VetBooks.ir Veterinary Ectoparasites: Biology, Pathology and Control VetBooks.ir Veterinary Ectoparasites: Biology, Pathology and Control Second Edition Richard Wall BSc, MBA, PhD David Shearer BvetMed, CertSAD, PhD, CBiol, MIBiol, MRCVS b Blackwell Science VetBooks.ir # 1997, 2001 by DISTRIBUTORS Blackwell Science Ltd Editorial Offices: Marston Book Services Ltd Osney Mead, Oxford OX2 0EL PO Box 269 25 John Street, London WC1N 2BS Abingdon 23 Ainslie Place, Edinburgh EH3 6AJ Oxon OX14 4YN 350 Main Street, Malden (Orders: Tel: 01235 465500 MA 02148 5018, USA Fax: 01235 465555) 54 University Street, Carlton Victoria 3053, Australia USA and Canada 10, rue Casimir Delavigne Iowa State University Press 75006 Paris, France A Blackwell Science Company 2121 S. State Avenue Other Editorial Offices: Ames, Iowa 50014-8300 (Orders: Tel: 800-862-6657 Blackwell Wissenschafts-Verlag GmbH Fax: 515-292-3348 KurfuÈ rstendamm 57 Web www.isupress.com 10707 Berlin, Germany email: [email protected] Blackwell Science KK Australia MG Kodenmacho Building Blackwell Science Pty Ltd 7±10 Kodenmacho Nihombashi 54 University Street Chuo-ku, Tokyo 104, Japan Carlton, Victoria 3053 (Orders: Tel: 03 9347 0300 Iowa State University Press Fax: 03 9347 5001) A Blackwell Science Company 2121 S. State Avenue A catalogue record for this title Ames, Iowa 50014-8300, USA is available from the British Library ISBN 0-632-05618-5 The right of the Author to be identified as the Author of this Work has been asserted in accordance with the Copyright, Library of Congress Designs and Patents Act 1988. Cataloging-in-Publication Data is available All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, For further information on recording or otherwise, except as permitted by the UK Blackwell Science, visit our website: Copyright, Designs and Patents Act 1988, without the prior www.blackwell-science.com permission of the publisher. First edition published by Chapman & Hall 1997 Second edition published by Blackwell Science 2001 Set in 10/12pt Times by DP Photosetting, Aylesbury, Bucks Printed and bound in Great Britain by Anthony Rowe Ltd, Chippenham The Blackwell Science logo is a trade mark of Blackwell Science Ltd, registered at the United Kingdom Trade Marks Registry VetBooks.ir Contents Preface to Second Edition ix 2.3 Life history 25 Acknowledgements x 2.4 Pathology 26 2.5 Classification 26 Chapter 1 The Importance and Diversity of 2.5.1 Astigmata 26 Arthropod Ectoparasites 1 2.5.2 Prostigmata 27 1.1 Introduction 1 2.5.3 Mesostigmata 27 1.2 Ectoparasite±host relationships 1 2.6 Recognition of mites of veterinary 1.3 Ectoparasite damage 2 importance 27 1.4 The evolution of ectoparasite±host 2.7 Astigmata (Sacroptiformes) 27 relationships 3 2.7.1 Sarcoptidae 27 1.5 A modern and growing problem? 