Canadian Special Publication of Fisheries and Aquatic Sciences 124 Guide to the Parasites of Fishes of Canada Edited by L Margolis and Z Kabata 11111111illyellfill Part IV Trematoda David L Gibson m Department ori Fisheries & Orean's Library rAu°Anur 22 1996 Ministere cles Perches et Oceans des OTTAWA c 3 1 ( LF cJ GUIDE TO THE PARASITES OF FISHES OF CANADA PART IV NRC Monograph Publishing Program R.H Haynes, OC, FRSC (York University): Editor, Monograph Publishing Program Editorial Board: W.G.E. Caldwell, FRSC (University of Western Ontario); P.B. Cavers (University of Western Ontario); G. Herzberg, CC, FRS, FRSC (NRC, Steacie Institute of Molecular Sciences); K.U. IngoId, OC, FRS, FRSC, (NRC, Steacie Institute of Molecular Sciences); M. Lecours (Université Laval); L.P. Milligan, FRSC (University of Guelph); G.G.E. Scudder, FRSC (University of British Columbia); E.W. Taylor, FRS (University of Chicago); B.P. Dan- cik, Editor-in-Chief, NRC Research Journals and Monographs (University of Alberta) Publishing Office: M. Montgomery, Director General, CISTI; A. Holmes, Director, Publishing Directorate; G.J. Neville, Head, Monograph Publishing Program; E.M. Kidd, Publication Officer. Publication Proposals: Proposals for the NRC Monograph Publishing Program should be sent to Gerald J. Neville, Head, Monograph Publishing Program, National Research Council of Canada, NRC Research Press, 1200 Montreal Road, Building M-55, Ottawa, ON K 1 A 0R6, Canada. Telephone: (613) 993-1513; fax: (613) 952-7656; e-mail: gerry.nevi lie@ nrc.ca . © National Research Council of Canada 1996 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the National Research Council of Canada, Ottawa, Ontario KlA 0R6, Canada. ISBN 0-660-16403-5 ISSN 0706-6481 NRCC No. 40083 Canadian Cataloging In Publication Data Main entry under title: Guide to the parasites of fishes of Canada (Canadian special publication of fisheries and aquatic sciences; no. 124) Partial contents: V. 4 Trematoda / David I. Gibson. Includes an abstract in French. Includes bibliographical references and an index. Issued by the National Research Council of Canada. ISBN 0-660-16403-5 1. Fishes — Parasites — Canada — Identification. I. Kabata, Z. 1924–. II. Margolis, L. 1927–. III. Gibson, David I. Trematoda IV. Canada. Dept. of Fisheries and Oceans. V. National Research Council Canada VI. Title. VII. Series. SH175.G81 1996 597'.0233'0971 C85-075080-6 rev. This publication is available from: Subscription Office, NRC Research Press, National Research Council of Canada, Ottawa, Ontario KlA 0R6, Canada. Enquiries: Telephone, 613-993-9084; fax, 613-952-7656 Correct citation for this publication: Gibson, D.I. 1996. Trematoda. In L. Margolis and Z. Kabata (ed). Guide to the parasites of fishes of Canada. Part 1V. Can. Spec. Publ. Fish Aquat. Sci. 124. 373 pp. Canadian Special Publication of Fisheries and Aquatic Sciences 124 Guide to the Parasites of Fishes of Canada Edited by L. Margolis and Z. Kabata Department of Fisheries and Oceans Pacific Biological Station Nanaimo, British Columbia V9R 5K6 Part IV Trematoda David I. Gibson Department of Zoology The Natural History Museum Cromwell Road London SW7 5BD United Kingdom ARC-CARC NRC RESEARCH PRESS Ottawa 1996 CONTENTS Guide to the Parasites of Fishes of Canada Part IV Preface vii Abstract/Résumé ix Introduction 1 TREMATODA 5 Key to the subclasses 5 ASPIDOGASTREA 5 Key to the orders 7 ASPIDOGASTRIDA 7 Aspidogastridae 7 STICHOCOTYLIDA 8 Rugogastridae 8 DIGENEA 10 Key to the superfamilies 10 STRIGEIDA Clinostomoidea 13 Diplostomoidea 16 Gymnophalloidea 38 Hemiuroidea 61 Schistosomatoidea 124 ECHINOSTOMIDA Echinostomatoidea 133 Paramphistomoidea 137 PLAGIORCHIIDA Allocreadioidea 139 Lepocreadioidea 191 Opisthorchioidea 222 Plagiorchioidea 247 Zoogonoidea 270 Host-parasite list 295 Acknowledgments 338 References 339 Index to parasites 359 Index to hosts 368 PREFACE In Part IV of this series, Guide to the Parasites of Fishes of Canada, the Trematoda are treated by a renowned helminthologist, Dr. David I. Gibson, from the Natural History Museum, London. He has studied these parasites of fishes from the "four corners" of the globe, including the Pacific and Atlantic coasts of Canada. We are indeed fortunate, as will be subsequent users of the Guide, that Dr. Gibson agreed to undertake the task of authoring this publication. The three previous parts* in this series dealt with the Monogenea, Turbellaria, Crustacea, Acanthocephala, and Cnidaria. Part IV provides practical keys for identification of the taxa of Trematoda represented in Canadian fishes down to the species level, accompanied by detailed line drawings. Although maintaining the general format of Parts I—III, in keeping with our policy of permitting the authors some degree of latitude in how they approached their task, Part IV has three departures from the general plan. Firstly, it discusses major taxonomic problems, when required, for genera and higher taxa on a global basis. The large number of species included in this work