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With a Phylogenetic Analysis of the Acanthuroidei (Pisces)

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With a Phylogenetic Analysis of the Acanthuroidei (Pisces) Morphology of Luvarus imperialis (Luvaridae), with a Phylogenetic Analysis of the Acanthuroidei (Pisces) JAMES C. TYLER, G. DAVID JOHNSON, IZUMINAKAMURA, and BRUCE B. COLLETTE SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 485 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 FRONTISPIECE.—Cleared and stained larval specimens of Luvarus imperialis, upper (MCZ 55291,10.5 mm SL), and Zanclus cornutus, lower( MCZ 62046,9.5 mm SL). SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 485 Morphology of Luvarus imperialis (Luvaridae), with a Phylogenetic Analysis of the Acanthuroidei (Pisces) James C. Tyler, G. David Johnson, Izumi Nakamura, and Bruce B. Collette SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1989 ABSTRACT Tyler, James C, G. David Johnson, Izumi Nakamura, and Bruce B. Collette. Morphology of Luvarus imperialis (Luvaridae), with a Phylogenetic Analysis of the Acanthuroidei (Pisces). Smithsonian Contributions to Zoology, number 485,78 pages, 50 figures, 1989.—The osteology of the rarely collected epipelagic fish Luvarus imperialis Rafinesque, the louvar, is redescribed, based for the first time on cleared and stained specimens of juveniles and young adults, as well as on dried skeletons of larger adults. Knowledge of the structure of fossil luvarids, from their first occurrence in the upper Paleocene to a very well-preserved specimen from the lower Eocene, is reviewed and their status re-evaluated. Earlier comparative anatomical studies are reviewed in historical perspective with regard to previously proposed relationships. It has been postulated most frequently that Luvarus is related to carangoid or scombroid fishes, the latter hypothesis having been the most widely accepted. With cladistic methodology and with much additional detailed information on the osteology and larval morphology of Luvarus and other groups, the carangoid and scombroid hypotheses are refuted. Luvarus is actually a highly specialized pelagic member of the typically reef-associated Acanthuroidei, the surgeon fishes and their allies. Scatophagids and ephippidids, respectively, are the first and second outgroups for the Acanthuroidei. Anatomical diagnoses of all acanthuroid genera are given. A cladistic analysis of the intra-relationships of the Acanthuroidei, based on 90 characters encompassing both larval and adult morphology, provides a highly corroborated hypothesis indicating the following phyletic sequence: Siganidae—Luvaridae—Zanclidac—Acanthuridae. 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 Morphology of Luvarus imperialis (Luvaridae), with a phylogenetic analysis of the Acanthuroidea (Pisces) / James C. lyicr ... [ct al.]. (Smithsonian contributions to zoology ; no. 485) Bibliography: p. 1. Louvar—Classification. 2. Louvar—Morphology. I. Tyler, James C, 1935- . II. Series. QL1.S54 no. 485 [QL638.L95] 591 s-dc20 [597'.58] 89-600070 Contents Page Introduction 1 Brief History of This Study 3 Methods 4 Material Examined 5 Acknowledgments 5 Anatomical Descriptions 6 79.2 mm SL 6 Skull 6 Gill Arches 14 Pectoral Girdle 17 Pelvic Girdle 18 Vertebral Column 19 Caudal Skeleton 20 Dorsal and Anal Fins 21 Scales 22 Olfactory Rosette 22 301 mm SL 22 485 mm SL 27 980 mm SL 27 1100 mm SL 29 1140 mm SL 29 1720 mm SL 29 1780 mm SL 31 Relationships 31 Fossil Record 31 Parasites 32 Diagnosis of the Family Luvaridae 32 Previously Proposed Relationships 34 Acanthuroid Characters 37 Character Analysis 42 Summary of Character Analysis 73 Evolution and Adaptations 73 Note 75 Literature Cited 76 Dedication This contribution is dedicated to the memory of Robert H. Gibbs, Jr., 30 July 1929 to 3 April 1988, our dear friend and esteemed colleague in the study of scombroid fishes. Morphology of Luvarus imperialis (Luvaridae), with a Phylogenetic Analysis of the Acanthuroidei (Pisces) James C. Tyler, G. David Johnson, Izumi Nakamura, and Bruce B. Collette Introduction Giinther (1866) provided a brief page-and-a-half description The louvar, Luvarus imperialis Rafinesque, 1810 (Figure 1), and a sketch of the lateral view of an entire dried skeleton of is an infrequently collected epipelagic fish of world-wide unspecified size in the Senckenberg Museum of Frankfurt. He distribution in tropical and temperate marine waters. The was impressed by "the feeble development of the whole monotypic Luvarus undergoes a remarkable and prolonged osseous structure" and inferred that Luvarus "is a deep-sea metamorphosis, various stages of which have received larval fish" from "a zone at a depth of perhaps a hundred fathoms." appellations (Hystricinella, Astrodermella, Luvarella, Figure He commented on the small mouth, curved haemal spines and 2), as described in detail by Roule (1924) and Roule and Angel elongate anal fin basal pterygiophores.the presence of 7 ribs on (1930). The most recent descriptions of larval Luvarus are by the 3rd to 9th vertebrae, the total of 22 vertebrae, and dorsal fin Leis and Richards (1984), Johnson and Washington (1987), pterygiophores "so much dilated above that their upper and Nishikawa (1987). Juveniles begin to resemble the adult by extremities appear to be united by one semiossified ligament, about 100 to 200 mm SL. Adults reach a size of over 1800 mm which extends from the parietal crest to the end of the dorsal." SL and 140 kg (for records see Bolin, 1940; Whitley, 1940; He noted that the pterygiophores of the anal fin are similar to Gotshall and Fitch, 1968). those of the dorsal fin, their united extremities "extending from Like many other strange and exotic epipelagic species such the end of the anal to the pubic bones, and forming a complete as the oar fish Regalecus and the giant ocean sunfish Mola, but feeble ring round and supporting the abdominal cavity." Luvarus has excited the imagination and moved the pen so Gunther's specimen apparently was at least a large juvenile or frequently that it has spawned a subset of ichthyological young adult, for the pelvic fin was proportionally small, being literature devoted to records of its occurrence and to its external "very short and coalesced, but slightly divergent behind, so as anatomy. Most of the early work was summarized by Roule to leave a narrow opening for the vent, which can be entirely (1924), Roule and Angel (1930), and Gregory and Conrad closed by the rudimentary.. .ventral fins." (1943). Recent regional papers include Gotshall and Fitch Giinther (1866) did not consider the classification or possible (1968) for the eastern Pacific, Buen (1957) for the north and relationships of Luvarus, but in his "Catalogue" (1860) he south western Atlantic, Paulin et al. (1982) for New Zealand, placed Luvarus (as Ausonia), without significant explanation, and Blache (1964) and Ibafiez (1981) for the eastern Atlantic. in the Coryphaenina, as one of five subdivisions of his James C. Tyler, Deputy Director, National Museum of Natural Scombridae. Shortly thereafter, Gill (1863), probably follow- History, Smithsonian Institution, Washington, D.C. 20560. G. David ing Gunther (1860), gave an equally vague allocation of Johnson, Division of Fishes, National Museum of Natural History, Luvarus as an aside to his brief classification of the Smithsonian Institution, Washington, D.C. 20560. Izumi Nakamura, Fisheries Research Station, Kyoto University, Maizuru, Kyoto 625, Scombroidae, stating that Luvarus, Diana, and Lampris Japan. Bruce B. Collette, Systematics Laboratory, National Marine represent distinct families related to Coryphaena, the latter of Fisheries Service, National Museum
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