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Infectious THIRD EDITION Diseases of Wild Mammals Infectious THIRD EDITION Diseases of Wild Mammals Edited by Elizabeth S. Williams and Ian K. Barker Iowa State University Press / Ames Elizabeth S. Williams,DVM,PhD,is a professor of veterinary sciences at the University of Wyoming. She has been specializing in the research of infectious diseases of wild animals since 1977. She also serves as a diagnos- tic pathologist. Ian K. Barker, DVM, PhD, is a professor of veterinary pathobiology in the Department of Pathology in the Ontario Veterinary College at the University of Guelph. His research focuses on infectious diseases of wild ani- mals,and he is an expert in alimentary tract diseases. ©2001,1981,1970 Iowa State University Press All rights reserved Iowa State 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 Copy- right 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 0-8138-2556-3/2001 $.10. Printed on acid-free paper in the United States of America First edition,1970 Second edition,1981 Third edition,2001 International Standard Book Number:0-8138-2556-3 Library of Congress Cataloging-in-Publication Data Infectious diseases of wild mammals / edited by Elizabeth S. Williams and Ian K. Barker.—3rd ed. p. cm. Includes bibliographical references (p. ). ISBN 0-8138-2556-3 (alk. paper) 1. Wildlife diseases. 2. Mammals—Diseases. 3. Communicable diseases in animals. I. Williams,Elizabeth S. II. Barker,Ian K. SF996.4 .I54 2000 639.9'64—dc21 00-047258 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 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii Caprine Herpesvirus Monika Engels . . . 173 Other Herpesviruses Anthony E. Castro 175 Part 1. Viral and Prion Diseases . . . . . . . . . 3 8. Poxvirus Infections Anthony J. Robinson 1. Rabies Charles E. Rupprecht,Klaus Stöhr, and Peter J. Kerr . . . . . . . . . . . . . . . . . . 179 and Courtney Meredith . . . . . . . . . . . . . . 3 9. Adenoviral Diseases Leslie W. Woods . . 202 2. Morbilliviral Diseases 10. Retrovirus Infections Michael Worley . . 213 Rinderpest Paul Rossiter . . . . . . . . . . . 37 11. Papillomavirus Infections John Peste des Petits Ruminants P. Sundberg,Marc Van Ranst, Paul Rossiter . . . . . . . . . . . . . . . . . . . . . 45 and A. Bennett Jenson . . . . . . . . . . . . . . . 223 Canine Distemper Elizabeth S. Williams 50 12. Pestivirus Infections Hana Van Campen, Feline MorbillivirusInfection Kai Frölich,and Martin Hofmann . . . . . . 232 Linda Munson . . . . . . . . . . . . . . . . . . . . . 59 13. Coronaviral Infections James Measles Linda Munson . . . . . . . . . . . . 63 F. Evermann and David A. Benfield . . . . 245 Morbillivirus Infections in Aquatic 14. Rodent-borne Hemorrhagic Fever Mammals Seamus Kennedy . . . . . . . . . 64 Viruses James N. Mills and James E. Childs . . . . . . . . . . . . . . . . . . . . . . . . . 254 3. Bluetongue,Epizootic Hemorrhagic Disease,and Other Orbivirus-Related 15. Orthomyxovirus and Paramyxovirus Diseases Elizabeth W. Howerth,David Infections Hana Van Campen and E. Stallknecht,and Peter D. Kirkland . . . 77 Greg Early . . . . . . . . . . . . . . . . . . . . . . . 271 4. Arbovirus Infections Thomas M. Yuill 16. Calicivirus Infections Cor Lenghaus, and Charles Seymour . . . . . . . . . . . . . . . 98 Michael J. Studdert,and Dolores Gavier-Widén . . . . . . . . . . . . . . . . . . . . . 280 5. Picornavirus Infections Gavin R. Thomson, Roy G. Bengis,and Corrie C. Brown . . . 119 17. Transmissible Spongiform Encephalopathies Elizabeth S. Williams,James 6. Parvovirus Infections Ian K. Barker and K. Kirkwood,and Michael W. Miller . . . . 292 Colin R. Parrish . . . . . . . . . . . . . . . . . . . 131 7. Herpesvirus Infections Part 2. Bacterial and Mycotic Diseases . . . . 303 Herpesviruses of Nonhuman Primates 18. Tularemia Torsten Mörner and Norval W. King . . . . . . . . . . . . . . . . . . . . 147 Edward Addison . . . . . . . . . . . . . . . . . . . 303 Malignant Catarrhal Fever Werner 19. Plague and Yersiniosis Peter W. Gasper P. Heuschele and Hugh W. Reid . . . . . . . 