JImzJbE"t､，５９(1)：125-140．March2S，１９９１

PhylogeneticStudyoftheFamilyFormicidaebasedon theSensillumStructuresontheAntennaeand

LabialPalpi(Hymenoptera,Aculeata）

YoshiakiHAsHIMoTo

DepartmentofGreenEnvironment，DivisionofEnvironmentalSciences，

GraduateSchoolofScienceandTechnology，ＫｏｂｅUniversity，

Rokkodai，Kobe，６５７Japan

AbstractThepresentstudyanalyzedthesensillumstructuresontheantennaeand

labialpalpiofFormicidaetotesthypothesesproposedforthehigher-1evelphylogeny

ofthisfamily・Sensillumstructuresexaminedwerethesensillabasiconica，sensilla

trichoideacurvata，sensillacoeloconica，sensillaampullacea，setifOrmsensillaand

button-likesensilla、Acladisticanalysisoftherelationshipsofninesubfamilieswas

madebasedonfifteenmorphologicalcharacters・Plesiomorphicandapomorphicstates

ofthecharactersweredeterminedbyoutgroupcomparisonwithnon-formicidaculeates・

Therecognizedpatternsinnestingrelationshipswere:((((Leptanillinae＋Cerapachyinae

＋AmblyoponiniofPonerinae＋ProceratiiniofPonerinae)＋(PoneriniofPonerinae＋

OdontomachiniofPonerinae)＋(EctatomminiofPonerinae＋Myrmicinae)＋Dorylinae）

＋(Pseudomyrmecinae＋Myrmeciinae))＋Dolichoderinae)＋Formicinae・Thecloserela-

tionshipbetweenCerapachyinaeandDorylinae，andthatbetweenProceratiiniand

EctatomminiofPonerinae，bothofwhichwereproposedbyearlierauthors，werenot

supportedThevalidityofPonerinaetobeplacedasamonophyletictaxonwasques-tioned

Keywords：Phylogeny；Formicidae；antennae；labialpalpi；sensilla．

Introduction

arecurrentlydividedintotensubfamilies：Myrmeciinae，

Formicinae，Dolichoderinae，Aneuretinae，Dorylinae，

TheFormicidae

Pseudomyrmecinae，Formicinae，Dolichoderlnae，AneuretInae，Doryllnae，

Leptanillinae,Cerapachyinae,PonerinaeandMyrmicinae(WHEELER＆WHEELER，

1972)．Severalauthorshavetriedtoclarifythephylogenyofthesesubfamilies， buttheconclusionsreachedbytheseauthorsvariedtoagreatextent(e､9.,WHEELER，

1928;BRowN，1954;TAYLOR，1978,etc.)．Itisnecessarytoscrutinizedifferent

hypothesesproposedforthehigher-1evelphylogenyofFormicidae・

Thescanningelectronmicroscope(SEM）offeredmanynewmorphological

charactersforsystematicstudyofinsects、sensillumstructureontheantennaeandotherappendagesareincludedinthesecharacters,whichcanprovideusefulin‐

formationtoreconstructtheirphylogeny,ashasbeendemonstratedbyKRzYwIEｃ

(1968),MEINEcKE（1975),BoHARTandMENKE（1976),WALTHER（1979),BARLIN

andVINsoN(1981),etc・Ａ１thoughneverhasthephylogeneticsignificanceofthesensillumstructuresbeenrealizedinFormicidae，thesestructuresareexpectedto

126 YoshiakiHAsFnMoTo

beofvalueinphylogeneticanalysisofthisfamily(HAsHIMoTo，1990)． Thepresentstudyanalyzedthesensillumstructuresontheantennaeandlabial

palpiofFormicidaetotesthypothesesaboutthehigher-1evelphylogenyofthis

familybynewcharacters・Particularemphasiswasplacedonthesubfamily

Ponerinae，becausethesubfamilyisextremelyheterogeneousandpoorlyde‐ lineatedonthetraditionaltaxonomiccriteria．

MaterialsandMethods

Taxaexamined：ThespeciesexaminedwerelistedinTable1．Ｂｅｃａｕｓｅｔｈｅ

ｗｏｒｋｅｒｉｓｂｙｆａｒｔｈｅｂｅｓｔknownandmostavailablecaste，workerspecimenswere investigate。、Femalespecimensofsomeotheraculeateswerealsoexaminedfor

theoutgroupcomparison、BecausethesubfamilyAneuretinaeｃａｎｎｏｔｂｅｅx-

amined，itwasnotincludedinthisanalysis、

ThehigherclassificationsystemofAculeataproposedbyBRoTHERs（1974） andtheformicidc1assihcationbyWHEELERandWHEELER（1985）wereusedas guidelines、

Phylogeneticmethod：Thephylogeneticanalysiswasmadeaccordingtothe Hennigianmethod（1966)，wherethephylogeneticrelationshipsofthetaxaare

analyzedbasedonsharedderivedfeatures(synapomorphiesLbutnotonshared primitivefeatures（symplesiomorphies)，noronindependentlyderivedfeatures・Todeterminecharacterpolarities，outgroupcomparisonwasadopted，Thisis summarizedasfollows：Ｗｈｅｎｔｗｏｏｒｍｏｒｅｓｔａｔｅｓｆｏｒａｇｉｖｅｎｃｈａｒａｃterwere observedwithinagroup，themostwidespreadstateinthecloserelativesofthe

groupshouldberegardedasaplesiomorphicstate(cfWILEY,1981)．Accordingly， thepolaritiesofcharacterstatesｗｅｒｅassignedbasedonthecomparisonwiththose foundintheotherAculeata・

