Atlas of Entomopathogenic Fungi Cordyceps tuberculata on moth (Sphingidae), Hirsutella versicolor on leafhopper (Cicadellidae), Hirsutella jonesisii on green planthopper Indonesia. Kenya. (Nephotettix), Indonesia. Erynia neoaphidis on Aphis (Aphididae), France. Conidiobolus obscurus on Acyrthosiphon pisum Entomophthora muscae on fly (Diptera), Denmark. (Aphididae), France. Erynia radicans on Dicyphus pallidus (Miridae), Switzerland. S porodiniel/a umbel/ata on cicada nymph Erynia gammae on lepidoptera larva (N octuidae), (Cicadidae), Papua New Guinea. Switzerland. Erynia eulieis on Coddis fly (Diptera), UK. Erynia eoniea on Simulium sp. (Diptera), UK. Nomuraea rileyi on rice looper (Lepidoptera), Indonesia. Fusarium eoeeophilum on pine-needle scales Cordyeeps variabilis on Coleoptera larva Cordyeeps unilateralis on Camponotus ant, Costa (Phenaeaspis), Honduras. (Elateridae), Canada. Rica. Entomophaga aulieae on lepidoptera larva, Canada. Cordyeeps sp. on adult beetle (Coleoptera), Brazil. 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), Metarhizium anisopliae var. majus on third instar Hymenostilbe sp. on Gryllidae, Ecuador. Indonesia. larvae of Oryctes rhinoceros, Philippines. Cordyceps locustiphila on Locustidae, Ecuador. Cordyceps sp. on Psychidae larva, Brazil. Erynia delphacis on green leafhopper (Nephotettix virescens), Indonesia. Cordyceps sp. oil cockchafer larva (Scarabidae), Tolypocladium cylindrosporum on mosquito larva Brazil. (Aedes aegyptii), USA. Gibellula species on spider (Salticidae), Brazil. Polycephalomyces ramosus on fly, the Netherlands. Cordyceps curculionum on weevil adult (Curculionidae), Brazil. Aschersonia cubensis on scale insect (Lecaniidae), Aschersonia aleyrodis on citrus whiteflies Gibellula pulchra on spider (Salticidae), Ecuador. Brazil. (Aleyrodiidae) USA. 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 IX Preface A knowledge ofi nsect diseases (whether or not it has to do with control) is off unda mental andfar-reaching importance in the study ofi nsect ecology. E. A. Steinhaus (1949). As biological control is becoming more acceptable as a practical hope that our book can go some way towards replacing this classic science and the dangers of the long-term use of chemical pesticides publication and thereby attain at least some of Cooke's earlier aspi are fully appreciated, there has been a resurgence of interest in em rations: 'In its present form I hope that the book will be welcome ploying fungal pathogens to combat insect pests. New production alike to the entomologist and the mycologist, and assist them in their and application techniques combined with a greater understanding respective studies'. of both fungal and insect ecology have meant that biological insecti We restrict the term 'entomopathogenic fungi' to those genera or cides can now compete on a more equal footing with traditional species which are proven pathogens of insects or for which circum chemical pesticides. Consequently, agrochemical companies are be stantial evidence exists concerning their pathogenicity. The looser ginning to take account of entomopathogens as both viable and eco term 'entomogenous fungi' is avoided here because this also relates nomic propositions and to invest more resources in their research to fungi growing on or colonising insect substrates, either faculta and development, particularly to improve the formulations of well tively or obligately, but which are strictly non-pathogenic. Hence documented pathogens, possibly by genetic manipulation, and to the commensal parasites of insects such as the Laboulbeniales are test more obscure or novel ones. It is for these reasons that an atlas excluded by definition from the true entomopathogenic fungi and or guide to the entomopathogenic fungi is considered necessary, not consequently will not be dealt with here. However, it is felt justified only to highlight and consolidate the many contributions of past my in including the obligate pathogens of spiders (araneopathogenic cologists but also to aid and hopefully to stimulate present and fu fungi) within any treatment of the entomopat:p.ogenic fungi since ture workers in the field, many of whom will come from disciplines many of them are related to those on insects and they exploit a simi other than mycology. These fungi have either been included within lar ecological niche and hence face the same problems of host pene general reviews of insect pathogens, and hence have received tration, colonisation and preservation in addition to sporulation on somewhat superficial treatment taxonomically, or conversely, they and dissemination from the host. Moreover, many of the metabolites have been described in mycological monographs essentially over necessary to achieve these ends are probably common to both groups specialised and unavailable to non-mycologists. This book, there offungi. Similarly, fungi also attack arthropods other than insects fore, is an attempt to combine the information relating to the taxo and spiders; some groups, for example, are common pathogens of nomy, ecology and physiology of this important group of organisms marine invertebrates. These mycopathogens will not be discussed in a format readily assimilable by agriculturists, biotechnologists, here with the exception of the crayfish pathogen Aphanomyces, be chemists and entomologists, but which at the same time can serve as cause of its relevance to recent progress in invertebrate immunology an identification guide useful to both practical field scientists and (see Chapter 4). The readers interested in these pathogens should specialist mycologists or pathologists in the laboratory. consult Lightner (1981). The only comparable 'popular' book in English solely devoted to The fungal genera Beauveria and Metarhizium are well known to the this subject was written by Cooke in 1892 under the evocative title majority of entomologists who would have come across these names Vegetable wasps and plant worms, and is essentially a mycological in the literature or assigned them to the fungi occurring on their spe treatise but skilfully presented to appeal to a wider audience. We cimens, depending on whether the 'mould' exhibited a white or a x green colouration. We wish to demonstrate to those who may be in volved 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 frame work upon which most research has been concentrated and that there exist many more genera which have received little or no atten tion 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 entomopatho genic fungi, increase the levels of knowledge, and thereby redirect or divert them into the mainstream of mycology: ' ... the study of ento mogenous fungi being a well known if somewhat esoteric, mycologi cal 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 Tar rant, 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 sup port and patience while working on the preparation of the book. SPRING 1988 THE AUTHORS