Entomology. Cedric Gillott University of Saskatchewan Saskatoon, Saskatchewan, Canada. Springer Science + Business Media, LLC

Transcription

1 Entomology

2 Entomology Cedric Gillott University of Saskatchewan Saskatoon, Saskatchewan, Canada Springer Science + Business Media, LLC

3 Gillott, Cedric. Entomology. Library of Congress Cataloging in Publication Data Bibliography: p. Includes index. 1. Entomology. I. Title. QL463.G ISBN ISBN (ebook) DOI / First Printing-September 1980 Second Printing- September 1982 Third Printing-October Springer Science+ Business Media New York Originally published by Plenum Press, New York in 1980 A Division of Plenum Publishing Corporation 233 Spring Street, New York, N.Y All rights reserved No part of this book may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher

4 Preface The idea of writing this book was conceived when, in the late 1960s, I began teaching a senior undergraduate class in general entomology. I soon realized that there was no suitable text for the class I intended to give. The so-called "general" or "introductory" texts reflected the traditional taxonomic approach to entomology and contained relatively little information on the physiology and ecology of insects. This does not mean that there were no books containing such information. There were several, but these were so specialized and detailed that their use in an introductory class was limited. I hold a strong belief that an undergraduate general entomology course should provide a balanced treatment of the subject. Thus, although some time should be devoted to taxonomy, including identification (best done in the laboratory, using primarily material which students themselves have collected, supplemented with specimens from the general collection), appropriate time should be given also to discussion of the evolution, development, physiology, and ecology of insects. In the latter category I include the interactions between insects and Man because it is important to stress that these interactions follow normal ecological principles. Naturally, the format of this book reflects this belief. The book has been arranged in four sections, each of which necessarily overlaps with the others. Section I (Evolution and Diversity) deals with the evolution of the Insecta both in relation to other arthropods (Chapter 1) and in terms of the individual orders within the class (Chapter 2). Chapter 3 serves two purposes: it provides a description of external structure, which remains the basis on which insects can be classified and identified, while stressing diversity by reference to mouthpart and limb modifications. In Chapter 4 the principles of classification and identification are discussed, and a key to the orders of insects provided. Diversity of form and habits is again emphasized in Chapters 5 to 10, which deal with the orders of insects. Section II (Anatomy and Physiology) deals with the homeostatic systems of insects; that is, those systems which serve to keep an insect "in tune" with its environment. The goal of these physiological systems is to enable insects to grow and reproduce optimally. The section begins with a discussion of the integument (Chapter 11), since this has had such a profound effect on the success of insects. Chapter 12 examines sensory systems, whose form and function are greatly influenced by the cuticular nature of the integument. In Chapter v

5 vi PREFACE 13 neural and chemical integration are discussed, and this is followed, in Chapter 14, by a discussion of muscle structure and function, including locomotion. Chapter 15 reveals the remarkable efficiency of the tracheal system in gaseous exchange. Chapter 16 deals with the acquisition and utilization of food, Chapter 17 with the structure and functions of the circulatory system, and Chapter 18 with nitrogenous waste removal and salt/water balance. In Section III are discussed reproduction (Chapter 19), embryonic development (Chapter 20), and postembryonic development (Chapter 21). The final section of the book (Ecology) examines those factors which affect the distribution and abundance of insects. In Chapter 22 abiotic (physical) factors in an insect's environment are discussed. Chapter 23 deals with biotic factors that influence insect populations and serves as a basis for the final chapter in which the specific interactions of insects and Man are discussed. As can be inferred from the opening paragraph of this Preface, the book is intended as a text for senior undergraduates taking their first class in entomology. Such students probably will have an elementary knowledge of insects, gleaned from an earlier class in general zoology, as well as a basic understanding of animal physiology and ecological principles. With such a background, students should have no difficulty in understanding the present text. On occasions, several of my colleagues have questioned the wisdom (ability?) of a single author attempting to cover the entire field of entomology; the more outspoken ones have plainly told me to "Get help!" I have resisted this advice, partly, I suppose, through selfishness, but mainly on the grounds that multiauthored treatises tend to lack continuity. However, to guard against errors of fact and interpretation resulting from my own ignorance, I have had each chapter reviewed by a specialist. During preparation of the book, there were moments when I wondered whether the finished product would ever see the light of day or, indeed, whether I should have undertaken the task in the first place! Now that the book is finished, I can truly say that its preparation has been a rewarding and pleasant experience. My hope is that its readers will now receive the same pleasure. April 1979 Cedric Gillott

