Inadequacy of Past Researches on the Oeningen and Radoboj Ants 5 Numerical Proportion of Known Fossil Ant Specimens to Those of Other Hymen- optera
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P- nj Omo ANTS X 2 3 Sto=c t:o |H V,4- _=CA "5 c o-a oOT3 "O2 o d C"^ .5>> ba o re E Cl [14
COLUMBIAUNIVERSITYBIOLOGICALSERIES.L\.
ANTSTHEIRSTRUCTURE,DEVELOPMENT
ANDBEHAVIOR
WILLIAMMORTONWHEELER,Pn.D.
deTrtpiTOI>S/St'ousTroXXda.v ys,/cat/xaXXocCTTIrCivf\aTTOvXetoock
THECOLUMBIAUNIVERSITYPRESS
1910Allrightsreserved
Copyrighted1910,by
THECOLUMBIAUNIVERSITYPRESS
PRESSOF
THENEWERAPRINTINGCOMPANY
LANCASTER.PA
TOMYWIFE
DORABAYEMERSON
PREFACE.
insects. foreigncontributions. viiPREFACE.
theNorthAmericanspecies. ofinterpretation. mologyinAmerica. ofNaturalHistory.Brueshasaidedmeinreadingtheproof.
BUSSEYINSTITUTION,
FORESTHILLS,BOSTON,MASS.,
October30,1909.
TABLEOFCONTENTS.
CHAPTERI.
ANTSASDOMINANTINSECTS.
PAGEII.TheDominanceofAnts,asshownby
andOtherAnimals2III.ProbableCauseoftheDominanceofAnts3
V.ComparisonofHumanandAntSocieties5
Organization.
VII.EconomicImportanceofAnts8
VIII.AntsasObjectsofBiologicalStudyII
CHAPTERII.
THEEXTERNALSTRUCTUREOFANTS.
I.GeneralDistinguishingCharacters13
II.TheSegmentationoftheBody14
III.TheIntegument15
IV.TheHead16
Mouth-parts.4.TheAntennae.5.TheEyes.
V.TheThorax20
xiTAIU.1'OFCOXTEXTS.
PACEVI.TheAbdomen26
4.TheTerminalSegmentsoftheMale.
CHAPTERIII.
THEINTERNALSTRUCTUREOFANTS.
I.TheAlimentaryTract31
II.TheGlandularSystem37
natorialOrgans39V.TheRespiratorySystem49
XT.TheMuscularSystem49
CHAPTERIV.
THEINTERNALSTRUCTUREOFANTS(CONCLUDED).
I.TheNervousSysteminGeneral51
II.TheBrain52
canceofthePedunculateBodies.III.TheVentralXerve-cord57
IV.TheSympatheticNervousSystem58
V.TheSense-organs59
MedianEyes,Stemmata,orOcelli.
CHAPTERV.
THEDEVELOPMENTOFANTS.
ThatofOtherHymenoptera67
11.TheNursingoftheBrood69
TABLEOFCONTENTS.xin
PAGEIII.TheEgg70
Development.
IV.TheLarva72
V.Pupation76
VI.ColorationoftheCallows79
VII.LengthofDevelopmentalPeriods80
VIII.LongevityofAdultAnts.81
IX.ResistanceofAntstoNoxiousInfluences83
CHAPTERVI.
4POLYMORPHISM.
I.DefinitionoftheTerm86
phisminHymenoptera90KnownamongAnts.
morphisminAnts91 disposition.V.TheThreeAspectsoftheProblem102
Phylogenetic.3.ThePsychological.
DevelopmentoftheWorker103
WorkerandRelatedTypes.5.Conclusion.
TABLEOPCONTENTS.
PACECHAPTERVII.
POLYMORPHISM(CONCLUDED).
DevelopmentoftheWorkerno
andwithThatoftheAuthor.VIII.ThePsychologicalViewoftheProblem117
stinctsinAnts. entiationinStructure119 changesPrecedeMorphologicalChanges.3.Conclusions122
CHAPTERVIII.
IThePresentStatusoftheStudyofAnts123
II.TheThreeView-pointsinAnt-study124
III.TheDevelopmentofAntTaxonomy124
IV.TheHistoryofAntEthology127
\Yriters.TABLEOFCONTENTS.xv
PAGEV.TheStudyofMyrmecophily128
VI.TheStudyofAntMorphology129
VII.TheDifficultiesofClassification129
i.TheThreeCastes.2.TheVariabilityofAnts.IX.QuadrinomialNomenclature131
X.ConspectusoftheClassificationofAnts134
CHAPTERIX.
THEDISTRIBUTIONOFANTS.
I.ModesofDissemination145
III.TheFaunisticDistributionofAnts147
IV.TheEthologicalDistributionofAnts156
CHAPTERX.
FOSSILANTS.
II.TheTertiaryAnts161
America.
TABLEOFCOXTEXTS.
PAGEIII.TheQuaternary,orPleistoceneAnts173
.174Ants174
CHAPTERXI.
THEHABITSOFANTSINGENERAL.
Food177
III.ProtectiveHabitsofAnts178
ofPreventingMixtureofAlienColonies.IV.ReproductionamongAnts182
berofIndividualsinaColony.CHAPTERXII.
ANT-NESTS.
I.GeneralArchitecturalCharacteristics192
II.MethodsofConstruction194
III.ChangeofAbode195
IV.ClassificationofAnt-nests198
V.NestsintheSoil199
?tc.CHAPTERXIII.
