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Entomology 3rd ed - C.Gillott

Contents


I. Evolution and Diversity
1
Arthropod
Evolution
1. Introduction................................................... 3
2. Arthropod Diversity......................................... 3
2.1. Onychophora, Tardigrada, and Pentastoma....... 4
2.2. Trilobita ................................................. 5
2.3. The Chelicerate Arthropods ......................... 6
2.4. The Mandibulate Arthropods........................ 8
3. Evolutionary Relationships of Arthropods.............. 14
3.1. The Problem............................................ 14
3.2. Theories of Arthropod Evolution................... 15
3.2.1. Mono- and Diphyletic Theories ............ 15

3.2.2. The Polyphyletic Theory .................... 17
3.3. The Uniramians........................................ 20
3.3.1. Myriapoda-Hexapoda Relationships ...... 20
4. Summary...................................................... 21
5. Literature...................................................... 22
2
Insect
Diversity
1. Introduction................................................... 25
2. Primitive Wingless Insects................................. 25
3. Evolution of Winged Insects .............................. 27
3.1. Origin and Evolution of Wings ..................... 27
3.2. Phylogenetic Relationships of the Pterygota ..... 33
3.3. Origin and Functions of the Pupa .................. 44
4. The Success of Insects...................................... 47
4.1. The Adaptability of Insects.......................... 47
4.2. The Importance of Environmental Changes...... 49

5. Summary...................................................... 53
6. Literature...................................................... 53
3
External
Structure
1. Introduction................................................... 57
2. General Body Plan .......................................... 57
3. The Head...................................................... 60
3.1. General Structure...................................... 60
3.2. Head Appendages ..................................... 64
3.2.1. Antennae........................................ 64
3.2.2. Mouthparts ..................................... 64
4. The Neck and Thorax....................................... 72
4.1. The Neck................................................ 73
4.2. Structure of the Thorax............................... 73
4.3. Thoracic Appendages................................. 75
4.3.1. Legs ............................................. 75
4.3.2. Wings ........................................... 79
5. The Abdomen................................................ 83
5.1. General Structure...................................... 83
5.2. Abdominal Appendages.............................. 84
5.2.1. External Genitalia............................. 85
5.2.2. Other Appendages ............................ 88
6. Literature...................................................... 89
4
Systematics
and
Taxonomy
1. Introduction................................................... 91
2. Naming and Describing Insects........................... 92
3. Classification................................................. 94
3.1. The History of Insect Classification ............... 96
4. Identification ................................................. 102
4.1. Key to the Orders of Insects ......................... 103
5. Literature...................................................... 111
5
Apterygote
Hexapods
1. Introduction................................................... 113
2. Collembola ................................................... 114
3. Protura......................................................... 118
4. Diplura ........................................................ 120
5. Microcoryphia ............................................... 122
6. Zygentoma.................................................... 123
6
Paleoptera
1. Introduction................................................... 127
2. Ephemeroptera............................................... 127
3. Odonata ....................................................... 136

7
The
Plecopteroid,
Blattoid, and
Orthopteroid
Orders
1. Introduction................................................... 147
2. Plecoptera..................................................... 147
3. Embioptera ................................................... 153
4. Dictyoptera ................................................... 156
5. Isoptera ........................................................ 163
6. Grylloblattodea .............................................. 173
7. Dermaptera ................................................... 175
8. Phasmida...................................................... 179
9. Mantophasmatodea.......................................... 182
10. Orthoptera .................................................... 184
11. Zoraptera ...................................................... 195
8
The
Hemipteroid
Orders
1. Introduction................................................... 199
2. Psocoptera .................................................... 199
3. Phthiraptera................................................... 203
4. Hemiptera..................................................... 210
5. Thysanoptera ................................................. 233
9
The
Panorpoid
Orders
1. Introduction................................................... 239
2. Mecoptera..................................................... 239
3. Diptera......................................................... 243
4. Siphonaptera.................................................. 264
5. Trichoptera.................................................... 268
6. Lepidoptera................................................... 276
10
The
Remaining
Endopterygote
Orders
1. Introduction................................................... 297
2. Megaloptera .................................................. 297
3. Raphidioptera ................................................ 299
4. Neuroptera .................................................... 301
5. Coleoptera .................................................... 305
6. Strepsiptera ................................................... 326
7. Hymenoptera ................................................. 330
II. Anatomy and Physiology
11
The
Integument
1. Introduction................................................... 355
2. Structure....................................................... 355
3. Cuticle Formation ........................................... 360
3.1. Preecdysis............................................... 360
3.2. Ecdysis .................................................. 363
3.3. Postecdysis ............................................. 363