6 Acknowledgments Though this book has single authorship, its preparation would not have been possible but for the contributions of numerous individuals and organizations, to whom I am most grateful. These include Miss Shirley Shepstone, who prepared all the original drawings and who redrew a large number of figures from their original sources; Mr. David Wong, Mr. Dennis Dyck, and Mr. Alex Campbell, who photocopied numerous figures; Mrs. Evelyn Peters, who typed the bulk of the manuscript, as well as Mrs. Denise Nowoselski, Mrs. Peggy Baird, and Mrs. Joan Ryan for their smaller, but nevertheless important, typographical contributions. I also thank the following individuals who reviewed specific chapters: Dr. G. G. E. Scudder (Chapters 1-3, 8), Dr. D. M. Lehmkuhl (Chapters 4, 6, 9, 22-24), Dr. D. K. McE. Kevan (Chapters 5, 7), Dr. G. E. Ball (Chapter 10), Dr. M. Locke (Chapter 11), Dr. R. Y. Zacharuk (Chapters 12, 13), Dr. P. J. Mill (Chapters 14,15), Dr. J. E. Steele (Chapter 16), Dr. J. C. Jones (Chapter 17), Dr. J. E. Phillips (Chapter 18), Dr. K. G. Davey (Chapter 19), and Dr. B. S. Heming (Chapters 20, 21). An especial debt of gratitude is owed to Dr. Scudder, who most willingly served also as general reviewer for the entire manuscript. Thanks are also extended to the large number of publishers, editors, and private individuals who allowed me to use material for which they hold copyright. In particular, I thank Mr. Charles S. Papp, who generously supplied me with negatives for a large number of his drawings. The source of each figure is acknowledged individually in the text. I am also grateful to Professor E. J. W. Barrington, former Head, Zoology Department, The University of Nottingham, for provision of facilities in his department, where preparation of this book began, during tenure of a sabbatical leave; and to the Plenum Publishing Corporation, especially Mr. Kirk Jensen (Editor), Mr. John Matzka (Managing Editor), and Mr. Geoffrey Braine (Production Editor), for their patience and assistance in seeing this project through to completion. And finally, the unceasing encouragement, assistance, and patience of my wife, Anne, must be acknowledged. To her fell such jobs as translating my hieroglyphics into a first typewritten draft, proofreading at all stages of the book's progress, and a variety of clerical work. It is to her that this book is dedicated. vii

7 Contents I. Evolution and Diversity 1 Arthropod Evolution 1. Introduction Arthropod Diversity Onychophora Trilobita The Chelicerate Arthropods The Mandibulate Arthropods Evolutionary Relationships of Arthropods The Problem Theories of Arthropod Evolution Mono- and Diphyletic Theories The Polyphyletic Theory Relationships within the Uniramia Summary Literature Insect Diversity 1. Introduction Monura, Microcoryphia, and Zygentoma Evolution of Winged Insects Origin and Evolution of Wings Phylogenetic Relationships of the Pterygota Origin and Function of the Pupa The Success of Insects The Adaptability of Insects The Importance of Environmental Changes Summary Literature ix

8 x 3 External Structure 1. Introduction General Body Plan The Head General Structure Head Appendages Antennae Mouthparts The Neck and Thorax The Neck Structure of the Thorax Thoracic Appendages Legs Wings The Abdomen General Structure Abdominal Appendages External Genitalia Other Appendages Literature Classification and Identification 5 Apterygote Hexapods 6 Paleoptera 7 The Orthopteroid Orders 1. Introduction Classification The History of Insect Classification Identification Key to the Orders of Insects Literature Introduction Collembola Protura Diplura Microcoryphia Zygentoma Introduction Ephemeroptera Odonata Introduction Plecoptera Embioptera Dictyoptera Isoptera Grylloblattodea

9 7. Dermaptera xi 8. Phasmida Orthoptera Zoraptera The Hemipteroid Orders 9 The Panorpoid Orders 10 The Remaining Endopterygote Orders 1. Introduction Psocoptera Phthiraptera Hemiptera Thysanoptera Introduction Mecoptera Trichoptera Lepidoptera Diptera Siphonaptera Introduction Megaloptera Raphidioptera Neuroptera Coleoptera Strepsiptera Hymenoptera II. Anatomy and Physiology 11 The Integument 1. Introduction Structure Cuticle Formation Preecdysis Ecdysis Postecdysis Coordination of Events Functions of the Integument Strength and Hardness Permeability Color Summary Literature

10 xii Sensory Systems 3. Introduction. ' Mechanoreception Sensory Hairs Proprioceptors Sound Reception Johnston's Organ Tympanal Organs Subgenual Organs Chemoreception Location and Structure of Sensilla Physiology of Chemoreception Humidity Perception Temperature Perception Photoreception Compound Eyes Form and Movement Perception Distance Perception Spectral Sensitivity and Color Vision Sensitivity to Polarized Light Simple Eyes Summary Literature Introduction Nervous System Nervous and 2.1. Central Nervous System Chemical Integration 2.2. Visceral Nervous System Physiology of Neural Integration Learning and Memory Endocrine System Neurosecretory Cells and Corpora Cardiaca Corpora Allata Molt Glands Other Supposed Endocrine Structures Pheromones Sex Pheromones Caste-Regulating Pheromones Aggregation Pheromones Alarm Pheromones Trail-Marking Pheromones Environmental,\Jeural, and Endocrine Interaction Summary Literature

