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The Cephalic Lateralis System of Cardinalfishes (Perciformes

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The Cephalic Lateralis System of Cardinalfishes (Perciformes THE CEPHALIC LATERALIS SYSTEM OF CARDINALFISHES (PERCIFORMES: APOGONIDAE) AND ITS APPLICATION TO THE TAXONOMY AND SYSTEMATICS OF THE FAMILY A DISSERTATION SUBMITTED TO THE GRADUATE DIVISION OF THE UNIVERSITY OF HAWAI'I IN PARTIAL FULLFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN ZOOLOGY (ECOLOGY, EVOLUTION AND CONSERVATION BIOLOGY) AUGUST 2004 By Laura M. Rodman Bergman Dissertation Committee: David Greenfield, Chairperson Robert Cowie Kenneth Kaneshiro Jack Randall Rebecca Cann ACKNOWLEDGMENTS I am grateful to all the curators and collection manages who allowed me to borrow and examine apogonid specimens: A. Suzumoto, O. Gon, B. Hutchings, S. Jewitt, and 1. Williams. I would particularly like to thank Alex Vagelli at New Jersey State Aquarium for the loan ofspecimens ofPterapogon (Pterapogon) kauderni. I would like to thank Sue Monden, the Department ofZoology illustrator, for allowing me to borrow one ofher microscopes with a camera lucida attachment during the illustration phase of this research. I am indebted to my advisor, Dave Greenfield for his support and confidence during the years it has taken me to complete this project. I am grateful to my committee members, Becky Cann, Rob Cowie, Ken Kaneshiro, and Jack Randall for their invaluable insight and advice. This research was sponsored, in part, by the Ecology, Evolution, and Conservation Biology Program's Graduate Teaching in K-12 Education Fellowship. 111 ABSTRACT The Apogonidae is one ofthe most speciose coral reeffish families. Members ofthis family are found on every coral reefin all tropical and subtropical waters worldwide and yet, despite this apparent ubiquity, the systematic relationships ofits species are poorly understood. Few studies have attempted to address the question ofthe family's evolutionary history and, as a result, little is known about the evolution ofthis major group ofcoral reeffish. This dissertation addresses this shortcoming by using characters derived from the skeleton, external morphology, and cephalic lateralis system to produce a cladistically derived hypothesis ofthe evolutionary relationships among the genera and subgenera ofthe family Apogonidae. The cephalic lateralis ofthe type species ofeach apogonid genus and subgenus was described and illustrated, and comments were made on the variability ofthis system within the supra-specific taxon to which the species belongs. The cephalic lateralis ofapogonids is a highly diverse system; much variation exists within and among the family's genera and subgenera. Cladistic analyses resulted in a number ofwell-resolved, but not well-supported (as indicated by the consistency and retention indices), equally parsimonious cladograms. Homoplasies aside, all ofthe cladograms suggest that the apogonid genera Apogon sensu lato and Pterapogon sensu lato are not monophyletic as they are currently defined. The results ofthis research highlight the need for revision ofthe family's classification in order more accurately reflect the group's phylogenetic history. tV TABLE OF CONTENTS ACKNOWLEDGEMENTS................................................................ 111 ABSTRACT.................................................................................. IV LIST OF TABLES........................................................................... V11 LIST OF FIGURES......................................................................... Vl11- CHAPTER 1. INTRODUCTION TO THE APOGONIDAE......................... 1 APOGONID CLASSIFICATION AND PHyLOGENY........................... 4 THE LATERALIS SYSTEM AND ITS PREVIOUS UTILIZATION IN FISH SySTEMATICS 24 CHAPTER 2. DESCRIPTIONS AND ILLUSTRATIONS OF THE CEPHALIC LATERALIS OF CARDINALFISHES................................................... 31 DESCRIPTION AND ILLUSTRATION OF THE CRANIAL LATERALIS SySTEMS 32 INSTITUTION ABBREVIATIONS.............................................. 32 METHODS 32 PRESENTATION OF DATA......... 33 SUBFAMILY APOGONINAE........................................................ 39 APOGON (APOGON) LACEPEDE, 1801....................................... 39 APOGON (BREPHAMIA) JORDAN IN JORDAN & JORDAN, 1922.. .... 47 APOGON (JAYDIA) SMITH, 1961................................................ 52 APOGON (LEIPDAMIA) GILL, 1863... 57 APOGON(OSTORHINCHUS)LACEPEDE, 1802 , 64 APOGON (PARONCHEILUS) SMITH, 1964......... 72 APOGON (PRISTIAPOGON) KLUNZINGER, 1870....................... 77 APOGON (PRISTICON) FRASER, 1972.......................................... 84 APOGON (YARICA) WHITLEY, 1930........................................... 91 APOGON (ZAPOGON) FRASER, 1972........... 96 APOGON (ZORAMIA) JORDAN, 1917[b].............................. 