3.4. Coordination of Events............................... 365
4. Functions of the Integument............................... 366
4.1. Strength and Hardness................................ 366
4.2. Permeability............................................ 367
4.3. Color..................................................... 369
4.4. Other Functions........................................ 370
5. Summary...................................................... 370
6. Literature...................................................... 371
12
Sensory
Systems
1. Introduction................................................... 373
2. Mechanoreception........................................... 374
2.1. Sensory Hairs .......................................... 374
2.2. Proprioceptors.......................................... 375
2.3. Signal Detection....................................... 378
3. Sound Reception............................................. 379
3.1. Johnston’s Organ ...................................... 380
3.2. Tympanal Organs...................................... 380
3.3. Subgenual Organs ..................................... 383
4. Chemoreception ............................................. 384
4.1. Location and Structure of Sensilla ................. 384
4.2. Physiology of Chemoreception..................... 386
5. Humidity Perception........................................ 387
6. Temperature Perception .................................... 388
7. Photoreception ............................................... 389
7.1. Compound Eyes ....................................... 389
7.1.1. Form and Movement Perception........... 392
7.1.2. Distance Perception .......................... 395
7.1.3. Spectral Sensitivity and Color Vision ..... 396
7.1.4. Sensitivity to Polarized Light............... 397
7.2. Simple Eyes ............................................ 398
8. Summary...................................................... 400
9. Literature...................................................... 401
13
Nervous and
Chemical
Integration
1. Introduction................................................... 405
2. Nervous System.............................................. 405
2.1. Central Nervous System ............................. 408
2.2. Visceral Nervous System............................ 411
2.3. Physiology of Neural Integration................... 411
2.4. Learning and Memory................................ 415
3. Endocrine System ........................................... 417
3.1. Neurosecretory Cells and Corpora Cardiaca ..... 417
3.2. Corpora Allata ......................................... 419
3.3. Molt Glands ............................................ 420
3.4. Other Endocrine Structures.......................... 420
4. Insect Semiochemicals ..................................... 421
4.1. Pheromones ............................................ 422

4.1.1. Sex Pheromones............................... 422
4.1.2. Caste-Regulating Pheromones.............. 425
4.1.3. Aggregation Pheromones.................... 425
4.1.4. Alarm Pheromones ........................... 426
4.1.5. Trail-Marking Pheromones ................. 427
4.1.6. Spacing (Epideictic) Pheromones.......... 428
4.2. Kairomones............................................. 429
4.3. Allomones .............................................. 429
5. Environmental, Neural, and Endocrine Interaction ... 430
6. Summary...................................................... 431
7. Literature...................................................... 432
14
Muscles and
Locomotion
1. Introduction................................................... 437
2. Muscles........................................................ 437
2.1. Structure ................................................ 438
2.2. Physiology.............................................. 441
3. Locomotion................................................... 443
3.1. Movement on or Through a Substrate ............. 443
3.1.1. Walking ......................................... 443
3.1.2. Jumping......................................... 447
3.1.3. Crawling and Burrowing .................... 448
3.2. Movement on or Through Water ................... 449
3.2.1. Surface Running .............................. 449
3.2.2. Swimming by Means of Legs............... 450
3.2.3. Swimming by Other Means................. 451
3.3. Flight .................................................... 452
3.3.1. Structural Basis................................ 452
3.3.2. Aerodynamic Considerations ............... 453
3.3.3. Mechanics of Wing Movements............ 458
3.3.4. Control of Wing Movements................ 460
3.3.5. Flight Metabolism............................ 462
3.4. Orientation.............................................. 464
4. Summary...................................................... 465
5. Literature...................................................... 466
15
Gas
Exchange
1. Introduction................................................... 469
2. Organization and Structure of the Tracheal System... 470
2.1. Tracheae and Tracheoles............................. 470
2.2. Spiracles ................................................ 473
3. Movement of Gases within the Tracheal System...... 474
3.1. Diffusion................................................ 475
3.2. Discontinuous Gas Exchange ....................... 476
3.3. Active Ventilation ..................................... 477
4. Gas Exchange in Aquatic Insects......................... 479
4.1. Closed Tracheal Systems ............................ 479
4.2. Open Tracheal Systems .............................. 481

6. Summary...................................................... 485
7. Literature...................................................... 485
16
Food Uptake
and
Utilization
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
5.1.1. Fat Body........................................ 504
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
17
The
Circulatory
System
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
7. Literature...................................................... 534
18
Nitrogenous
Excretion and
Salt and
Water
Balance
1. Introduction................................................... 537
2. Excretory Systems........................................... 537
2.1. Malpighian Tubules—Rectum...................... 537
2.2. Other Excretory Structures .......................... 539
3. Nitrogenous Excretion...................................... 541
3.1. The Nature of Nitrogenous Wastes................. 541
3.2. Physiology of Nitrogenous Excretion ............. 543
3.3. Storage Excretion ..................................... 545
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
Reproduction
1. Introduction................................................... 561
2. Structure and Function of the Reproductive System.. 561
2.1. Female................................................... 532
2.2. Male...................................................... 565
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
3.2. Male...................................................... 579
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

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
Embryonic
Development
1. Introduction................................................... 597
2. Cleavage and Blastoderm Formation .................... 597
3. Formation and Growth of Germ Band................... 598
4. Gastrulation, Somite Formation, and Segmentation .. 602
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
Postembryonic
Development
1. Introduction................................................... 623
2. Growth......................................................... 624
2.1. Physical Aspects....................................... 624
2.2. Biochemical Changes during Growth ............. 626
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
6.2. Factors Initiating and Terminating
Molt Cycles............................................. 643
7. Polymorphism................................................ 645
8. Summary...................................................... 649
9. Literature...................................................... 650
IV. Ecology
22
The Abiotic
Environment
1. Introduction................................................... 655
2. Temperature .................................................. 655
2.1. Effect on Development Rate ........................ 655
2.2. Effect on Activity and Dispersal.................... 657
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
The Biotic
Environment
1. Introduction................................................... 691
2. Food and Trophic Relationships.......................... 691
2.1. Quantitative Aspects.................................. 691
2.2. Qualitative Aspects ................................... 694
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

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
5.2.4. Fungi ............................................ 716
5.2.5. Protozoa ........................................ 717
5.2.6. Nematodes ..................................... 718
6. Summary...................................................... 718
7. Literature...................................................... 720
24
Insects and
Humans
1. Introduction................................................... 725
2. Beneficial Insects ............................................ 726
2.1. Insects Whose Products Are Commercially
Valuable ................................................. 727
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

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