This book attempts to summarize what we know about inseet-plant relationships without beeoming too involved with untestable hypo- theses. It is not intended to be eomprehensive and we have deliberately excluded diseussion of aquatie organisms and fungi. Our definition of inseet herbivores is intentionally broad. It includes all inseets whieh feed on plants, although we have emphasized those whieh feed primarily on the photosynthetie tissues. Some referenee is made to seed predation but pollination eeology is excluded. We thank Ors P.H. Smith and M. Luxton for their helpful eommenb on the manuseript but we accept full responsibility for any mistakes whieh may remain. Finally, we thank the various publishers and a uthor~ who gave us permission to use copyright material. 7 I Introduction The net primary produetion of the 300000 speeies of vaseular plant whieh inhabit the dry land surfaee of the earth has been estimated at 9 about 115 x 10 t per annum. This represents a massive resouree potentially available for exploitation by the herbivorous inseets, which themselves probably number in exeess of 500000 species.

INSECTS PROVIDE an ideal medium in which to study all the problems of physiology. But if this medium is to be used to the best advantage, the principles and peculiarities of the insect's organization must be first appreciated. It is the purpose of this book to set forth these principles so far as they are understood at the present day. There exist already many excellent text-books of general ento mology; notably those of Imms, Weber, and Snodgrass, to mention only the more recent. But these authors have necessarily been preoccupied chiefly with describing the diversity of form among insects; discussions on function being correspondingly condensed. In the present work the emphasis is reversed. Struc ture is described only to an extent sufficient to make the physiological argument intelligible. Every anatomical peculiarity, every ecological specialization, has indeed its physiological counterpart. In that sense, anatomy, physiology and ecology are not separable. But regarded from the standpoint from which the present work is written, the endless modifications that are met with among insects are but illustrations of the general principles of their physiology, which it is the aim of this book to set forth. Completeness in such a work is not possible, or desirable; but an endeavour has been made to illustrate each physiological characteristic by a few concrete examples, and to include sufficient references to guide the student to the more important sources. The physiology of insects is to some the handmaid of Economic Entomology."

'A remarkable field guide ... one of those books that come along every now and then and cause a revolution.' Michael McCarthy Independent This concise guide is a companion to the main Field Guide to the Moths of Great Britain and Ireland by the same expert authors, but is in a condensed form with artwork opposite the species descriptions and lay-flat binding for ease of use in the field. It includes brief but comprehensive field descriptions of all the macro-moths in Great Britain, Ireland, the Isle of Man and the Channel Islands, and this second edition has been thoroughly revised and updated to reflect the latest advances in taxonomy. Featuring more than 1,700 superbly detailed colour artworks and covering nearly 900 species, this portable guide is an essential addition to every moth-lover's field kit.

Cockroaches are ideal subjects for laboratory investigation at all educational levels. Compared with many other laboratory animals, cockroaches are easily and inexpensively maintained and cultured and require relatively little space. They are hardy and are readily available. The purpose of this book is to provide background material and experimental leads for utilizing cockroaches in the teaching laboratory and in designing research projects. The level of difficulty of the experiments varies according to the depth of understanding desi red by the instructor. In most cases at least a part of each experiment or technique can be incorporated into the laboratory component of elementary, high school or college curriculum. Sections of the lab book are appropriate for courses in Animal Behavior, Entomology, Organismic Biology and Insect Physiology. Aside from this main purpose, the book also provides a wealth of experimental ideas and techniques for a scientist at any level of education. Lawrence, Kansas June 15, 1981 W. J. B. ACKNOWLEDGEMENTS. Virtually all graduate students who have worked on cockroach research in my laboratory have knowingly or unknowingly contributed to this book. The most important contribution was from Sandy Jones McPeak, who encouraged me to finish the project. Segments of various chapters were conceived, developed or reviewed by Michael D. Breed, Sandy Jones McPeak, Michael K. Rust, Coby Schal, Thomas R. Tobin, W. Alexander Hawkins, Gary R. Sams and Chris Parsons Sams.

