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M E D I C A L INSECTS A N D A R A C H N I D S

THE NATURAL HISTORY MUSEUM

M E D I C A L INSECTS A N D A R A C H N I D S

Edited by R I C H A R D P. L A N E and ROGER W. CROSSKEY

Department of Entomology, The Natural History Museum

(British Museum (Natural History)) London, UK

m SPRINGER-SCIENCE+BUSINESS MEDIA, B.V.

First edition 1993 © 1993 Springer Science+Business Media Dordrecht Originally published by Chapman & Hall in 1993 Softcover reprint of the hardcover 1st edition 1993 Typeset in 10/12pt Palatino by Expo H o l d i n g , Malaysia

ISBN 978-94-010-4679-4 ISBN 978-94-011-1554-4 (eBook) DOI 10.1007/978-94-011-1554-4

A p a r t f rom any fair dealing for the purposes of research or private study, or criticism or review, as permitted under the U K Copyr ight Designs and Patents Act , 1988, this publication may not be reproduced, stored, or transmitted, i n any form or by any means, without the prior permission i n wr i t ing of the publishers, or i n the case of reprographic reproduction only i n accordance w i t h the terms of the licences issued by the Copyr ight Licensing Agency i n the U K , or i n accordance w i t h the terms of licences issued by the appropriate Reproduction Rights Organizat ion outside the U K . Enquiries concerning reproduction outside the terms stated here should be sent to the publishers at the L o n d o n address printed on this page.

The publisher makes no representation, express or i m p l i e d , w i t h regard to the accuracy of the information contained i n this book and cannot accept any legal responsibility or l iabi l i ty for any errors or omissions that may be made.

A catalogue record for this book is available f rom the Brit ish Library

Library of Congress Cataloging-in-Publication data available

The use of the term 'man' is not indicative of a specific gender and should be taken to refer to both men and w o m e n throughout.

Contents

Contributors xi Preface xiii

1 General introduction 1 R. P. Lane and R. W. Crosskey

Classification and nomenclature 1 Species and species complexes 12 Distribution and zoogeographical regions 16 Vectors and vector status 19 Collection, storage and handling of specimens 23 References 27

2 Introduction to the arthropods 30 R. P. Lane

Structure 30 Classification 32 The principal arthropod groups 33 Further reading 46

Part One Diptera 49

3 Introduction to the Diptera 51 R. W. Crosskey

Morphology 52 Taxonomy 63 Biology 72 References 75

4 SandfLies (Phlebotominae) 78 R. P. Lane

Recognition and elements of structure 79 Classification and identification 89 Biology 100 Medical importance 103 Control 107

vi Contents

Collecting, preserving and rearing material 111 References 113

5 Mosquitoes (Culicidae) 120 M. W. Service

Recognition and elements of structure 121 Classification and identification 139 Biology 173 Medical importance 196 Control 208 Collecting, preserving and rearing material 216 References 221

6 BlackfLies (Simuliidae) 241 R. W. Crosskey


7 Biting midges (Ceratopogonidae) 288 John Boorman


8 Horse-flies, deer-flies and clegs (Tabanidae) 310 John E. Chainey

Recognition and elements of structure 310 Classification and identification 316 Biology 323 Medical importance 325 Control 327

Collecting, preserving and rearing material References

9 Tsetse-flies (Glossinidae) A. M. Jordan

Recognition and elements of structure Classification and identification Biology Medical importance Control Collecting, preserving and rearing material References

10 Stable-flies and horn-flies (bloodsucking Muscidae) R. w. Crosskey

Recognition and elements of structure Classification and identification Biology Medical importance Control Collecting, preserving and rearing material References

11 House-flies, blow-flies and their allies (calyptrate Diptera) R. W. Crosskey and R. P. Lane

Recognition of calyptrates and elements of structure Identification of medically important calyptrate flies Biology of synanthropic flies Medical importance of adult calyptrate flies Control References

