SECOND Parasitic EDITION Diseases of Wild Mammals
Parasitic SECOND EDITION Diseases of Wild Mammals Edited by William M. Samuel Margo J. Pybus A. Alan Kocan Iowa State University Press / Ames
William M. Samuel, Ph.D., is a professor in the Department of Biological Sciences at the University of Alberta. He is also the coordinator for the Challenge Grants in Biodiversity Program, which provides research funds to graduate students studying conservation biology and ecology. Margo J. Pybus, Ph.D., is a wildlife disease specialist with the Alberta Natural Resources Service Fisheries and Wildlife. She has over 20 years of experience in field and experimental investigation of disease and parasites in free-ranging wildlife. A. Alan Kocan, Ph.D., is a professor of pathobiology in the College of Veterinary Medicine at Oklahoma State University. His major research interests are parasitic and infectious diseases of wild and domestic animals, with recent emphasis on tick-transmitted diseases. 2001, 1971 Iowa State University Press All rights reserved Iowa Sate University Press 2121 South State Avenue, Ames, Iowa 50014 Orders: 1-800-862-6657 Office: 1-515-292-0140 Fax: 1-515-292-3348 Web site: www.isupress.com Authorization to photocopy items for internal or personal use, or the internal or personal use of specific clients, is granted by Iowa State University Press, provided that the base fee of $.10 per copy is paid directly to the Copyright Clearance Center, 222 Rosewood Drive, Danvers, MA 01923. For those organizations that have been granted a photocopy license by CCC, a separate system of payments has been arranged. The fee code for users of the Transactional Reporting Service is 0-8138-2978-X/2001 $.10. Printed on acid-free paper in the United States of America First edition, 1971 Second edition, 2001 Library of Congress Cataloging-in-Publication Data Parasitic diseases of wild mammals / edited by William M. Samuel, Margo J. Pybus, and A. Alan Kocan. 2nd ed. p. cm. Rev. ed. of: Parasitic diseases of wild mammals / John William Davis. 1st ed. 1971. Includes bibliographical references and index (p. ). ISBN 0-8138-2978-X (alk. paper) 1. Wildlife diseases. 2. Mammals Parasites. 3. Veterinary parasitology. I. Samuel, William M. II. Pybus, Margo J. III. Kocan, A. Alan. IV. Davis, John William. Parasitic diseases of wild mammals. SF996.4 P37 2001 639.9'64 dc21 00-047257 The last digit is the print number: 9 8 7 6 5 4 3 2 1
C O N T E N T S Preface............................... PART ONE: ECTOPARASITES........ 3 1. Lice (Phthiraptera) Lance A. Durden... 3 2. Biting Flies (Class Insecta: Order Diptera) Sandra A. Allan................... 18 3. Bot Flies and Warble Flies (Order Diptera: Family Oestridae) Douglas D. Colwell.. 46 4. Ticks (Class Arachnida: Order Acarina) Sandra A. Allan................... 72 5. Sarcoptes scabiei and Sarcoptic Mange Set Bornstein, Torsten Mörner, and William M. Samuel................. 107 PART TWO: ENDOPARASITES........ 121 6. Liver Flukes Margo J. Pybus......... 121 7. Taeniasis and Echinococcosis Arlene Jones and Margo J. Pybus................ 150 8. Gastrointestinal Strongyles in Wild Ruminants Eric P. Hoberg, A. Alan Kocan, and Lora G. Rickard................ 193 9. Extrapulmonary Lungworms of Cervids Murray W. Lankester................ 228 10. Lungworms of Marine Mammals Lena N. Measures........................ 279 11. Baylisascaris procyonis and Related Species Kevin R. Kazacos.................. 301 12. Filarioid Nematodes Roy C. Anderson.. 342 13. Dioctophymatosis Lena N. Measures... 357 14. Hepatic Capillariasis David M. Spratt and Grant R. Singleton................. 365 vii 15. Trichinella spp. and Trichinellosis Terry A. Dick and Edoardo Pozio............. 380 PART THREE: PROTOZOANS......... 397 16. Enteric Protozoans Amebiasis, Balantidiasis, and Enteric Trichomoniasis A. Alan Kocan........ 397 Giardia and Giardiasis Merle E. Olson and Andre G. Buret................. 399 Cyclospora, Eimeria, Isospora, and Cryptosporidium spp. Donald W. Duszynski and Steve J. Upton................. 416 17. Tissue-inhabiting Protozoans Opportunistic Amoebae David T. John.. 460 Hepatozoon spp. and Hepatozoonosis Thomas M. Craig.................. 462 Besnoitia spp. and Besnoitiosis Frederick A. Leighton and Alvin A. Gajadhar....... 468 Toxoplasmosis and Related Infections Jitender P. Dubey and Klaus Odening... 478 18. Blood-inhabiting Protozoans Blood-inhabiting Protozoan Parasites A. Alan Kocan..................... 520 Piroplasms (Theileria spp., Cytauxzoon spp., and Babesia spp.) A. Alan Kocan and Kenneth A. Waldrup................ 524 Contributors........................... 537 Index............................... 539 v
