Ecology of insects PDF
Preview Ecology of insects
Speight-TheInsects 9781405131148_1_pretoc FinalProof page 1 23.4.2008 6:43pm Ecology of Insects Speight-TheInsects 9781405131148_1_pretoc FinalProof page 2 23.4.2008 6:43pm Speight-TheInsects 9781405131148_1_pretoc FinalProof page 3 23.4.2008 6:43pm Ecology of Insects C O N C E P T S A N D A P P L I C A T I O N S Martin R. Speight Mark D. Hunter Allan D. Watt A John Wiley & Sons, Ltd., Publication Speight-TheInsects 9781405131148_1_pretoc FinalProof page 4 23.4.2008 6:43pm Thiseditionfirstpublished2008,©2008byMartinR.Speight,MarkD.HunterandAllanD.Watt BlackwellPublishingwasacquiredbyJohnWiley&SonsinFebruary2007.Blackwell’spublishingprogramhasbeen mergedwithWiley’sglobalScientific,TechnicalandMedicalbusinesstoformWiley-Blackwell. 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Setin9/11Photina bySPiPublisherServices,Pondicherry,India PrintedinSingaporebyCOSPte 1 2008 Speight-TheInsects 9781405131148_2_toc FinalProof page 5 22.4.2008 10:06am CONTENTS Prefacetosecondedition viii Forewordtofirstedition x 1 Anoverviewofinsectecology 1 1.1 Introduction 1 1.2 Historyofecologyandentomology 1 1.3 Ecologicalassociations 2 1.4 TheInsecta 2 1.5 Fossilhistoryandinsectevolution 5 1.6 Habitsofinsects 10 1.7 Numbersofinsects:speciesrichness 11 1.8 Variationsinspeciesnumber 14 1.9 Thenumberofinsects:abundance 22 1.10 Insectsandhumans 29 1.11 Conclusion 32 2 Insectsandclimate 33 2.1 Introduction 33 2.2 Temperature 33 2.3 Daylength(photoperiod) 40 2.4 Rainfall 41 2.5 Wind 47 2.6 Climatechange 54 2.7 Conclusion 60 3 Insectherbivores 61 3.1 Thetroublewithplantsasfood 61 3.2 Feedingstrategiesofherbivorousinsects 64 3.3 Plantdefenses 65 3.4 Planthybridzonesanddefenseagainstherbivores 94 3.5 Whenisadefensenotadefense? 95 3.6 Costsofresistanceandtolerancetoherbivores 95 3.7 Overcompensation 98 3.8 Plantdefenseunderelevatedcarbondioxide 100 Speight-TheInsects 9781405131148_2_toc FinalProof page 6 22.4.2008 10:06am vi Contents __________________________________________________________________________ 4 Resourcelimitation 103 4.1 Theimportanceofresourcelimitationoninsectpopulations 103 4.2 Competitionforlimitedresources 106 4.3 Thenicheconcept 108 4.4 Theoreticalapproachestothestudyofcompetition 109 4.5 Competitionamonginsectsinexperimentalandnaturalpopulations 113 4.6 Competitionamonginsectsotherthanherbivores 131 5 Naturalenemiesandinsectpopulationdynamics 135 5.1 Introduction 135 5.2 Thevarietyofnaturalenemies 135 5.3 Theimpactofnaturalenemiesoninsectpopulations 138 5.4 Modelingpredator–preyinteractions 148 5.5 Synthesis:combiningtheimpactsofnaturalenemiesandotherfactorsoninsect populationdynamics 163 5.6 Conclusion 178 6 Evolutionaryecology 180 6.1 Introduction 180 6.2 Lifehistorystrategies 180 6.3 Sexualstrategies:optimizingreproductivepotential 185 6.4 Lifehistoryvariationswithregion 194 6.5 Coevolution 196 6.6 Sequestrationofplantsecondarymetabolites 229 6.7 Demeformationandadaptivegeneticstructure 235 6.8 Extremewaysoflife 239 7 Physiologicalecology 250 7.1 Introduction 250 7.2 Foodandfeedinginjuveniles 250 7.3 Foodandfeedinginadults 256 7.4 Metabolism 258 7.5 Respiration 262 7.6 Excretion 266 7.7 Waterrelations 268 7.8 Development:ecdysisandpupation 271 7.9 Reproduction 274 7.10 Conclusion 279 8 Insectsinecosystems 280 8.1 Whatisecosystemecology? 280 8.2 Afewfundamentalsofecosystemecology 280 8.3 Insectsandtheterrestrialcarboncycle 283 8.4 Leaf-shreddinginsectsandstreamecosystems 288 8.5 Insectdefoliatorsandthecyclingofnutrients 291 8.6 Insects,plantcommunitystructure,andsuccession 295 8.7 Conclusion 302 9 Biodiversity 303 9.1 Introduction 303 Speight-TheInsects 9781405131148_2_toc FinalProof page 7 22.4.2008 10:06am __________________________________________________________________________ Contents vii 9.2 Measuringbiodiversity 303 9.3 Patternsininsectdiversity 317 10 Insectconservation 342 10.1 Introduction 342 10.2 Statusandtrendsininsectdiversity 342 10.3 Threatstoinsects 353 10.4 