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Acanthuroidei: Siganidae) •».«L"WHB' vn«74MV /ir, ^/j" -w irjur- Relationships of the Fossil and Recent Genera of Rabbitfishes (Acanthuroidei: Siganidae) R • - 5Vf^> ES C. TYLt and fDREF.BAN ->: m ^ 1 •"- . *6$B O PALEO * i 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 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. I. Michael Heyman Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY • NUMBER 84 Relationships of the Fossil and Recent Genera of Rabbitfishes (Acanthuroidei: Siganidae) James C. Tyler and Alexandre F. Bannikov SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1997 ABSTRACT Tyler, James C, and Alexandre F. Bannikov. Relationships of the Fossil and Recent Genera of Rabbitfishes (Acanthuroidei: Siganidae). Smithsonian Contributions to Paleobiology, number 84, 35 pages, 21 figures, 1 table, 1997.—Four genera of fossil siganid fishes of early Eocene to early Oligocene age are recognized in addition to the single Recent genus. The osteological features of these five genera are described and illustrated. A phylogenetic analysis utilizing PAUP indicates that the genera have the following phyletic sequence convention: Ruffoichthys Sorbini (two species from the middle Eocene of Italy)—Eosiganus, new genus (one new species from the middle Eocene of Russia)—Siganopygaeus Danilchenko (one species from the early Eocene of Turkmenistan)—Protosiganus Whitley (one species from the early Oligocene of Switzerland)—Siganus Forsskal (27 Recent species in the Indo-Pacific). OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Annals of the Smithsonian Institution. SERIES COVER DESIGN: The trilobite Phacops rana Green. Library of Congress Cataloging-in-Publication Data Relationships of the fossil and recent genera of rabbitfishes (Acanthuroidei: Siganidae) / James C. Tyler and Alexandre F. Bannikov. p. cm.—(Smithsonian contributions to paleobiology ; no. 84) Includes bibliographical references. 1. Siganidae, Fossil—Europe. 2. Paleontology—Eocene. 3. Paleontology—Oligocene. 4. Animals, Fossil— Europe. 5. Siganidae. I. Bannikov, A. F. (Aleksandr Fedorovich) II. Title. III. Series. QE701 .S56 no. 84 [QE852.P4] 560 s—dc21 [567.7] 97-21827 ® 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 1 Acknowledgments 2 Systematic Descriptions of the Genera of Siganidae 2 jRuffoichthys Sorbini, 1983 3 Ruffoichthys bannikovi Tyler and Sorbini, 1991, and Ruffoichthys spinosus Sorbini, 1983 3 fEosiganus, new genus 8 Eosiganus kumaensis, new species 8 fSiganopygaeus Danilchenko, 1968 13 Siganopygaeus rarus Danilchenko, 1968 13 tProtosiganus Whitley, 1935 16 Protosiganus glaronensis (Wettstein, 1886) 16 Siganus Forsskal, 1775 20 Siganus species 20 Analysis of Characters 25 Analytical Protocols 25 Characters Used in Phylogenetic Analysis 28 1. Number of Pelvic-Fin Spines 28 2. Procumbent Spine on First Pterygiophore of Spiny Dorsal Fin 29 3. Number of Anal-Fin Spines 29 4. Number of Supernumerary Dorsal-Fin Spines 29 5. Length of First Supernumerary Dorsal-Fin Spine 29 6. Association of Postcleithrum with First Anal-Fin Pterygiophore 30 7. Number of Dorsal-Fin Spines 30 8. Number of Dorsal-Fin Rays 30 9. Number of Anal-Fin Rays 30 10. Number of Supraneurals 31 11. Shape of Teeth 31 12. Ornamentation of Procumbent Spine 32 Other Characters 32 Vacant Interneural Space 32 Size of Uroneural 32 Subocular Shelf 32 Longest Dorsal-Fin Spine 33 Conclusion 33 Literature Cited 34 in Relationships of the Fossil and Recent Genera of Rabbitfishes (Acanthuroidei: Siganidae) James C. Tyler and Alexandre F. Bannikov Introduction five genera based on the 12 informative characters that can be determined in the Recent and most of the fossil taxa. Siganid rabbitfishes are one of the more prominent herbivo­ METHODS.