entomology 3rd ed - c.gillott

Entomolog y Entomology Th ir d E d i t i o n Entomo l og y Entomo l og y Th ir d E d i t i on C e d ric Gi ll ot t Universit y of Saskatchewa n S askatoon, Saskatchewan, Canada A C.I.P. Cata l o g ue recor df or t hi s b oo ki sava il a bl e f rom t h eL ib rar y o f Con g ress. ISBN-10 1-4020-3182-3 ( PB ) I S BN- 13 9 7 8 - 1 - 4020 - 3182 - 3( PB ) I S BN- 10 1 - 4020 - 3184 -X ( HB ) ISBN-13 978-1-4020-3184-7 ( HB ) ISBN-10 1-4020-3183-1 (e-book ) ISBN-13 978-1-4020-3183-0 ( e- b oo k) Published by Springer , P.O. Box 17 , 3300 AA Dordrecht , The Netherlands . www.springeronline.com P rinted on acid- f ree pape r Cover ima g e : Bee fli es an d a bli ster b eet l e f ee di n g on po ll en o f Ec hi nace a (courtesy of Jason Wolfe and Tyler Wist) A ll R igh ts Reserve d  C 2 005 Springe r N o part of this work may be reproduced, stored in a retrieval system, or transmitte d i nan yf orm or by an y means, e l ectron i c, mec h an i ca l ,p h otocop yi n g ,m i cro fil m i n g , recor di n g or ot h erw i se, w i t h out wr i tten perm i ss i on f rom t h ePu bli s h er, w i t h t h e except i o n of any material supplied specifically for the purpose of being entere d an d execute d on a computer s y stem, f or exc l us i ve use by t h e purc h aser o f t h ewor k. Printed in the Netherlands . C ontents Preface . xv A cknowledgment s x vi i I . Evolution and Diversit y 1 A rthro p o d Evolutio n 1. Intro d uct i on . 3 2. Art h ropo d D i vers i t y 3 2 .1. On y chophora, Tardi g rada, and Pentastoma . 4 2. 2 . Trilobit a 5 2 .3. The Chelicerate Arthro p ods . 6 2 .4. T h e Man dib u l ate Art h ropo ds 8 3. Evo l ut i onary Re l at i ons hi ps o f Art h ropo d s . 1 4 3 .1. The Proble m 14 3 .2. Theories of Arthro p od Evolutio n . 15 3 .2.1. Mono- and Diphyletic Theorie s . 1 5 3 .2.2. T h ePo l yp h y l et i cT h eor y 17 3 .3. T h eUn i ram i an s . 2 0 3 .3.1. M y riapoda-Hexapoda Relationships 2 0 4 . S ummar y 21 5 . Literature 22 2 In sec t Di vers i t y 1. Intro d uct i on . 2 5 2. Pr i m i t i ve W i n gl ess Insects . . 2 5 3. Evolution of Win g ed Insects . 27 3.1. Ori g in and Evolution of Win gs 2 7 3.2. P h y l ogenet i cRe l at i ons hi ps o f t h e Pterygota 3 3 3.3. Or i g i nan d Funct i ons o f t h e Pup a . 4 4 4 .T h e Success o f Insects 4 7 4 .1. The Adaptabilit y of Insect s 47 4 .2. The Importance of Environmental Chan g es 49 v vi C ontents 5 . Summar y 5 3 6 . Literatur e 53 3 E xterna l S tructur e 1. Introductio n 5 7 2 . General Bod y Plan . 57 3 .T h e Hea d 60 3 .1. G enera lS tructure . 60 3 .2. Hea d Appen d a g es . 6 4 3 . 2 . 1 . Antenna e 64 3.2.2. Mouth p arts . . 6 4 4 .T h e Nec k an d T h orax . 7 2 4.1. T h e Nec k 7 3 4.2. Structure o f t h eT h ora x . 73 4.3. Thoracic Appenda g es . 7 5 4 . 3 . 1 .Le g s 7 5 4 .3.2. W i ng s . 79 5 . The Abdomen . 83 5 .1. General Structure . 83 5.2. Abdominal Appenda g es 84 5 .2.1. External Genitalia 85 5 .2.2. Other Appendages 88 6 . Literatur e 8 9 4 S ystematics a nd T axonomy 1 . Intro d uct i o n 91 2 . Nam i ng an d Descr ibi ng Insects . 92 3 .C l ass ifi cat i o n . 94 3 .1. The Histor y of Insect Classification 9 6 4. Identificatio n . 102 4.1. Key to t h eOr d ers o f Insect s 1 03 5 . Literatur e 111 5 A ptery g ot e H exapo d s 1 . Intro d uct i o n 11 3 2 .Co ll em b o l a 114 3 . Protura 118 4 .Di p lur a . 120 5. Microcoryphia . 12 2 6 . Zygentoma 12 3 6 P aleopter a 1 . Intro d uct i o n 12 7 2 .Ep h emeroptera . 