Circulatory System

5. Gas Exchange in Endoparasitic Insects 484

6. Summary 485

7. Literature 485

1. Introduction 487

2. Food Selection and Feeding 487

3. The Alimentary System 489

3.1. Salivary Glands 489

3.2. Foregut 491

3.3. Midgut 492

3.4. Hindgut 496

4. Gut Physiology 496

4.1. Gut Movements 496

4.2. Digestion 498

4.2.1. Digestive Enzymes 498

4.2.2. Factors Affecting Enzyme Activity 499

4.2.3. Control of Enzyme Synthesis and

Secretion 500

4.2.4. Digestion by Microorganisms 501

4.3. Absorption 502

5. Metabolism 503

5.1. Sites of Metabolism 503

5.1.2. Mycetocytes 504

5.2. Carbohydrate Metabolism 505

5.3. Lipid Metabolism 506

5.4. Amino Acid and Protein Metabolism 506

5.5. Metabolism of Insecticides 507

6. Summary 509

7. Literature 510

1. Introduction 515

2. Structure 515

3. Physiology 519

3.1. Circulation 519

3.2. Heartbeat 520

4. Hemolymph 521

4.1. Plasma 522

4.1.1. Composition 522

4.1.2. Functions 524

4.2. Hemocytes 524

4.2.1. Origin, Number, and Form 524

4.2.2. Functions 526

5. Resistance to Disease 530

5.1. Wound Healing 530

5.2. Immunity 530

5.2.1. Resistance to Host Immunity 532

6. Summary 533 xi

7. Literature 534


18 1. Introduction 537

2. Excretory Systems 537

Nitrogenous 2.1. Malpighian Tubules—Rectum 537

. 2.2. Other Excretory Structures 539

Excretionand 3. Nitrogenous Excretion 541

Salt and 3.1. The Nature of Nitrogenous Wastes 541

Water 3.2. Physiology of Nitrogenous Excretion 543

3.3. Storage Excretion 545

Balance 4. Salt and Water Balance 546

4.1. Terrestrial Insects 546

4.2. Freshwater Insects 550

4.3. Brackish-Water and Saltwater Insects 551

5. Hormonal Control 554

6. Summary 556

7. Literature 556

III. Reproduction and Development

19 1- Introduction 561

2. Structure and Function of the Reproductive System.. 561

Reproduction 2-1- Female 532

3. Sexual Maturation 568

3.1. Female 568

3.1.1. Vitellogenesis 569

3.1.2. Vitelline Membrane and Chorion Formation 570

3.1.3. Factors Affecting Sexual Maturity in the Female 572

4. Mating Behavior 581

4.1. Mate Location and Recognition 581

4.2. Courtship 582

4.3. Copulation 583

4.3.1. Insemination 584

4.4. Postcopulatory Behavior 586

5. Ovulation 587

6. Sperm Use, Entry into the Egg, and Fertilization 587

6.1. Sperm Use 587

6.2. Sperm Entry into the Eggs 588

6.3. Fertilization 588

xii 7. Oviposition 589

7.1. Site Selection 589


7.2. Mechanics and Control of Oviposition 590

7.3. Oothecae 590

8. Summary 591

9. Literature 593

20 1. Introduction 597

2. Cleavage and Blastoderm Formation 597

Embryonic 3. Formation and Growth of Germ Band 598

4. Gastrulation, Somite Formation, and Segmentation.. 602

Development 5. Formation of Extra-Embryonic Membranes 605

6. Dorsal Closure and Katatrepsis 606

7. Tissue and Organ Development 607

7.1. Appendages 607

7.2. Integument and Ectodermal Derivatives 608

7.3. Central Nervous System 609

7.4. Gut and Derivatives 611

7.5. Circulatory System, Muscle, and Fat Body 611

7.6. Reproductive System 612

8. Special Forms of Embryonic Development 612

8.1. Parthenogenesis 613

8.2. Polyembryony 614

8.3. Viviparity 614

8.4. Paedogenesis 617

9. Factors Affecting Embryonic Development 617

10. Hatching 619

11. Summary 619

12. Literature 621

21 1. Introduction 623

2. Growth 624

Postembryonic Physical Aspects 624

2.2. Biochemical Changes during Growth 626

Development 3. Forms of Development 627

3.1. Ametabolous Development 628

3.2. Hemimetabolous Development 628

3.3. Holometabolous Development 628

3.3.1. The Larval Stage 630

3.3.2. Heteromorphosis 631

3.3.3. The Pupal Stage 631

4. Histological Changes During Metamorphosis 634

4.1. Exopterygote Metamorphosis 634

4.2. Endopterygote Metamorphosis 634

5. Eclosion 639

6. Control of Development 639

6.1. Endocrine Regulation of Development 640 xiii

6.2. Factors Initiating and Terminating

Molt Cycles 643 Contents

7. Polymorphism 645

8. Summary 649

9. Literature 650

IV. Ecology

22 1. Introduction 655

2. Temperature 655

The Abiotic 2.1. Effect on Development Rate 655

2.2. Effect on Activity and Dispersal 657

nvironment 2.3. Temperature-Synchronized Development and

Emergence 658

2.4. Survival at Extreme Temperatures 659

2.4.1. Cold-Hardiness 659

3. Light 662

3.1. Daily Influences of Photoperiod 662

3.1.1. Circadian Rhythms 663

3.2. Seasonal Influences of Photoperiod 666

3.2.1. Nature and Rate of Development 667

3.2.2. Reproductive Ability and Capacity 668

3.2.3. Diapause 668

4. Water 674