This volume deals mainly with the biology of the American cockroach, Periplaneta americana (1.). Contributors were urged to emphasize recent findings, including unpublished data when possible, a goal that would not have been feasible if it were not for the two previously published books on the basic biology of cockroaches, The Biology o/the Cockroach (1968) by D. M. Guthrie and A. R. Tindall and The Cockroach, Volume 1 (1968) by P. B. Cornwell. Those topics not included in The American Cockroach, such as external morphology, are well covered in the two preceding books. In addition, these books provided a broad background upon which contributors to The American Cockroach have been able to build with recent trends, new and established concepts and integration. Although this book deals primarily with the American cockroach, many chapters offer a comparative approach in sections where the more recent and exciting research has been accomplished on other species. Most contributors place the cockroach in perspective with regard to its appropriateness or inappropriateness for various types of biological investigations. Many questions are realistically left unanswered when no acceptable or obvious solution is apparent; an invitation to new researchers to consider the cockroach as an experimental subject.

As editor of this volume I am not going to emphasize the economic im portance of the Bruchids of pulses, nor how necessary it is to know the conditions of infestation of crops in order to achieve an efficient protection; both points are testified by F.A.O.'s sponsorship, as well as of the International Organization for Biological Control. On the other hand I would like to stress the scientific interest of the study of 'domestic' bruchids. It raises questions which require that present basic entomological knowledge be extended. I am not going to review all questions raised but I will emphasize those I am most aware of. Some bruchid species have been able to colonize habitats differing totally; some differing in their latitude, and thus their basic periodicities; others differing in their degree of complexity (stocks of seeds): What are the respective roles played by polymorphism and plasticity in that exceptional capacity? What differences are there between the populations living on wild plants in dispersal areas and those living in stocks? What are the alterations brought about by the drastic selection pressures in populations having later colonized habitats ecologically similar to the original habitats? What factors determine the degree of specificity of trophic relationships, and the possibilities of extension of the niche? What influences are exerted by the other elements of the original bio cenoses upon the behavioural evolution of these species? Such are a few fundamental problems that can be tackled directly through"

Courses on the invertebrates have two principal aims: (1) to introduce students to the diversity of animal life and (2) to make them aware that organisms are marvellously integrated systems with evolutionary pasts and ecological presents. This text is concerned exclusively with the second aim and assumes that the reader will already know something about the diversity and classification of invertebrates. Concepts of whole-organism function, metabolism and adaptation form the core of the subject-matter and this is also considered in an ecological setting. Hence, the approach is multi-disciplinary, drawing from principles normally restricted to comparative morphology and physiology, ecology and evolutionary biology. Invertebrate courses, as with all others in a science curriculum, also have another aim - to make students aware of the general methods of science. And these I take to be associated with the so-calledhypothetico deductive programme. Here, therefore, I make a conscious effort to formulate simple, some might say naive, hypotheses and to confront them with quantitative data from the real world. There are, for example, as many graphs in the book as illustrations of animals. My aim, though, has not been to test out the principles of Darwinism, but rather to sharpen our focus on physiological adaptations, given the assumption that Darwinism is approximately correct. Whether or not I succeed remains for the reader to decide."

It was gratifying to be invited to prepare a third edition of this book, which first appeared in 1951. Preliminary discussions with the publishers, however, revealed a considerable challenge in the present high costs of printing, so that changes and some improvements were clearly necessary to justify the venture. It was immediately apparent that the chapter on chemical control measures would have to be substantially re-written, because of the great changes in usage due to resistance and the regulations introduced to prevent environmental pollution. Also, I decided to expand the scope of the book by increased coverage of the pests of continental Europe and North America, including some new figures and keys in the Appendix. These two undertakings resulted in considerable expansion in length, with about 370 new references and 250 additional specific names in the Index. In order to avoid too alarming an increase in price, I decided to sacrifice three chapters from the earlier editions: those dealing with the structure and classification of insects, their anatomy and physiology, and their ecology. Readers who require basic biological information on insects should buy one of the various short introductions to entomology available.

