Atlas of Entomopathogenic Fungi

Cordyceps tuberculata on moth (Sphingidae), Indonesia.

Erynia neoaphidis on Aphis (Aphididae), France.

S porodiniel/a umbel/ata on cicada nymph (Cicadidae), Papua New Guinea.

Hirsutella versicolor on leafhopper (Cicadellidae), Kenya.

Conidiobolus obscurus on Acyrthosiphon pisum (Aphididae), France.

Erynia gammae on lepidoptera larva (N octuidae), Switzerland.

Hirsutella jonesisii on green planthopper (Nephotettix), Indonesia.

Entomophthora muscae on fly (Diptera), Denmark.

Erynia radicans on Dicyphus pallidus (Miridae), Switzerland.

Erynia eulieis on Coddis fly (Diptera), UK.

Fusarium eoeeophilum on pine-needle scales (Phenaeaspis), Honduras.

Entomophaga aulieae on lepidoptera larva, Canada.

Erynia eoniea on Simulium sp. (Diptera), UK.

Cordyeeps variabilis on Coleoptera larva (Elateridae), Canada.

Cordyeeps sp. on adult beetle (Coleoptera), Brazil.

Nomuraea rileyi on rice looper (Lepidoptera), Indonesia.

Cordyeeps unilateralis on Camponotus ant, Costa Rica.

Cordyceps militaris on Lepidoptera purpa, Netherlands.

Robert A. Samson Harry C. Evans Jean-Paul Latge

Atlas of Entornopathogenic Fungi

Springer-Verlag Berlin Heidelberg GmbH

Cordyceps dipterigena on robberfly (Asilidae), Indonesia.

Cordyceps locustiphila on Locustidae, Ecuador.

Cordyceps sp. oil cockchafer larva (Scarabidae), Brazil.

Metarhizium anisopliae var. majus on third instar larvae of Oryctes rhinoceros, Philippines.

Cordyceps sp. on Psychidae larva, Brazil.

Tolypocladium cylindrosporum on mosquito larva (Aedes aegyptii), USA.

Hymenostilbe sp. on Gryllidae, Ecuador.

Erynia delphacis on green leafhopper (Nephotettix virescens), Indonesia.

Gibellula species on spider (Salticidae), Brazil.

Aschersonia cubensis on scale insect (Lecaniidae), Brazil.

Polycephalomyces ramosus on fly, the Netherlands.

Aschersonia aleyrodis on citrus whiteflies (Aleyrodiidae) USA.

Cordyceps curculionum on weevil adult (Curculionidae), Brazil.

Gibellula pulchra on spider (Salticidae), Ecuador.

Robert A. Samson Centraalbureau voor Schimmelcultures, Baarn, The Netherlands

Harry C. E vans CAB International Institute ofBiological Control, Silwood Park, Ascot, United Kingdom

Jean-Paul Latge Institut Pasteur, Paris, France

Sole distribution rights outside the Netherlands granted to Springer-Verlag Berlin Heidelberg GmbH

ISBN 978-3-662-05892-3 ISBN 978-3-662-05890-9 (eBook) DOI 10.1007/978-3-662-05890-9

© Springer-Verlag Berlin Heidelberg 1988 Originally published by Springer-Verlag Berlin Heidelberg New York in 1988 Softcover reprint of the hardcover lst edition 1988

Preface

A knowledge of insect diseases (whether or not it has to do with control) is of funda­mental andfar-reaching importance in the study of insect ecology. E. A. Steinhaus (1949).

As biological control is becoming more acceptable as a practical science and the dangers of the long-term use of chemical pesticides are fully appreciated, there has been a resurgence of interest in em­ploying fungal pathogens to combat insect pests. New production and application techniques combined with a greater understanding of both fungal and insect ecology have meant that biological insecti­cides can now compete on a more equal footing with traditional chemical pesticides. Consequently, agrochemical companies are be­ginning to take account of entomopathogens as both viable and eco­nomic propositions and to invest more resources in their research and development, particularly to improve the formulations of well­documented pathogens, possibly by genetic manipulation, and to test more obscure or novel ones. It is for these reasons that an atlas or guide to the entomopathogenic fungi is considered necessary, not only to highlight and consolidate the many contributions of past my­cologists but also to aid and hopefully to stimulate present and fu­ture workers in the field, many of whom will come from disciplines other than mycology. These fungi have either been included within general reviews of insect pathogens, and hence have received somewhat superficial treatment taxonomically, or conversely, they have been described in mycological monographs essentially over­specialised and unavailable to non-mycologists. This book, there­fore, is an attempt to combine the information relating to the taxo­nomy, ecology and physiology of this important group of organisms in a format readily assimilable by agriculturists, biotechnologists, chemists and entomologists, but which at the same time can serve as an identification guide useful to both practical field scientists and specialist mycologists or pathologists in the laboratory. The only comparable 'popular' book in English solely devoted to

this subject was written by Cooke in 1892 under the evocative title Vegetable wasps and plant worms, and is essentially a mycological treatise but skilfully presented to appeal to a wider audience. We