—The methodology used for the phylogenetic rous components of the modern-day Indo-Pacific ichthyofauna. hypotheses is given under "Analytical Protocols" at the They are associated with coral reefs, the surrounding grass beginning of the "Analysis of Characters" section. flats, and other algae-rich environments, such as mangroves and rocky shores. There is a single extant genus (Siganus), but Abbreviations for the repositories of materials are as follows: it is exceptionally speciose (27 species that differ mostly in ANSP, Academy of Natural Sciences of Philadelphia; coloration and proportions). This genus is unique among MCSNM, Museo Civico di Storia Naturale di Milano; Recent teleosts in having the palatine divided into two separate MCSNV, Museo Civico di Storia Naturale di Verona; MCZ, ossifications (palatine and prepalatine) and in having each Museum of Comparative Zoology, Harvard University; NMB, pelvic fin with two spines (outer and inner, between which are Naturhistorisches Museum Basel; PIN, Paleontological Insti­ three rays). Along with the description of a new fossil species tute, Moscow; USNM, National Museum of Natural History, in the morphologically primitive Eocene genus Ruffoichthys Smithsonian Institution (collections of the former United States (palatine probably composed of two separate ossifications but National Museum). no inner pelvic-fin spine), Tyler and Sorbini (1991) compared Specimen length is always standard length (SL). Vacant that genus with the two other fossil genera then known (the interneural spaces are given the number of the preceding neural Eocene Siganopygaeus and Oligocene Protosiganus), but they spine (e.g., the first space is between the first and second did not analyze their phylogenetic relationships. We herein neurals, as in Baldwin and Johnson, 1993), and the terms describe a new genus of Eocene siganid (Eosiganus), rede- supraneural (as in Mabee, 1988) and epineural (as in Patterson scribe all of the other fossil taxa on the basis of our examination and Johnson, 1995) are used rather than predorsal bone and of all of the type materials, and utilize PAUP (Phylogenetic epipleural intermuscular bone; all three of these terms are Analysis Using Parsimony, Version 2.4.1, written by D.L. changes from those used in Tyler et al. (1989) for siganids and Swofford, then of the Illinois Natural History Survey, now of other acanthuroids. the Smithsonian Institution) to analyze the relationships of the For ease of comparison, descriptive features of the species are given in the same order, beginning with the few that are unique to either all or some siganids (palatine composed of two James C. Tyler, National Museum of Natural History, Smithsonian Institution (MRC-106), Washington, D.C. 20560. Alexandre F. separate ossifications, two spines in each pelvic fin, deep Bannikov, Paleontological Institute, Russian Academy of Sciences, notches on sides of teeth). Profsoyuznaya 123, 117647 Moscow, Russia. All five genera of siganids have 10 abdominal and 13 caudal Reviewers: Carole Baldwin, Department of Vertebrate Zoology, vertebrae, and these data are not repeated. Most other families National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Antony S. Harold, Department of Ichthyol­ of acanthuroids (luvarids, zanclids, acanthurids) have a more ogy and Herpetology, Royal Ontario Museum, Toronto, Ontario, specialized condition of one fewer abdominal vertebra, Canada M5S 2C6. Robert L. Shipp, Department of Marine Sciences, 9+13=22, but the fossil kushlukiids (sister group of luvarids) University of South Alabama, Mobile, Alabama 36688. have a secondarily increased number of vertebrae 1 On— 19— SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY 20=29-30. Scatophagids are like siganids in having 10+13=23, The relationships of Gazolaichthys as a basal member of the itself a specialized reduction of one caudal vertebra
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