Conservationandrestoration 365 10.5 Prospectsforinsectconservation 378 11 Insectsanddiseases 379 11.1 Introduction 379 11.2 Diseasesandpathogens 379 11.3 Pathogenspread 380 11.4 Vectorecology 384 11.5 Vectordispersal 387 11.6 Pathogentransmission 389 11.7 Effectsonthevector 392 11.8 Epidemiology:thespreadofdisease 397 11.9 Humanactivitiesandvectors 399 11.10 Vectorcontrol 403 11.11 Casestudies 409 11.12 Conclusion 427 12 Insectpestmanagement 429 12.1 Introduction 429 12.2 Theconceptof‘‘pest’’ 429 12.3 Whypestoutbreaksoccur 430 12.4 Ecologicalpestmanagement 443 12.5 Biologicalcontrol 450 12.6 Chemicalcontrol 475 12.7 Integratedpestmanagement 497 References 514 Taxonomicindex 597 Subjectindex 602 Colourplatesectionappearsbetweenpp.310and311 Speight-TheInsects 9781405131148_3_posttoc FinalProof page 8 22.4.2008 10:07am PREFACE TO SECOND EDITION Thefirsteditionofthisbookwaspublishedin1999. our second edition reflects that acceptance. For In the 9 years since then, many changes have ecologists,climatechangerepresentsoneofthemost occurred in the worlds of ecology and entomology. important topics for debate and research that has First, and very sadly, the author of the foreword of arisen this century. Entomologists, too, have a clear our first edition, Professor Sir Richard Southwood, role to play in predicting changes in the distribution died in 2005. Dick was an eminent taxonomist, a andabundanceofplantpestsanddiseasevectors.As classicalecologistandpopulationdynamisist,agreat air currents, precipitation, and temperature change, diplomat and leader, and a guiding light to many many insect species will be lost from current geo- students,academics,andauthorsalike. graphicareasandgainedinothers.InChapter2,we Second, major advances have been made in our describe how speckled wood butterflies are moving understandingoftheecologyandevolutionofinsects, northwards in the UK while skipper butterflies are especially in the fields of molecular evolution and moving northward in the USA. What about insect systematics. We have completely rewritten much of diversity? Despite uncertainty about the future thissecondeditiontotakeaccountofrecentchanges climate, there is concern among some scientists that inthesefields.Forexample,whowouldhavethought biodiversity losses could be serious (see Chapter 10). back in 1999 that the comfortable and to many, Changing distributions of pest species are already satisfying, place in the Arthropoda occupied by the apparent. The western corn rootworm and the insectswouldbeconsideredverydifferentlessthana Coloradopotatobeetlearemovingfurtherandfurther decadelater?Whenthefirsteditionwaspublished,the intonorthernEurope(seeChapter12),whileremote Insecta sat neatly in the Hexapoda as part of the areasinAlaskaareexperiencingrangeexpansionsof subphylum Uniramia, along with their clear close agriculturalandforestpests. relativestheMyriapoda(surely,insectsarejustmilli- There have also been major advances in the pedesorcentipedeswithfewerlegs?).However,tech- applied entomological fields of disease epidemiology niquesinmolecularbiologyhaverevealedpreviously and pest management since the first edition of this unrecognizedrelationshipsamongtaxa,detailsabout book. Emerging diseases vectored by insects such as evolutionary pathways, and resulting taxonomic WestNilevirushavegainedprominence,whereasold associationsthathaveoverthrownourlong-accepted ones,suchasmalariaandbarleyyellowdwarfvirus, comforts. As we explain in detail in Chapter 1, the have become even more deleterious despite many Insectanowbelonginthemajorgroupknownasthe attempts at control. Chapter 11 describes in detail Pan-Crustacea along with the true Crustacea as a how both plant and animal (including human) sister group. The unfortunate Myriapods appear to diseases have continued to proliferate because of the becloselyrelatedtono-one. actions of insects as vectors or predispositioning Thepastdecadehasseenalmostuniversalaccept- agents,andhowkeeneffortsarebeingmadetoreduce