dictated that certain measures be taken to keep the volume within a manageable size. This led to the second and third departures from the basic plan: figures are provided for only one species in a genus, rather than all species, and full descriptions are not provided for individual species. These omissions are compensated for by detailed generic descriptions and inclusion of important distinguishing characters in the keys to species. The first three parts in this series were published by the Canadian Department of Fisheries and Oceans. Publication responsibilities have now been assumed by the National Research Council of Canada to whom we express our appreciation for continuing this series within its monograph publication program. Financial assistance for publication was provided by the Pacific Biological Station, Department of Fish- eries and Oceans. The Editors * Part I. Beverley-Burton, M. 1984. Monogenea and Turbellaria. Part II. Kabata, Z., F. Rafi, and E.L.Bousfield. 1988. Crustacea. Part III. Arai, H. P. 1989. Acanthocephala; and Arai, M. N. 1989. Cnidaria. vii ABSTRACT GIBSON, D.I. 1996. Trematoda. In: L. Margolis and Z. Kabata (ed.) Guide to the parasites of fishes of Canada. Part IV. Can. Spec. Publ. Fish. Aquat. Sci. 124: 373 pp. More than 80 keys leading to the identification of more than 220 species of trematode parasites of Cana- dian fishes are provided. Diagnoses of more than 80 family-group taxa and 130 genera are given. Each genus is illustrated. For each species of trematode a Canadian host list, locality data, and any relevant remarks are included. In addition, a host-parasite list is given plus parasite and host indices. RÉSUMÉ GIBSON, D.I. 1996. Trematoda. Dans L. Margolis et Z. Kabata (éd.) Guide to the parasites of fishes of Canada. Part IV. Can. Spec. Publ. Fish Aquat. Sci. 124: 373 pp. On y retrouve plus de 80 clefs qui permettent d'identifier quelque 220 espèces de trématodes parasites des poissons du Canada. Le livre comprend la description de plus de 80 taxums de famille/groupe et de 130 genres. Chaque genre y est illustré. À chaque espèce de trématode est associée une fiche signalétique, con- tenant des données sur l'emplacement et des remarques particulières à l'espèce. De plus, on y retrouve une liste des paires hôte-parasite ainsi qu'un index des hôtes et des parasites. ix INTRODUCTION The first record of trematode parasites in Canadian Synotzym(s). These are other names that have been fishes appears to be that of Wright (1879), who used for the species in the past: they may be totally recorded the presence of metacercariae of different names that have been synonymised with Distomum gracile (Leidy), which is probably the present species or the same epithet with a dif- Clinostomum complanatum (Rud.), from perch ferent combination of generic name. These syn- Perca flavescens probably in Ontario. The first spe- onymy lists are usually not complete, tending to cialist who carried out detailed work on the trema- include only those cited in publications that relate tode fauna of fishes was, however, Stafford, who to the Canadian fauna. between 1900 and 1907, after training in Germany, produced a series of papers on the trematodes of Site. This is the position or organ in or on the fish marine and freshwater fishes of eastern Canadian where the parasite has been reported in Canadian waters. Subsequently, although numerous authors fishes. In some cases, where this information is not added information, it was not until the second half given, its expected site is given in brackets. of the 20th century that a knowledge of this fauna has been built up. Great advances were made, espe- Host(s). These are cited using currently accepted cially during the 1960s, 1970s, and early 1980s, nomenclature. The numbers in parentheses follow- after which a noticeable decline in faunistic studies ing the host name are the numbers assigned to the became apparent. Although parts of the Canadian relevant record, i.e., the author(s) responsible for fauna are now relatively well known, particularly in recording the parasite from the particular host. relation to freshwater and commercial marine Question marks associated with these numbers fishes, other areas, and particularly in the deep-sea indicate that the record is questionable. and Arctic regions, are less so. Distribution. The distribution of the records of the The arrangement of the text follows largely that of various species of parasite in Canadian waters is other contributions in the series. The classification indicated by abbreviations. Marine records are (see below) is based at the superfamily level, with listed under three geographical regions: Atlantic definitions of the various taxa down to the generic (At1), Pacific (Pac), and eastern Arctic (EArc). level. Keys, usually dichotomous, to the superfami- Freshwater records are indicated in relation to the lies, families, subfamilies (where included), genera, various Provinces or Territories of Canada: Alberta and species are included.