Insect Behavior: 2nd Edition

518 Pages·2010·12.59 MB·English

by  Robert W. Matthews, Janice R. Matthews (auth.)

Insect Behavior
·
Robert W. Matthews Janice R. Matthews
Insect Behavior
Second Edition
123
RobertW.Matthews JaniceR.Matthews
UniversityofGeorgia UniversityofGeorgia
Dept.Entomology Dept.Entomology
AthensGA30602 AthensGA30602
USA USA
[email protected] [email protected]
ISBN978-90-481-2388-9 e-ISBN978-90-481-2389-6
DOI10.1007/978-90-481-2389-6
SpringerDordrechtHeidelbergLondonNewYork
LibraryofCongressControlNumber:2009926821
©SpringerScience+BusinessMediaB.V.2010
Nopartofthisworkmaybereproduced,storedinaretrievalsystem,ortransmittedinanyformorby
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Coverdesign:BoekhorstDesignBV
Cover photo credit: The male Eastern Tiger Swallowtail, Papilio glaucus, shown here is one of the
mostrecognizablebutterfliesinNorthAmerica;femalesofthisspeciesaredimorphic,appearinginboth
yellowandblackforms.Likemanyotherbutterflies,swallowtailsareattractedtothearomaticflower
clustersofLantanafornectar,butdonotlayeggsonitssomewhattoxicfoliage.Theauthorstookthis
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SpringerispartofSpringerScience+BusinessMedia(www.springer.com)
Preface to the Second Edition
This book is for all who are interested in the biological sciences. Like the course
thatoriginallyinspiredit,thetextisdesignedforuseatseniorundergraduatelevel
for college and university students, so we trust that it will find readership among
those who have had some basic introduction to entomology and animal behavior.
However,wealsohopeitwillproveusefultonewcomerswhomaybeapproaching
behavioralstudyfromotherperspectives.
Thisisoursecondedition,bornanewafterthirtyyears.Muchhaschangedduring
thattime,especiallyinthebreadthanddepthofafieldthat(likeus)wasfairlyyoung
back in 1978. New technologies are allowing scientists to shape—and answer—
questions in ways that once could not even have been envisioned. Insect behavior
researchnowhaswings,andispoisedtotakeoff.However,atthisjuncture,weall
mustalsotakecarenotloseanawarenessofourroots.Thus,aswritersintroducing
thisexcitingfieldofstudytothenextgenerationofscientistsandinsectenthusiasts,
wehavetriedtostrikeabalancebetweennewideasandold,andbetweenmodern
developmentsandhistoricalinsights.
Ourobjectivesinwritingthiseditionremainthesameastheyhavealwaysbeen.
The first of these has been to help readers understand how a number of major
behavioral systems function. Thus, this is not an encyclopedia, but an introduc-
tiontofundamentalconceptsandprocessesasseenfromacomparativeevolutionary
viewpoint.Wehavenotdocumentednumerousstringsofexamplesmerelyfor‘com-
pletenessofcoverage’butinsteadhavetriedtogiveaflavorofthediversityofways
in which insects approach similar life tasks. Because the Internet and excellent
search engines have made access to information sources easy and nearly instan-
taneous, we have not burdened readers with a cumbersome citations in the text;
searching on key terms, aided when necessary by references associated with fig-
ure credits at the book’s end, will provide entry into additional literature for those
interestedinfurtherpursuingsubjectswecanbutintroduce.
Our second objective has been to help readers gain insights into accessible
waysinwhichbehavioralresearchcanbeconducted.Wheneverpossible,wehave
included discussions of important experiments and investigations, rather than pre-
sentingarhetoricofconclusions.Selectedprinciplesareinterwovenwithcasestud-
ies of specific situations, presenting actual examples in a manner compatible with
thedynamic,open-endedfieldandlaboratoryexperiencesinwhichtheyhavearisen.
v
vi PrefacetotheSecondEdition
Likeanywritersofageneraltextbook,werecognizeadeepobligationtomany
others—to those of whose work we write, to other authors whose ideas we use,
toourownteacherswhohaveshapedourperspectivesandinterests,andtoourstu-
dents,friendsandcolleagueswiththeirmanystimulatingandinvaluablesuggestions
andcriticisms.
We also thank the many scientists and journal editors who have freely granted
permission for the use of published material. Many colleagues have generously
provided us with photographs, which are acknowledged in the credits list; special
thanksareduetoDouglasW.Whitmanforcontributingnumerousnewimagesand
toRobertE.SilbergliedandCarlW.Rettenmeyer,bothnowdeceased,whosepho-
tographictalentcontinuestoenhancethisedition.Wealsoareespeciallygratefulto
JoanW.KrispynandPaulH.Matthewsfornumerousoriginaldrawings.
