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PHYLOGENETIC RELATIONSHIPS and TAXONOMIC REVISION of the BLENNIID FISH GENERA Cirripectes and Sear T Ichthys

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PHYLOGENETIC RELATIONSHIPS and TAXONOMIC REVISION of the BLENNIID FISH GENERA Cirripectes and Sear T Ichthys PHYLOGENETIC RELATIONSHIPS AND TAXONOMIC REVISION OF THE BLENNIID FISH GENERA Cirripectes AND Sear t ichthys By JEFFREY TAYLOR WILLIAMS A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 1986 ACKNOWLEDGEMENTS During my work on this project, many people have assisted me in various ways. I thank all of them. In particular, I thank W. F. Smith-Vaniz, E. Bohlke, W. Saul, and staff (Academy of Natural Sciences of Philadelphia); P. C. Heemstra (J. L. B. Smith Institute of Ichthyology); T. Iwamoto, W. N. Eschmeyer, L. Dempster, and staff (California Academy of Sciences); R. Winterbottom and staff (Royal Ontario Museum); C. R. Gilbert, G. Burgess and J. B. Miller (Florida State Museum); K. Hartel (Museum of Comparative Zoology, Harvard); M. L. Bauchot and M. Desoutter (Museum National d'Histoire Naturelle); A. G. Gerberich, K. A. Bruwelheide, L. P. Norrod , S. L. Jewett, J. R. Gomon, and Victor G. Springer (Smithsonian Institution), and the staff at all other museums that provided specimens, radiographs, and/or other technical assistance. I thank V. G. Springer, W. F. Smith-Vaniz, and H. Bath for their gracious hospitality while I was visiting museums. For providing color transparencies of specimens, I thank J. E. Randall, V. G. Springer, R. Winterbottom, and B. Carlson. C. R. Gilbert and V. G. Springer generously gave their time to discuss my project. I gratefully acknowledge their help. Financial support for this study was provided in part by the Department of Zoology and Florida State Museum, University of Florida; the California Academy of Sciences; and the National Science Foundation 11 (Predoctoral Improvement Grant DEB 8207313). I thank my parents for their support throughout the study. Finally, I thank my wife, Karen, for her support, editorial comments, and seemingly limitless understanding. in TABLE OF CONTENTS PAGE ACKNOWLEDGEMENTS " ABSTRACT v i INTRODUCTION 1 HISTORICAL TREATMENT OF THE BLENNIIDAE 6 METHODS 9 Taxonomic Methods 9 Methods of Phylogenetic Analysis 10 PHYLOGENETIC ANALYSIS 12 Family Blenniidae 15 Salariini-"Blenniini" Group 18 "Blenniini" 19 Salariini 20 Alticus Group 28 Node I Group Genus Istiblennius 32 Node II Group 32 Genus Entomacrodus 36 Node III Group 36 Genus Stanulus 37 Node IV Group 38 Genus Pereulixia - 45 Node V Group 49 Genus Exal lias 54 Node VI Group 54 Genus Cirripectes > Node VII Group 61 Genus Ophioblennius 61 Genus Scartichthys 62 DISCUSSION OF PHYLOGENETIC ANALYSIS OF GENERA 63 PHYLOGENETIC RELATIONSHIPS OF CIRRIPECTES SPECIES 68 Node VIII Group 68 Node IX Group Node X Group 73 LV , Node XI Group 76 Node XII Group 76 Node XIII Group 77 Node XIV Group 80 Node XV Group 80 Node XVI Group 81 Node XVII Group 82 Cirripectes SWAINSON 87 Key to the Species of Cirripectes 109 Cirripectes alboapicalis (Ogilby) 143 Cirripectes auritus Carlson 149 Cirripectes castaneus (Valenciennes) 155 Cirripectes chelomatus Williams and Mauge 165 Cirripectes filamentosus (Alleyne and Macleay) 170 Cirripectes fuscoguttatus Strasburg and Schultz 180 Cirripectes gi^berti new species 184 Cirripectes hutchinsi new species 189 Cirripectes imitator Williams 195 Cirripectes jenningsi Schultz 199 Cirripectes kuwamurai Fukao 203 Cirripectes obscurus (Borodin) 205 Cirripectes perustus Smith 209 Cirripectes polyzona (Bleeker) . 213 Cirripectes quagga (Fowler and Bail) 219 Cirripectes randalli new species. 226 Cirripectes springeri new species.,,, 230 Cirripectes stigmaticus Strasburg and Schultz 234 Cirripectes vanderbilti (Fowler) 241 Cirripectes variolosus (Valenciennes) 246 Scartichthys JORDAN AND EVERMANN 254 Key to the Species of Scartichthys 260 Scartichthys crapulatus new species 265 Scartichthys gigas (Steindachner) 267 Scartichthys variolatus (Valenciennes) 272 Scartichthys viridis (Valenciennes) 276 Exallias JORDAN AND EVERMANN 282 Exallias brevis (Kner) 283 HISTORICAL ZOOGEOGRAPHY 291 APPENDIX 302 LITERATURE CITED 307 BIOGRAPHICAL SKETCH 314 Abstract of Dissertation Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy PHYLOGENETIC RELATIONSHIPS AND TAXONOMIC REVISION OF THE BLENNIID FISH GENERA Cirripectes AND Scartichthys By Jeffrey Taylor Williams May 1986 Chairman: Carter R. Gilbert Major Department: Zoology The blenniid fish tribe Salariini comprises a group of marine shore fishes found circumglobally in tropical and subtropical seas. Members of the tribe are quite similar ecologically, with most being herbivores or deposit feeders and inhabiting rocky or coralline areas. The salariin genera Cirripectes , Scartichthys , Exallias , Ophioblennius , and