5 Guide to the suborders of Acari 28 1.6 An introduction to arthropod Guide to species and families of structure and function 6 veterinary importance 28 1.6.1 Arthropod segmentation 6 2.7.2 Psoroptidae 34 1.6.2 The arthropod exoskeleton 6 Guide to the identification of life-cycle 1.6.3 Jointed legs 8 stages of Psoroptes mites 36 1.6.4 Spiracles and gas exchange 8 2.7.3 Knemidocoptidae 40 1.6.5 The arthropod circulatory 2.7.4 Listrophoridae 41 system 9 2.7.5 Astigmatid mites of minor 1.6.6 The arthropod nervous system 10 veterinary interest 43 1.6.7 Digestion and absorption 11 2.8 Prostigmata (Trombidiformes) 44 1.6.8 Arthropod sense organs 12 2.8.1 Demodicidae 44 1.6.9 Arthropod reproduction 13 2.8.2 Cheyletiellidae 45 1.6.10 Arthropod size 13 2.8.3 Trombiculidae 47 1.7 Patterns of arthropod development 14 2.8.4 Psorergatidae 48 1.7.1 Moulting 14 2.8.5 Prostigmatid mites of minor 1.7.2 Simple and complex life-cycles 14 veterinary interest 49 1.8 The classification of diversity 16 2.9 Mesostigmata (Gamesid mites) 50 1.9 The origins of arthropods 16 2.9.1 Macronyssidae 50 1.10 Living arthropod groups 17 2.9.2 Dermanyssidae 52 1.10.1 Arachnids 18 2.9.3 Mesostigmatid mites of minor 1.10.2 Insects 19 veterinary interest 53 1.10.3 Other living arthropod classes 20 Further reading and references 54 1.11 Arthropod distributions 21 Further reading and references 22 Chapter 3 Ticks (Acari) 55 3.1 Introduction 55 Chapter 2 Mites (Acari) 23 3.2 Morphology 55 2.1 Introduction 23 3.2.1 Ixodidae 55 2.2 Morphology 23 3.2.2 Argasidae 57 v VetBooks.ir vi Contents 3.3 Life history 58 4.8.1 Tabanidae (horse flies, deer 3.3.1 Ixodidae 58 flies and clegs) 101 3.3.2 Argasidae 60 4.9 Nematocera 104 3.4 Pathology 61 4.9.1 Simuliidae (black flies) 104 3.4.1 Cutaneous effects of tick feeding 61 4.9.2 Ceratopogonidae (biting midges) 107 3.4.2 Systemic effect: vectors of disease 61 4.9.3 Culicidae (mosquitoes) 108 3.4.3 Systemic effects: tick paralysis 64 4.9.4 Psychodidae (sand flies) 110 3.4.4 Other systemic effects 65 4.10 Other Diptera of veterinary interest 111 3.5 Classification 65 4.10.1 Eye gnats 111 3.6 Recognition of ticks of veterinary Further reading and references 112 importance 65 Guide to tick identification 66 Chapter 5 Myiasis 114 3.7 Ixodidae 67 5.1 Introduction 114 3.7.1 Ixodes 67 5.2 Morphology 114 3.7.2 Dermacentor 71 5.3 Life history 115 3.7.3 Haemaphysalis 74 5.4 Pathology 116 3.7.4 Rhipicephalus 75 5.5 Classification 116 3.7.5 Boophilus 76 5.6 Recognition of dipterous agents of 3.7.6 Amblyomma 77 myiasis 117 3.7.7 Hyalomma 78 Guide to third-stage larvae causing 3.8 Argasidae 78 myiasis in domestic animals 117 3.8.1 Argas 78 Guide to genera of adult Diptera 3.8.2 Otobius 80 causing myiasis in domestic animals 119 3.8.3 Ornithodoros 80 5.7 Oestridae 121 Further reading and references 81 5.7.1 Oestrinae 121 5.7.2 Gasterophilinae 123 Chapter 4 Adult Flies (Diptera) 83 Guide to the third-stage larvae 4.1 Introduction 83 of the most important 4.2 Morphology 83 Gasterophilus species 124 4.3 Life history 85 5.7.3 Hypodermatinae 126 4.4 Pathology 86 5.7.4 Cuterebrinae 129 4.5 Classification 87 5.8 Calliphoridae 130 4.5.1 Cyclorrhapha 87 5.8.1 Cochliomyia 130 4.5.2 Brachycera 88 5.8.2 Chrysomya 132 4.5.3 Nematocera 88 5.8.3 Lucilia 134 4.6 Recognition of flies of veterinary 5.8.4 Phormia and Protophormia 137 importance 88 5.8.5 Calliphora 138 4.7 Cyclorrhapha 88 5.8.6 Cordylobia 139 Guide to families of adult Diptera of 5.9 Sarcophagidae 140 veterinary importance 89 