4.1. Terrestrial Insects 674

4.2. Aquatic Insects 677

5. Weather 678

5.1. Weather and Insect Abundance 678

5.2. Migration 679

5.2.1. Categories of Migration 681

6. Summary 686

7. Literature 688

23 1. Introduction 691

2. Food and Trophic Relationships 691

The Biotic 2.1. Quantitative Aspects 691

, . 2.2. Qualitative Aspects 694

nvironment 3. Insect-Plant Interactions 694

3.1. Herbivores 694

3.2. Insect-Plant Mutualism 697

3.3. Detritivores 701

4. Interactions between Insects and Other Animals 702

4.1. Intraspecific Interactions 702

4.1.1. Underpopulation 702

4.1.2. Overpopulation 703

xiv 4.2. Interspecific Interactions 705

4.2.1. Competition and Coexistence 705


4.2.2. Predator-Prey Relationships 709

4.2.3. Insect-Insect Mutualisms 711

5. Insect Diseases 711

5.1. Epizootics 712

5.2. Types of Pathogens 713

5.2.1. Bacteria 713

5.2.2. Rickettsias 715

5.2.3. Viruses 715

5.2.5. Protozoa 717

5.2.6. Nematodes 718

6. Summary 718

7. Literature 720

24 1. Introduction 725

2. Beneficial Insects 726

Insects and 2.1. Insects Whose Products Are Commercially

Valuable 727

Humans 2.2. Insects as Pollinators 728

2.3. Insects as Agents of Biological Control 728

2.4. Insects as Human Food 731

2.5. Soil-Dwelling and Scavenging Insects 733

2.6. Other Benefits of Insects 735

3. Pest Insects 736

3.1. Insects That Affect Humans Directly 736

3.2. Pests of Domesticated Animals 740

3.3. Pests of Cultivated Plants 740

3.4. Insect Pests of Stored Products 743

4. Pest Control 743

4.1. Legal Control 746

4.2. Chemical Control 746

4.3. Biological Control 753

4.3.1. Microbial Control 757

4.4. Genetic Control 766

4.5. Cultural Control 769

4.6. Integrated Pest Management 770

5. Summary 775

6. Literature 776

Index 783


The strongly favorable reception accorded previous versions of this book, together with the not infrequent urgings of colleagues and students, encouraged me to take on the task of preparing a third edition of Entomology. My early retirement, in 1999, freed up the time necessary for a project of this size, and for the past 2 years my effort has been almost entirely focused in this direction. Obviously, all chapters have been updated; this includes not only the addition of new information and concepts (some of which are highlighted below), but also the reduction or exclusion of material no longer considered 'mainstream' so as to keep the book at a reasonable size.

My strong belief that an introductory entomology course should present a balanced treatment of the subject still holds and is reflected in the retention of the format of earlier editions, namely, arrangement of the book into four sections: Evolution and Diversity, Anatomy and Physiology, Reproduction and Development, and Ecology.

Section I (Evolution and Diversity) has again undergone a great reworking, mainly because the last decade has seen the uncovering of significant new fossil evidence, and the application of molecular and cladistic analyses to extant groups. As a result, ideas both on the relationships of insects to other arthropods and on the higher classification of many orders have changed drastically. However, as in previous editions, I have stressed that most phylogenies are not 'embedded in stone' but represent the consensus based on existing information; thus, they are liable to refinement as additional data are forthcoming. Chapter 1 discusses the evolution of Insecta in relation to other arthropods, emphasizing the ageless debate on whether arthropods form a monophyletic or polyphyletic group, and the relationship of insects to other hexapodous arthropods. Evolutionary relationships within the Insecta are considered in Chapter 2, together with discussion of the factors that contributed to the overwhelming success of the group. Chapter 3 serves two purposes: It provides a description of external structure, which remains the principal basis on which insects can be classified and identified, while stressing diversity with reference to mouthpart and appendage modifications. In Chapter 4 the principles of classification and identification are outlined, and a key to the orders of insects is provided. Diversity of form and habits is again emphasized in Chapters 5 to 10, which deal with the orders of insects, including the Mantophasmatodea, established only in 2002. For many orders, new proposed phylogenies are presented, and the text has undergone significant rearrangement to reflect modern ideas on the classification of these taxa.

