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The Indo-Pacific Blenniid Fish Genus Ecsenius

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The Indo-Pacific Blenniid Fish Genus Ecsenius The Indo-Pacific Blenniid Fish Genus Ecsenius VICTOR G. SPRINGER SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 465 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 465 The Indo-Pacific Blenniid Fish Genus Ecsenius Victor G. Springer SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1988 ABSTRACT Springer, Victor G. The Indo-Pacific Blenniid Fish Genus Ecsenius. Smithsonian Contributions to Zoology, number 465, 134 pages, 68 figures, 14 color plates, 30 tables, 1988.—Ecsenius McCulloch is defined and its major specializations illustrated. It is not possible to propose its sister group unequivocally within the tribe Salariini. Ecsenius comprises 12 species groups, most hypothesized to be monophyletic based on a few heavily weighted characters; a few groups are possibly paraphyletic. The groups contain from one to nine species. Interrelationships of some groups, and some species within some groups, are hypothesized based on heavily weighted characters and/or distribution patterns. Ecsenius comprises 46 species, 20 here described as new. The new species are: E. aequalis (Great Barrier Reef; Trobriand Islands), E. alleni (Rowley Shoals and Scott Reef, Western Australia), E. australianus (Great Barrier Reef), E. axelrodi (Hermit Islands; New Britain; Solomon Islands), E. bathi (Java; Komodo), E. dentex (Gulf of Aqaba), E. dilemma (Philippines), E. fijiensis (Fiji: Viti Levu south to Vatoa, Lau Islands), E. kurti (Philippines), E. lubbocki (Phuket, Thailand, Indian Ocean), E. monoculus, (Philippines; Dot du Sud, Viet Nam; Moluccas), E. oculatus (Christmas Island, Indian Ocean; Western Australia), E. pardus (Fiji), E. paroculus (Phuket, Thailand; Malacca, Malaysia; Mentawei, Bawean, Seribu islands, Indonesia), E. portenoyi (Rotuma; American Samoa), E. sellifer (Umboi Island and Trobriand Islands, Papua New Guinea; Solomon Islands; Palau Islands), E. stictus (Great Barrier Reef), E. taeniatus (Basilaki and Goodenough islands, western Papua New Guinea), E. tessera (New Caledonia; New Hebrides), and E. tigris (Osprey and Bougainville reefs, Coral Sea). Ecsenius minutus Klausewitz, formerly synonymized with E. nalolo Smith, is resurrected. Several of the species have two or three strikingly different color-pattern forms, which are not associated with size or sex. A key is given to all species, all are illustrated in black-and-white halftones, and most are illustrated live in color photographs. Distribution maps are provided for all the species groups and species. Of the 12 species groups, all the species within each of five groups, containing from three to nine species, are allopatric. Two of these five groups, one with five and one with eight species, are hypothesized to form a clade. Of the 13 species in this clade, 12 are allopatric. One of the five species in one group is broadly sympatric with six of the eight species in the other group. The overall distribution of the two groups, and the distributions of the species in each of them, are used as the basis for a speculative vicariance biogeographic discussion, in which the tectonic history of India is proposed as the primary physical barrier that isolated biotas in the western Indian Ocean and allowed them to diverge. Species within a group usually exhibit sharply delimited distributions with respect to each other. For instance, one species of a group may have an extremely broad distribution with islandless voids of hundreds of kilometers separating its populations. Populations of the same species, however, may be separated from populations of another species in the same group at other islands by distances of only a few tens of kilometers. Such distribution patterns appear to be better explained by the tectonic history of the pertinent areas than by dispersal. 