11 14 Muscles and Locomotion 15 Gaseous Exchange 16 Food Uptake and Utilization 1. Introduction xiii 2. Muscles Structure Physiology Locomotion Movement on or through the Substrate Walking Jumping Crawling and Burrowing Movement on or through Water Surface Running Swimming by Means of Legs Swimming by Other Means Flight Structural Basis Aerodynamic Considerations Mechanics of Wing Movements Control of Wing Movements Orientation Summary Literature Introduction Organization and Structure of Tracheal System Tracheae and Tracheoles Spiracles Movement of Gases within Tracheal System Diffusion Passive (Suction) Ventilation Active Ventilation Gaseous Exchange in Aquatic Insects Closed Tracheal Systems Open Tracheal Systems Gaseous Exchange in Endoparasitic Insects Summary Literature Introduction Food Selection and Feeding The Alimentary System Salivary Glands Foregut Midgut Hindgut

12 xiv 4. Gut Physiology Gut Movements Digestion Digestive Enzymes Factors Affecting Enzyme Activity Control of Enzyme Synthesis and Secretion Digestion by Microorganisms Absorption Metabolism Sites of Metabolism Fat Body Carbohydrate Metabolism Lipid Metabolism Amino Acid and Protein Metabolism Metabolism of Insecticides Summary Literature The Circulatory System 18 Nitrogenous Excretion and Salt and Water Balance 1. Introduction Structure Physiology Circulation Heartbeat Hemolymph Plasma Composition Functions Hemocytes Number and Form Functions Resistance to Disease Wound Healing Immunity Summary Literature Introduction Excretory Systems Malpighian Tubules-Rectum Other Excretory Structures Nitrogenous Excretion The Nature of Nitrogenous Wastes Physiology of Nitrogenous Excretion Storage Excretion Salt and Water Balance Terrestrial Insects Brackish-Water and Saltwater Insects

13 4.3. Freshwater Insects xv 4.4. Hormonal Control Summary Literature III. Reproduction and Development 1. Introduction..., Structure and Function of the Reproductive Reproduction System Female Male Sexual Maturation Female Vitellogenesis Vitelline Membrane and Chorion Formation Factors Affecting Sexual Maturity in the Female Male Mating Behavior Mate Location and Recognition Courtship Copulation Insemination Postcopulatory Behavior Entry of Sperm into the Eggs and Fertilization Oviposition Site Selection Mechanics and Control of Oviposition Oothecae Summary Literature Embryonic Development Introduction Cleavage and Blastoderm Formation Formation and Growth of Germ Band Gastrulation, Somite Formation, and Segmentation Formation of Extraembryonic Membranes.., Dorsal Closure and Katatrepsis Tissue and Organ Development Appendages Integument and Ectodermal Invaginations Central Nervous System Gut and Derivatives

14 xvi 7.5. Circulatory System, Muscle, and Fat Body Reproductive System Special Forms of Embryonic Development Parthenogenesis Polyembryony Viviparity Paedogenesis Factors Affecting Embryonic Development " Hatching Summary Literature Postembryonic Development 1. Introduction Growth Physical Aspects Biochemical Changes during Growth Forms of Development Ametabolous Development Hemimetabolous Development Holometabolous Development The Larval Stage Heteromorphosis The Pupal Stage Histological Changes during Metamorphosis Exopterygote Metamorphosis Endopterygote Metamorphosis Adult Emergence Control of Development Endocrine Regulation of Development Factors Initiating and Terminating Molt Cycles Polymorphism Summary Literature IV. Ecology 22 The Abiotic Environment 1. Introduction., Temperature., Effect on Developmental Rate Effect on Activity and Dispersal Temperature-Synchronized Development and Emergence Survival at Extreme Temperatures Cold-Hardiness

15 3. Light Daily Influences of Photoperiod Circadian Rhythms Seasonal Influences of Photoperiod Nature and Rate of Development Reproductive Ability and Capacity Diapause Water Terrestrial Insects Aquatic Insects Wind Migration Summary Literature xvii 23 The Biotic Environment 24 Insects and Man 1. Introduction Food Quantitative Aspects Qualitative Aspects Insect-Plant Interactions Interactions between Insects and Other Animals Intraspecific Interactions Underpopulation Overpopulation Interspecific Interactions Competition and Coexistence Predator-Prey Relationships Insect Diseases Epizootics Types of Pathogens Bacteria Rickettsias Viruses Fungi Protozoa Nematodes Summary 6. Literature Introduction Beneficial Insects Insects Whose Products are Commercially Valuable Insects as Pollinators Insects as Agents of Biological Control Insects as Food

16 xviii 2.5. Soil-Dwelling and Scavenging Insects Other Benefits of Insects Pest Insects Insects Affecting Man Directly Pests of Domesticated Animals Pests of Cultivated Plants Insect Pests of Stored Products Pest Control Legal Control Chemical Control Biological Control Genetic Control Cultural Control Integrated Control Summary Literature Author Index Subject Index