The Familial Phylogeny of the Tetraodontiformes (Acanthopterygii: Pisces) As Evidenced by Their Comparative Myology
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The Familial Phylogeny of the Tetraodontiformes (Acanthopterygii: Pisces) as Evidenced by Their Comparative Myology RICHARD WINTERBOTTOM m 4 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 155 SERIAL PUBLICATIONS OF THE SMITHSONIAN INSTITUTION The emphasis upon publications as a means of diffusing knowledge was expressed by the first Secretary of the Smithsonian Institution. In his formal plan for the Insti- tution, Joseph Henry articulated 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 keynote of basic research has been adhered to over the years in the issuance of thousands of titles in serial publications under the Smithsonian imprint, com- mencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Annals of Flight Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Studies in History and Technology In these series, the Institution publishes original articles and monographs dealing with the research and collections of its several museums and offices and of professional colleagues at other institutions of learning. These papers report newly acquired facts, synoptic interpretations of data, or original theory in specialized fields. These pub- lications are distributed by mailing lists to libraries, laboratories, and other interested institutions and specialists throughout the world. Individual copies may be obtained from the Smithsonian Institution Press as long as stocks are available. S. DILLON RIPLEY Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 155 The Familial Phylogeny of the Tetraodontiformes (Acanthopterygii: Pisces) as Evidenced by Their Comparative Myology Richard Winterbottom SMITHSONIAN INSTITUTION PRESS City of Washington 1974 ABSTRACT Winterbottom, Richard. The Familial Phylogeny of the Tetraodontiformes (Acanthopterygii: Pisces) as Evidenced by Their Comparative Myology. Smith- sonian Contributions to Zoology, number 155, 201 pages, 185 figures, 1974.—The myology of 46 species of the order Tetraodontiformes is described, the species representing the 11 families initially recognized for the purposes of this study. The data gathered from the descriptions has been interpreted phylogenetically, using only those characters of a shared specialized (synapomorph) nature. This analysis suggests that the Triacanthodidae and Triacanthidae are sister groups, and separated off first from the ancestral stock. The remaining families are divided into two lines. On the one hand, the Balistidae form the sister group of the Monacanthidae, and together form the sister group of the Aracanidae and Ostraciidae. In the other lineage, the Triodontidae separated off first, fol- lowed by the Molidae. The Diodontidae form the sister group of the Tetra- odontidae plus Canthigasteridae. In the final analysis, such fossils as are considered pertinent to the under- standing of tetraodontiform phylogeny are considered. The information pro- vided by these fossils, combined with the results of the myological interpretations, suggest the following systematic changes in the hierarchical classification of tetraodontiforms in order to bring it into line with the proposed phylogeny: (1) that two suborders be provisionally recognized, the first containing the Triacanthodidae and Triacanthidae, the second consisting of two superfamilies; (2) that a superfamily Balistoidea be recognized to contain the families Balistidae, fSpinacanthidae, and Ostraciidae; (3) that the families Monacanthidae and Aracanidae be treated as subfamilies of the Balistidae and Ostraciidae respec- tively, and that the fSpinacanthinae of Tyler (1968) be removed from the Triacanthodidae and placed as the sister group of the Balistidae, with equal rank (as fSpinacanthidae); (4) that a superfamily Tetraodontoidea be formed to contain the families tEoplectidae, Triodontidae, Tetraodontidae, Diodontidae, and Molidae, and, should -fZignoichthys prove to be further along the gymnodont line than fEoplectus, as hinted by Tyler (1972) and tentatively preferred here, the fZignoichthyidae; (5) that the family Canthigasteridae be reduced to sub- familial rank within the family Tetraodontidae. and that the fEoplectinae of Tyler (1972) be transferred from the Triacanthodidae to the superfamily Tetrao- dontoidea and raised to familial rank, in keeping with its phylogenetic position. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SI PRFSS NUMBER 4823. SERIES COVER DESICN: The coral Montastrea cavernosa (Linnaeus) . Library of Congress Cataloging in Publication Data Winterbottom, Richard, 1944— The familial phylogeny of the Tetraodontiformes. (Smithsonian contributions to zoology, no. 155) 1. Tetraodontiformes—Anatomy. 2. Phylogeny. 3. Muscles. 4. Fishes—Evolution. I. Title. II. Series: Smithsonian Institution. Smithsonian contributions to zoology, no. 155. QL1.S54 no. 155 [QL639] 591'.08s [597'.58] 73-2684 For sale by the Superintendent of Documents, U.S. Government Printing Office Washington, D.C. 20402 - Price $2.80 (paper cover) Contents Page Introduction 1 Acknowledgments .. 3 Materials, Methods, and Terminology 4 Myological Descriptions of Representative Triacanthodids 5 Subfamily Triacanthodinae 6 Triacanthodes anomalus (Schlegel) 6 Tydemania navigatoris Weber 11 Macrorhamphosodes platycheilus Fowler 12 Halirriochirurgus centriscoides Alcock 12 Subfamily Hollardiinae 12 Parahollardia lineata (Longley) 12 Hollardia hollardi Poey 13 Summary of Triacanthodidae 13 Myological Descriptions of Representative Triacanthids 13 Trixiphichthys weberi (Chaudhuri) 13 Triacanthus biaculeatus (Bloch) 18 Tripodichthys oxycephalus (Bleeker) 18 Summary of Triacanthidae 18 Myological Descriptions of Representative Balistids 18 Balistes vetula Linnaeus 19 Balistapus undulatus (Mungo Park) 24 Melichthys niger (Bloch) . 24 Rhinecanthus aculeatus (Linnaeus) 25 Odonus niger (Riippell) 25 Summary of Balistidae 25 Myological Descriptions of Representative Monacanthids 25 Pervagor melanocephalus (Bleeker) 26 Stephanolepsis auratus (Castlenau) 31 Paramonacanthus barnardi Fraser-Brunner 31 Chaetoderma spinosissimus (Quoy and Gaimard) 31 Paraluteres prionurus (Bleeker) . 31 Cantherhines pardalis (Riippell) 32 Oxymonacanthus longirostris (Bioch and Schneider) 32 Pseudalutarius nasicornis (Schlegel) 33 Ahiterus monoceros (Linnaeus) 33 Aluterus heudeloti Hollard 34 Aluterus schoepfi (Walbaum) 34 Anacanthus barbatus Gray 34 Summary of Monacanthidae 35 Myological Descriptions of Representative Aracanids 35 Kentrocapros aculeatus (Houttuyn) 35 Capropygia unistriata Kaup 39 ui Page Myological Descriptions of Representative Ostraciids 40 Ostracion tuberculatus Linnaeus 40 Lactoria cornuta (Linnaeus) 44 Tetrosomus concatenatus (Bloch) 44 Acanthostracion quadricornis (Linnaeus) 45 Summary of Ostraciidae 45 Myological Description of the Representative Triodontid 45 Triodon macropterus Lesson 45 Myological Descriptions of Representative Tetraodontids 50 Lagocephalus lunaris (Bloch and Schneider) 50 Amblyrhynchotes honckenii (Bloch) 56 Colomesus psittacus (Bloch and Schneider) 56 Xenopterus naritus (Richardson) 57 Arothron immaculatus (Bloch) 57 Tetraodon schoutedeni Pellegrin 58 Summary of Tetraodontidae 58 Myological Descriptions of Representative Canthigasterids 58 Canthigaster margaritatus (Riippell) 58 Canthigaster valentini (Bleeker) 63 Myological Descriptions of Representative Diodontids 63 Diodon hystrix Linnaeus 63 Chilomycterus orbicularis (Bloch) 68 Chilomycterus schoepfi (Walbaum) 68 Summary of Diodontidae 68 Myological Descriptions of Representative Molids 69 Mola mola Linnaeus 69 Ranzania laevis (Pennant) 72 Phylogenetic Significance of Observed Myological Variation 73 Interpretation 74 Discussion 90 Alternative Interpretations 94 A New Classification of the Tetraodontiformes 97 Literature Cited 100 List of Abbreviations 104 Figures 1-185 106-201 IV The Familial Phylogeny of the Tetraodontiformes (Acanthopterygii: Pisces) as Evidenced by Their Comparative Myology Richard Winterbottom Introduction changes appear to be of a grosser and less subtle nature (and thus more obvious) than those This paper represents an attempt to elucidate the involving other facets of the functional systems. familial phylogeny of the order Tetraodontiformes, Since this paper represents one of the first using a comparative myological approach. Past attempts at detailed phylogenetic interpretation classifications, usually based on external characters, from a purely myological standpoint, it seems neces- osteology or both, have often been vague concern- sary to outline certain factors influencing myology. ing the precise interrelationships of the families, The evolutionary mechanism of muscular change although certain family groupings have been almost may be by the subdivision of new sections from universally accepted. Since the aim is to resolve the existing muscles, the reduction and ultimate loss familial phylogeny, the eleven families recognized of a muscle, or change in fiber direction, since by Tyler (1970a) are provisionally accepted rather these alterations