5.9.1 Wohlfahrtia 140 4.7.1 Muscidae 92 Further reading and references 141 4.7.2 Fanniidae 97 4.7.3 Hippoboscidae (keds and forest Chapter 6 Fleas (Siphonaptera) 143 flies) 98 6.1 Introduction 143 4.7.4 Glossinidae (tsetse flies) 99 6.2 Morphology 144 4.7.5 Cyclorrhaphous flies of minor 6.3 Life history 146 veterinary interest 100 6.4 Pathology 148 4.8 Brachycera 101 6.5 Classification 149 VetBooks.ir Contents vii 6.6 Recognition of fleas of veterinary 8.2.4 Deep skin scraping (deep importance 149 epidermal examination) 181 6.7 Pulicidae 149 8.2.5 Collection of free-living 6.7.1 Ctenocephalides 149 ectoparasites 181 Guide to the flea species of veterinary 8.2.6 Biopsy and histopathology 181 importance 150 8.3 The chemical control of ectoparasites ± 6.7.2 Spilopsyllus 154 ectoparasiticides 181 6.7.3 Echidnophaga 155 8.3.1 Ectoparasiticides: early 6.7.4 Pulex 156 compounds 181 6.7.5 Xenopsylla 156 8.3.2 Ectoparasiticides: neurotoxins 182 6.8 Ceratophyllidae 157 8.3.3 Ectoparasiticides: insect growth 6.8.1 Ceratophyllus 157 regulators 184 6.8.2 Nosopsyllus 158 8.3.4 Repellents 185 6.9 Flea species of minor veterinary 8.3.5 Desiccants 185 interest 160 8.4 Mode of ectoparasiticide Further reading and references 160 application 185 8.4.1 Topical preparations 185 Chapter 7 Lice (Phthiraptera) 162 8.4.2 Systemic preparations 185 7.1 Introduction 162 8.4.3 Environmental preparations 186 7.2 Morphology 162 8.5 Problems with chemical control 186 7.3 Life history 164 8.5.1 Poisoning and environmental 7.4 Pathology 164 contamination 186 7.5 Classification 165 8.5.2 Resistance 187 7.6 Recognition of lice of veterinary 8.6 Non-chemical control of importance 166 ectoparasites 187 Guide to the genera of lice of 8.6.1 Physical control 187 veterinary interest 166 8.6.2 Barriers 188 7.7 Amblycera 168 8.6.3 Biological control 188 7.7.1 Menoponidae 168 8.6.4 Vaccination 188 7.7.2 Boopidae 169 8.6.5 Trapping 189 7.7.3 Gyropidae 169 8.6.6 Sterile insect technique 189 7.8 Ischnocera 170 8.6.7 Modelling and forecasting 189 7.8.1 Philopteridae 170 8.7 Cattle 190 7.8.2 Trichodectidae 172 8.7.1 Mites 190 7.9 Anoplura 174 8.7.2 Ticks 192 7.9.1 Haematopinindae 174 8.7.3 Flies 193 7.9.2 Linognathidae 176 8.7.4 Myiasis 194 7.9.3 Polyplacidae 177 8.7.5 Fleas 195 Further reading and references 178 8.7.6 Lice 195 8.8 Sheep 196 Chapter 8 The Diagnosis and Control of 8.8.1 Mites 196 Ectoparasite Infestation 179 8.8.2 Ticks 199 8.1 Introduction 179 8.8.3 Flies 199 8.2 Diagnosis of ectoparasite infestation 179 8.8.4 Myiasis 200 8.2.1 Hair examination 180 8.8.5 Fleas 202 8.2.2 Acetate strip examination 180 8.8.6 Lice 202 8.2.3 Superficial skin scraping 8.9 Horses 202 (epidermal surface examination) 180 8.9.1 Mites 202 VetBooks.ir viii Contents 8.9.2 Ticks 204 8.15 Guinea-pigs 230 8.9.3 Flies 205 8.15.1 Mites 230 8.9.4 Myiasis 207 8.15.2 Flies 231 8.9.5 Fleas 208 8.15.3 Myiasis 231 8.9.6 Lice 208 8.15.4 Fleas 231 8.10 Pigs 208 8.15.5 Lice 232 8.10.1 Mites 208 8.16 Mice and rats 232 8.10.2 Ticks 210 8.16.1 Mites 232 8.10.3 Flies 210 8.16.2 Flies 233 8.10.4 Myiasis 210 8.16.3 Myiasis 233 8.10.5 Fleas 211 8.16.4 Fleas 233 8.10.6 Lice 211 8.16.5 Lice