Preface And Acknowledgments For First Edition

Insects are extremely successful animals and they affect many aspects of our lives, despite their small size. All kinds of natural and modified, terrestrial and aquatic, ecosystems support communities of insects that present a bewildering variety of life-styles, forms and functions. Entomology covers not only the classification, evolutionary relationships and natural history of insects, but also how they interact with each other and the environment. The effects of insects on us, our crops and domestic stock, and how insect activities (both deleterious and beneficial) might be modified or controlled, are amongst the concerns of entomologists.

The recent high profile of biodiversity as a scientific issue is leading to increasing interest in insects because of their astonishingly high diversity. Some calculations suggest that the species richness of insects is so great that, to a near approximation, all organisms can be considered to be insects. Students of biodiversity need to be versed in entomology.

We, the authors, are systematic entomologists teaching and researching insect identification, distribution, evolution and ecology. Our study insects belong to two groups - scale insects and midges - and we make no apologies for using these, our favourite organisms, to illustrate some points in this book.

This book is not an identification guide, but addresses entomological issues of a more general nature. We commence with the significance of insects, their internal and external structure, and how they sense their environment, followed by their modes of reproduction and development. Succeeding chapters are based on major themes in insect biology, namely the ecology of ground-dwelling, aquatic and plant-feeding insects, and the behaviours of sociality, predation and parasitism, and defence. Finally, aspects of medical and veterinary entomology and the management of insect pests are considered.

Those to whom this book is addressed, namely students contemplating entomology as a career, or studying insects as a subsidiary to specialized disciplines such as agricultural science, forestry, medicine or veterinary science, ought to know something about insect system-atics - this is the framework for scientific observations. However, we depart from the traditional order-by-order systematic arrangement seen in many entomological textbooks. The systematics of each insect order are presented in a separate section following the ecological-behavioural chapter appropriate to the predominant biology of the order. We have attempted to keep a phylogenetic perspective throughout, and one complete chapter is devoted to insect phylogeny, including examination of the evolution of several key features.

We believe that a picture is worth a thousand words. All illustrations were drawn by Karina Hansen McInnes, who holds an Honours degree in Zoology from the Australian National University, Canberra. We are delighted with her artwork and are grateful for her hours of effort, attention to detail and skill in depicting the essence of the many subjects that are figured in the following pages. Thank you Karina.

This book would still be on the computer without the efforts of John Trueman, who job-shared with Penny in second semester 1992. John delivered invertebrate zoology lectures and ran lab classes while Penny revelled in valuable writing time, free from undergraduate teaching. Aimorn Stewart also assisted Penny by keeping her research activities alive during book preparation and by helping with labelling of figures. Eva Bugledich acted as a library courier and brewed hundreds of cups of coffee.

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The following people generously reviewed one or more chapters for us: Andy Austin, Tom Bellas, Keith Binnington, Ian Clark, Geoff Clarke, Paul Cooper, Kendi Davies, Don Edward, Penny Greenslade, Terry Hillman, Dave McCorquodale, Rod Mahon, Dick Norris, Chris Reid, Steve Shattuck, John Trueman and Phil Weinstein. We also enjoyed many discussions on hymenopteran phylogeny and biology with Andy. Tom sorted out our chemistry and Keith gave expert advice on insect cuticle. Paul's broad knowledge of insect physiology was absolutely invaluable. Penny put us straight with springtail facts. Chris' entomological knowledge, especially on beetles, was a constant source of information. Steve patiently answered our endless questions on ants. Numerous other people read and commented on sections of chapters or provided advice or helpful discussion on particular entomological topics. These people included John Balderson, Mary Carver, Lyn Cook, Jane Elek, Adrian Gibbs, Ken Hill, John Lawrence, Chris Lyal, Patrice Morrow, Dave Rentz, Eric Rumbo, Vivienne Turner, John Vranjic and Tony Watson. Mike Crisp assisted with checking on current host-plant names. Sandra McDougall inspired part of Chapter 15. Thank you everyone for your many comments which we have endeavoured to incorporate as far as possible, for your criticisms which we hope we have answered, and for your encouragement.