157 and Rowena P. Watson . . . . . . . . . . . . . . 313 Pseudorabies (Aujeszky’s Disease) 20. Pasteurellosis Michael W. Miller . . . . . 330 David E. Stallknecht and Elizabeth 21. Mycobacterial Diseases W. Howerth . . . . . . . . . . . . . . . . . . . . . . . 164 Mycobacterium bovisInfections Richard Elephant Herpesvirus Infections S. Clifton-Hadley,Carola M. Sauter-Louis, Laura K. Richman and Richard Ian W. Lugton,Ronald Jackson,Peter J. Montali . . . . . . . . . . . . . . . . . . . . . . . . 170 A. Durr,and John W. Wilesmith . . . . . . . . 340 v vi / Contents Paratuberculosis and Other Mycobacterial CampylobacterInfection Torsten Diseases Elizabeth S. Williams . . . . . . . 361 Mörner . . . . . . . . . . . . . . . . . . . . . . . . . . 488 22. Brucellosis E. Tom Thorne. . . . . . . . . . . . 372 Dermatophilosis Frederick A. Leighton 489 23. Anthrax C. Cormack Gates,Brett Elkin, ErysipelothrixInfection Frederick and Dan Dragon . . . . . . . . . . . . . . . . . . . 396 A. Leighton . . . . . . . . . . . . . . . . . . . . . . . 491 24. Diseases due to Mycoplasmas Kevin Fusobacterium necrophorum Whithear . . . . . . . . . . . . . . . . . . . . . . . . . 413 Infection Frederick A. Leighton . . . . . . 493 25. Chlamydiosis of Koalas Richard HelicobacterInfection Torsten Mörner 496 Whittington . . . . . . . . . . . . . . . . . . . . . . . 423 Leptospirosis Frederick A. Leighton 26. Lyme Borreliosis Richard N. Brown and Thijs Kuiken . . . . . . . . . . . . . . . . . . 498 and Elizabeth C. Burgess . . . . . . . . . . . . 435 Listeriosis Torsten Mörner . . . . . . . . . . 502 27. Order Rickettsiales Salmonellosis Torsten Mörner . . . . . . . 505 Anaplasmosis William R. Davidson Shigellosis Torsten Mörner . . . . . . . . . 507 and Will L. Goff . . . . . . . . . . . . . . . . . . . 455 StaphylococcusInfection Gary Wobeser 509 Ehrlichioses William R. Davidson, Jacqueline E. Dawson,and Sidney Tyzzer’s Disease Gary Wobeser . . . . . . 510 A. Ewing . . . . . . . . . . . . . . . . . . . . . . . . . 466 29. Mycotic Diseases Kathy Burek . . . . . . . 514 Heartwater Nancy D. Kock . . . . . . . . . . 477 Contributors . . . . . . . . . . . . . . . . . . . . . . . . . . . 533 Salmon Poisoning Disease William Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 537 J. Foreyt . . . . . . . . . . . . . . . . . . . . . . . . . 480 28. Miscellaneous Bacterial Infections Actinomycesand Arcanobacterium Infections Gary Wobeser . . . . . . . . . . . 487 P R E F A C E Nearly 20 years have passed since the publication zoonotic agents,such as those causing the rodent- of the second edition of Infectious Diseases of borne hemorrhagic fevers and Lyme borreliosis in Wild Mammals, an important resource for, and people, have little or no impact on the wildlife inspiration to, a generation of wildlife biologists that serve as the reservoir for human infection. and veterinarians. All those concerned with dis- Some well-recognized pathogens of domestic eases of wildlife are greatly indebted to the late animals, such as canine distemper virus and par- John Davis,and to Lars Karstad and Dan Trainer, voviruses, are emerging in significance or for consolidating knowledge in this field and increasingly recognized in free-ranging wildlife. making it so readily accessible. Their work Diseases such as brucellosis,bovine tuberculosis, helped establish the widespread and growing rinderpest, and foot-and-mouth disease arguably recognition of the influence of disease on popula- are more strongly associated with domestic ani- tions of wild mammals, consideration of disease mals than with wildlife,but they pose some of the in the fields of wildlife management and ecology, knottiest epidemiologic,management,and ethical and greater appreciation of disease as a compo- problems where wild animals are affected. Some nent in the relationship among wild mammals, agents, such as the myxoma virus, rabbit cali- humans,and domestic animals. civirus, and Salmonella, have been exploited as To us has fallen the humbling task of regener- biologic agents of vertebrate pest control, with ating and updating this work, in collaboration