An analysis of the particular topics in biological control programmes all over the world indicates an increased interest in the utilization of aphid parasites. Besides the so-called traditional biological control of introduced aphids, there appears a situation which could perhaps be called a renaissance of biocontrol, i. e. the utilization of biotic agents against insecticide-resistant populations of aphid pests. The rapid increase in the amount of information as well as the necessity of syntheti cal papers are the well-known features in today's entomology. This requirement is much more topical in the groups where brief summarized information is needed for research workers in applied branches. The author has been well aware of all these difficulties and requirements owing to his own experience both in basic and applied research. Several years ago he decided to summarize our knowledge on the aphid parasites of the world by elaborating synthetical studies on the particular zoogeographical areas. These papers have been intended to represent annotated reviews of the parasite fauna, distribution, biologies and utilization in aphid pest management, with keys to genera and species, hos- parasite catalogue, and a list of references added. Naturally, these studies are only relatively updated, with respect to a certain deadline from which the research may be further continued. At present, the Far East Asian and the Mediterranean areas have been reviewed in this or at least in a similar manner.

This Naturalists' Handbook aims to attract more people to the study of solitary wasps by describing the ecology, distribution and natural history of these insects, including all relevant research in one convenient volume. Contents include an overview of the natural history of the solitary wasp, guidelines on identification, and advice on techniques and approaches to study. Further reading, a systematic checklist of genera and an alphabetical checklist of species and their distributions are included. Detailed keys to the identification of the species form the centre of the book and the text is accompanied by clear illustrations throughout, making this an invaluable practical guide for anyone seeking to broaden their knowledge of these fascinating, diverse creatures. Smaller, gentler and less intimidating than the black and yellow social wasps, the solitary wasps are attractive because of their bright colours and their fascinating behaviour. A female wasp will construct a nest, excavating it from wood or sand or building it from mud. She provisions the nest with prey, hunting down a suitable creature, perhaps a caterpillar or a fly, which she will paralyse before dragging it home to the nest. She lays her egg on the paralysed prey, and the larva when it hatches feeds on the prey. On a sunny day it is easy to observe the apparently purposeful behaviour of female wasps as they prepare their nests and stock them with food for the next generation. This book is a digital reprint of ISBN 0-85546-295-7 (1995). Naturalists' Handbooks encourage and enable those interested in natural history to undertake field study, make accurate identifications and to make original contributions to research.

This title offers a guide to the world of arthropods, covering many insect orders, including beetles, flies, stick insects, dragonflies, ants and wasps, as well as microscopic creatures. It provides a fascinating overview of insects and spiders, including their habitats and classification, all shown in over 195 beautiful photographs and illustrations. All aspects of insect life are covered, such as the way insects defend themselves and how they are able to jump, leap and fly. It describes cryptic coloration, and the way insects can use camouflage to blend into their background and escape attack from predators. It offers various methods of feeding are discussed, from biting and chewing to lapping, sucking, piercing and filter feeding, according to their different mouthparts. It outlines their useful role in pollination of crops, production of honey, and removing insect pests. In the arthropoda phylum, insects are one of the most successful species, and spiders are one of the largest groups. This book studies how they organize their lives. The first section provides information of every aspect of insect life: evolution, anatomy, life cycles, flight and social organization. The last section describes the 30 orders within the class Insecta, demonstrating the huge variety of insects, from microscopic creatures to giant stick insects and large beetles. Typical features of insects in each order are highlighted. With expert text, illustrations and clear photographs, this guide will be enjoyed by all who take an interest in natural history.