101 APOGONICHTHYS BLEEKER, 1854[a]......................................... 107 ARCHAMIA GILL, 1863 ,............. 112 ASTRAPOGONFOWLER, 1907................................................... 118 CERCAMIA RANDALL & SMITH, 1988....................................... 124 CHEILODIPTERUSLACEPEDE, 1801.......................................... 129 CORANTHUS SMITH, 1961...... 136 FOA JORDAN & EVERMANN IN JORDAN & SEALE, 1905... 142 FOWLERIA JORDAN & EVERMANN, 1903......... 148 GLOSSAMIA GILL, 1863........................................................... 154 HOLAPOGONFRASER, 1973.................................................... 159 LACHNERATUS FRASER & STRUHSAKER, 1991........................... 165 NEAMIA SMITH & RADCLIFFE IN RADCLIFFE & RADCLIFFE, 1912 170 PHAEOPTYXFRASER & ROBINS, 1970................................... 174 PTERAPOGON (PTERAPOGON) KOUMANS, 1933... 180 PTERAPOGON (QUINCA) MEES, 1966.......................................... 185 RHABDAMIA (BENTUVIAICHTHYS) SMITH, 1961........................ 190 V RHABDAMIA (RHABDAMIA) WEBER, 1909................................. 196 RHABDAMIA (VERULUX) FRASER, 1972.................................... 201 SIPHAMIA WEBER, 1909.......................................................... 207 SPHAERAMIA FOWLER & BEAN, 1930....................................... 212 VINCENTIA CASTELNAU, 1872............... 218 SUBFAMILY PSEUDAMINAE...................................................... 224 GYMNAPOGONRRREGAN, 1905............................................... 224 PAXTON BALDWIN & JOHNSON, 1999....................................... 229 PSEUDAMIA BLEEKER, 1865................................................... 234 PSEUDAMIOPS SMITH, 1954.................................................... 239 CHAPTER 3: PHYLOGENY OF CARDINALFISHES............................... 244 METHODS FOR CLADISTIC ANALySIS......................................... 247 ASSESSMENT OF THE MONOPHYLY OF APOGON...................... 247 CLADISTIC ANALYSIS OF FRASER'S DATA FOR THE APOGONIDAE...................................................................... 248 RESULTS...... 249 CHARACTERS COMPILED FROM FRASER'S (1972) MONOGRAPH. 249 ASSESSMENT OF RELATIONSHIPS WITHNG APOGON................. 273 CLADISTIC ANALYSIS OF THE APOGONDIAE........................... 273 DISCUSSION............................................................................ 277 ASSESSMENT OF THE RELATIONSHIPS WITHIN APOGON........... 277 CLADISTIC ANALYSIS OF FRASER'S DATA FOR THE APOGONIDAE...................................................................... 279 IMPLICATIONS FOR THE CLASSIFICATION OF THE FAMILy....... 293 CHAPTER 4. EVOLUTION OF THE APOGONID CEPHALIC LATERALIS.. 299 METHODS FOR THE CLADISTIC ANALYSIS OF THE COMBINED DATA SET............ 300 RESULTS... 301 CHARACTERS COMPILED FROM THE DESCRIPTIONS OF THE CEPHALIC LATERALIS SySTEM............................................... 301 SHARED CHARACTERS...................................................... 301 VARIABLE CHARACTERS................................................... 303 CLADISTIC ANALYSIS OF THE COMBINED DATA SET................ 315 DISCUSSION.. ...... .... ..... ................... ... ... ........................... ....... 318 COMMENTS ON THE USEFULNESS OF CEPHALIC LATERALIS CHARACTERS IN THE CLADISTIC ANALYSIS OF THE APOGONIDAE...................................................................... 318 EVOLUTION OF THE CEPHALIC LATERALIS SYSTEM IN THE APOGONIDAE...................................................................... 320 CLADISTIC ANALYSIS OF THE COMBINED DATA SET......... 326 CHAPTER 5. CONCLUSIONS.......................................................... 338 LITERATURE CITED..................................................................... 349 VI LIST OF TABLES Table Page 1. Classification ofthe Apogonidae....................... 7-8 Vll LIST OF FIGURES Figure Page 1. General structure ofthe apogonid canal system fishes.. ....... ....... 37 2. Illustration ofarbitrary canal divisions.................................. 38 3. Canal System ofApogon (Apogon) imberbis.......................... 45-46 4. Canal System ofApogon (Brephamia) parvulus........................ 50-51 5. Canal System ofApogon (Jaydia) truncatus............................ 56-57 6. Canal System ofApogon (Lepidamia) kalosoma....................... 63-64 7. Canal System ofApogon (Ostorhinchus)jleurieu..................... 70-71 8. Canal System ofApogon (Paroncheilus)fragilis...................... 76-77 9. Canal System ofApogon (Pristiapogon)fraenatus.................... 83-84 10. Canal System ofApogon (Pristicon) trimaeulatus..................... 89-90 11. Canal System ofApogon (Yarica) hyalosoma.......................... 94-95 12. Canal System ofApogon (Zapogon) evermanni........................ 100-101 13. Canal System ofApogon (Zoramia) leptacanthus....................
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