12 Diptera causing myiasis in man Martin J. R. Hall and Kenneth G. V. Smith

Recognition and elements of structure Review of dipterous families associated with human myiasis Maggot therapy Control Collecting, preserving and rearing material References

Contents vii

327 328

333

334 340 365 372 379 381 382

389

389 391 396 398 399 400 400

403

403 407 419 423 425 426

429

430 431 459 459 462 464

Vlll Contents

Part Two Other insects 471

13 Cockroaches (Blattaria) 473 N. R. H. Burgess

Recognition and elements of structure 473 Classification and identification 474 Biology 479 Medical importance 480 Control 481 References 482

14 Bedbugs and kissing-bugs (bloodsucking Hemiptera) 483 C. J. Schofield and W. R. Dolling

Recognition and elements of structure 484 Classification and identification 485 Biology of the Triatominae 502 Medical importance 506 Control 508 Collecting, preserving and rearing material 509 References 510

15 Lice (Anoplura) 517 Joanna Ibarra

Recognition and elements of structure 518 Biology 521 Medical importance 523 Control 525 References 527

16 Fleas (Siphonaptera) 529 Robert E. Lewis


17 Insects of minor medical importance 576 Kenneth G. V. Smith

Diptera 576 Hymenoptera 581

Coleoptera Lepidoptera Thysanoptera (thrips) Incidental insects Insects and hygiene References

Part Three Arachnids

18 Ticks and mites (Acari) M. R. G. Varma

Recognition, structure and classification of the Acari Review of the suborder Ixodida (ticks) Biology of ticks Medical importance of ticks Introduction to mites: recognition and structure Classification and identification of mites Biology and medical importance of mites Control of ticks and mites Collecting, preserving and rearing material of ticks and mites References

19 Spiders and scorpions (Araneae and Scorpiones) J. L. Cloudsley-Thompson

Recognition and elements of structure Classification and identification Biology Medical importance Control Collecting and preserving material References

Scientific names index

Subject index

Contents ix

583 585 588 589 589 589

595

597

597 600 616 623 631 632 634 648 649 654

659

660 666 670 672 678 679 680

683

706

Contributors

Mr John Boorman 6 Beckingham Road, Guildford, Surrey GU2 6BN

Dr N. R. H. Burgess Department of Military Entomology, Royal Army Medical College, Millbank, London SW1P 4RJ

Mr John E. Chainey Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD

Professor J. L. Cloudsley-Thompson Department of Biology, Medawar Building, University College London, Gower Street, London WC1E 6BT

Dr R. W. Crosskey Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD

Mr W. R. Dolling Brook Farmhouse, Elstronwick, Hull HU12 9BP

Dr Martin J. R. Hall Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD

Mrs Joanna Ibarra Community Hygiene Concern, 32 Crane Avenue, Isleworth, Middlesex TW7 7JL

Dr A. M. Jordan Tsetse Research Laboratory, University of Bristol, Langford House, Langford, Bristol BS18 7DU

xii Contributors

DrR. P. Lane Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD

Professor Robert E. Lewis Department of Entomology, Iowa State University, Ames, Iowa 50011, USA

Dr C. J. Schofield 7 rue Maclonay, Pregnin, St. Genis - Pouilly, 01630 France

Professor M. W. Service Vector Biology and Control Group, Liverpool School of Tropical Medicine, Pembroke Place, Liverpool L3 5QA

Mr Kenneth G. V. Smith 70 Hollick Wood Avenue, London N12 OLT

Professor M. G. R. Varma c/o Department of Medical Parasitology, London School of Hygiene and Tropical Medicine, Keppel Street, London WC1E 7HT

Preface

Surprising though it seems, the world faces almost as great a threat today from arthropod-borne diseases as it did in the heady days of the 1950s when global eradication of such diseases by eliminating their vectors with synthetic insecticides, particularly DDT, seemed a real possibility. Malaria, for example, still causes tremendous morbidity and mortality throughout the world, especially in Africa. Knowledge of the biology of insect and arachnid disease vectors is arguably more important now than it has ever been. Biological research directed at the development of better methods of control becomes even more important in the light of the partial failure of many control schemes that are based on insecticides -although not all is gloom, since basic biological studies have contributed enormously to the outstanding success of international control programmes such as the vast Onchocerciasis Control Programme in West Africa.

