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Teleostei: Siluriformes) Monophyly and Phylogenetic Diagnosis of the Family Cetopsidae, with Synonymization of the Helogenidae (Teleostei: Siluriformes) MARIO C.C. DE PINNA and RICHARD P. VARI W9\ I SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 571 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 ZOOLOGY • NUMBER 571 Monophyly and Phylogenetic Diagnosis of the Family Cetopsidae, with Synonymization of the Helogenidae (Teleostei: Siluriformes) Mario C.C. de Pinna and Richard P. Vari SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1995 ABSTRACT de Pinna, Mdrio C.C., and Richard P. Vari. Monophyly and Phylogenetic Diagnosis of the Family Cetopsidae, with Synonymization of the Helogenidae (Teleostei: Siluriformes). Smithsonian Contributions to Zoology, number 571, 26 pages, 20 figures, 1995.—The Neotropical catfishes currently recognized as the families Cetopsidae and Helogenidae are hypothesized to be sister groups. The two taxa share numerous synapomorphies, including a lap joint between the opercle and interopercle, a dorsal displacement of the attachment of the interoperculo-mandibular ligament on the interopercle, and a strongly depressed palatine. Some of these derived characters are unique within Siluriformes. On the basis of this hypothesis of relationships, cetopsids and helogenids are united into an expanded Cetopsidae, itself subdivided into the monophyletic subfamilies Cetopsinae and Helogeninae. The Cetopsinae is synapomor- phically diagnosable by, among other characters, an expanded and often ramified cartilaginous posterior process of the basipterygium, an anterior curvature of the third epibranchial, and the articulation of the palatine with the neurocranium only by means of its anterior cartilage. Synapomorphies supporting the monophyly of the Helogeninae include a reduction in the degree of ossification between the mesethmoid cornua and the remainder of the neurocranium, a large paired fontanel in frontal and sphenotic, the posterior expansion of the third basibranchial, and the lack of contact of the sphenotic with the supraoccipital. Phylogenetic diagnoses are provided for the redefined Cetopsidae, Cetopsinae, and Helogeninae. The external morphology of juvenile Helogenes and a number of unusual juvenile features, possibly unique to the genus, are described for the first time. 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 Pinna, Mario C.C. de. Monophyly and phylogenetic diagnosis of the Family Cetopsidae, with synonymization of the Helogenidae (Teleostei: Siluriformes) / Mario C.C. de Pinna and Richard P. Vari. p. cm. — (Smithsonian contributions to zoology ; no. 571) Includes bibliographic references. I. Cetopsidae—Classification. 2. Cetopsidae—Phylogeny. 3. Helogeneidae—Classification. I. Van, Richard P. II. Series. QL1.S54 no. 571 [QL638.C37) 591 s-dc20 [597'.52] 94-48553 ® 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 Acknowledgments 2 Material and Methods 3 Institutional Abbreviations 3 Anatomical Abbreviations 4 Character Evidence 4 Cetopsidae Clade 4 Cetopsinae Clade 7 Helogeninae Clade 13 Discussion 15 New Classificatory Scheme 17 Family CETOPSIDAE Bleeker, 1858, new usage 18 Subfamily CETOPSINAE Bleeker, 1858 18 Subfamily HELOGENINAE Regan, 1911, new rank 19 Comparisons with Previous Hypotheses and Classifications 19 Description of the Juvenile of Helogenes 20 Resumo 23 Literature Cited 24 111 Monophyly and Phylogenetic Diagnosis of the Family Cetopsidae, with Synonymization of the Helogenidae (Teleostei: Siluriformes) Mario C.C. de Pinna and Richard P. Vari Introduction known about helogenid biology indicates they are free-living generalized predators of allochthonous terrestrial insects (Vari The families Helogenidae and Cetopsidae (sensu Greenwood and Ortega, 1986:15) and other small invertebrates. The et al., 1966) are two small groups of Neotropical freshwater species- and generic-level taxonomy of helogenids recently catfishes, which are comprised of four and approximately 15 was revised by Vari and Ortega (1986). currently recognized species, respectively. The phylogenetic relationships of both families to other siluriforms always have The Cetopsidae, with about five nominal genera, also form a been obscure, in part due to the poor overall knowledge of rather distinctive group of catfishes (Figure 2). Cetopsids phylogenetic relationships among familial-level catfish taxa. display far greater intrafamilial morphological variation than Although recognized as distinct entities for over eight decades, does the Helogenidae and a significantly greater range of neither the Cetopsidae nor the Helogenidae have been maximum size among the included species (30 to 300 mm SL). demonstrated to be monophyletic on the basis of synapomor- The limited available information on cetopsid biology indicates phic characters, a situation they share, however, with the that they range from free-living insect eaters (Saul, 1975; majority of catfish families. Baskin et al., 1980) to semi-predatory scavengers and The Helogenidae, with a single valid genus Helogenes flesh-eaters (sometimes erroneously termed "parasitic") known Gunther, is a group of distinctive small catfishes (maximum in the Brazilian Amazon as "candiru ac,u." The flesh-eating size 73 mm SL) with rather uniform morphology and cetopsids, notorious for their voracious habits (Magalhaes, pigmentation (Figure 1). Helogenids inhabit small, clear and 1931; Goulding, 1980:193-194), are often inappropriately blackwater streams of the Guyanas and the Amazon and lumped with some semi-parasitic trichomycterids under the Orinoco basins, being most often captured in shallow-water common name of "candiru." Cetopsids have a wider distribu- microhabitats of blackwater rain forest streams. The little tion than helogenids, occurring in the Amazon, Essequibo, Orinoco, Sao Francisco, and Parana-Paraguay basins, the Mdrio C.C. de Pinna, Departamento de Zoologia, Instituto de rivers of the Pacific slope of Ecuador and Colombia, and Biociencias, Universidade de Sao Paulo, Caixa Postal 11294, Sao drainages of the Caribbean versant of Colombia and northwest- Paulo, SP 05422-970, Brazil. Richard P. Vari, Department of Vertebrate Zoology, National Museum of Natural History, Smith- ern Venezuela. Their known habitats range from small rain sonian Institution, Washington, D.C. 20560. forest creeks, similar to those inhabited by helogenids, to large Review Chairman: Stanley H. Weitzman, National Museum of Natural rivers. In the absence of a thorough revision of cetopsids, the History, Smithsonian Institution. taxonomy of the group remains unsettled both at the species Reviewers: Scott A. Schaefer, Department of Ichthyology, Academy of and generic levels (Ferraris and Brown, 1991). Although two Natural Sciences of Philadelphia; and Carl. J. Ferraris, Jr., Department of Ichthyology, California Academy of Sciences. revisionary studies of cetopsids have been completed in recent SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY FIGURE 1.—Representative example of the Cetopsidae, subfamily Helogeninae, Helogenes marmoratus, USNM 273057, Guyana, Potaro River. FIGURE 2.—Representative example of the Cetopsidae, subfamily Cetopsinae, Pseudocetopsis amphiloxa, USNM 305348, Colombia, Choco, Rfo Truando system. years (de Oliveira, 1988; Milani de Amal, 1991), neither has morphology prompted us to provide a brief description of the been published. external anatomy
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