127 3 . Odonata 13 6 v ii C ontents 7 The P leco p teroid, B lattoid , an d O rthoptero id O rder s 1 . Introduction 147 2. P l eco p ter a . 147 3. Em bi opter a . 1 53 4 .D i ct y opter a 1 5 6 5 . Iso p tera . 1 6 3 6. Gr y lloblattodea 1 7 3 7. Derma p ter a 17 5 8 .P h asm ida 17 9 9. Mantop h asmato d e a . 182 10. Orthoptera 184 11 . Zora p tera . 19 5 8 T h e H em i ptero id O rder s 1. Intro d uct i on . 1 99 2. Psocoptera 1 99 3. Phthiraptera . 203 4 . Hemi p ter a . 210 5 . Thysanopter a 233 9 The Panor p o id Order s 1 . Intro d uct i on . 239 2 . Meco p ter a 239 3. Di p tera . 243 4 .S iph ona p ter a 2 64 5 . Trichopter a . 2 68 6 . Lepidopter a . 27 6 10 T h e R ema i n i n g E n d optery g ote O rder s 1. Intro d uct i on . 2 97 2. Mega l optera 2 97 3. Rap hidi optera . 299 4 . Neuro p tera 301 5 . Coleo p tera 305 6. Stre p si p ter a 3 2 6 7. Hymenopter a 330 I I. Anatomy and Phys i olo g y 11 T h e I nte g ument 1. Intro d uct i on . 355 2. Structure 3 55 3. Cuticle Formation 3 60 3 .1. Preecd y sis 360 3 .2. Ec d ys i s . 363 3 .3. Postec d ys is 363 viii C ontents 3 .4. Coordination of Event s . 36 5 4. Functions of the Inte g umen t 3 66 4.1. Stren g th and Hardness 366 4.2. Permea bili t y . 36 7 4. 3 . C o l o r . 3 69 4.4. Other Functions . 3 7 0 5. Summar y 3 7 0 6 . Literatur e 3 7 1 12 S ensory S ystems 1 . Intro d uct i o n 3 7 3 2 . Mec h anorecept i o n . 37 4 2.1. Sensor y Hair s 3 7 4 2.2. Pro p rioce p tors 3 7 5 2.3. S i gna l Detect i on . 3 7 8 3 . Soun d Recept i on 37 9 3 .1. Jo h nston’s Organ . 380 3 .2. T y mpanal Or g ans . 380 3 .3. Sub g enual Or g ans . . 383 4 .C h emorece p t i on 384 4.1. Locat i on an dS tructure o fS ens ill a 384 4.2. P h ys i o l ogy o f C h emorecept i o n 38 6 5 . Humidit y Perceptio n . 38 7 6 .Tem p erature Perce p tion . . . 388 7 . Photorece p tio n . 389 7 .1. Compoun d Eye s 38 9 7.1.1. Form an d Movement Percept i on . 3 9 2 7.1.2. Distance Perceptio n 3 95 7.1.3. Spectral Sensitivit y and Color Vision 39 6 7.1.4. Sensitivity to Polarized Ligh t 39 7 7 .2. S i mp l e Eye s 3 98 8. Summary 4 00 9 .L i teratur e 401 1 3 N ervous an d Chemica l I nte g ration 1. Introductio n 4 05 2 . Nervous Syste m . 4 05 2.1. Centra l Nervous System . 4 08 2.2. V i scera l Nervous S y ste m 411 2.3. Ph y siolo gy of Neural Inte g ratio n 411 2.4. Learnin g and Memor y 41 5 3 .En d ocr i ne System 417 3 .1. Neurosecretory Ce ll san d Corpora Car di ac a . 41 7 3 .2. Corpora A ll at a . 41 9 3 .3. Molt Gland s 420 3 .4. O ther Endocrine S tructures . 420 4. Insect S em i oc h em i ca l s 421 4.1. P h eromone s 422 ix C ontents 4.1.1. Sex Pheromones 422 4.1.2. Caste-Re g ulatin g Pheromones 425 4.1.3. A gg re g ation Pheromone s 42 5 4.1.4. A l arm P h eromones . 42 6 4.1. 5 . Trail-Marking Pheromone s 427 4.1. 6 . Spacin g (Epideictic) Pheromone s 428 4. 2 . Kairomones . 429 4 . 3 . Allomones 429 5 . Environmental, Neural, and Endocrine Interaction 43 0 6 . Summar y . 4 3 1 7 . L i terature . 432 14 Muscles a n d Loco m o ti on 1 . Introduction . 437 2. Musc l es . 4 3 7 2 .1. S tructur e 4 38 2 .2. P hy s i o l o gy 44 1 3. Locomotion 443 3 .1. Movement on or Throu g h a Substrat e 443 3 .1.1. Walking 44 3 3 .1.2. Jump i ng 44 7 3 .1.3. Craw li n g an d Burrow i n g . 44 8 3 .2. Movement on or Throu g h Wate r . 449 3 .2.1. Surface Runnin g . . 449 3 .2.2. Swimming by Means of Legs . 4 50 3 .2.3. Sw i mm i ng b yOt h er Means . 4 51 3.3. F li g h t 4 5 2 3 .3.1. Structural Basis . 4 5 2 3 .3.2. Aerod y namic Consideration s . 45 3 3 .3.3. Mechanics of Wing Movements 4 58 3 .3.4. Contro l o f W i ng Movement s 4 60 3 .3. 