SEMobservation：TheantennaeandlabialpalpiwereeｘａｍｉｎｅｄｂｙＳＥＭ、Thesampleswerecleanedbyanultrasonic-washerinthechlorofOrm-methanol(２：１)， driedinair，mountedonstubsandthensputteredwithgoldToshowtheinner aspectofthecuticularstructures，ｔｈｅｓａｍｐｌｅｓｗｅｒｅcutwitharazorbladeand

thecellularmaterialweredigestedwithl０％KOHbeforethegoldcoatingwas applied

Thetermino10gyfOrsensillummorpho10gyfollowedafterWALTHＥＲ(1979)． AbbreviationsusedfOrfigureswere：ａ:aperture;ａｍ:sensillaampullacea;ｂ：

sensillabasiconica；ｂｒ：bristle；ｂｕ：button-likesensilla；ｃｈ：sensillachaetica；ＣＯ： sensillacoeloconica；ｄ：duct；ｅｌ：encirclingledge；ｇ：groove；ｉｍ：internalbasal membrane;ｍｌ:middleledge；ｐ：sensillaplacodea;ｓ：setiformsensilla；ｓｏ：socket； t：sensillatrichoideacurvata．

PhylogeneticStudyofFormicidae 127

TableLListoftaxaexamined．

Vespoidea：ＦｏｒｍicifOrmers Formicidae

Formicinae：AcrOpyga6aod[Joe"s応TERAYAMA,Ａ"OpﾉoﾉBpjs/o"gゎａｓ(JERDoN),QJmpo"０t"ｓノiabe1℃jFoREL,Ｃ/(Zpo"jc"sMAYR,Ｃ､〃jppo"jc"sWImELER,ＥＣ肺Opmrsp.,ＦＷｍ〃んkmjWHEELER，Ｒノ(Jpo"jcaMoTscHuLsKY，Ｌ“伽ノヲmws（FABRIcIus)，Ｌ〃ｊｇｅ７（LINNAEus)，

Ｍｾﾉ｡pﾉioro"ｓｓｐ.，OecOpﾉiy肋ｓａｍａｍｇｄ伽FABRIcIus，Ｐｍｍ”c"伽ｂｏ"巾o"ｊ“（FoREL)，

ＰﾉZZgjoにｐｊｓｓｐ.，ＰＣかelg"ｓｓａｍ"〃YANo，ＰＣ()、αc肱ｄ/ｖｅｓＦ・SMITH，Ｐ７℃"o/Ｅｐなｓｐ.，〃o伽ｍｓｓｐ，

Dolichoderinae：Borhrjo"Uﾉｍｚｅｘｓｐ.，ＤＣﾉﾉcﾉiode'ｗｓｂ〃berc"ﾉ、"sMAYR，Djdom)ﾉ、?eｘノノoj

FoREL,TtJpj"o"!αｊ"`ｊｃ""zFoREL,712cﾉi"o"!)rmexgjbbos"s(WHEELER） Myrmeciinae：Ｍ)ｗ７ｚｅｃｊａｇ"ﾉCsα(FABRIcIus),Ｍｓｐ・

Pseudomyrmecinae：T1ermpo"e、αﾉﾙrbom"sWALxER，ＴＸｓｐ、

Dorylinae：肥"ｊα"MzeWcﾋﾞps(F・SMITH),Ａ/”meTERAYAMA,ＤＣび/"ssp、Leptanillinae：Leprα"ﾉﾉﾉZzﾉ(Zpo"/αzBARoNIURBANI

Cerapachyinae：αr(ZpacﾉZysbjMFoREL,Ｃ､ｓα"rerjFoREL,Ｃｓｐ・

Myrmicinae：Ａｐﾉiae"og“rer/iJmeﾉﾉαJ(F・SMITH)，QJmjoco"q【ｿﾉUJ〃"dZJ(MAYR)，Ｃ、α"ﾉUJC"ｓ

ｓｐ.,ｏｅｍａｒｏｇ印rermans冴加"MFoREL,印ｉか〃MexzzmelwsBRowN,KWdrな、"/伽BRowN，ＭＢｓ９ｏｒａｃｊｃ"〃"ｓ（Fr・SMITH)，Ｍ)ｗ?Tecj"α９ｍｍ〃ｃＭ７〃jppo"jaJWHEELER，Ｍ)′｢〃αＪ

ノ“e"sjsFoREL，Ｏﾉﾉgo”ｍｚｅｘｓα"rerノFoREL，〃"'“/'ｗｍａｃｕｚ"j"αBRowNetBoIsvERT，Ｐﾉiejdoだ〃ノビノノSANTscHI，ＰﾉiejcjOgao〃ｓｐ.，Ｐｒ伽oﾉｱZyl・ｍｅｘｐ""ge""sMAYR，Ｓ〃ﾉﾉi伽｢"、αノCZpo"ｊｃｚＪ（ITO)，soに"OpsLssp.，Ｓ/'wmjgelUAsにw〃CAMERoN，〃mmorj"ｍｃａａＳＰ加加（LINNAEus)，リノＭＥＭｏｙｍｅｍｅびjWHEELER