Pereulixia form a monophyletic group closely related to the genera Stanulus , Entomacrodus , and Istiblennius . A phylogenetic analysis supports the hypothesis that each of the following genera is the sister genus to a combination of all preceding genera: Scartichthys , Ophioblennius , Cirripectes , Exallias , Pereulixia , least the Stanulus , Entomacrodus , and Istiblennius (including at J_. gibbifrons species complex). A hypothesis of the relationships among the species of Cirripectes is illustrated in a cladograra. VI . The genera Cirripectes and Scartichthys are taxonomically revised. Cirripectes occurs throughout the Indo-Pacific region and comprises the following 20 species: C_. alboapicalis (Australia to Easter Island), C_. auritus (western Indian Ocean to Line Islands), C_. castaneus (Indian Ocean and Red Sea to western Pacific Ocean), C_. chelomatus (Coral Sea to Tonga Islands), C. f ilamentosus (western Indian Ocean to western Pacific Ocean), C_. fuscoguttatus (Pacific Plate), C_. gilberti new species (Indian Ocean), C. hutchinsi new species (western Australia), C_. imitator (Taiwan to Japan and Ogasawara [Bonin] Islands), £. jenningsi (Gilbert Islands to Tuamotu Archipelago), C_. kuwamurai (Japan), C_. obscurus (Hawaiian Islands), C_. perustus (western Indian Ocean to Gilbert Islands), C. polyzona (western Indian Ocean to Line Islands), C. quagga (western Indian Ocean to Pitcairn Island Group), C_. randalli new species (Mauritius and Cargados Carajos Shoals), C. springeri new species (Philippines to Solomon Islands), C_. stigmaticus (western Indian Ocean to Samoa), C. vanderbilti (Hawaiian Islands and Johnston Island), and _C. variolosus (Pacific Plate). Scartichthys is restricted to the eastern Pacific Ocean and comprises four species: S. crapulatus new species (northern Chile and vicinity of Valparaiso), S^. gigas (Panama to northern Chile), S_. variolatus (San Felix, San Ambrosio, and Juan Fernandez Islands), and S_. viridis (vicinity of Valparaiso to southern Peru). Area cladograms derived from the phyiogenetic analyses are analyzed. Eleven geographic patterns indicated by the cladogram are discussed Vll ; INTRODUCTION The blenniid fish tribe Salariini comprises a group of marine shore fishes found circumglobally in tropical and subtropical seas. Members of the tribe are quite similar ecologically; most are herbivores or deposit feeders and inhabit rocky or coralline areas. The objectives of the present study are to 1 ) test the hypotheses that the genera Cirripectes , Scartichthys , Ophioblennius , Pereulixia and Exallias comprise a monophyletic group and are each monophyletic 2) if the first hypotheses are confirmed, form phylogenetic hypotheses of interrelationships of the species of Cirripectes ; 3) taxonomically revise Cirripectes and Scartichthys , and provide complete species descriptions of and distribution maps for each of the included species; and 4) analyze the distribution patterns of these genera and their included species. Smith-Vaniz and Springer (1971) gave a synopsis of the Salariini to provide a foundation for future work on the tribe. They pointed out several genera that needed additional study and provided a preliminary hypothesis of the relationships of the 24 genera they recognized in the tribe. Their method was not cladistic, but they did establish the monophyletic nature of the tribe Salariini. Although they did not cladistically analyze the members of the tribe, they provided a dendrogram (Smith-Vaniz and Springer, 1971: fig. 51) of the genera showing their interrelationships. This dendrogram suggests that the genera Cirripectes , Scartichthys , Exallias , Pereulixia , and Ophioblennius are more closely related to each other than to other members of the tribe. The genus Cirripectes Swainson is widespread throughout the Indo-Pacif ic . It is a common inhabitant of coral and rocky reefs and is usually common in heavy surge areas. Specimens of Cirripectes are common in museum collections, but complex nomenclatural problems and the lack of adequate keys to the species have resulted in numerous misidentif ications. Smith-Vaniz and Springer (1971) listed 24 nominal species of Cirripectes , and four additional species have since been described (Table 1). There has been no comprehensive revision of the genus and all published keys to species have been of a regional nature (Chapman, 1951, for the Indo-Australian Archipelago; Fukao , 1984, for Japan; Schultz and Chapman, 1960, for the Marshall and Mariana Islands; Smith, 1959, for the western Indian Ocean). As a result, numerous populations from different geographic areas have been described as new species, even though these populations are actually members of the same widespread species. There is also sexual polychroraatism in some species and, in many instances,
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