xvi The chapters in Section II (Anatomy and Physiology) deal with the homeostatic systems of insects; that is, those systems that keep insects 'in tune' with their environment, enabling

Preface them to develop and reproduce optimally. The section begins with a discussion of the integument (Chapter 11), as this has had such a profound influence 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 13, where neural and chemical integration are discussed, new sections on kairomones and allomones have been included. Chapter 14 considers muscle structure and function, including locomotion. In this chapter the section on flight has been significantly revised, especially with respect to recent proposals for the generation of lift using non-steady-state aerodynamics. Chapter 15 reveals the remarkable efficiency of the tracheal system in gaseous exchange, and Chapter 16 deals with the acquisition and utilization of food. Chapter 17 describes the structure and functions of the circulatory system, including the immune response of insects about which much has been learned in the past decade. New to this chapter is a section on how parasites and parasitoids are able to defend themselves against the host insect's immune system. Chapter 18 concludes this section with a discussion of nitrogenous waste removal and salt/water balance.

In Section III reproduction (Chapter 19), embryonic development (Chapter 20), and postembryonic development (Chapter 21) are discussed. Chapter 19 includes additional information on behavioral aspects of reproduction (courtship, mate guarding and sexual selection), as well as sperm precedence. Chapter 21 has been revised to provide an updated account of the endocrine regulation of development and molting.

Section IV (Ecology) examines those factors that affect the distribution and abundance of insects. In Chapter 22 abiotic (physical) factors in an insect's environment are considered. Chapter 23 deals with the biotic factors that influence insect populations and serves as a basis for the final chapter, in which the specific interactions of insects and humans are discussed. Of all of the chapters, Chapter 24 has received the most drastic overhaul; such has been the 'progress' (and the costs of such progress) in the battle against insect pests.

As may be inferred from the opening paragraph of this Preface, the book is intended as a text for senior undergraduates taking their first course in entomology. Such students probably will have an elementary knowledge of insects acquired from an earlier course in general zoology, as well as a basic understanding of animal physiology and ecological principles. With such a background, students should have no difficulty understanding the text.

Preparation of the third edition has benefited, not only from both published and unsolicited reviews of previous editions, but also from my solicitation of comments on the content of specific chapters from experts in those areas. Of course, any errors that remain, and I hope these are extremely few, are my responsibility. I have enjoyed preparing this third edition, for it has given me, once again, the opportunity to delve into aspects of entomology that are well outside the range of an 'insect sexologist'. For example, I never cease to be impressed by the remarkable discoveries and insights of those entomologists who deal with fossil insects, by those who develop integrated strategies for the management of insect pest populations, and by the patience and dedication (and imagination—see Chapter 4, Section 2) of insect taxonomists. Hopefully, readers of the new edition will receive the same enjoyment.

Cedric Gillott Professor Emeritus

University of Saskatchewan, Saskatoon, Saskatchewan, Canada


Though the book has single authorship, its preparation would not have been possible but for my colleagues, too numerous to mention individually, who provided information and answered specific questions that improved the book's content and currency. To these people I am most grateful.

Mr. Dennis Dyck and Mrs. Shirley Brodsky are thanked for their considerable assistance with preparation of the original figures. For this edition, all figures were converted into electronic format and, when necessary, reworked by Mr. Dyck to achieve greater uniformity of style.

Thanks are also extended to a large number of publishers, editors, and private individuals who allowed me to use materials for which they hold copyright. The source of each figure is acknowledged individually in the text.

I am grateful to the University of Saskatchewan, which granted me the facilities necessary to bring this project to fruition. I specifically acknowledge the assistance given by staff in the Library's inter-library loans department; so numerous were my requests for material that I felt, at times, as though they were my personal assistants! The confidence, patience, and assistance of Kluwer Academic Publishers, especially Zuzana Bernhart (Publishing Editor, Life Sciences), Ineke Ravesloot (Assistant to the Publishing Editor), and Tonny van Eekelen (Production Supervisor, Books) are also appreciated.

Finally, the enormous help given me by my wife, Anne, is acknowledged. To her fell the major task of proofreading to ensure that the revised text was coherent, figures were correctly numbered, labeled and cited, reference lists were accurate, and tables were complete. She also checked copyright approvals and assisted in preparation of the index. It is to her that this book is dedicated.

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