Ponerinae：Amblyoponini：Ａｍｂｂ'Opo"ｅα"ｓｒｍﾉﾊﾞsERIcHsoN，Ａｓｊﾉvelrrjj（WHEELER） Proceratiini：〃ocem加加ノ(Zpo"に"mSANTscHI，Ｐ・ｗａｒ“eｊ（WHEELER)，Ｐ、ノroノFoREL，Djscorhy”ａｓ｡"rerノFoREL，、．ｓp・Ectatommini：Ｇ"αmprogemAscosrara（EMERY)，Ｒﾉﾛyr〃Opo"ｅｍｐ"、"だａ（EMERY)，Ｒ・ｓｐ・Ponerini：Ｂｍｃﾉ！〕lpo"ｅｍｃｈｊ"e"血（EMERY)，Ｏ〕lpmpo"ｅｍｍｅｒｊ(WHEELER),Ｄｊａａｚｍｍａｓｐ.,ｍｒｏｍｏｍ)ﾉrmexﾉαvα""sMAYR,HDlpOpo"era〃収po"α(SANTscHI)，Ｏdo"rOpo"ｅｍｒｍ"sve応α(Fr､SMITH),ＰＣ"e、ﾉ〃o"jcaWHEELER,Ｐ・sazbmWHEELER，７ｍcﾉiymesOp"ｓｐﾉﾉOsjor(WHEELER)，ＴＸｓ/zα/mFoREL，T1rzJpeZjOpe/ｒａｓｐ、Odontomachini：Odo"romac〃ｓｍｏ"rjcMzEMERY，０．ｓp､，Ａ〃oc/zer"ｓｓｐ・

Vespoidea：VespifOrmes

Tiphiidae：ＭＥｵﾉiocﾉiα/qpo"jcaYAsuMATsu,７Zp"ｊＭｚ伽rjaraALLENetJAYNEsMutillidae：Ｍｲが肋ｅ”Opaeα〃kadoCAMERoN,Ｍ)ｗｍｓα〃igrq/､Sc〃αYAsuMATsuPompilidae：Ａ"Opjmssamarje"sjF(PALLAs),qlpﾉio"o'U2xao応αﾉﾊﾞ,(LEPELETIER）Scoliidae：armosco"α〃Zeﾉﾋz"osomα/tzFcjarα(SMITH),Ｍ２ｇａｃα”somerJsgrossα〃zα“ｍｚｍｚＪ

（BETEREM）

Masaridae：町e"dOm“αｒＪｓｃｏｑｗｊﾉﾉErrjRoHwER

Eumenidae：Djscoeﾉﾉ"MJpo"ｊｃ"sPEREz,Ｅ"ｍｅ"assp

Vespidae：ＰＣ肋raschj"e"sjMJ"花"α肋PEREz，P70veSpasp，

Sphecoidea

Sphecidae：ＡｍｍＯｐｈ加地/臼ｓｒａＦ・SMITH,Ａ叩"にｘｄ伽eaor(THuNBERG）

Colletidae：ＣＭｅｒａｓｐａｔｅ肋/"ｓＰEREzAndrenidae：Ａ"cﾉ7℃"α加放adOSTRANDetYAsuMATsu

Halictidae：Ｌ“jogﾉｏ３,"ｍｓｐ、Melittidae：Ａ化"r、ﾉ(Zpo"jcaYAsuMATsuetHIRAsImlA

Megachilidae：Ａ〃rhjd"msepremqpJ"ommLEPELETIER

Anthophoridae：Ｘ)ﾉﾉocOpα(Zppe"djc"ﾉUmJcjrc"'、ﾉoﾉUJ"sSMITHApidae：〃jgo"αsp

Bethyloidea

BethyUdae：Ｃ２ｐｈａﾉO"ｏｍｍｇα"icMz(AsHMEAD） Chrysididae：Ｃ/z'”jss"α"g〃ｊｅ"sjsSMITH

Cleptidae：αどpressp、Dryinidae：HtIpﾉogo"αｍｐ"ｓ(Zpjcα/isPERKINs

YoshiakiHAsHIMoTo 1２８

Table２．Distributionofcharacter-statesamongsubfamiliesinFormicidae・

Characternumbersandcharacterstatesaredenotedintext．

１２３４５６７８９１０１１１２１３１４１５ CharacterNo．

００００１１１

０００００１１

０００００００

０００００００

０００００００

０

１

１

１

１

１

１ ００１１１１１

０１０００００

ａ（Ｕ刊，、）〈Ｕ〈Ｕ（０

１．１

００１１０００

０１０００００

００００００１

Formicinae

Dolichoderinae

Myrmeciinae

Pseudomyrmecinae

Dorylinae

Leptanillinae

Cerapachyinae

Ponerinae

Amblyoponini Proceratiini

Ponerini

Odontomachini

Ectatommini

Myrmicinae

Ｏ０００００ｂ

ｌ

０００００００

１１１１２２

１１１１１１

００００００

００００００

００１１００

０１００００

０００００１

１１１１１１

１１００００

〈Ｕ（Ｕａａｎ〉〈Ｕ

１．１

００００００

００００００

００００００

１１１１１１

１０００１１

Results

Thestructuralfeaturesoffourtypesofantennalsensilla(sensillabasiconica，

sensillatrichoideacurvata,sensillacoeloconica,andsensillaampullacea)andtwo

typesoflabialsensilla(setiformsensillaandbutton-1ikesensilla)wereanalyzedin theformicidsandinotheraculeates、

