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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\.

ANTS

THEIRSTRUCTURE,DEVELOPMENT

ANDBEHAVIOR

WILLIAMMORTONWHEELER,Pn.D.

deTrtpiTOI>S/St'ousTroXXda.v ys,/cat/xaXXocCTTIrCivf\aTTOvARISTOTLE

Xetoock

THECOLUMBIAUNIVERSITYPRESS

1910

Allrightsreserved

Copyrighted1910,by

THECOLUMBIAUNIVERSITYPRESS

PRESSOF

THENEWERAPRINTINGCOMPANY

LANCASTER.PA

TOMYWIFE

DORABAYEMERSON

PREFACE.

insects. foreigncontributions. vii

PREFACE.

theNorthAmericanspecies. ofinterpretation. mologyinAmerica. ofNaturalHistory.

Brueshasaidedmeinreadingtheproof.

BUSSEYINSTITUTION,

FORESTHILLS,BOSTON,MASS.,

October30,1909.

TABLEOFCONTENTS.

CHAPTERI.

ANTSASDOMINANTINSECTS.

PAGE

II.TheDominanceofAnts,asshownby

andOtherAnimals2

III.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

xi

TAIU.1'OFCOXTEXTS.

PACE

VI.TheAbdomen26

4.TheTerminalSegmentsoftheMale.

CHAPTERIII.

THEINTERNALSTRUCTUREOFANTS.

I.TheAlimentaryTract31

II.TheGlandularSystem37

natorialOrgans39

V.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

PAGE

III.TheEgg70

Development.

IV.TheLarva72

V.Pupation76

VI.ColorationoftheCallows79

VII.LengthofDevelopmentalPeriods80

VIII.LongevityofAdultAnts.81

IX.ResistanceofAntstoNoxiousInfluences83

CHAPTERVI.

4

POLYMORPHISM.

I.DefinitionoftheTerm86

phisminHymenoptera90

KnownamongAnts.

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

PAGE

V.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.

PAGE

III.TheQuaternary,orPleistoceneAnts173

.174

Ants174

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

PAGE

VII.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.

PAGE

II.EarlyObservations268

Forel,Andre,EmeryandOthers269

dologeton275 noplus276 ...278

CHAPTERXVII.

THERELATIONSOFANTSTOVASCULARPLANTS.

III.TheAntsDwellingonPlants302

TABLEOFCONTENTS.xix

PAGE

SouthAmerica.

IV.OtherRelationsofAntstoPlants315

InjurioustoAnts.

CHAPTERXVIII.

THEFUNGUS-GROWINGANTS.

andTheirFungi318

II.CharacteristicsoftheTribeAttii319

III.ReviewoftheLiterature320

IV.Belt'sObservationsonAttaccphalotcs321

V.Moeller'sObservations324

Cyphomyrmex.

Huber.)329

OriginoftheHabit.

CHAPTERXIX.

HOPPERSANDCATERPILLARS.

II.RelationsofAntswithAphids340

3.TheAbdominalSiphonsofAphids.

TABLEUI;CONTENTS.

PAGE

III.RelationsofAntswithCoccids347

\.RelationsofAntswithfree-hoppers^o

HomopterainGeneral351

VII.TheEulgoridae,etc356

IX."Trophobiosis

"360

CHAPTERXX.

HONEYAXTS.

I.DescriptionoftheHoney-storingHabit361

II.TheSpeciesofHoneyAnts363

III.TheCausesofRepletion374

mentofRepletes375 ...376

CHAPTERXXI.

PERSECUTEDANDTOLERATEDGUESTS.

I.ExtranidalandIntranidalSymbiosis378

II.MyrmecophilesinGeneral379

III.TheSynechthrans382

IY.TheSynoeketes383

TABLEOFCONTEXTS.xxi

CHAPTERXXII.

TRUEGUESTS,Ecro-ANDENTOPARASITES.

PAGE andParasitestotheAnts398

II.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.

PAGE

I.Introductory471

Polyergusrufescens.

III.TheAmericanAmazons474

lucidus.

IV.TheFoundingofAmazonColonies486

CHAPTERXXVII.

ficationandDefinition488

II.TheDegenerateSlave-makers489

6.Strongylognathustcstaceus.

2,Harpagoxenusamericanus.

III.ThePermanentSocialParasites495

SociologicalAnalogies.

CHAPTERXXVIII.

THESENSATIONSOFANTS.

