Systematics of the Carlia “fusca” Lizards (Squamata: Scincidae) of New Guinea and Nearby Islands George R. Zug Bishop Museum Bulletin in Zoology 5 Bishop Museum Press Honolulu, 2004 Cover: Published by Bishop Museum Press 1525 Bernice Street Honolulu, Hawai‘i 96817-2704, USA Copyright ©2004 Bishop Museum All Rights Reserved Printed in the United States of America ISSN 0893-312X Zug — Carlia “fusca” Lizards from New Guinea and Nearby Islands v TABLE OF CONTENTS Acknowledgments ......................................................................................................................... vii Abstract ........................................................................................................................................ viii Introduction ................................................................................................................................... 1 Carlia: An Analysis for Species Relationships ........................................................................... 1 Characters and Taxa .................................................................................................................. 2 Phylogenetic Analysis................................................................................................................ 8 New Guinea Carlia “fusca” ....................................................................................................... 9 Materials and Methods.................................................................................................................. 9 Characters Examined ................................................................................................................ 10 Mensural characters ............................................................................................................. 10 Meristic characters............................................................................................................... 12 Color pattern ........................................................................................................................ 12 General analytical protocols ................................................................................................ 12 Character Analysis: Results ........................................................................................................ 13 Sample Size and Variation ........................................................................................................ 13 Body Size and Proportions ....................................................................................................... 14 Size at sexual maturity ........................................................................................................ 14 Sexual dimorphism .............................................................................................................. 16 Head Scalation .......................................................................................................................... 17 Interparietal and prefrontals ................................................................................................ 17 Supraoculars, supraciliaries, and eyelid scales ................................................................... 17 Temporals and loreals ........................................................................................................ 17 Labials ................................................................................................................................. 17 Auricular lobes and ear shape ............................................................................................ 18 Body and Limb Scalation.......................................................................................................... 18 Dorsal and midbody scale rows .......................................................................................... 18 Keeling ................................................................................................................................ 18 Precloacal scales .................................................................................................................. 18 Subdigital lamellae .............................................................................................................. 18 Evaluation of Quantitative Characters....................................................................................... 18 Color pattern ............................................................................................................................. 19 Juvenile ................................................................................................................................ 20 Adult ................................................................................................................................... 23 Color pattern resume .......................................................................................................... 27 Geographic Patterns of Variation.................................................................................................. 27 Decisions on speciation in Carlia “fusca” ............................................................................... 35 Taxonomy .................................................................................................................................... 39 History and decisions ............................................................................................................... 39 Taxa recognized .................................................................................................................. 41 Carlia Gray........................................................................................................................ 41 Carlia fusca (Duméril & Bibron) ................................................................................. 42 Carlia pulla (Barbour) .................................................................................................. 44 Sepik populations ........................................................................................................... 45 Carlia mysi new species ................................................................................................. 45 Carlia ailanpalai new species ........................................................................................ 47 Carlia eothen new species ............................................................................................ 48 Carlia luctuosa (Peters & Doria) ................................................................................. 50 Highlands populations ................................................................................................... 52 vi BISHOP MUSEUM BULLETIN IN ZOOLOGY 5 (2004) Taxa recognized (continued) Carlia aenigma new species ............................................................................................... 53 Carlia aramia new species.................................................................................................. 54 Carlia longipes (Macleay) ................................................................................................. 56 Merauke population ........................................................................................................... 56 Carlia diguliensis (Kopstein) ............................................................................................. 57 Agats to Ayamura ................................................................................................................ 58 Carlia beccarii (Peters & Doria) ....................................................................................... 59 Kai unicolor ......................................................................................................................... 59 Carlia babarensis (Kopstein) ............................................................................................ 60 Carlia leucotaenia (Bleeker) ............................................................................................ 61 Carlia tutela new species ................................................................................................... 63 Biogeography of Carlia fusca Complex .................................................................................... 64 Literature Cited ............................................................................................................................. 70 Appendix .................................................................................................................................... 74 Table A. Specimens Examined, Carlia species phylogenetic analysis .................................... 74 Table B. Gazetteer of Sample Localities, Carlia “fusca” analysis........................................... 75 Table C. Discriminant Function Analysis Sample Sets ........................................................... 77 Table D. Specimens Examined, Carlia “fusca” analysis ......................................................... 78 Table E. Specimens Examined, but not part of the 76 sample localities ................................. 82 Zug — Carlia “fusca” Lizards from New Guinea and Nearby Islands vii ACKNOWLEDGMENTS This systematic investigation study arose from field observations made in Papua New Guinea dur- ing a 1971–1972 research trip sponsored by the Smithsonian Research Awards program. Other Smithsonian and National Museum of Natural