Ｔｈｅmorphologicalcharacteristicsandcharacterstatesofthesesensillum

structuresweredescribedasbelow,where(0)meansplesiomorphiccondition,(1)，

(１ａ)，（1ｂ)ａｎｄ(2)apomorphicconditons,ａｎｄ(Ｏｔｏｌ)，（Ｏｔｏｌａ)，（Ｏｔｏｌｂ）ｏｒ

(Ｏｔｏ２）transformationseries・ThedistributionofthecharacterstateswithinFormicidaewasshowniｎＴａｂｌｅ２．

Antennae

Se"sﾉﾉﾉZJb“jco"ｊｃａ，

Thistypeofsensillaalwaysconsisｔｓｏｆｔｗｏｐａｒｔｓ，apegandasocketwhich

isthemembraneconnectingthepegwiththeantennalcuticle(WALTHER，1981；

MARTIN１，１９８６ａ)(Figs1and6)．

Characterl(Ｏｔｏｌａ;Ｏｔｏｌｂ）

Pegstoutandalmoststraightwithablunttip-(O)(Fig.２),slender,narrowed

apicallyandbasally,withtheapexpointing-(1ａ)(Fig.３),orthinandgreatly

bent-(1ｂ)(Fig.４)．

Character2(Ｏｔｏｌ）

PhylogeneticStudyofFormicidae 1２９

Pegshort（＜20瓜、)－(0),ｏｒlong（＞30瓜、)－(1)(Fig.５)．Character3(Ｏｔｏｌ）

Socketflushwiththeantennalsurface-(O),orgreatlyelevatedabovethesur-

臼ﾄｽ羅耀７鰯露

３－ 》■■》口髭一

Figs，１－８．Thesensillabasiconica,－１－５.Pegsofthesensillabasiconica:１，TXPﾉｾﾉﾛﾉ、応"､/α

（Tiphiidae);２，，０ﾉﾌﾉﾉ"ssp.(Dorylinae,Formicidae);３，Ｃ'yprOpo"esα"reri(Ponerinae,For-

micidae);４，αｍｐａｃﾉzjAssum花ｒｊ(Cerapachyinae,Formicidae);５，Ｐﾉｶejdo/ePjeﾉﾉ(Myrmicinae，

Formicidae)．－６－８．Socketsofthesensillabasiconica：６，７１ＧｍＩＰＯ"e、α肋bom"ｓ

（Pseudomyrmecinae，Formicidae)，longitudinalsectionofthesocket；７，〃jdO'7U′ｍｚｅｘｊｒｏｊ

（Dolichoderinae,Formicidae);８，internalbasalmembraneofthesensillabasiconica．α、‐

Ｐａｃﾉ！)Ｆｂｊｒｏｊ(Cerapachyinae,Formicidae),pegbaselongitudinallydissectedtoshowinternal

basalmembrane．（Scalebar＝2.5瓜、）

YoshiakiHAsHIMoTo 130

face－(1)(Fig.６)．

Character4(OtoD

SocketHat-(0),orwithraisedrim-(1)(Fig.７)．

Character5(Ｏｔｏｌ）

Internalbasalmembraneabsent-(0),orpresent-(1)(Fig.８)．

Se"sﾉﾉhJかjchojdeac"ｒｗＪｒａ・

Thissensillumisacurvedhair，，，possessinganencirclingandamiddle 66

cuticularledges(Fig.１１)．Sincetheseledgesarefeaturescharacteristictothepore Plaf尼sensilla（Fig.１０)，thesensillatrichoideacurvatacanberegardedasahomo10gueofsensillaplacodea（WALTHER，1981；MARTINI＆SCHMIDT，1984； MARTIN1,1986ｂ)．

Character6(OtoD

Platetype-(O)(Fig.８),orhairtype-(1)(Fig.１１)． Character7(Ｏｔｏ２）

Thebasalapertureofthehairextenｄｅｄｆｒｏｍｔｈｅｐｒｏｘｉｍａｌｅｎｄｎｅａｒｔｏｔｈｅ upperinnercornerofcurvedhairapparatuses-(O）（Fig.１２)，showedin-termediateinthestatebetween(0)ａｎｄ(2)－(1),orneverextendedbeyondthe

base-(2)(Fig.１３)．

Character8(OtoD

Hairinsertedintotheantennalcuticledeeply-(O)(Fig.１４),orshallowly-(1)

（Fig.１５)．

Se"sjﾉﾉﾋzcoe/OCC"にα・

Thesensillacoeloconicaarepegswithlongitudinalgrooveinthechamber

belowtheantennalsurface(Fig.１８)．

Character9(OtoD

Openingsmall(diameter＜3.0〃、)－(O)(Fig.１６)，ｏｒlarge（＞4.0,ｕｍ)－(1)（Fig.１７)．

CharacterlO(Ｏｔｏｌａ)(Ｏｔｏｌｂ）

Chambertube-form-(O)(Fig.１９)，fUnnel-form-(1ａ)(Fig.２０)，ｏｒｂｏｔｈ－

（1bl

Characterll(Ｏｔｏｌ）

Surfacegroovesofthepegnotextendingbasallybeyondthemidpoint-(O） （Fig.１８),orreachingnearlythebase-(1)(Fig.１９.）