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hope that our book can go some way towards replacing this classic publication and thereby attain at least some of Cooke's earlier aspi­rations: 'In its present form I hope that the book will be welcome alike to the entomologist and the mycologist, and assist them in their respective studies'. We restrict the term 'entomopathogenic fungi' to those genera or species which are proven pathogens of insects or for which circum­stantial evidence exists concerning their pathogenicity. The looser term 'entomogenous fungi' is avoided here because this also relates to fungi growing on or colonising insect substrates, either faculta­tively or obligately, but which are strictly non-pathogenic. Hence the commensal parasites of insects such as the Laboulbeniales are excluded by definition from the true entomopathogenic fungi and consequently will not be dealt with here. However, it is felt justified in including the obligate pathogens of spiders (araneopathogenic fungi) within any treatment of the entomopat:p.ogenic fungi since many of them are related to those on insects and they exploit a simi­lar ecological niche and hence face the same problems of host pene­tration, colonisation and preservation in addition to sporulation on and dissemination from the host. Moreover, many of the metabolites necessary to achieve these ends are probably common to both groups offungi. Similarly, fungi also attack arthropods other than insects and spiders; some groups, for example, are common pathogens of marine invertebrates. These mycopathogens will not be discussed

here with the exception of the crayfish pathogen Aphanomyces, be­cause of its relevance to recent progress in invertebrate immunology (see Chapter 4). The readers interested in these pathogens should consult Lightner (1981). The fungal genera Beauveria and Metarhizium are well known to the majority of entomologists who would have come across these names in the literature or assigned them to the fungi occurring on their spe­cimens, depending on whether the 'mould' exhibited a white or a

x

green colouration. We wish to demonstrate to those who may be in­volved with any aspect of arthropod population dynamics, be it the ecologist, the practical agriculturist or the pesticide sales executive, that the range and variety of entomopathogenic fungi encompasses considerably wider dimensions than this relatively narrow frame­work upon which most research has been concentrated and that there exist many more genera which have received little or no atten­tion but which conceivably may contain species exploitable by man as sources ofbiocontrol agents or of useful metabolites. At the very least we shall dispel some of the myths surrounding entomopatho­genic fungi, increase the levels of knowledge, and thereby redirect or divert them into the mainstream of mycology: ' ... the study of ento­mogenous fungi being a well known if somewhat esoteric, mycologi­cal by-way' (Ainsworth, 1981).

Acknowledgements We are very grateful to the following individuals who provided data and material for illustrations: Stanizlaw Balazy, Mildred Blackwell, Paul Brey, Jorgen Eilenberg, Brian Federici, Dave Malloch, Dan Molloy, Jacqie Pendland, David Perry, Chris Prior, Clay McCoy, Steve Moss, Bert Orr, J. Pillai, Michiel Rombach, J. P. Skou, Kenneth Soderhiill, George Soares, Tony Sweeney, Christine Tar­rant, Alain Vey, John Webster and Neil Wilding. A special word of thanks should be expressed to Siegfried Keller for his generous loan of the beautiful colour slides of the Entomophthorales. We wish to thank Richard Hall, Drion Boucias and Kenneth Soderhiill for reviewing the manuscripts of some chapters; Mrs. Ans Spaapen - de Veer for invaluable help with the typing, and Francis Snippe-Claus and Ellen Mul for assisting with the drawings and the photography respectively. Finally we are indebted to our families for their sup­port and patience while working on the preparation of the book.

SPRING 1988 THE AUTHORS

Contents

Preface IX

I Introduction I

2 Taxonomy of entomopathogenic fungi 5

3 Illustrations of common fungal pathogens 17

4 Fungal pathogenesis 128

5 Natural control: ecology and biology 140

6 Biotechnology (J. P. Latge and R.Moletta) 152

7 Biological control: past, present and future 165

Bibliography 173

Index 185

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