Athens,Georgia RobertW.Matthews
JaniceR.Matthews
Contents
1 TheHistoryandScopeofInsectBehavior . . . . . . . . . . . . . . 1
1.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.1.1 WhatIsInsectBehavior? . . . . . . . . . . . . . . . . 1
1.1.2 InsectBehavior’sBiologicalContext . . . . . . . . . . 3
1.1.3 HistoricalFoundations . . . . . . . . . . . . . . . . . 6
1.1.4 TheWatershedYears . . . . . . . . . . . . . . . . . . 9
1.1.5 TheRiseofEthology . . . . . . . . . . . . . . . . . . 13
1.2 ConceptualFrameworks . . . . . . . . . . . . . . . . . . . . . 15
1.2.1 EvolutionbyNaturalSelection . . . . . . . . . . . . . 15
1.2.2 GeneticsandBehavior . . . . . . . . . . . . . . . . . 17
1.2.3 TheComparativeApproach . . . . . . . . . . . . . . . 24
1.2.4 ConceptualPitfalls . . . . . . . . . . . . . . . . . . . 25
1.3 Phylogeny’sRole . . . . . . . . . . . . . . . . . . . . . . . . . 28
1.3.1 MicroevolutionandMacroevolution . . . . . . . . . . 28
1.3.2 PhylogeneticSystematicsandCladistics . . . . . . . . 33
1.3.3 BehaviorandSpeciation . . . . . . . . . . . . . . . . 36
1.4 QuestionsandPerspectives . . . . . . . . . . . . . . . . . . . . 38
1.4.1 ProximateandUltimateAnalyses. . . . . . . . . . . . 40
1.4.2 TypesofApproach . . . . . . . . . . . . . . . . . . . 41
2 ProgrammingandIntegratingBehavior . . . . . . . . . . . . . . . 45
2.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
2.2 Nerve-BasedCoordination . . . . . . . . . . . . . . . . . . . . 46
2.2.1 TheInsectNervousSystem . . . . . . . . . . . . . . . 47
2.2.2 SimpleReflexesandRepeatedMotorPatterns . . . . . 50
2.2.3 EthologicalExplanations . . . . . . . . . . . . . . . . 56
2.3 LifeinaStimulus-RichWorld . . . . . . . . . . . . . . . . . . 59
2.3.1 SensoryTuningandFiltering . . . . . . . . . . . . . . 60
2.3.2 MemoryandLearning. . . . . . . . . . . . . . . . . . 68
2.3.3 InsectIntelligence . . . . . . . . . . . . . . . . . . . . 81
2.4 Hormone-BasedCoordination . . . . . . . . . . . . . . . . . . 83
2.4.1 ClocksandReiterativeRhythms . . . . . . . . . . . . 86
2.4.2 GatedRhythms . . . . . . . . . . . . . . . . . . . . . 88
vii
viii Contents
3 SpatialAdjustment . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
3.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
3.2 Locomotion . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
3.2.1 TerrestrialandAquaticLocomotion . . . . . . . . . . 96
3.2.2 AerialLocomotion . . . . . . . . . . . . . . . . . . . 98
3.3 Orientation . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100
3.3.1 LocomotoryResponses . . . . . . . . . . . . . . . . . 101
3.3.2 PostureandPosition. . . . . . . . . . . . . . . . . . . 103
3.3.3 OrientationtoRadiantEnergy . . . . . . . . . . . . . 105
3.3.4 MagneticFieldOrientation . . . . . . . . . . . . . . . 108
3.3.5 OrientationtotheEvidenceofOthers’Presence . . . . 109
3.4 Thermoregulation. . . . . . . . . . . . . . . . . . . . . . . . . 110
3.4.1 DormancyandThermotolerance . . . . . . . . . . . . 110
3.4.2 RegulationofHeatGain . . . . . . . . . . . . . . . . 113
3.4.3 HeatProduction . . . . . . . . . . . . . . . . . . . . . 114
3.5 Migration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
3.5.1 SeasonalMigration . . . . . . . . . . . . . . . . . . . 117
3.5.2 MigrationUnderEphemeralConditions . . . . . . . . 122
3.5.3 DispersalandNavigation . . . . . . . . . . . . . . . . 124
4 ForagingandFeeding . . . . . . . . . . . . . . . . . . . . . . . . . 131
4.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
4.1.1 FoodRecognitionandAcceptance . . . . . . . . . . . 134
4.1.2 RegulationofFeeding . . . . . . . . . . . . . . . . . . 137
4.2 ForagingStrategies . . . . . . . . . . . . . . . . . . . . . . . . 140