It is a sine qua non for proper understanding of the epidemiology and successful vector control of any human disease transmitted by an arthropod that all concerned with the problem - medical entomologist, parasitologist, field technician - have a good basic understanding of the arthropod's biology. Knowledge will be needed not only of its direct relationship to any parasite or pathogen that it transmits but also of its structure, its life history and its behaviour - in short, its natural history. Above all, it will be necessary to be sure that it is correctly identified.

Close attention to biological details often reveals the presence of species complexes of vectors, which if they are to be analysed for correct identification of component sibling species, require the application of sophisticated techniques extending beyond the ubiquitous morphological approach to identification. Study of such complexes leads to understanding of speciation mechanisms, and has begun to take medical entomology into evolutionary biology. In a similar manner, there is no clear division between medical entomology and veterinary entomology - very often the same groups of insects, mites or ticks transmit parasites to domestic animals. Clearly, medical entomology does not stand in isolation.

Unfortunately, while medical entomology is comparatively rich in works which deal with particular groups of arthropods at the level needed by specialists, it is rather less well supplied with more general literature which broadly summarizes information and meets the needs of students in tropical medicine and allied fields.

The book we present here, under the short title Medical Insects and Arachnids, brings to fruition a project we initiated several years ago when it became clear to us that there was no up-to-date and comprehensive introduction to arthropods important in human health. The march of events in medical entomology over the past 20 years has outdated the well-known book Insects and Other Arthropods of Medical Importance that was edited by our colleague Mr K. G. V. Smith and published by the British Museum (Natural History) in 1973 - one need think only of how the importance of sibling species complexes in arthropod-borne vectors of

xiv Preface

human diseases has come to be fully realized and of the varied approaches to vector taxonomy and identification this has brought in its train.

This book is rooted in group-by-group systematic coverage, and on this account we have (as editors) prepared the chapters in generally similar format so that the structure and biology of the different groups can be easily compared. So far as is practicable, we have balanced the chapters to reflect the biological complexity and proved medical importance of the groups concerned. Control is covered only briefly, with the focus mainly on the biological rationale for the interventions adopted.

We have paid particular attention to the identification keys that are included (revising manuscripts where necessary) because we are conscious of how difficult students find keys when these are written in the jargon of taxonomists. The keys are presented in a completely uniform style and in what we believe is more than usually friendly language, avoiding unnecessary technical terms.

As part of the essential aim of the book, stress is laid on 'systematic' aspects of medical entomology, that is to say on the structure and biology of each medically related group of arthropods and upon how the important families, genera and species can most easily be identified. Special mention is made of non-morphological taxonomic techniques, and the scientific background to methods and concepts applicable in medical entomology is summarized in a general introduction. (We have omitted the names of those who described the genera, species and subspecies mentioned in the text because these do not form part of the scientific names concerned and serve no purpose in a book of this kind.)

Major effort has been applied to the illustrations, many of which have been specially drawn or modified to meet the needs of the text. Labelling of illustrations is to a uniform style and in full, thereby obviating the need (which we have always found irksome) to refer to lengthy legends or to memorize abbreviations. Roman numerals have been everywhere avoided as we see no point in imposing this unnecessary system on those who might not be familiar with it; larval instars and abdominal segments, for example, are labelled in arabic numbers.

Many medical entomologists working in the field without access to libraries depend on literature obtained in photocopy or through library loan. Correct bibliographic citation is therefore essential. To meet this need, we have verified all the references cited in the book. The references provide leads for readers wanting more information on morphology, taxonomy, biology, vector/parasite relations, vector control and faunistics. (In connection with the last, we observe that we did not anticipate the rapid demise of the Soviet Union while this book was being completed and the now- historical abbreviation USSR is used.)

