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Detritivores of the South American Fish Family Prochilodontidae (Teleostei: Ostariophysi: Characiformes): a Phylogenetic and Revisionary Study

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Detritivores of the South American Fish Family Prochilodontidae (Teleostei: Ostariophysi: Characiformes): a Phylogenetic and Revisionary Study Detritivores of the South American Fish Family Prochilodontidae (Teleostei: Ostariophysi: Characiformes): A Phylogenetic and Revisionary Study RICARDO M.C. CASTRO and RICHARD P. VARI W9\ I SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 622 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. Lawrence M. Small Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 622 Detritivores of the South American Fish Family Prochilodontidae (Teleostei: Ostariophysi: Characiformes) A Phylogenetic and Revisionary Study Ricardo M. C. Castro and Richard P. Vari Smithsonian Contributions and Studies Series AN IMPRINT OF SMITHSONIAN BOOKS Washington, D.C. 2004 ABSTRACT Castro, Ricardo M.C., and Richard P. Vari. Detritivores of the South American Fish Family Prochilodontidae (Teleostei: Ostariophysi: Characiformes): A Phylogenetic and Revisionary Study. Smithsonian Contributions to Zoology, number 622, 189 pages, 71 figures, 23 tables. 2004.—The investigation of the inter- and intrafamilial relationships of the Prochilodontidae was investigated based on information from osteological and soft anatomical systems. Synapo- morphies uncovered during this study provide additional evidence that (1) the clade formed by the Prochilodontidae plus Curimatidae is the sister group to the clade that consists of the Anos- tomidae plus Chilodontidae, (2) supports the hypothesis of the monophyly of the Prochilodon- tidae, Ichthyoelephas Posada Arango (1909), Prochilodus Agassiz (in Spix and Agassiz, 1829), and Semaprochilodus Fowler (1941), (3) demonstrates that Prochilodus is the sister group to the clade formed by Ichthyoelephas plus Semaprochilodus, and (4) partially resolves interspe- cific relationships. Recognized genera of the Prochilodontidae are Ichthyoelephas, Prochilodus, and Semapro- chilodus. Pacu Agassiz (in Spix and Agassiz, 1829) and Chilomyzon Fowler (1906) are syn- onyms ofProchilodus. Ichthyoelephas includes two species: /. humeralis (Giinther, 1859); and/, longirostris (Stein- dachner, 1879). Ichthyoelephas patalo Posada Arango (1909) and /. longirostris neglectus Dahl (1971) are synonyms of/, longirostris. Prochilodus consists of thirteen species: P. argenteus Agassiz (in Spix and Agassiz, 1829); P. brevis Steindachner (1874); P. britskii Castro (1993); P. costatus Valenciennes (in Cuvier and Valenciennes, 1850); P. hartii Steindachner (1874); P. lacustris Steindachner (1907); P. linea- tus (Valenciennes, 1836); P. magdalenae Steindachner (1879); P. mariae Eigenmann (1922); P. nigricans Agassiz (in Spix and Agassiz, 1829); P. reticulatus Valenciennes (1850); P. rubrotae- niatus Jardine (1841); and P. yimboides Kner (1859). Semaprochilodus contains six species: S. brama (Valenciennes, in Cuvier and Valenciennes, 1850); S. insignis (Jardine, 1841); S. kneri (Pellegrin, 1909); S. laticeps (Steindachner, 1879); S. taeniurus (Valenciennes, in Humboldt and Bonpland, 1817); and S. varii Castro (1988). Semaprochilodus squamilentus Fowler (1941) is a synonym of 5. brama. Prochilodus thera- ponura Fowler (1906) and P. amazonensis Fowler (1906) are synonyms of Semaprochilodus insignis. Keys are provided to the genera and species of prochilodontids. Neotypes are designated for Prochilodus argenteus and P. nigricans. Lectotypes are designated for Prochilodus affinis, P. asper, P. brevis, P. cearensis, P. hartii, P. humeralis, P. kneri, P. lacustris, P. longirostris, P. asper var. magdalenae, P. oligolepis, P. reticulatus, P. scrofa, and P. vimboides. The phylogenetic biogeography of the Prochilodontidae indicates that the family dates back minimally to approximately 12 million years ago, with higher level intrafamilial cladogenic events also dating to at least that time period; these dates are congruent with data from the fos- sil record for more encompassing groups within the Characiformes. Major shifts of overall bauplan characterize the Prochilodontidae and the major clades within the family, but there is relatively little interspecific diversity. The Prochilodus bauplan has apparently been static for at least 12 million years. 