4.2.1 Herbivory . . . . . . . . . . . . . . . . . . . . . . . . 141
4.2.2 ActiveSearch . . . . . . . . . . . . . . . . . . . . . . 141
4.2.3 TrappingandAmbush . . . . . . . . . . . . . . . . . . 146
4.2.4 ParasitesandParasitoids . . . . . . . . . . . . . . . . 148
4.2.5 TheftandKleptoparasitism . . . . . . . . . . . . . . . 152
4.2.6 InsectAgriculture . . . . . . . . . . . . . . . . . . . . 154
4.2.7 NestSymbionts:BecomingaHousePet . . . . . . . . 157
4.3 CoevolutionandtheArmsRace . . . . . . . . . . . . . . . . . 164
4.3.1 Attack,Defense,andCounterattack. . . . . . . . . . . 165
4.3.2 EmployingMercenariesforProtection . . . . . . . . . 170
4.3.3 TheTommyTuckerSyndrome:FoodinReturn
forServices . . . . . . . . . . . . . . . . . . . . . . . 173
4.4 FeedingasaCommunalActivity . . . . . . . . . . . . . . . . . 177
4.4.1 SimpleGroupsandFeedingAggregations . . . . . . . 177
4.4.2 SocialFeedingBehaviors . . . . . . . . . . . . . . . . 181
5 Defense:ASurvivalCatalogue . . . . . . . . . . . . . . . . . . . . 185
5.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185
5.2 DefenseMessages . . . . . . . . . . . . . . . . . . . . . . . . 186
5.3 PassiveMessages . . . . . . . . . . . . . . . . . . . . . . . . . 187
5.3.1 Crypsis:‘I’mNotHere!’ . . . . . . . . . . . . . . . . 187
Contents ix
5.3.2 SystemicDefenses:‘I’mNoxious!’ . . . . . . . . . . . 194
5.3.3 Mimicry:‘I’mSomeoneElse!’ . . . . . . . . . . . . . 197
5.3.4 AposematicDefenses:‘I’mDangerous!’ . . . . . . . . 202
5.4 ActiveMessages . . . . . . . . . . . . . . . . . . . . . . . . . 204
5.4.1 Attack:‘I’mTurningtheTables!’ . . . . . . . . . . . . 204
5.4.2 Startle:‘I’mNotWhatYouThought!’ . . . . . . . . . 209
5.4.3 GroupActions:‘We’reinThisTogether!’ . . . . . . . 213
6 ChemicalCommunication . . . . . . . . . . . . . . . . . . . . . . . 217
6.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
6.2 MechanismsofChemicalCommunication . . . . . . . . . . . . 217
6.2.1 OdorCreationandReception . . . . . . . . . . . . . . 218
6.2.2 CommunicationThroughChemistry . . . . . . . . . . 223
6.3 TheFunctionsofChemicalCommunication . . . . . . . . . . . 227
6.3.1 FindingandChoosingMates . . . . . . . . . . . . . . 228
6.3.2 Assembly,Aggregation,andRecruitment . . . . . . . 231
6.3.3 AlarmandAlert . . . . . . . . . . . . . . . . . . . . . 240
6.3.4 Host-Marking . . . . . . . . . . . . . . . . . . . . . . 242
6.3.5 Recognition . . . . . . . . . . . . . . . . . . . . . . . 246
6.4 TheInformationContentofPheromones . . . . . . . . . . . . 249
6.4.1 PhysiologicalAdjustments:TheQ/KRatio . . . . . . . 249
6.4.2 PheromonesasLanguage:SyntaxandLexicon. . . . . 251
6.4.3 ExploitationandCode-Breaking . . . . . . . . . . . . 255
6.4.4 TheChemicalChannelandOtherSignalModes . . . . 256
6.5 ChemicalCommunicationandInsectControl . . . . . . . . . . 258
7 VisualCommunication . . . . . . . . . . . . . . . . . . . . . . . . . 261
7.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . 261
7.2 Bioluminescence . . . . . . . . . . . . . . . . . . . . . . . . . 262
7.2.1 ThePhysiologyofInsectLightProduction . . . . . . . 263
7.2.2 BioluminescenceasaCommunicationMethod. . . . . 264
7.3 LightReception. . . . . . . . . . . . . . . . . . . . . . . . . . 268
7.3.1 ReceptorsandFormPerception . . . . . . . . . . . . . 268
7.3.2 VisualAcuityandFlickerVision . . . . . . . . . . . . 272
7.3.3 PolarizedLightPerception . . . . . . . . . . . . . . . 275
7.3.4 ColorVision . . . . . . . . . . . . . . . . . . . . . . . 277
7.4 FunctionsofVisualCommunication . . . . . . . . . . . . . . . 279
7.4.1 AggregationandDispersion . . . . . . . . . . . . . . 281
7.4.2 Alarm . . . . . . . . . . . . . . . . . . . . . . . . . . 283