5 . Flight Metabolis m 4 6 2 3 .4. Orientation 4 64 4 . S ummar y 46 5 5 . Literatur e . 4 66 15 G a s E xc h an g e 1. Intro d uct i on . 4 69 2. Or g an i zat i on an d Structure o f t h e Trac h ea l S y stem 47 0 2. 1 . Tracheae and Tracheoles . 4 7 0 2 .2. S p iracles 4 7 3 3. Movement o f Gases w i t hi nt h e Trac h ea l System 474 3 .1. D iff us i o n . 47 5 3 .2. D i scont i nuous Gas Exc h an ge . 47 6 3 .3. Active Ventilation 4 7 7 4 . Gas Exchan g e in Aquatic Insect s 4 7 9 4 .1. C l ose d Trac h ea l Systems 479 4 .2. Open Trac h ea l Systems . 4 81 x C ontents 5 . Gas Exchan g e in Endoparasitic Insects . 484 6 . Summar y 48 5 7. Literatur e 4 85 16 F ood U p tak e a n d U t i l i zat i on 1. Introductio n 487 2 .Foo d Se l ect i on an d Fee di n g . 4 8 7 3 .T h eA li mentary System . . 48 9 3 .1. Salivar y Glands 489 3 . 2 . Fore g ut . 491 3 .3. Midgu t 492 3 .4. H i n d gut . 4 9 6 4 . Gut P h ys i o l og y 4 96 4 . 1 . G ut Movement s 4 96 4.2. Di g estio n 498 4 .2.1. Digestive Enzymes . 498 4 .2.2. Factors A ff ect i ng Enzyme Act i v i ty 4 99 4 .2.3. Contro l o f Enzyme Synt h es i san d S ecret i o n . 5 00 4 .2.4. Di g estion b y Microor g anism s . 501 4.3. Absor p tio n 50 2 5 . Metabolism 503 5 .1. S ites of Metabolism. . . 503 5 .1.1. Fat Bod y . 5 0 4 5 .1.2. M y cetoc y tes . 504 5 .2. Carboh y drate Metabolis m 505 5 .3. Li p id Metabolism . 506 5 .4. Amino Acid and Protein Metabolism . 506 5 . 5 . Metabolism of Insecticides . 5 0 7 6 . Summar y 509 7. Literatur e 5 1 0 17 The Circulator y S ystem 1. Introductio n 5 1 5 2 . S tructur e . 5 1 5 3 .P h ys i o l og y . 5 1 9 3 .1. C i rcu l at i o n 5 1 9 3 .2. Heartbea t 520 4 . Hemol y mph 5 2 1 4.1. P l asm a 5 2 2 4 .1.1. Compos i t i on 5 22 4. 1 . 2 . Funct i on s . 5 24 4 . 2 . Hemoc y te s 52 4 4 .2.1. Ori g in, Number, and Form 5 2 4 4 .2.2. Funct i on s . 5 2 6 5 . Resistance to Disease . 530 5 .1. Wound Healin g . 5 3 0 5 .2. Immunit y . 530 5 .2.1. Resistance to Host Immunit y 532 xi C ontents 6 . Summar y 5 33 7 . Literature . 534 1 8 Ni trogenou s E x c r e ti o n a n d S alt an d W ater B a l ance 1 . Introduction . 53 7 2 . Excretor ySy stems 537 2 .1. Ma l p i g hi an Tu b u l es—Rectum 537 2 .2. Ot h er Excretory Structures . 5 3 9 3. N i tro g enous Excret i on . 5 41 3 .1. The Nature of Nitro g enous Wastes 54 1 3 .2. Ph y siolo gy of Nitro g enous Excretio n . 5 4 3 3 .3. Storage Excret i on . 5 4 5 4 . S a l tan d Water Ba l anc e 5 4 6 4 .1. Terrestr i a l Insects . 5 4 6 4 .2. Freshwater Insect s 550 4 . 3 . Brackish-Water and S alt w ater Insect s 551 5 . Hormonal C ontrol . 554 6 . Summar y 556 7. L i teratur e . 55 6 I II. Re p roduction and Develo p ment 19 R e p roductio n 1 . Introduction . 561 2. Structure and Function of the Reproductive System 5 61 2 .1. Fema l e 53 2 2. 2 . Male . 5 6 5 3. Sexual Maturation 568 3 .1. Female 568 3 .1.1. V i te ll ogenes i s 56 9 3 .1.2. V i te lli ne Mem b rane an dCh or i on F ormat i on . 5 70 3 .1.3. Factors Affectin g Sexual Maturit y in th e Fe m a l e . 5 72 3 .2. Ma le . 579 4 . Mat i ng Be h av i o r . 581 4 .1. Mate Locat i on an d Reco g n i t i on 5 8 1 4 .2. Courtship 58 2 4 .3. Co p ulation 583 4. 3 .1. Insemination 58 4 4 .4. Postcopu l atory Be h av i or . . 586 5 . Ovulatio n . 5 8 7 6. Sperm Use, Entr y into the E gg , and Fertilization 5 87 6 .1. S p erm Us e . 587 6 .2. Sperm Entry into the Egg s 588 6 . 3 . Fertilizatio n 588 [...]... 