The 2nd International Symposium on Trichoptera was held at the University of Reading, England, 25-29 July 1977. It attracted 68 participants from 22 countries, which was a gratifying response to the circulation of about 250 workers on caddis flies. It was H. MALICKY who appreciated the need for a specialized meeting of this kind and organized the 1st International Symposium on Trichoptera, which was held at Lunz am See, Austria, 16-20 September 1974. This volume of Proceedings includes 38 papers; all except one were presented and discussed in the sessions listed in the programme. The papers were given in a lecture theatre of the Palmer Building, and demonstrations were laid out in a laboratory of the Department of Zoology where members met for their mprning and afternoon breaks. Members were accommodated in St Patrick's Hall, one of the University Halls of Residence. They were the guests of the University at an informal reception on 25 July. On the afternoon of 27 July an excursion was made to the River Lambourn at Bagnor near NeWbUry. This chalk stream has been the subject of an ecological study by a team from the Department of Zoology since 1970. The excursion was also an opportunity to see something of the local caddis fauna, and to do some collecting. The final session on 29 July, under the chairmanship of G. B. WIGGINS, was followed by a discussion on future plans."

In his preface to early editions of this book, the late Dr. A. D. Imms said that he intended it to be an elementary account of entomology as a branch of general biology. He had especiaHy in mind the needs of university students of zoology and agriculture, as weH as those intending later to specialize in entomology, and he suggested that the book might also interest teachers of advanced biology in schools. These general aims and the balance between the different aspects of the subject have changed linIe in this and in our previous revision. We have, however, tried to bring the present edition up to date on the lines of our revised tenth edition of Imms' General Textbook 0/ Entomology, published in 1977. The text has been entirely re-set and eleven illustrations have been replaced by new figures. The same orders of insects are recognized as in the last edition, but the sequence in which the Endopterygote groups appear has been changed to reflect more accurately their probable evolutionary relationships. Many small changes and some addi tions have been made in the physiological sections, the chapter on the origin and phylogeny of insects has been rewritten, and a new bibliography provides a selection of modern references for the in tending specialist. It has been our object to make these alterations without materially increasing the length of the book or its level of difficulty."

There has been a large increase in the commercial use of integrated crop/pest management methods for pest and disease control on a wide range of crops throughout the world since the first edition of this book. The completely revised second edition of the bestselling Biological Control in Plant Protection: A Color Handbook continues the objective of providing a handbook with profiles and full-color photographs of as many examples of biological control organisms from as wide a global area as possible. It is designed to help readers anticipate and recognize specific problems of pest management and then resolve them using the natural enemies of pests-parasites, predators, and pathogens. The authors first describe the impact of predator-prey relationships on host plant species in arable, orchard, and protected environments. The main sections of the book include profiles of pests, beneficial arthropods (insects and mites), and beneficial pathogens (bacteria, fungi, viruses, and nematodes), featuring a tabular pest identification guide. Descriptions of biocontrol organisms are divided into four sections: species characteristics, lifecycle, crop/pest associations, and influences of growing practices. The text is illustrated throughout with color photographs of the highest quality. This revised edition helps readers more fully understand the concepts and practice of biological control and integrated pest management. All chapters have been updated and expanded, and more than 300 new photographs have been added. The second edition covers new beneficial organisms and pest profiles, and it includes a new chapter on the practical aspects and application of biological control. It also contains a new final chapter that puts biological control in perspective, discussing interactions that occur when using biocontrol for population management as well as some of the possible mechanisms of biocontrol.

There has been a considerable upsurge in interest in insect bio chemistry and physiology in recent years and this has been reflected in a notable expansion in the number of original papers in this field. Whereas insect physiology has tended to receive ample attention from reviewers, the same has not always been true for the more biochemical aspects of insect research. This book is a venture to help redress the balance. No attempt has been made to cover all aspects of insect biochemistry, but rather a few topics have been selected which seemed to us to merit a review at the present time. One reason for this increased interest in insect biochemistry is perhaps the growing realization that insects can be very useful organisms to act as model systems for the experimental study of general biochemical principles. One remembers, for instance, that Keilin's perceptive observations on the flight muscles of living bees and wax moths led to his discovery of the cytochromes. The fundamental unity of biochemistry has long been accepted as a dogma by the faithful and the insect kingdom provides no exception to it. The main biochemical processes in insects are being revealed as essentially the same as in other life forms but, as so often found in comparative biochemistry, there are interesting variations on the central theme."