ANT-NESTS(CONCLUDED).
VI.NestsintheCavitiesofPlants207
CarpenterAnts;(b~)GallAnts.
TABLEOFCONTENTS.xvn
PAGEVII.SuspendedNests213
Description;(b)Manufacture.
VIII.NestsinUnusualSituations221
IX.AccessoryStructures222
Nests.3.Tents,orPavilions.
CHAPTERXIV.
THEPONERINEANTS.
IV.TheCastesofthePonerinae230
V.NestingandFeedingHabits232
VI.TheThreeTypesofPonerineLarvae...233
VIII.CarefortheBrood237
IX.TheHatchingoftheCallows238
X.DeportationandHuntingHabits240
XII.PhytogenyoftheGroup243
CHAPTERXV.
THEDRIVERANDLEGIONARYANTS.
I.TheDorylinaeasaGroup246
II.DescriptionoftheCastes248
III.DifficultiesofNomenclature248
IV.TheGenusDoryhts249
V.TheGenusjEnictns253
VI.TheGeneraEcitonandCheliomyrmex255
VII.TheProblemsoftheDorylinae265
T.DomesticEconomy.2.Dichthadiigynes.
'I'ABLEOFCONTENTS.I'll.\ITERXVI.
THE11AKVKSTINGANTS.
PAGEII.EarlyObservations268
Forel,Andre,EmeryandOthers269
dologeton275 noplus276 ...278CHAPTERXVII.
THERELATIONSOFANTSTOVASCULARPLANTS.
III.TheAntsDwellingonPlants302
TABLEOFCONTENTS.xix
PAGESouthAmerica.
IV.OtherRelationsofAntstoPlants315
InjurioustoAnts.
CHAPTERXVIII.
THEFUNGUS-GROWINGANTS.
andTheirFungi318II.CharacteristicsoftheTribeAttii319
III.ReviewoftheLiterature320
IV.Belt'sObservationsonAttaccphalotcs321
V.Moeller'sObservations324
Cyphomyrmex.
Huber.)329
OriginoftheHabit.
CHAPTERXIX.
HOPPERSANDCATERPILLARS.
II.RelationsofAntswithAphids340
3.TheAbdominalSiphonsofAphids.
TABLEUI;CONTENTS.
PAGEIII.RelationsofAntswithCoccids347
\.RelationsofAntswithfree-hoppers^oHomopterainGeneral351
VII.TheEulgoridae,etc356
IX."Trophobiosis
"360CHAPTERXX.
HONEYAXTS.
I.DescriptionoftheHoney-storingHabit361
II.TheSpeciesofHoneyAnts363
III.TheCausesofRepletion374
mentofRepletes375 ...376CHAPTERXXI.
PERSECUTEDANDTOLERATEDGUESTS.
I.ExtranidalandIntranidalSymbiosis378
II.MyrmecophilesinGeneral379
III.TheSynechthrans382
IY.TheSynoeketes383
TABLEOFCONTEXTS.xxi
CHAPTERXXII.
TRUEGUESTS,Ecro-ANDENTOPARASITES.
PAGE andParasitestotheAnts398II.TheSymphilesStructuralAdaptations398
Antenna?.
III.TypicalSymphiles402
Lomechusinae.(a)SummaryofLifeHistory.(b)
IV.Ectoparasites412
5.TheChalcididse.
V.Entoparasites419
ClassesofParasites421
VII.MyrmecophagsandMyrmecoids422
CHAPTERXXIII.
THECOMPOUNDNESTS.
I.SocialSymbiosis423
MixedColonies.
TABLEOFCONTENTS.
PACE thtimxglacialis.CHAPTERXXIV.
THETEMPORARYSOCIALPARASITES.
I.AntParasitisminGeneral437
QueensinAlienColonies,andTheirPhylogem.
Parasitism44r
IV.GeneralConclusions449
CHAPTERXXV.
III.TheFacultativeSlave-makers454
theDuloticInstincts.TABLEOFCONTENTS.xxiii
CHAPTERXXVI.
THEAMAZONS,OROBLIGATORYSLAVE-MAKERS.
PAGEI.Introductory471
Polyergusrufescens.
III.TheAmericanAmazons474
lucidus.IV.TheFoundingofAmazonColonies486
CHAPTERXXVII.
ficationandDefinition488II.TheDegenerateSlave-makers489
6.Strongylognathustcstaceus.
2,Harpagoxenusamericanus.
III.ThePermanentSocialParasites495
SociologicalAnalogies.
CHAPTERXXVIII.
THESENSATIONSOFANTS.
TheSensesasaBasisforitsStudy505
xxivTAlil.HOFCONTENTS. PAGEII.SensePerceptioninAnts508
Ultra-violetandRontgenRays.
andVibrationsintheLivesofAnts517CHAPTERXXIX.
THEINSTINCTIVEBEHAVIOROFANTS.
7.RegulationinInstinct.
DivinatorySympathy529
CHAPTERXXX.
THEPLASTICBEHAVIOROFANTS.
TypesofPlasticBehavior531
5.Docility.
III.TheNatureofMemoryinAnts531)
TABLEOFCONTEXTS.xxv
PAGE IV 7 soninginAnts540APPENDICES.
C.AListofDescribedNorthAmericanAnts561
D.MethodsofExterminatingNoxiousAnts573
E.Literature578
CHAPTER1.
ANTSASDOMINANTINSECTS.
allexerunt."Nylander,Europse,"1846.
1908.2I ANTS. mentalstimuli. animalsmanincluded.