We benefited from discussions concerning published and unpublished views on insect phylogeny (and fossils), particularly with Jim Carpenter, Mary Carver, Niels Kristensen, Jarmila Kukalova-Peck and John Trueman. Our views are summarized in the phylogenies shown in this book and do not necessarily reflect a consensus of our discussants' views (this was unattainable).

Our writing was assisted by Commonwealth Scientific and Industrial Research Organization (CSIRO) providing somewhere for both of us to work during the many weekdays, nights and weekends during which this book was prepared. In particular, Penny managed to escape from the distractions of her university position by working in CSIRO. Eventually, however, everyone discovered her whereabouts. The Division of Entomology of the CSIRO provided generous support: Carl Davies gave us driving lessons on the machine that produced reductions of the figures, and Sandy Smith advised us on labelling. The Division of Botany and Zoology of the Australian National University also provided assistance in aspects of the book production: Aimorn Stewart prepared the SEMs from which Fig. 4.7 was drawn, and Judy Robson typed the labels for some of the figures.

Chapter 1


Charles Darwin inspecting beetles collected during the voyage of the Beagle. (After various sources, especially Huxley & Kettlewell 1965 and Futuyma 1986.)

Curiosity alone concerning the identities and lifestyles of the fellow inhabitants of our planet justifies the study of insects. Some of us have used insects as totems and symbols in spiritual life, and we portray them in art and music. If we consider economic factors, the effects of insects are enormous. Few human societies lack honey, provided by bees (or specialized ants). Insects pollinate our crops. Many insects share our houses, agriculture, and food stores. Others live on us, our domestic pets, or our livestock, and yet more visit to feed on us where they may transmit disease. Clearly, we should understand these pervasive animals.

Although there are millions of kinds of insects, we do not know exactly (or even approximately) how many. This ignorance of how many organisms we share our planet with is remarkable considering that astronomers have listed, mapped, and uniquely identified a comparable diversity of galactic objects. Some estimates, which we discuss in detail below, imply that the species richness of insects is so great that, to a near approximation, all organisms can be considered to be insects. Although dominant on land and in freshwater, few insects are found beyond the tidal limit of oceans.

In this opening chapter, we outline the significance of insects and discuss their diversity and classification and their roles in our economic and wider lives. First, we outline the field of entomology and the role of entomologists, and then introduce the ecological functions of insects. Next, we explore insect diversity, and then discuss how we name and classify this immense diversity. Sections follow in which we consider past and some continuing cultural and economic aspects of insects, their aesthetic and tourism appeal, and their importance as foods for humans and animals. We conclude with a review of the conservation significance of insects.


Entomology is the study of insects. Entomologists, the people who study insects, observe, collect, rear, and experiment with insects. Research undertaken by entomologists covers the total range of biological disciplines, including evolution, ecology, behavior, anatomy, physiology, biochemistry, and genetics. The unifying feature is that the study organisms are insects. Biologists work with insects for many reasons: ease of cul-turing in a laboratory, rapid population turnover, and availability of many individuals are important factors. The minimal ethical concerns regarding responsible experimental use of insects, as compared with vertebrates, are a significant consideration.

Modern entomological study commenced in the early 18 th century when a combination of rediscovery of the classical literature, the spread of rationalism, and availability of ground-glass optics made the study of insects acceptable for the thoughtful privately wealthy. Although people working with insects hold professional positions, many aspects of the study of insects remain suitable for the hobbyist. Charles Darwin's initial enthusiasm in natural history was as a collector of beetles (as shown in the vignette for this chapter). All his life he continued to study insect evolution and communicate with amateur entomologists throughout the world. Much of our present understanding of worldwide insect diversity derives from studies of nonprofessionals. Many such contributions come from collectors of attractive insects such as butterflies and beetles, but others with patience and ingenuity continue the tradition of Henri Fabre in observing close-up activities of insects. We can discover much of scientific interest at little expense concerning the natural history of even "well known" insects. The variety of size, structure, and color in insects (see Plates 1.1-1.3, facing p. 14) is striking, whether depicted in drawing, photography, or film.

A popular misperception is that professional entomologists emphasize killing or at least controlling insects, but in fact entomology includes many positive aspects of insects because their benefits to the environment outweigh their harm.

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