This book could not have been prepared without the keen cooperation of our 13 fellow contributors, and we take this opportunity to thank them not only for their contributions but for allowing us to modify their texts when this seemed desirable for consistency and balance between chapters. Others have helped us behind the scenes, and it is our pleasure to acknowledge assistance received from Dr Anne Baker (on mites) and Mr Paul Hillyard (on spiders), from librarians Julie Harvey and Kathy Martin who obtained literature not immediately to hand in the British Museum (Natural History), and from Jan Townsend and Angel Hathaway who typed parts of the text. Professor M. W. Service provided us with some helpful bibliographic information.

Preface xv

Some illustrations for which the British Museum (Natural History) does not own copyright have appeared in previous literature, and we are grateful to the following publishers for permission to reproduce the figures indicated: American Museum of Natural History (Fig. 14.2a); Blackwell Scientific Publications (Figs 5.2d, 5.7b, 5.9, 5.12, 5.16b,d, 5.20); CAB International (Fig. 5.1Of); Canada Agriculture (Figs l1.5a, 12.1.2b,c, 17.2b); Commonwealth Scientific and Industrial Research Organization, Canberra (Figs 3.9a, 3.9d, 11.5b,c originally drawn by T. Binder); Edward Arnold (Fig. 5.14); Entomological Society of America (Fig. 5.25); Faune de Madagascar (Fig. 5.22); Oxford University Press (Fig. 5.29); Unwin Hyman's Ltd (Fig. 5.lOc); Walter Reed Biosystematics Unit (Figs 5.8, 5.lOa,b,d,e, 5.11, 5.13a,b, 5.16a,c, 5.17, 5.19, 5.24, 5.26); World Health Organization (Fig. 5.30). Figs 4.2, 4.6, 6.3, 8.4 and 1O.2b,d,e are reproduced courtesy of the Wellcome Trust from Boris Jobling's Anatomical Drawings of Biting Flies (1987); Figs 11.5a, 12.12b,c and 17.2c are from Agriculture Canada's Manual of Nearctic Diptera, volume 1 (1981), and are reproduced with the permission of the Minister of Supply and Services, Canada, 1992. Mr M. Smith kindly supplied the photograph used for Fig. 15.3.

Acknowledgements made by individual chapter contributors are as follows. J. L. Cloudsley-Thompson (Araneae and Scorpiones) thanks Maureen Hakney and Helen Wilson for deciphering his handwriting and typing his manuscript; R. E. Lewis (Siphonaptera) thanks Mr F. G. A. Smit for the splendid job done by him in the flea chapter published previously in the book edited by K. G. V. Smith (see above); C. J. Schofield and W. R. Dolling (Hemiptera) thank Professor Randall T. Schuh and Professor Carl W. Schaeffer for advice on higher classification of bugs, and Dr H. Brailovsky for helpful discussion of recently described triatomine species; M. W. Service (Culicidae) thanks Professor A. N. Clements, Professor P. S. Corbet, Dr. E. L. Peyton and Dr H. Townson for comments on draft text, Professors L. de Vos, G. Josens and B. Vray for the photograph used in Fig. 5.4, Dr M. Coetzee and Dr R. Hunt for the photograph used in Fig. 5.31, Dr L. P. Lounibos for the photograph used in Fig. 5.28, and Dr. R. A. Ward for information on the number of world mosquito species.

We conclude this preface with appreciative thanks tei our wives for their forbearance in tolerating long periods when we appeared to be 'incommunicado' while in the throes of editing and preparing this book. Special thanks go to Peggy Crosskey, and to her sister Jean Johns, who shouldered the considerable task of compiling the draft index to scientific names.

Richard P. Lane and Roger W. Crosskey

British Museum (Natural History), London, January 1992

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