Se"sﾉﾉhJα"ZpMZJcea，

Thesensillaampullacearesemblethecoeloconicsensilla,butaredistinctinthe chamberbeingmoredeeplysituated(Fig.２１)anddifferentsurfacestructureofthe sensehairinthechamber(Fig.２４)．

PhylogeneticStudyofFormicidae 1３１

Figs、９－１５．－９－１０.Thesensillaplacodea:９，DiFcoeﾉﾉ"s/叩o"jczｲs(Eumenidae);10,助ｍｅ"ｅｓｓｐ.,inneraspect、－１１－１５.Thesensillatrichoideacurvata：11,Polye堰"ｓｓａｍ"〃(For-micinae,Formicidae),longitudinalsectionofhair;12-13,hairbaseaperturesofthesensilla

trichoideacurvata,12,Ctm1po"omMIpo"ｊｃ"s(Formicinae,Formicidae),hairbasedissected toshowtheaperture;13,Ａpﾉme"og“rer/bｍｅ/ｊｃａ(Myrmicinae,Formicidae),hairremovedto

showtheaperture)；14-15,hairinsertionofthesensillatrichoideacurvata,14,07ｍPC"o'"ｓ

ノ(ＪＰＯ"ｊｃ"３（Formicinae，Formicidae)；１５，αｍｐａｃﾉDAFｂ伽ノ（Cerapachyinae，Formicidae）（Scalebar＝2ｓ似、）

YoshiakiHAsmMoTo 1３２

弓匡溺

Figs，１６－２４．－１６－２０．Thesensillacoeloconica：１６－１７，externalopeningsofthesensilla

coeloconica，１６，Ｌ“ｊ"ｓ〃Jger（Formicinae，Formicidae)；１７，７１ec〃o、)w7zexgjbbos"ｓ

（Dolichoderinae,Formicidae)；18-19,longitudinalsectionsofinnerchamberofthesensilla

coeloconicashowingthepeg，18,CremarogmrermaLs"、"raj(Myrmicinae,Formicidae)；１９，

０ｺlptOpo"esamerj(Ponerinae,Formicidae),notethatpeg-surfacegroovesrunfromthetip

neartothebase;20,ｍnerviewsofthesensillacoeloconiＣａ,ＭＥﾉOp/joro"ｓｓｐ．（Formicinae，

Formicidae)．－２１－２４．Innerviewsofthesensillaampullacea：２１，Ａ、p此xdissecror

（Sphecidae)；２２，Ｌ“伽〃jger（Formicinae，Formicidae)；２３－２４，Pro“、伽、ノ(Zpo"jc"ｍ

（Ponerinae,Formicidae),24,chamberdissectedtoshowinnerhair・（Scalebar＝Ｚ５Ｊｕｍ）

PhylogeneticStudyofFormicidae 1３３

Figs、２５－３１．－２５－２６．Distributionpatternofsensilla：２５，basalsegmentsofantennalfUniculus，Mm77eα"α９ｍ〃"ｊｃＭｚ〃仰o"jａＪ（Myrmicinae，Formicidae)，、otethatthe

segmentshavebristlesonly;26,thebristlesdissectedtoshowthebase,Ｓ〃rhZsrlwmaﾉ(Ipo"ｊａＪ（Myrmicinae,Formicidae),notethatthebristleshavenoaperturesatthebaseofcuticular

apparatus､－２７－３LLabial-palpsensilla:27-28,Ｍｲ〃αｅ"rOpaeα〃ｋａａｏ(Mutillidae)，２８，close-upofthetipoflabialpalp，notebutton-likesensillum；２９，Ｍｙｒｍｅｃｊａｇ"ﾉCsα

（Myrmeciinae，Formicidae)；３０，OyPrOPo"Ｃｓα"ｒｅｒｊ（Ponerinae，Formicidae)；３１，月o‐Ce、伽ｍｗａｍｓｅｊ(Ponerinae,Formicidae),close-upofthetipoflabialpalpshowingbutton‐likesensillum．（Scalebar＝2.s似、）