P R E F A C E When the first edition of Parasitic Diseases of Wild Mammals was published in 1971, the field of wildlife parasitology was, for the most part, in the stage of documenting the presence and identity of parasites in mammalian hosts. Few had challenged the idea that parasites were so well adapted to their hosts that they seldom caused disease (but see coverage of lungworms and elaeophorosis in the first edition). Nonetheless, Parasitic Diseases of Wild Mammals served many years as a reference book for parasitologists, veterinarians, professors, senior students, wildlife biologists, public health workers, and others. With chapters arranged in similar and uniform format, the first edition provided current information on 17 selected parasites, or related groups of parasites, of wild mammals. The field of wildlife parasitology, particularly as it relates to mammals, has changed dramatically since publication of the first edition, moving well beyond the case report and parasite survey stage of development. Much information has accrued on the ecology and epidemiology/epizootiology of parasites of wild mammals, as well as the significance of parasites in conservation biology. This information explosion has clearly illuminated the complexity of the host-parasite relationship for those few parasites of wild mammals studied in detail and has revealed that some parasites are able to cause severe disease and death of hosts in certain situations. Reasons for this information explosion are many and include establishment of 1. strong research programs on specific parasites with disease-causing propensities, 2. teaching/research curricula at the university level, 3. data-gathering and disseminating facilities around the world (for example, Canadian Cooperative Wildlife Health Centre, U.S. National Wildlife Health Center, Southeastern Cooperative Wildlife Disease Study) and disease monitoring programs in Scandinavia and France that are specific to wildlife disease, and 4. growth and maturation of the Wildlife Disease Association and its major publication, the Journal of Wildlife Diseases. In summary, today it is commonplace to use integrated approaches to better understand host-parasite interactions and the inherent role of parasitic organisms within natural ecosystems. This book updates information on several important parasites or parasitic groups covered in the first edition and adds information on groups not covered previously. It will become immediately obvious that the second edition is long overdue. As one brief example, Trichinella is no longer considered a monospecific zoonotic organism, but rather comprises a suite of different isolates, ecotypes, subspecies, or species infecting mammals worldwide. Virtually all aspects of the epidemiology of trichinosis have changed in recent years. The second edition of Parasitic Diseases of Wild Mammals will serve as a benchmark reference for parasites or parasitic groups 1. with members that are significant pathogens of mammals (for example, see Chapter 9 on lungworms and Chapter 5 on sarcoptic mange), 2. that normally do not have pathogenic tendencies, or are difficult to assess, but nonetheless have potential to cause problems (examples include Chapter 8 on gut nematodes of ungulates, Chapter 7 on taeniid tapeworms, and Chapter 14 on hepatic capillariasis), 3. with zoonotic importance (examples include Chapter 17 on giardiasis, Chapter 15 on trichinellosis, and Chapter 11 on Baylisascaris), 4. that we might temptingly refer to as emerging diseases (examples include crytosporidians in Chapter 18 and new forms in Chapter 22, on toxoplasmosis), and 5. that are conspicuous organisms that evoke management concerns or public interest (examples include Chapter 4 on ticks, and Chapter 6 on liver flukes). This volume presents up-to-date summaries of five groups of ectoparasites (lice, biting flies, bot flies, ticks, and mange mites), ten groups of helminths or endoparasites (liver flukes, taeniid tapeworms, gastrointestinal nematodes of ruminants, lungworms of terrestrial and marine mammals, baylisascarids, filarioid nematodes, kidney worms, hepatic nematodes, and Trichinella), and nine groups of protozoans including enteric forms (Amoeba, Giardia, coccidia), tissueinvaders (amoebae, hepatozoons, Besnoitia, and Toxoplasma and relatives), and blood-inhabiting forms (trypanosomes and relatives, piroplasms). The format is, for the most part, uniform, although in some cases authors had freedom to arrange the presentation to best present the information. vii
viii / Preface It is hoped that the contents here will be worthwhile for both novices and experts in wildlife parasitology and also for those with other expertise in need of a reference on parasitic diseases of wild mammals. The editors acknowledge the Wildlife Disease Association, the membership of which initiated the idea for this volume. We also acknowledge the many members who contributed to this volume. Royalties that will accrue from sales go to the Wildlife Disease Association.