TheSensesasaBasisforitsStudy505

xxivTAlil.HOFCONTENTS. PAGE

II.SensePerceptioninAnts508

Ultra-violetandRontgenRays.

andVibrationsintheLivesofAnts517

CHAPTERXXIX.

THEINSTINCTIVEBEHAVIOROFANTS.

7.RegulationinInstinct.

DivinatorySympathy529

CHAPTERXXX.

THEPLASTICBEHAVIOROFANTS.

TypesofPlasticBehavior531

5.Docility.

III.TheNatureofMemoryinAnts531)

TABLEOFCONTEXTS.xxv

PAGE IV 7 soninginAnts540

APPENDICES.

C.AListofDescribedNorthAmericanAnts561

D.MethodsofExterminatingNoxiousAnts573

E.Literature578

CHAPTER1.

ANTSASDOMINANTINSECTS.

allexerunt."Nylander,

Europse,"1846.

1908.
2I ANTS. mentalstimuli. animalsmanincluded.

ANTSASDOMINANTINSECTS.3

minatedenvironment.

4ANTS.

1Andhowevermuch

theseubiquitousinsects. man'smostdangerousenemiesareothermen." .L\'TSASDOMINANTINSECTS.5 nowengrossingtheattentionofbiologists. *likeTopinard, withtheharvestingants."

Co.,1890,p.IQ2etscq.

AN'IS.

towhichtheantshaveneverattained. inmind: itslifeasanactivememberofthecommunity.

ANTSASDOMIX.1XTINSECTS.7

tionoftheactivitiesofitsfellows. problemsoflifeandmind.

8ANTS.

ofdestroyingnoxiousants.

ANTSASDOMINANTINSECTS.9

entomologists. ioJ.VTS. cephalotes itsbodyandleftasaligature.

ANTSASDOMINANTINSECTS.n

leaf-fungi. i-ANTS. moreseriousandphilosophicalstudent.

CHAPTERII.

THEEXTERNALSTRUCTUREOFAXTS.

KalKa\ov.

"DePartibus

Animalium,"I,5.

ANTS.

Nassonow.

THEEXTERNALSTRUCTUREOFANTS.15

scales. i6AMS. legs(I'idcinfra,p.24).In desertstheseinsectseasily becomecoveredwiththe drysoilorsandandhave toremoveitfromtheir bodiesandlimbsbymeans ar,ofthestrigils.Theseor- gansarethenthrustalong theammochaetseinmuch thesamewayasweclean acombbymeansof threads.Theclypeusand mandiblesofmanyantsare alsofringedwithunusually longhairs(clypealand mandibularammochaetse) whichareemployedinre-

FIG.2.Ammocha?t?eofdesertants.(Ori-

ginal.)A,HeadofMessorpcrgandeiinprofile;

B,ventralaspectofsame;C,headofMyrmeco-

cystnsbicolorinprofile;D.ventralaspectof same;a.clypeal;b,mandibular;c.gular;d, mentalammochsetae.

THEEXTERNALSTRUCTUREOFAXIS.i/

3 iSANTS. ofridges,thefrontalcarinaeor laminae,justmesialtotheinser- tionsoftheantennae.Asmall, usuallytriangular,medianregion, thefrontalarea,canbeeasily seeninthemiddlelinejustback oftheclypeus,andoftenthereis animpressedline,thefrontal groove,extendingbackfromthi- areaoverthemiddleofthefront.

Thefrontalregionpasseswith-

outdefiniteboundaryintothe vertexandtemples,theformer extendingposteriorly,thelatter lyingaboveandbehindtheeyes.

Theshortregionbetweenthe

vertexandthenarrowopening, orforamenthroughwhichthe alimentarytractandnervous systempassintothethorax,may becalledtheocciput.The cheeks,orgenae,comprisethe portionsofthecraniumanterior totheeyesandlateraltothe frontalcarinae.Theventralpor- tionofthehead,boundedin frontbythelabium,onthesides bythecheeksandextendingto theoccipitalforamen,isthe throat,orgula.Itiswell-de- velopedintheantsandisusually dividedintotwoequalhalvesby alongitudinalsuture.

FIG.4.Externalstructureofhead

inMyrmicanibraworker.(Janet.)A,

Dorsalaspectofhead;B,anterior

aspect;a.mandible;b,clypeus;c, frontalarea;d,frontalgroove;e, frontalcarina;/,vertex;g,occiput; h,temple;/,baseofantennalscape; k,cheek:/,eye;ni,lateralocellus;n, medianocellus;o,tentorialpit;p,quotesdbs_dbs14.pdfusesText_20