YoshiakiHAsHIMoTo 134

Characterl2(Ｏｔｏｌ）

Theinternalchamberflask-form-(0)(Fig.２２),ortube-form-(1)(Fig.２３)．

PZJrre'〃ｑ/・se"sﾉﾉﾉ、α肋伽rjo"・

Characterl3(Ｏｔｏｌ）

Thesensillaofalltypesshowedratherhomogenousdistribution-(O)，ｏｒ

concentratedinapicalsegements-(1)(Fig.２５)．

Labialpalpi

Sa肋r"zse"sﾉﾉﾉﾛ・

Theseareslenderhairspointingsharply(Fig.２７)．

Characterl4(OtoD

Thepalpuscoveredwiththesetiformsensilladensely-(0）（Fig.２９)，or

sparsely-(1)(Fig.３０)．

Ｂ"がoﾙﾉﾉｋｅｓｅ"sﾉﾉﾉ[7．

Ｔｈｉｓｓｅｎｓｉｌｌｕｍｉｓａｐｅｇｓｅｔｉｎａｓｏｃｋｅtonthetipoflastlabial-palpsegment

(Fig.２８)。

Characterl5(OtoD

Thepegshort（＜５〃、)－(O)(Fig.３０),ｏｒlong（＞５〃、)－(1)(Fig.３１)．

Discussion

Thesensillumstructuresstudiedhereprovidedsynapomorphicstatesforthe

followingnestedpatternsofrelationshipsamongtheformicidsubfamilies、

EctatomminiofPonerinaeandMyrmicinaeweregroupedtogetherbasedo、

thelongpegofthesensillabasiconica(character2)andsmallhair-baseapertureof

thesensillatrichoideacurvata(character7)．ThoughAmblyoponiniofPonerinae

alsohadthelongpegsofthesensillabasiconica(seeTable2),ｔｈｉｓｍａｙｂｅｃｏｎ‐

vergentwiththetwotaxamentionedabovebecauseacloserelationshipbetween

Amblyoponiniand(Ectatommini＋Myrmicinae)wascontradictedbythecharac-

terstatesofothersensillumstructures（characters7andl5)．Poneriniand

OdontomachiniofPonerinaeweregroupedtogetherbyauniquepegshapeof

thesensillabasiconica(characterl）andauniquesurfacegrooveconditioninthe

pegsofthesensillacoeloconica(characterll)．Leptanillinae,Cerapachyinae,and

AmblyoponiniandProceratiiniofPonerinaeweregroupedtogetherbythe

elongatedpegofthelabialbutton-likesensilla（characterl5)．Althoughrela‐

tionshipsamongthesethreegroups（Ectatommini＋Myrmicinae，Ponerini＋

Odontomachini，Leptanillinae＋Cerapachyinae＋Amblyoponini＋Proceratiini）

PhylogeneticStudyofFormicidae 1３５

werenotclarihedinthisstudy，thesegroupsandDorylinaeweregroupedbya synapomorphy，ノ.ａ，ｓｍｏｏｔｈlabialpalpiwithreducedhairs（characterl4)．MyrmeciinaeandPseudomyrmecinaeformedagroupbasedonthestrongly elevatedsocketofthesensillabasiconica(character3),whichisthesistergroupof Dorylinae＋Leptanillinae＋Cerapachyinae＋Ponerinae＋Myrmicinae,basedonthe hairsinsertedshallowlyintotheantennalcuticleofthesensillatrichoideacurvata

(character8)．Dolichoderinaewasthesistergroupofthemonophyleticcompo-nentconsistingoftheabovesevensubfamiliesbasedonthetransformationseries

ofthehair-baseapertureofthesensillatrichoideacurvata(character7),andthese

togetherconstitutedthesistergroupofFormicinaebasedonthehair-likeplates

(ｊ､ａ，thesensillatrichoideacurvata)（character6)．Asthesensillaplacodeaof

Hymenopterahavebeenbelievedtohaveevolvedfromhair-typesensilla(VoGEL，

1923),thesensillatrichoideacurvatawerepreviouslyinterpretedasprecursorsof

thesensillaplacodea(WAcKER，1925)．Thishypothesis,however,mustassume

independentoriginofsensillaplacodeawithineachlineageofAculeata，because

theplatesensillawerewelldevelopedintheotheraculeateswhichmustbeearlier

derivedgroupsthanFormicidae，suchasBethyloideaandSphecoidea（Ｃｆ・

BROTHERS,1974)．A1ternatively,ifthesensillaplacodeaevolvedonceintheAculeata，

thehair-1ikeplateofFormicidaｅｍｕｓｔｂｅｔｈｅｒｅｓｕｌｔｓｏｆｔｈｅｒｅｖｅｒｓａｌevolution，

Thus，themostparsimoniousinterpretationfortherestricteddistributio、ofthe

sensillatrichoideacurvataonlywithinFormicidaewouldbethatthesesensillaare

derivativesoftheplateofsensillaplacodea，ｊ､e,，thehair-1ikeplatewasanapo-

morphicstateintheFormicidae，

Thebasicconceptforphylogenyofformicidsubfamilieshadbeentreatedin

threedifferentmanners(e､9.,WHEELER,1928;BRowN,1954;TAYLOR,1978)．Based

ontheabovenestedpatternsofrelationshipsamongthesubfamilies，thebasic

conceptsforformicidphylogenywerescrutinizedWHEELER(1928)hypothesized

thattheponerinesasthemostprimitivegroupfromwhichtheothersubfamilies

directlydiverged・DespitenoconvincingevidencegivenbyWHEELER(10c・Cit.)，

REＩＤ(1941)haspointedoutthatponerinesdisplaymostdiversethoraciccharac-

ters,whichprobablysupportstheWHEELER,shypothesis、However,thereexistsno

reasonwhyanancestralgroupshouldbemostdiversifiedinmorphology，Even

though,ｆｏｒexample,thefossilrecordsuggeststhattheｍｙrmicinesareconsidered

asthemostrecentlyderivedgroupoftheformicids(cfBRowN,1973),theydisplay

byfarthegreatestdiversityinmorpho1ogysuchasofmouthpart(GoTwALD,1969）

andlarvalstructures(WHEELER＆WHEELER,1976)．Thesensillumcharactersdid

notsupportWHEELER(1928),becauseacceptingthehypothesisofponerineorigin

forotherformicidsrequirednumbersofunlikelyreversionsincharacter-state

evolutionofthesensillumstructures(charactersl,２，７，８，１１，１４ａｎｄｌ５)．

ＢＲｏｗＮ(1954)proposedthatthefOrmicidssplitearlyintheirevolutioninto

twophylands：thePoneroidphylandincludingPonerinae，Myrmicinae，Dory-

1inae，LeptanillinaeandCerapachyinae，ａｎｄtheMyrmecioidphylandincluding

YoshiakiHAsHIMoTo 1３６

Myrmeciinae,Pseudomyrmecinae,DolichoderinaeandFormicinae・WILsoN,ｅｒａﾉ．

(1967),supportingBRowN(1954),emphasizedapossibilitythattheformicidsmay bederivedtwiceinevolution・ThoughBRowN（1954)ａｎｄＷＩＬｓｏＮｅｒａﾉ.（1967）

