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Istiblennius, Blenniella, and Paralticus, New Genus

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Istiblennius, Blenniella, and Paralticus, New Genus * The Indo-West Pacific Blenniid Fish Genus Istiblennius Reappraised: A Revision of Istiblennius, Blenniella, and Paralticus, New Genus VICTOR G. SPRINGER and JEFFREY T. WILLIAMS I SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 565 SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of "diffusing knowledge" was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to the Marine Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Folklife Studies Smithsonian Studies in Air and Space Smithsonian Studies in History and Technology In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world of science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs'submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. Robert McC. Adams Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 565 The Indo-West Pacific Blenniid Fish Genus Istiblennius Reappraised: A Revision of Istiblennius, Blenniella, and Paralticus, New Genus Victor G. Springer and Jeffrey T. Williams SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1994 ABSTRACT Springer, Victor G., and Jeffrey T. Williams. The Indo-West Pacific Blenniid Fish Genus Istiblennius Reappraised: A Revision of htiblennius, Blenniella, and Paralticus, New Genus. Smithsonian Contributions to Zoology, number 565, 193 pages, 73 figures, 45 tables, 1994.—Istiblennius Whitley, as most recently defined (Smith-Vaniz and Springer, 1971), comprises 2 monophyletic groups of species, which we recognize as separate genera, Blenniella Reid (9 species) and Istiblennius (14 species). A cladistic analysis supports the hypothesis that Blenniella and Istiblennius form a clade. The 14 species of Istiblennius include 2 new species: /. pox, from southernmost Red Sea, Persian Gulf, Gulf of Oman, and /. spilotus, from most of East African coast, Comoro Islands, Madagascar, Gulf of Oman, and Pakistan. A monotypic new genus, Paralticus, is described for Salarias amboninensis Bleeker, which had been assigned most recently (Smith-Vaniz and Springer, 1971) to Praealticus Schultz and Chapman. Although superficially similar to Istiblennius species, P. amboinensis is a member of a clade that also comprises Praealticus, Alticus, and Andamia. Most of the species we revise exhibit statistically significant meristic sexual dimorphism, as well as noticeable geographic variation. All species are illustrated, including both sexes, individual and geographic variants, and ophioblennius stage prejuveniles (for all 5 species for which these stages are known). Distribution maps for all species are presented. Cladistic analyses are given for a subset of the genera of the tribe Salariini, to which the 3 genera we revise belong, and for the species of Blenniella and Istiblennius. The generic level analysis corroborates a major portion of the analysis proposed by Williams (1990). Because of character variability, we consider the results of the generic analyses highly tentative. Discussions of the biogeography of the genera and species are presented. The Philippine Islands are noted as a general area of endemism for fishes and an explanation is proposed. The pattern of east-west disjunctly distributed species, or sister species, first noted by Springer and Williams (1990), is modified slightly and shown to be common one for fishes. The western portion of the disjunct distributions appears to be restricted to the western Indian Ocean south of the equator and west of the mid-Indian Ocean Ridge. Some species that are not disjunctly distributed are limited in distribution to the same area of the Indian Ocean as the western disjunct portions of other species. The area of disjunction almost always includes, at least, the Gulf of Thailand and the southern half of the South China Sea. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus). Library of Congress Cataloging-in-Publication Data Springer, Victor Gruschka, 1928- The Indo-West Pacific blenniid fish genus Istiblennius reppraised: a revision of Istiblennius, Blenniella, and Paralticus, new genus / Victor G. Springer and Jeffrey T. Williams p. cm.—(Smithsonian contributions to zoology ; no. 565) Includes bibliographical references. 1. Istiblennius. 2. Blenniella. 3. Paralticus. 4. Blenniidae-Classification. I.Williams, Jeffrey T. II. Title. III. Series Q11.S54 no. 565 [QL638.B6] 591 s—dc20 94-15323 [59T.58] CIP © The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48—1984. Contents Page Introduction 1 Methods 2 Acknowledgments 9 Joint Nomenclatural Discussion of Blenniella, Istiblennius, and Paralticus 9 Key to Species of Blenniella, Istiblennius, and Paralticus 17 Blenniella 23 Blenniella cyanostigma 23 Blenniella caudolineata 31 Blenniella interrupta 35 Blenniella leopardus 40 Blenniella bilitonensis 43 Periophthalmus Species Complex . 49 Blenniella paula 65 Blenniella periophthalmus 65 Blenniella gibbifrons 68 Blenniella chrysospilos 78 Istiblennius 85 Lineatus Species Complex 86 Istiblennius pox, new species 96 Istiblennius steindachneri 97 Istiblennius lineatus 98 Istiblennius rivulatus 100 Istiblennius edentulus 103 The Istiblennius enosimae Problem 118 Istiblennius meleagris 126 Istiblennius dussumieri 131 Istiblennius zebra 139 Istiblennius bellus 143 Istiblennius muelleri 149 Istiblennius unicolor 152 Istiblennius spilotus, new species 155 Istiblennius flaviumbrinus 158 Istiblennius colei 161 Paralticus, new genus 164 Paralticus amboinensis 164 Phylogenetic Analysis 167 Intergeneric Relationships 170 Blenniella Intrarelationships 173 Istiblennius Intrarelationships 175 Biogeographic Discussion 177 Literature Cited 189 in "The secret of life is to have a task...And the most important thing is—it must be something you cannot possibly do!" Henry Moore, sculptor The Indo-West Pacific Blenniid Fish Genus Istiblennius Reappraised: A Revision of Istiblennius, Blenniella, and Paralticus, New Genus Victor G. Springer and Jeffrey T. Williams Introduction specimens, and thousands of repeat counts made for purposes The original purpose of our study was to revise the genus of verification. Numerous radiographed specimens were Istiblennius Whitley as currently accepted and as essentially counted independently by one of us and an assistant, and where defined by Smith-Vaniz and Springer (1971). Springer began counts for the same specimen differed, they were repeated one work on the revision as a secondary project shortly after 1971, or more times to rectify the differences. and progressed slowly until 1984, when he began phasing it in It was not until we had essentially completed the routine as his primary project and Williams joined in the effort. A examination and analysis of our material, prepared the species preliminary assessment of museum holdings of Istiblennius descriptions, and begun to prepare summary tables of charac- specimens indicated that it was the most abundantly repre- ters (Tables 1-3), that we became convinced we were dealing sented genus of Salariini, if not all blenniids, and that the time with at least three clearly different groups of species. Further required to examine the material and to record and analyze data examination of the species, including skeletal preparations of taken from it, conceivably could extend beyond that remaining all but one species, convinced us that the groups should be to the senior author. recognized generically. We also found indications that the genera probably formed a polyphyletic group. For our study, we recorded and analyzed detailed data from The obvious next step in our study would have been a over 9000 specimens, which emanated from 35 museums and phylogenetic analysis of all currently recognized genera (28) one private collection, almost all of which we visited. We and species (estimated at over 200) of Salariini. Fortunately, estimate that we encountered many thousands more museum Williams (1990) had already hypothesized characters for a specimens of Istiblennius and Blenniella on which we recorded node that split
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