny of the Iguanidae, 1 1 5. Skull of Braehylophus vitiensis, 22 6. Skull and mandible of Braehylop hus vitiensis, 23 7. Posteroventral views of iguanine premaxillae, 24 8. Dorsal views of the preorbital portions of iguanine skulls, 25 9. Dorsal views of the skulls of Cyclura cornuta and Sauromaliis obesus, 11 10. Posterodorsal views of the anterior orbital regions oi Brachylophus fasciatm and Conolophus pallidus, 28 11. Dorsal view of the skull of Amblyrhynchus cristatus, 29 12. Ventral views of iguanine frontals, 31 13. Dorsal views of the parietals in an ontogenetic series of Iguana iguana, 34 14. Lateral view of the skull of Ctenosaura similis, 36 15. Lateral views of the posterolateral comers of iguanine skulls, 38 16. Posterodorsal views of disarticulated right palatines of Iguana delicatissima and Conolophus subcristatus, 40 17. Posterodorsal views of the right orbits of five iguanines and Morunasaurus annularis, 41 18. Ventral view of the skull of Iguana delicatissima, 43 19. Anterolateral views of the left orbitosphenoids in an ontogenetic series of Iguana iguana, 45 20. Ventral views of the posterior portion of the palate and anterior portion of the braincase of Sauromalus varius and Amblyrhynchus cristatus, 46 21. Ventral views of iguanine neurocrania, 47 22. Lateral views of the right mandibles of Iguana delicatissima and Amblyrhynchus cristatus, 50 23. Lingual views of the left mandibles of three iguanines, 51 24. Lateral views of the right mandibles of Conolophus pallidus and Cyclura cornuta, 52 vu viii List of Illustrations 25. Lateral views of the right mandibles of Iguana delicatissima, Sauromalus obesus, and Amblyrhynchus cristatus, 53 26. Lateral views of the right mandibles of Dipsosaurus dorsalis, Brachylophus vitiensis, and Iguana iguana, 55 27. Medial views of the left mandibles of Iguana delicatissima and Conolophus subcristatus, 56 28. Dorsal views of the posterior ends of the right mandibles in ontogenetic series of Ctenosaura hemilopha and Amblyrhynchus cristatus, 57 29. Dorsal views of the posterior ends of the right mandibles in an ontogenetic series of Dipsosaurus dorsalis, 58 30. Lingual views of left maxillary teeth of four iguanines and Basiliscus plumifrons, 62 31. Hypothetical character phylogeny for the iguanine pterygoid tooth patch, 65 32. Corneal view of the left scleral ring of Ctenosaura similis, 67 33. Ventral views of the iguanine hyoid apparati, 68 34. Twentieth presacral vertebra of Brachylophus vitiensis, 70 35. Lateral views of the twentieth presacral vertebrae of Sauromalus obesus and Ctenosaura pectinata, 11 36. Dorsolateral views of the twentieth presacral vertebrae of Dipsosaurus dorsalis and Sauromalus obesus, 73 37. Dorsal views of caudal vertebrae of Dipsosaurus dorsalis from different regions of the tail, 76 38. Lateral views of the ninth caudal vertebrae of Dipsosaurus dorsalis and Iguana iguana, 79 39. Presacral and sacral vertebrae and ribs of Dipsosaurus dorsalis in ventral view, 80 40. Pectoral girdles of three iguanines, 82 41. Dorsal views of the pelvic girdles of Sauromalus obesus and Ctenosaura pectinata, 86 42. Bones of the anterior limb of Brachylophus fasciatus, 87 43. Right hind limb skeleton of Brachylophus fasciatus, 88 44. Right
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