didnotexplicitlystatethecharacterdatadividingthePoneroidandMyrmecioid

phylands，itismostprobablefromtheirdiscussionthatthesetwophylandswere

establishedbytheoverallsimilaritiesoftheprimitivemembersofeachofthe

phylands，whichimpliesthattheymightbebaseｄｏｎｔhesymplesiomorphicstates・

Becausesymplesiomorphieshavenovaluefordeterminingthephylogeneticrela-

tionships,thetwophylandsproposedbyBRowN(1954)mayormaynotrepresent

phylogeneticreality，Thepresentstudyindicatedthatthemembersofboth

phylandssharedthefimdamentalhomogeneousstructuresinthesensillaandthat

alltheformicidsstudiedhereweregroupedtogetherbythepossessionofthesensilla

trichoideacurvata(seeTable2,character6)．ThedeepsplitproposedbyBRowN

(1954)intheformicidevolutionwas,therefore,notfoundinthesensillumstruc-tures・

TAYLOR(1978),modifyingBRowN(1954),transferredMWmecjaofMyrmeci-

inaebuttheothermyrmeciinegenusjVOrho"Zyrmeaa，andPseudomyrmecinae

fromtheMyrmecioidphylandtothePoneroidphylandHeproposedthathis

PoneroidandMyrmecioidphylands(whichwasrenamed``Formicioid,，)represent

separateandtwomajorlineagesintheformicidphylogeny・Thismodificationwas

basedlargelyoneitherfUsedorseparatescleritesofthe4thabdominalsegment・

ThemembersofPoneroidphylandandａｌｓｏＭ)w7zecjaofMyrmeciinaeand

PseudomyrmecinaehadfUsedsclerites，whereastheywereunfUsedintheother

membersofMyrmecioidphyland・However，theunfusedconditioninthe

memberｓｏｆｔｈｅＦｏｒｍｉｃｉｏｉｄｐｈｙｌａｎｄｍａｙｂｅｓｙmplesiomorphic，becausethisisa

conditionoccurringinmostaculeates・ThoughjVOrﾉio"ＺＷｍｅａａｗｅｒｅｎｏｔａｖａｉｌ－

ａｂｌｅｉｎｔｈisstudy，themonophylyofFormicinae＋Dolichoderinae，ａｔleast，was

notsupportedbythesensillumcharacters、Instead,thisstudyindicatedthatthese

subfamilieswerederivedindependentlyfromthemainformicidstockatveryearly

time，becausetheysharednoapomorphiccharactersandborerelativelyfew，but

unique,sensillumcharactersinderivedstates(seeTable2,characters4９＆１０)．

Consequently,eachofthethreephylogeniesfortheformicidsubfamilieswasnot

supportedbythisstudy・

Thepresentstudycanalsoshednewlightonsomeproblemspuzzlingformicid

systematists、WHEELER(1928)relatedPseudomyrmecinaeandMyrmicinaebased

ontwosegmentalelementsofthepetiole・However，twosegmentedpetioleoc-

curredindependentlyinPseudomyrmecinae，Myrmicinae，Myrmeciinae，some

Dorylinae，Leptanillinae，CerapachyinaeandsomePonerinae、ＢＲｏｗＮ（1954）

relatedPseudomyrmecinaetoMyrmeciinaeinsteadoftoMyrmicinae，buthedid

notprovideanyconvincingevidenceforthisrelationship、ThoughEINsNER

(1957)，supportingBRowN（1954)，indicatedthattheproventriculusofPseudo-

myrmecinaewassimilartothatofMyrmeciinae，ｔｈｅｓｉｍｉｌａｒｉｔｙｍａｙｂｅｄｕｅｔｏ

PhylogeneticStudyofFormicidae 137

symplesiomorphybecausetheirproventriculusisthesametypefoundinmost otherAculeata・ThesensillumcharactercorroboraｔｅｄＢＲｏｗＮ(1954),asindicatedabove(character3)．

ThecerapachyineshavebeenconsideredtobeancestraltoDorylinae,which servesasatransitionaltaxonfromPonerinaetoDorylinae(EMERY1895;WHEELER

＆WHEELER，1985),thoughREID（1941）didnotsupportthisviewbasedonthe

thoracicstructures，Thepresentstudysupportedneitherthecerapachyineorigin

ofthedorylinesnortheintermediateplacementofcerapachyinesbetween

PonerinaeandDorylinae・ＴｈｅｓｅnsiUumstructuresofcerapachyinesand

ponerinesweremorespecializedthanthoseofthedorylines・Particularly，ｔｈｅ

cerapachyineshaduniquelymodifiedpegsofthesensillabasiconica(characterl)，

aconditionthatthedorylinescouldnotariseRelatednessofcerapachyineswith

someponerines，suchasAmblyoponiniandProceratiini，wasproposedbythe

thoraciccharacters(REID,1941)andthemouthpartcharacters(GoTwALD,1969)． Thiswassupportedalsobythesensillumcharactersasincharacterl5・

