The Prosobranch Snail Family Hydrobiidae (Gastropoda: Rissooidea): Review of Classification and Supraspecific Taxa
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The Prosobranch Snail Family Hydrobiidae (Gastropoda: Rissooidea): Review of Classification and Supraspecific Taxa ALANR KABAT and ROBERT HERSHLE SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 547 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 FoUdife 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 o^ 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 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 547 The Prosobranch Snail Family Hydrobiidae (Gastropoda: Rissooidea): Review of Classification and Supraspecific Taxa Alan R. Kabat and Robert Hershler SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1993 ABSTRACT Kabat, Alan R., and Robert Hershler. The Prosobranch Snail Family Hydrobiidae (Gastropoda: Rissooidea): Review of Classification and Supraspecific Taxa. Smithsonian Contributions to Zoology, number 547, 94 pages, 4 tables, 1993.—A nomenclatural analysis is provided for the 75 family level names and 725 generic-level names in the fresh- and brackish-water prosobranch gastropod family Hydrobiidae. For the 725 generic-level taxa, 405 are nomenclaturally available and in "current usage"; 51 are junior homonyms; 65 are commonly acknowledged junior synonyms; 24 are nomina nuda or otherwise invalid; and 180 are errors or emendations. The Hydrobiidae are redefined and differentiated from other rissooidean families, and a review of the classification of this family is presented. 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 Kabat, Alan R. The prosobranch snail family Hydrobiidae (Gastropoda: Rissooidea): review of classification and supraspecific taxa / Alan R. Kabat and Robert Hershler. p. cm.—(Smithsonian contributions to zoology ; no. 547) Includes bibliographical references (p.) and indexes. 1. Hydrobiidae—Classification. 2. Hydrobiidae—-Nomenclature. I. Hershler, Robert II. Title. III. Series. QU.S54no. 547 [QL430.5.H9] 591 s—dc20 93-11450 [594'.32] CIP ® 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 Review of Hydrobiid Classification 2 What is a Hydrobiid? 5 Diagnosis of the Family Hydrobiidae 5 Family-group Names 6 Materials and Methods 8 Acknowledgments 9 Genus-group Names 9 Literature Cited 61 Appendix I: Geographical Index 85 Appendix II: Stratigraphical Index 89 Index to Species 91 in The Prosobranch Snail Family Hydrobiidae (Gastropoda: Rissooidea): Review of Classification and Supraspecific Taxa Alan R. Kabat and Robert Hershler Introduction due to the high diversity and broad distribution of the group, The prosobranch family Hydrobiidae is a cosmopolitan which has not been revised at the genus level since the group of small aquatic1 snails found in permanent nonmarine compilation of Wenz (1939). Important nomenclatural prob- habitats (a few taxa occur in brackish coastal waters), ranging lems also remain unresolved, including the status of the family from small seeps to large rivers and inland "seas." These snails name Hydrobiidae Troschel, 1857, which is a junior homonym comprise a major biotic component of inland waters owing to of Hydrobiidae Mulsant, 1844 (Coleoptera): a submission will their ubiquity and diversity, which has been critically estimated have to be made to ICZN (International Commission on at over 100 genera (Taylor and Sohl, 1962; Ponder, 1988) and Zoological Nomenclature) to remove this homonymy (per about 1000 Recent species (Boss, 1971). We suspect that the ICZN, Article 55b; see Newton and Thayer, 1990). (ICZN, group is larger than this. The Hydrobiidae is one of the largest Opinion 1664 (1992), specified "that HYDROBIIDAE Tro- of the more than 20 families of small-sized, predominantly schel should remain in use, even though a junior homonym, in aquatic snails placed in the superfamily Rissooidea. Hydrobiids the absence of a Commission ruling resolving its homonymy.") have a long fossil record extending to the Early Carboniferous Additionally, progress in revising the family has been slow (Knight et al., 1960; Solem and Yochelson, 1979), although due to the nature of these snails and difficulties in evaluating Ponder (1988) recently suggested (without justification) that their phylogenetic relationships. Hydrobiids generally have Paleozoic shells attributed to the family instead belong to the smooth, nondescript shells, which offer few characters useful to Ampullariidae (Ampullarioidea).2 The higher systematics of the systematist other than general form, whose range of the Hydrobiidae is in a confused state as the validity, rank, and variation in the family matches that of the entire Rissooidea relationships of nominal supra-specific taxa remain suspect. (Ponder, 1988, fig. 1). Furthermore, evolutionary convergence Widely divergent classifications for the family have been in shell form amongst hydrobiid-like (or hydrobioid3) gastro- proposed in recent years, some of which were based on study of pods appears to have occurred frequently (Davis, 1979). The local (albeit diverse) subsets of the family seemingly without implications of this for evaluation of empty shells (fossil or consideration of extra-limital taxa, and none of which was based on phylogenetic analysis. (The exemplary treatment of 1 rissooidean phylogeny of Ponder (1988) does not extend below Falniowskia neglectissima apparently is terrestrial (Falniowski and the family level.) Monophyly of the family has not been clearly Steffek, 1989) and at least one species, Fonscochlea zeidleri, is amphibious (Ponder et al., 1989). established (see below) and the distinction between hydrobiids 2 and other rissooidean families (such as Pomatiopsidae; Davis et Ponder's discussion of the earliest appearance of the family focused on supposed late Paleozoic records from Africa and South America. Since then, an al., 1985) remains uncertain. This unsettled situation is partly additional record from the Upper Permian of China has been postulated (Yu and Zhu. 1990), strengthening the case for a Paleozoic origin for these snails. Alan R. Kabat and Robert Hershler, Department of Invertebrate 3 An informal, adjectival term for rissooidean snails resembling hydrobiids Zoology, NHB MRC 118, National Museum of Natural History, in general features of head/foot, operculum, genitalia, fecal pellets, and Smithsonian Institution, Washington, D.C. 20560. pigmentation (Davis, 1979:7). SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY otherwise) are obvious and systematic treatment of these (at trunk-like, foot with omniphoric groove, progression step-like, least at higher taxonomic levels) probably will remain largely ctenidium rudimentary-absent. Estuarine, freshwater, terres- conjectural barring a discovery of phylogenetically important trial, sometimes amphibious. (Abbott, 1958; Marcus and information resident in shell structure. Unfortunately, the Marcus, 1965.) proliferation of new higher taxa based on shells alone has not Barleeidae: Shell with inner chitinous layer, foot some- abated in recent years. times with posterior mucous gland, osphradium enlarged, The dimunitive size of hydrobiids (usually <8 mm standard oesophageal pouches sometimes present, penis sometimes with shell height) renders anatomical study of these animals prostatic tissue, oviduct glands with simple histology. Marine. difficult, and may explain why many Recent taxa still are being (Ponder, 1983.) described bereft of non-conchological details beyond penis, Bithyniidae: Shell often colored, operculum calcareous, operculum, and radula. Those who choose to dissect can be neck with nuchal lobe; pallial cavity with epitaenia, food frustrated, as hydrobiid anatomy is simple