seem as appropriate now as the original balance was when Dr A. D. Imms' textbook was first published over fifty years ago. There are 35 new figures, all based on published illustrations, the sources of which are acknowledged in the captions. We are grateful to the authors concerned and also to Miss K. Priest of Messrs Chapman & Hall, who saved us from many errors and omissions, and to Mrs R. G. Davies for substantial help in preparing the bibliographies and checking references. London O.W.R. May 1976 R.G.D. Part I ANATOMY AND PHYSIOLOGY Chapter I INTRODUCTION Definition of the Insecta (Hexapoda) The insects are tracheate arthropods in which the body is divided into head, thorax and abdomen. A single pair of antennae (homologous with the anten nules of the Crustacea) is present and the head also bears a pair of mandibles and two pairs of maxillae, the second pair fused medially to form the labium. The thorax carries three pairs of legs and usually one or two pairs of wings. The abdomen is devoid of ambulatory appendages, and the genital opening is situated near the posterior end of the body. Postembryonic development is rarely direct and a metamorphosis usually occurs."

seem as appropriate now as the original balance was when Dr A. D. Imms' textbook was first published over fifty years ago. There are 35 new figures, all based on published illustrations, the sources of which are acknowledged in the captions. We are grateful to the authors concerned and also to Miss K. Priest of Messrs Chapman & Hall, who saved us from many errors and omissions, and to Mrs R. G. Davies for substantial help in preparing the bibliographies and checking references. O. W. R. London R. G. D. May 1976 Part III THEORDERSOFINSECTS THE CLASSIFICATION AND PHYLOGENY OFINSECTS The classification of insects has passed through many changes and with the growth of detailed knowledge an increasing number of orders has come to be recognized. Handlirsch (1908) and Wilson and Doner (1937) have reviewed the earlier attempts at classification, among which the schemes of Brauer (1885), Sharp (1899) and Borner (1904) did much to define the more distinctive recent orders. In 1908 Handlirsch published a more revolutionary system, incorporating recent and fossil forms, which gave the Collembola, Thysanura and Diplura the status of three independent Arthropodan classes and considered as separate orders such groups as the Sialoidea, Raphidioidea, Heteroptera and Homoptera. He also split up the old order Orthoptera, gave its components ordinal rank and regrouped them with some of the other orders into a subclass Orthopteroidea and another subclass Blattaeformia.

Diseases of insects are important factors reducing the number of insects in natural populations. They are density dependent and they act especially under conditions of outbreaks and close contact of susceptible hosts. They are im portant allies of man in his effort to replace toxic chemicals with non-toxic, biological means in the conservation of the environment of mankind. The importance of diseases as reducing factors has often been disregarded in studies on insect ecology and population dynamics. Most cases of mortality in carefully watched experiments have simply been characterized as "natural mortality," neglecting the fact that there is no "natural" mortality before the insects have finished their life assignment, the deposition of their eggs. The environmental factors including the pathogens are responsible for any other than natural mortality of the larvae, pupae and young adults of the pests. It is rather difficult to distinguish the different symptoms of a disease in insects. Therefore they are not noticed and their influence on populations is overlooked. We feel that a series of photographs of different stages of a collection of insect diseases could help such workers in a general orientation and we are sure from own experience that each specialist needs some broader oriented guide for the fields bordering his own homeland."

entiated free-living organism (larva), which is The success of the Insecta as a class (nearly extensively destroyed and rebuilt into a mor 1 million spp.; phylum Arthropoda) is largely phologically different form (adult) suitable for due to their adaptability to profoundly different ecological niches. Insects have attracted the life in a different ecological niche, is controlled attention of scientists both as useful model by a single genome. This is probably the most systems for the study of many basic biological dramatic reorganization of a growing animal phenomena, and also for the rational develop known. Certain carefully selected insect ment of new methods of controlling the pest material can, thus, provide suitable model species. As a class, insects have played an systems for developmental studies. important role in the elucidation of numerous The majority of the individual metabolic basic biochemical phenomena. For example, reactions occurring in insects are similar to work on the genetic control of eye pigment those found in other groups of organisms."