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 cavarnosa (Linnaeus). Library of Congress Cataloging-in-Publication Data Springer, Victor Gruschka, 1928- The Indo-Pacific blenniid fish genus Ecsenius. (Smithsonian contributions to zoology ; no. 465) Bibliography: p. Supt of Docs, no.: SI 1.27:465 1. Blenniidae—Classification. 2. Blenniidae—Geographical distribution. 3. Fishes—Classification. 4. Fishes— Geographical distribution. I. Title. II. Series. QL1.S54 no.465 [QL638.B6] 591 s 88-600023 [597'.58] Contents Page Introduction 1 Methods 2 Acknowledgments 7 Ecserdus McCulloch 8 Description and Relationships 8 General Biogeographical Comments 12 Key to the Species of Ecsenius 21 Pulcher Group 27 Ecsenius aroni Springer 28 Ecsenius gravieri (Pellegrin) 28 Ecsenius pulcher (Murray) 31 Stigmatura Group 32 Ecsenius lividanalis Chapman and Schultz 32 Ecsenius melarchus McKinney and Springer 34 Ecsenius midas Starck 37 Ecsenius stigmatura Fowler 39 Frontalis Group 41 Ecsenius frontalis (Ehrenberg) 41 Bicolor Group 44 Ecsenius bicolor (Day) 44 Ecsenius namiyei (Jordan and Evermann) 48 Lineatus Group 50 Ecsenius lineatus Klausewitz 50 Oculus Group 53 Ecsenius monoculus, new species 57 Ecsenius oculatus, new species 60 Ecsenius oculus Springer 62 Ecsenius pardus, new species 63 Ecsenius paroculus, new species 64 Ecsenius portenoyi, new species 65 Ecsenius sellifer, new species 66 Ecsenius tessera, new species 67 Yaeyamaensis Group 67 Ecsenius dentex, new species 69 Ecsenius minutus Klausewitz 71 Ecsenius nalolo Smith 72 Ecsenius stictus, new species 74 Ecsenius yaeyamaensis (Aoyagi) 75 Opsifrontalis Group 76 Ecsenius alleni, new species 81 Ecsenius australianus, new species 82 Ecsenius axelrodi, new species 84 Ecsenius bathi, new species 86 Ecsenius dilemma, new species 88 Ecsenius fijiensis, new species 90 Ecsenius fourmanoiri Springer 93 Ecsenius opsifrontalis Chapman and Schultz 94 iii iv SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Ecsenius tigris, new species 98 Isos Group 100 Ecsenius isos McKinney and Springer 101 Ecsenius lubbocki, new species 103 Ecsenius trilineatus Springer 104 Prooculis Group 105 Ecsenius bandanus Springer 106 Ecsenius bimaculatus Springer 107 Ecsenius collettei Springer 109 Ecsenius prooculis Chapman and Schultz 110 Ecsenius taeniatus, new species 112 Pictus Group 113 Ecsenius pictus McKinney and Springer 113 Mandibularis Group 115 Ecsenius aequalis, new species 115 Ecsenius kurti, new species 117 Ecsenius mandibularis McCulloch 119 Ecsenius schroederi McKinney and Springer 120 Appendix I: Speculative Discussion of the History of the Present-day Distribution of the Oculus-Yaeyamaensis Clade 122 Appendix II: Interrelationships and Biogeography of the Species of the Yaeyamaensis Group 127 Literature Cited 132 Color Plates follows page 134 The Indo-Pacific Blenniid Fish Genus Ecsenius Victor G. Springer Introduction Ecsenius in numbers of species. There is even a reasonable possibility that some of the Ecsenius species I recognize will This is the fourth study I have published (Springer, 1971, warrant subdivision when more complete geographical samples 1972; McKinney and Springer, 1976) on the systematics of are available (e.g., E. opsifrontalis, E. lineatus), or when more Ecsenius. In the first study I recognized 18 species, but the is known about their genetics (e.g., E.frontalis). There is also material actually comprised 26 species. Of the eight species a good possibilty that undescribed species of Ecsenius for that I did not recognize nomenclaturally, seven were described which no material is available lurk in various parts of as variations within other species of Ecsenius, and one Indonesia, the New Hebrides, and islands off eastern and (Ecsenius minutus Klausewitz), known from a single juvenile western Australia—areas
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