varying degrees of success and safety. Others, with an international field of chapter authors. Our such as poxviruses and Leptospira,are functional goal has been to summarize knowledge in the or candidate recombinant vectors for the delivery field of wildlife diseases relevant to wild mam- of antigens to immunize wild mammals. The sig- mals, in a form that will be useful to students in nificance for wild mammals of agents such as wildlife biology or veterinary medicine; wildlife Chlamydia and its relatives, and the retroviruses biologists and managers; veterinarians dealing and mycoplasmas, seems relatively restricted, with free-living and captive wildlife; and epi- taxonomically and/or geographically, while oth- demiologists and public health professionals con- ers,such as the lyssaviruses,the poxviruses,Sal- cerned with wildlife zoonoses. monella, and Leptospira, affect a wide array of The format of the third edition follows that of wild mammals on most continents. its predecessors,with chapters arranged by taxon Knowledge of diseases included in the last edi- of infectious agent. They deal with agents estab- tion has greatly expanded, and a number of new lished or suspected as pathogens in wild mam- agents that fall within the scope of this volume mals, and/or transmissible between wild mam- have emerged in the past two decades. Accord- mals and domestic animals or people. Many are ingly,we have added major treatments of problems capable of causing devastating diseases, such as such as rodent-borne hemorrhagic fevers, Lyme tularemia, plague, or rabies, among wild mam- borreliosis, transmissible spongiform encepha- mals; almost all of these are capable of infecting lopathies,and calicivirus infections. Consideration domestic animals or people, as well. Other of the implications for wildlife of agents such as vii viii / Preface the parvoviruses, canine distemper virus, and the Selected on the basis of their expertise and famil- rickettsiae has been expanded. And, of necessity, iarity with the wildlife ramifications of the topic some topics found in the second edition have been assigned, the authors span the disciplines of truncated,consolidated,or eliminated. wildlife biology,veterinary medicine,epidemiol- Within the limits of the space allotted, chapter ogy,and microbiology. Years of experience in the authors provide an entrée to the historical litera- field and in the laboratory, on all continents ture on an agent or disease,a summary of current except Antarctica, have been distilled onto these knowledge on the etiology, pathogenesis, immu- pages. The editors found it exciting to foster inter- nity, and diagnosis, and discussions of implica- national collaborations on authorship of chapters, tions of the agent or disease for captive or free- often by means of electronic communication,and ranging wild mammals, domestic animals, and to participate as they developed and evolved. Like people. Relevant current information on the biol- the editors, many authors had to cope with the ogy and epidemiology of pathogens gained by effects of economic rationalism and downsizing molecular techniques has been incorporated, but on their workplace. Chapter authorship was an act the rapid expansion of such knowledge, and the of dedication, for little reward, often carried out inevitable lag in bringing a work such as this to to difficult deadlines under trying circumstances. press, no doubt already will have dated chapters Sadly,two of our contributors,Drs. Charles Sey- on some very active topics. mour and Werner Heuschele, died during the Despite the expansion in size of this volume, preparation of their chapters. To the chapter space limitations forced elimination of most com- authors goes credit for the quality of the content prehensive reviews,lists of citations,and detailed of this book; the editors assume responsibility for treatment of clinical signs and comparative diag- any shortcomings. nosis. Of necessity, many chapters are surveys We also wish to acknowledge colleagues who rather than full reviews of the topic, and readers read drafts of chapters or who