JImzJbE"t､，５９(1)：125-140．March2S，１９９１

PhylogeneticStudyoftheFamilyFormicidaebasedon theSensillumStructuresontheAntennaeand LabialPalpi(,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＋)＋） ＋(＋))＋)＋・Thecloserela- tionshipbetweenCerapachyinaeandDorylinae，andthatbetweenProceratiiniand EctatomminiofPonerinae，bothofwhichwereproposedbyearlierauthors，werenot supportedThevalidityofPonerinaetobeplacedasamonophyletictaxonwasques- tioned Keywords：Phylogeny；Formicidae；antennae；labialpalpi；sensilla．

Introduction

TheFormicidae arecurrentlydividedintotensubfamilies：Myrmeciinae， Pseudomyrmecinae，Formicinae，Dolichoderlnae，，Doryllnae，Formicinae，Dolichoderinae，Aneuretinae，Dorylinae， Leptanillinae,Cerapachyinae,PonerinaeandMyrmicinae(WHEELER＆WHEELER， 1972)．Severalauthorshavetriedtoclarifythephylogenyofthesesubfamilies， buttheconclusionsreachedbytheseauthorsvariedtoagreatextent(e､9.,WHEELER， 1928;BRowN，1954;TAYLOR，1978,etc.)．Itisnecessarytoscrutinizedifferent hypothesesproposedforthehigher-1evelphylogenyofFormicidae・ Thescanningelectronmicroscope(SEM）offeredmanynewmorphological charactersforsystematicstudyofinsects、sensillumstructureontheantennae andotherappendagesareincludedinthesecharacters,whichcanprovideusefulin‐ formationtoreconstructtheirphylogeny,ashasbeendemonstratedbyKRzYwIEｃ (1968),MEINEcKE（1975),BoHARTandMENKE（1976),WALTHER（1979),BARLIN andVINsoN(1981),etc・Ａ１thoughneverhasthephylogeneticsignificanceofthe sensillumstructuresbeenrealizedinFormicidae，thesestructuresareexpectedto 126 YoshiakiHAsFnMoTo

beofvalueinphylogeneticanalysisofthisfamily(HAsHIMoTo，1990)． Thepresentstudyanalyzedthesensillumstructuresontheantennaeandlabial palpiofFormicidaetotesthypothesesaboutthehigher-1evelphylogenyofthis familybynewcharacters・Particularemphasiswasplacedonthesubfamily ，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kmj WHEELER，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rjaraALLENetJAYNEs Mutillidae：Ｍｲが肋ｅ”Opaeα〃kadoCAMERoN,Ｍ)ｗｍｓα〃igrq/､Sc〃αYAsuMATsu Pompilidae：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/"ｓＰEREz Andrenidae：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"sSMITH Apidae：〃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 1２８ YoshiakiHAsHIMoTo

Table２．Distributionofcharacter-statesamongsubfamiliesinFormicidae・ Characternumbersandcharacterstatesaredenotedintext．

CharacterNo． １２３４５６７８９１０１１１２１３１４１５ ００００１１１ ０００００１１ ０００００００ ０００００００ ０００００００ ００１１１１１ ０１０００００ １．１ ００１１０００ ０１０００００ ００００００１ Ｏ０００００ｂ ０００００００ Formicinae ０ ａ（Ｕ刊，、）〈Ｕ〈Ｕ（０ Dolichoderinae １ Myrmeciinae １ Pseudomyrmecinae １ Dorylinae １ Leptanillinae １ Cerapachyinae ｌ １ Ponerinae １１１１２２ １１１１１１ ００００００ ００００００ ００１１００ ０１００００ ０００００１ １１１１１１ １１００００ 〈Ｕ（Ｕａａｎ〉〈Ｕ ００００００ ００００００ ００００００ １１１１１１ Amblyoponini １０００１１ Proceratiini Ponerini １．１ Odontomachini Ectatommini Myrmicinae

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・Thedistributionofthecharacterstateswithin FormicidaewasshowniｎＴａｂｌｅ２．

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、） 130 YoshiakiHAsHIMoTo

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，，，possessinganencirclingandamiddle66 cuticularledges(Fig.１１)．Sincetheseledgesarefeaturescharacteristictothepore Plaf尼sensilla（Fig.１０)，thesensillatrichoideacurvatacanberegardedasa homo10gueofsensillaplacodea（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"ｅｓ ｓｐ.,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、） 1３２ YoshiakiHAsmMoTo

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，basalsegmentsofantennal fUniculus，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、） 134 YoshiakiHAsHIMoTo

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 1３６ YoshiakiHAsHIMoTo

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),asindicated above(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 1３８ YoshiakiHAsHIMoTo 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：，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）