7.4.3 SexualSignals . . . . . . . . . . . . . . . . . . . . . . 284
7.4.4 MultimodalSignaling . . . . . . . . . . . . . . . . . . 288
8 Mechanocommunication . . . . . . . . . . . . . . . . . . . . . . . . 291
8.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . 291
8.2 ProducingandSendingSignals . . . . . . . . . . . . . . . . . 292
8.2.1 SoundCreation . . . . . . . . . . . . . . . . . . . . . 293
x Contents
8.2.2 DistanceandSubstrate . . . . . . . . . . . . . . . . . 295
8.3 ReceivingSignals. . . . . . . . . . . . . . . . . . . . . . . . . 296
8.3.1 Vibration . . . . . . . . . . . . . . . . . . . . . . . . 298
8.3.2 Hearing . . . . . . . . . . . . . . . . . . . . . . . . . 303
8.3.3 CommunicationbyTouch . . . . . . . . . . . . . . . . 305
8.4 TheAcousticChannel . . . . . . . . . . . . . . . . . . . . . . 307
8.4.1 ParametersofInsectSong. . . . . . . . . . . . . . . . 307
8.4.2 SongSynchronies . . . . . . . . . . . . . . . . . . . . 308
8.4.3 ActiveAcoustics . . . . . . . . . . . . . . . . . . . . 311
8.4.4 SoundasaCommunicationMethod . . . . . . . . . . 313
8.5 FunctionsofInsectCommunicativeSounds . . . . . . . . . . . 314
8.5.1 Protest,Alarm,andAggression . . . . . . . . . . . . . 314
8.5.2 AposematicSoundsandAcousticMimicry. . . . . . . 319
8.5.3 SexualSignals . . . . . . . . . . . . . . . . . . . . . . 321
8.5.4 SocialSounds . . . . . . . . . . . . . . . . . . . . . . 327
9 ReproductiveBehavior . . . . . . . . . . . . . . . . . . . . . . . . . 341
9.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . 341
9.2 CourtshipandMating . . . . . . . . . . . . . . . . . . . . . . 342
9.2.1 ThePhysiologyofMatingBehavior . . . . . . . . . . 344
9.2.2 ReproductionModes . . . . . . . . . . . . . . . . . . 346
9.2.3 ComplexityandPlasticity . . . . . . . . . . . . . . . . 349
9.2.4 PollinationandMaleReproductiveBehavior . . . . . . 354
9.3 CourtshipandConflict . . . . . . . . . . . . . . . . . . . . . . 359
9.3.1 Dimorphism,SexualSelection,andMateChoice . . . 361
9.3.2 IntrasexualCompetition . . . . . . . . . . . . . . . . . 369
9.3.3 TerritorialityandDominance . . . . . . . . . . . . . . 371
9.3.4 NuptialGifts. . . . . . . . . . . . . . . . . . . . . . . 372
9.4 MatingSystemsandParentalInvestment . . . . . . . . . . . . 380
9.5 OvipositionBehavior . . . . . . . . . . . . . . . . . . . . . . . 382
9.5.1 SelectingaSiteorHost . . . . . . . . . . . . . . . . . 383
9.5.2 ReproductiveRatesandEnergyAllocation . . . . . . . 387
10 ParentalBehaviorsandSocialLife . . . . . . . . . . . . . . . . . . 389
10.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . 389
10.2 SocialOrganization . . . . . . . . . . . . . . . . . . . . . . . . 390
10.2.1 AggregationsandSimpleGroups . . . . . . . . . . . . 390
10.2.2 Parent-OffspringInteractions . . . . . . . . . . . . . . 394
10.2.3 SolitaryandCommunalNesters . . . . . . . . . . . . 400
10.3 TheInsectSocialRegister . . . . . . . . . . . . . . . . . . . . 407
10.3.1 TheAnts . . . . . . . . . . . . . . . . . . . . . . . . . 410
10.3.2 TheEusocialWasps . . . . . . . . . . . . . . . . . . . 416
10.3.3 TheBees. . . . . . . . . . . . . . . . . . . . . . . . . 420
10.3.4 TheTermites . . . . . . . . . . . . . . . . . . . . . . 423
10.3.5 LesserKnownCandidates. . . . . . . . . . . . . . . . 427
10.4 ImplicationsandCorrelatesofSocialLife . . . . . . . . . . . . 428
Contents xi
10.4.1 TheEcologyofParentalCare . . . . . . . . . . . . . . 430
10.4.2 ParadoxesofInsectSociality . . . . . . . . . . . . . . 434
10.4.3 InterspecificSocialInteractions . . . . . . . . . . . . . 440
Credits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 445
Plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 463
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 503