provided technical will need to consult the second edition and other advice on various topics. Iowa State University sources cited for fuller historical and current Press, in particular Gretchen Van Houten, has information. The number of illustrations and the been very helpful and extremely accommodating number and scope of tables also were limited in over the course of the preparation of this edition. deference to textual material. Nonetheless, We greatly appreciate the tolerance and encour- between these covers is a vast amount of infor- agement to carry this project through that were mation,covering a wide array of infectious agents offered by our spouses, Tom Thorne and Susy affecting virtually all orders of mammals. Refer- Carman. ence lists lead to the wider literature, and the Lastly, we wish to recognize the Wildlife Dis- index will allow the reader to find and consolidate ease Association and its many dedicated mem- information affecting particular families or gen- bers, who, over the course of nearly 50 years, era of mammals, which may be scattered among have shared in the study and dissemination of several chapters. Walker’s Mammals of the World, knowledge of diseases of wildlife,in the interests fifth edition, by R.M. Nowak (Johns Hopkins of animal and human health and welfare, and Press,1991),was used as the authority for mam- environmental conservation. The Wildlife Dis- malian nomenclature, except where another ease Association supported the preparation of this authority is cited by chapter authors. volume and is the beneficiary of the royalties that We gratefully acknowledge the effort that our will accrue from its use. many authors dedicated to their contributions. Infectious THIRD EDITION Diseases of Wild Mammals I PART Viral and Prion Diseases 1 RABIES CHARLES E. RUPPRECHT,KLAUS STÖHR, AND COURTNEY MEREDITH Synonyms:al kalabe,beshenstvo,derriengue,hari, book, significant advances in biotechnology have hydrophobia (humans),kalevet,kuang cheng,lyssa, occurred, particularly in the diagnostic arena with the mal de calderas,oulo fato,polar madness,rabhas, development of monoclonal antibodies (Mabs) and rage,rabia,rabbia,raiva,thao,tollwut. molecular typing techniques that have shed consider- able light on the epidemiology of rabies. In addition,in the vaccine field,there has been substantial application INTRODUCTION. Rabies is an acute fatal viral of oral immunization toward wildlife rabies control. encephalomyelitis. The etiologic agents belong to the genus Lyssavirus, in the family Rhabdovirus. Described for at least four millennia, it is one of the HISTORY AND HOSTS oldest recognized infectious diseases. A clinical entity indistinguishable from classic rabies has been reported Africa. Circumstantial evidence indicts Africa as the on all inhabited continents. Warm-blooded vertebrates cradle of lyssavirus evolution, because nowhere else are susceptible to experimental infection, but only are at least four serotypes or genotypes present (Rup- mammals are important in the epidemiology of rabies. precht et al. 1991; King et al. 1994). Besides rabies The etiologic agents are neurotropic viruses that pri- virus,only two European bat lyssaviruses and a newly marily replicate in the central nervous system (CNS) described Australian virus have become established and pass to the salivary glands and saliva. Transmission outside of Africa, perhaps by virtue of bat flight and principally occurs by the bites of infected carnivores subsequent zoogeographic spread. Classic serotype/ and bats. As a major zoonosis,rabies has considerable genotype 1 rabies virus, found mainly in dogs Canis public health,veterinary,and economic impact,and it familiaris,has achieved virtual worldwide distribution, should be of concern to all professionals that deal primarily with human assistance (Smith and Seidel directly with the health of free-ranging or captive mam- 1993),whereas the other viruses by comparison may be mals. Since the publication of the first edition of this viewed somewhat as biologic curiosities. 3

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