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 coral Montastrea cavernosa (Linnaeus). Library of Congress Cataloging-in-Publication Data Castro, Ricardo M.C. Detritivorcs of the South American fish family Prochilodontidae (Teleostei:Ostariophysi:Characiformes): a phy- logenetic and revisionary study / Ricardo M.C. Castro and Richard P. Vari. p. cm. — (Smithsonian contributions to zoology ; no. 622) Includes bibliographical references (p.). I. Prochilodontidae—Phylogeny. 2. Prochilodontidae—Classification. I. Vari, Richard P. II. Title. III. Series. QL1.S54 no. 622 [QL638.P786] 590s-dc22 [597'.48] 2003067250 © 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 and Material 5 Morphometrics 5 Meristics 5 Institutional Abbreviations 7 Osteological Preparations 7 Terminology 9 Acknowledgments 9 Character Descriptions and Analysis 10 Branchial Arches 10 Gill Filaments 12 Dorsal Portions of Branchial Arches 12 First Infrapharyngobranchial 12 First Epibranchial 13 First through Third Epibranchials 13 Second Epibranchial 14 Third Infrapharyngobranchial 14 Third Epibranchial 15 Fourth Upper Pharyngeal Tooth Plate 15 Fourth Upper Pharyngeal Tooth Plate and Fourth Infrapharyngobranchial .... 15 Fourth Epibranchial 15 Fifth Epibranchial 16 Fifth Upper Pharyngeal Tooth Plate 16 Epibranchial Organ 17 Ventral Portions of Branchial Arches 17 First Basibranchial 17 First through Fourth Ceratobranchials 17 Fourth Ceratobranchial 18 Fifth Ceratobranchial 18 Hyoid Apparatus 18 Basihyal 18 Dorsal Hypohyal 18 Anterior Ceratohyal 19 Interhyal 19 Branchiostegal Rays 19 Urohyal 20 Oral Dentition 20 Maxillary Dentition 20 Premaxillary and Dentary Dentition 20 Replacement Tooth Rows 23 Tooth Form 23 Lips 23 Upper Jaw 24 Soft Tissues 25 Bones 26 Premaxilla 26 Maxilla 28 iv SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Lower Jaw 30 Dentary 30 Coronomeckelian Bone 35 Angulo-articular 36 Retroarticular 37 Primordial Ligament 37 Infraorbital Series 37 First Infraorbital 37 Second Infraorbital 39 Third Infraorbital 40 Fourth Infraorbital 41 Fifth Infraorbital 41 Sixth Infraorbital 42 Supraorbital 42 Suspensorium 43 Palatine 43 Ectopterygoid 44 Quadrate 44 Mesopterygoid 45 Metapterygoid 45 Symplectic 46 Opercle 47 Preopercle 47 Interopercle 48 Suprapreopercle 48 Cranium 49 Mesethmoid 49 Vomer 49 Lateral Ethmoid 50 Orbitosphenoid 50 Parasphenoid 51 Frontal 52 Parietal 52 Sphenotic 53 Exoccipital 53 Myology 53 Pigmentation 54 Head 54 Body 54 Fins 55 Procumbent Dorsal-Fin Spine 55 Scales 56 Phylogenetic Reconstruction 58 MonophylyoftheProchilodontidae, Curimatidae, Anostomidae, and Chilodontidae Clade 58 Monophyly of the Prochilodontidae and Curimatidae Clade 59 Monophyly of the Curimatidae 60 Monophyly and Intrafamilial Phylogeny of the Prochilodontidae 60 Monophyly of the Prochilodontidae 60 Monophyly of Prochilodus 62 Interspecific Relationships within Prochilodus 63 Monophyly of the Semaprochilodus and Ichthyoelephas Clade 64 Monophyly of Semaprochilodus 64 Interspecific Relationships within Semaprochilodus 65 NUMBER 622 Monophyly of Ichthyoelephas 66 Homoplasies 67 Bauplan Shifts and Evolutionary Stasis 68 Family Prochilodontidae 70 Key to the Genera of the Prochilodontidae 71 Chave de Identificacao dos Generos de Prochilodontidae 71 Genus Ichthyoelephas Posada Arango, 1909 71 Key to the Species of Ichthyoelephas 72 Chave de Identificacao das Especies de Ichthyoelephas 72 Ichthyoelephas humeralis (Giinther, 1859) 73 Ichthyoelephas
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