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... proposed 2 Arthropod Diversity Arthropods share certain features with which they can be defined These features are: segmented body covered with a chitinous exoskeleton that may be locally hardened and is periodically shed, tagmosis (the grouping of segments into functional units, for example, head, thorax, and abdomen in insects), presence of preoral segments, paired jointed appendages on a varied number... 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... heart enclosed within a pericardium, nervous system comprising dorsal brain and ventral ganglionated nerve cord, muscles almost always striated, and epithelial tissue almost always non-ciliated 3 4 CHAPTER 1 Though the “true” arthropods fit readily within this definition, three small groups, the Onychophora, Tardigrada, and Pentastoma, whose members are soft-bodied, wormlike animals with unjointed appendages,... lived near or on the sea floor While some species preyed upon small, soft-bodied animals, the majority were scavengers However, like earthworms, a few smaller trilobites took in mud and digested the organic matter from it On the basis of X-ray studies of pyritized trilobite specimens, which show that trilobites possess a combination of chelicerate and crustacean characteristics, Cisne (1974) concluded... Shear, in Fortey and Thomas, 1998) Some 3000 species of chilopods (centipedes) (Figure 1.7A) have been described (Lewis, 1981) They are typically active, nocturnal predators whose bodies are flattened dorsoventrally The first pair of trunk appendages (maxillipeds) are modi ed into poison claws that are used to catch prey In most centipedes the legs increase in length from the anterior to the posterior of... formed from the same five original segments A third difficulty of the Snodgrass scheme is the implied homology of the sevento nine-segmented biramous appendage of Crustacea with the five-segmented, uniramous appendage of Insecta Supporters of the Mandibulata concept, for example, Matsuda (1970), derived the insect leg from the ancestral crustacean type by proposing that the extra segments were incorporated... the major arthropod groups and related phyla Hatched lines ending in a question mark indicate arthropod fossils not easily assigned to existing taxa (Onychophora-Myriapoda-Hexapoda) In some schemes the Trilobita are included as a sister group of the Chelicerata in the phylum Arachnomorpha; in others they are ranked as an independent phylum Manton, especially, argued that there are fundamental differences... 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. .. readers interested in this should consult Tiegs and Manton (1958) Nevertheless, a few very early schemes should be noted to show how ideas changed as new information became available The first monophyletic scheme for arthropod evolution was devised by Haeckel (1866).* Though believing that arthropods had evolved from a common ancestor, he divided them into the Carides (Crustacea, which included Xiphosura, . t h eL ib rar y o f Con g ress. ISBN-10 1-4 02 0-3 18 2-3 ( PB ) I S BN- 13 9 7 8 - 1 - 4020 - 3182 - 3( PB ) I S BN- 10 1 - 4020 - 3184 -X ( HB ) ISBN-13 97 8-1 -4 02 0-3 18 4-7 ( HB ) ISBN-10 1-4 02 0-3 18 3-1 (e-book ) ISBN-13. factors that affect the distribution and abundance o f insects. In Chapter 22 abiotic (ph y sical) factors in an insect’s environment are considered . Cha p ter 23 deals with the biotic factors. 70 3 x i v C ontents 4.2. Interspeci c Interactions . 70 5 4 .2.1. Com p etition and Coexistence 705 4 .2.2. Predator-Pre y Relationships . 7 09 4 .2. 3 . Insect-Insect Mutua li sm s 71 1 5 . Insect