EctatomminiofPonerinaewasconsideredastheancestralrelativeof

Myrmicinae，becauseMyrmicinaeretainedsomeprimitivecharactersofthe

ectatomminesandafossilgenusAgroeco"2〕ﾉﾉmexmayprovidedatransitional

for、(BRowN，1954)．However,noapomorphicstateswerefoundtoindicatea

sister-grouprelationshipbetweenthem・Thepresentstudyprovidedtwosynapo-

morphiesforMyrmicinae＋Ectatommini(characters2and7)． ＢＲｏｗＮ（1958）afIiliatedProceratiiniwithEctatommini，basedonthemor-

phologicalresemblancesobservedin月ocem伽mofProceratiiniandHCrero‐

po"emofEctatommini・However,WHEELERandWHEELER(1976)disagreedwith

him（1958)．TheyindicatedthatlarvaeofProceratiinidiffbredfromthoseof

HaerOpo"ｅｍａｎｄｆｒｏｍｔｈｏｓｅｏｆａｎｙｏｔｈｅｒｋｎｏｗｎｅctatommines・Although

Haempo"emhavenotbeenexamined，thepresentresultwasnotconsistentwith

BRwoN（1958）becausethesensillumstructuresofProceratiinidifIeredgreatly

fromthoseofEctatommini，particularlyinthestructuresofthesensillaampul-

1acea;innootherformicidtaxatheductlesschamberwasfound(characterl2)．

ThoughtheductlesschamberwascommoninAculeata,theconditioninProcera‐

tiinisuggesteditssecondaryderivation,ratherthanaretainedplesiomorphicstate，

becausethechamberofProceratiinihadstillmarkedlynarrowpart，unlikethe ductlesschamberofotheraculeates・

OdontomachiniofPonerinaeisthemostdistinctiveandeasilyrecognizedof

alltheFormicidaebyitsunusualheadandmandibles・Thoughthedistinctiveness

oftheheadandmandiblestendstoobscuretheirphylogeneticrelationshiptothe otherformicids,ＢＲｏｗＮ(1976)relatedOdontomachinitoPonerinibasedonsiｍｉ‐

laritiesobservedinthelarvae・ThesensillumcharacterofadultOdontomachini

andPonerinisupporteｄＢＲｏｗＮ（1976),asincharacterslandlL

Basedonthesensillacharacters,thefollowingsubfamiliesofFormicidaewere

delimited：FormicinaewascharacterizedbythecharacterlO，Dolichoderinaeby

YoshiakiHAsHIMoTo 1３８

thecharacters4and9，MyrmeciinaebythecharacterlO，Cerapachyinaebythe

charactersｌａｎｄ５，andMyrmicinaebythecharacterl3,thoughmorecomparative

datawereneededtodeterminethatthesestructuresareavailablethroughouttaxa・

ItshouldbenotedthatthevalidityofPonerinaewasquestionedinthisstudy， becausethesensillumcharactersindicatedthatexaminedmembersofthissub-

familyshouldbedividedintogroupssomeofwhicharecloselyrelatedtoother

subfamilies;group（１）AmblyoponiniandProceratiiniwhichconstituteamono-

phyleticcomponenttogetherwithCerapachyinaeandLeptanillinae，group（２）

PoneriniandOdontomachini，group（３）Ectatomminiwhichisthesistergroup

ofMyrmicinae・PonerinaeisthemostheterogeneousgroupofFormicidae，and

thecharacterstatesdelimitingthissubfamilymaybeplesiomorphicforFormicidae

(cfDALLAToRRE，1893；ＲＥID，1941；WHEELER＆WHEELER，1964；GoTwALD，

1969)．Sinceextensivevariationsinmorphologyandbehaviorexist,CLARK(1951）

evendividedtheponerinesintoseveralsubfamilies：Amblyoponinae，Dis-

cothyrinae，OdontomachinaeandPonerinae、AlthoughCLARK'sclassification

wasnotsupportedbythisstudyandthefinalconclusioｎｏｆｔａｘｏｎｏｍｉｃｓtatusfor

ponerinesmustawaitforfUrtherinvestigation，Iamtemptedtoconsider

Ponerinaeasasubfamilyunwarranted

Thepresentresultssuggestedthattheexistingconceptsofphylogenyand

highertaxaofFormicidaeshouldbechanged，Amorerealisticphylogenycanbe

approachedbytestingandretestingofnewcharacterssuchassensillumcharacters

usedinthisstudy．

Acknowledgments

IexpressmysinceregratitudetoDr．Ｍ・TAKEDA，forhiscriticalreadingof

EnglishtextandDr.Ｔ､NAITo，forhisreadingthroughthemanuscript・Iwishto

thankProf、ＳＭｏＭｏｌ，KobeUniversityforhisencouragement，Ｉａｍｍｕｃｈｉｎ‐

debtedtothefoUowinggentlemenfortheirkindnessinofferingvaluablematerial

andusefUlinformation：Ｍ・AzuMA，Ｒ､Ｈ、CRozIER，Ｍ、KuBoTA,Ｋ・ＭＡｓｕＫｏ，Ｋ・

OGATA，Ｒ､Ｗ・TAYLOR，Ｍ・TERAYAMA，ＪＲ・WALTHER，ＳＫ・ＹＡＭＡＮＥ，ａｎｄＫ・

ＹＡＭＡｕｃＨＬ

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(ReceivedJune7，1990;AcceptedDecemberl8,1990）