ance by the scientific community of the role that or even prevent disease incidence by insect popul- humans are playing in global climate change and ation management. Newer techniques in vector Speight-TheInsects 9781405131148_3_posttoc FinalProof page 9 22.4.2008 10:07am _________________________________________________________ Prefacetosecondedition ix management are based largely in molecular biology. andcolleaguesintheproductionofthisnewedition. An example is the manipulation of the mosquito Two anonymous reviewers provided a wealth of genometorenderitimmunetothemalarialprotozoa comments. Generations of Oxford undergraduates Plasmodium, thus removing the ability of mosquitoes read some of the chapters, in particular numerous to vector the pathogenic agent. Similarly, modern (in fact annual) updates of Chapter 12. Graduate crop pest management is becoming more and more students at the University of Georgia helped to reliant on genetic techniques. While continuing remove(unwanted)bugsfromseveralchapters.Clive to refine and improve the efficiency of traditional Hambler, Peter Henderson, Angela Speight, Mary- methods in biological and chemical control, pest Carol Hunter, and Katy Watt were always there managers are now increasingly exploring ways to for us with advice, support, and in some cases, manipulate crop genomes to outwit insect pests. refreshments. Peter Holland, David Ferrier, Charlie Chapter 12 discusses the pros and cons of genetic Gibson, Dave Thompson, Charles Godfray, Angela modification of crops to express genes of the highly Douglas, and George McGavin advised us on aspects efficient insecticidal bacterium, Bacillus thuringiensis. of ecology and entomology. Jane Andrew, Rosie Likeitornot,thistechnologyhasavitalroletoplay Hayden,andWardCooperfromBlackwellPublishing inthefutureofglobalfoodproduction. gottheproducttothebookshelves. The power of evolution by natural selection Gratefulacknowledgementismadetoallpublishers continues to structure much of what we see in the who granted us permission to use tables, graphs, biological world, and we now know considerably and photos from journals, books, and websites. In moreaboutitsstrengthsandlimitationsthanwedid particular, we thank Wiley-Blackwell Publishing, a decade ago. As we discuss in Chapter 6, it seems Elsevier, Springer Science and Business Media, that that precise matching of insect herbivore and Ecological Society of America, Entomological Society plant phylogenies might be rarer than we once of America, Cambridge University Press, National thought. At the same time, students of insect evolu- AcademyofSciencesoftheUnitedStatesofAmerica, tion continue to provide us with wonderful new American Association for the Advancement of discoveries(orolddiscoveriesrecentlybroughttothe Science, Inter-Research Science Centre, Japanese notice of authors such as ourselves). One of the Society of Applied Entomology and Zoology, Annual mostexcitingnewinclusionsinthesecondeditionof ReviewsOrganisation,AmericanMedicalAssociation, this book is the story of the barnacle predator, Ingenta Publishers, Company of Biologists Ltd., Oedoparena, described in Chapter 6. This insect has and Utah State University. We have tried very hard managed to get its own back on what we are now to seek permissions from all concerned. If we have toldareitsclosestrelatives,theCrustacea.Whilenot failedonanyoccasion, weapologize,andwillrectify beingmarineinthetruesenseoftheterm,thelarvae thesituationinfutureeditions.Allphotosweretaken of this insect are able to kill and eat a truly marine byMartinSpeightunlessotherwiseacknowledged. animal, when said animal has the audacity to try to invadearealmoftheinsects,inthiscase,thelittoral MartinSpeight,Oxford zoneoftheNorthAmericanintertidal. MarkHunter,Michigan We would like to acknowledge the assistance, AllanWatt,Edinburgh advice, help, and support of our families, friends, August2007
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