Taxonomic Characterization of Some Live-stem Inhabiting Azteca (Hymenoptera: Formicidae) in Costa Rica, with Special Reference to the Ants of Cordia (Boraginaceae) and Triplaris (Polygonaceae)

26 Pages·1996·8.6 MB·English

by  J T Longino

J.HYM.RES.
Vol.5,1996,pp.131-156
Taxonomic Characterization ofSome Live-stem InhabitingAzteca
(Hymenoptera: Formicidae) in Costa Rica, with Special Reference to
the Ants of Cordia (Boraginaceae) and Triplaris (Polygonaceae)
JohnT.Longing
TheEvergreenStateCollege,Olympia,WA98505
—
Abstract. In themorphologicalspacedefinedbyqueenheadlengthandheadwidth,seven
CostaRicanspeciesorspeciescomplexesintheantgenusAztecahaverelativelynarrow,subrec-
tangularheads(headlength21.3timesheadwidth),andallofthemsharecharacteristicnesting
bqeuheaevnioarndinwloirvkeesrt-ebmass.eTdhiedseentsipfeiccaiteisonangduisdpeesciaersecporomvpildeexde.sAarseubtsaextonoofmtihceaslelyspcehcaireasctaerreiczeodm,maonnd
inhabitantsofthespecializedantplantsCordiaalliodora(Boraginaceae)andTriplarismelacnodcndron
(Polygonaceae).AztecalongicepsisanobligateinhabitantofT.mdaenodendron,butisknownonly
fromtwomid-elevationPacificslopesites.InthePacificlowlandsT. melaenodendron isusually
inhabitedbyeitherA.beltiiorPseudomyrmexviduus,twospeciesthatarenotobligateinhabitants
Aofztpeacratipciuttliaerrianctompplalnetxs,cbounttaiinnsstetahdemcaoymmbeonfooubnldigianteaivnahriaebtiytaonftsdiofffeCroerntdiaantalpliloadnotra.speAciegse.neTrhael
descriptionofantcommunitycompositioninCordiaandTriplarisantplants,anddiscussionof1)
theadaptivesignificanceofqueencharactersinAzteca,2)problemsofspeciesdefinitionsasre-
vealedbythisstudy,3)possiblemechanismsgeneratingcomplexcharacterdistributionsintheA.
Tpiattxioerniocmoimcplcehxa,ngaensda4r)e:thAeztceocnatrabeslttiiingEmroelresyo1f89r3e,ginoneawlfsalautn.as[=anldaetgaloWbhaelerleevrisi1o9n4s2anreewprosyvni.d,ed=.
sSttoaltl.i;FAozrteelca19n1i2grniceawnssyFno.r]e;lAz1t8e9c9a,cnoerdwincstoalta.;FoArzetlec1a92p0a,trnueelwisstFaotr.e;lAz1t9e0c8a,jnureuwemsitsat.F;orAezlte1c9a04p,itntieerwi
Forel1899[=emarginatisquaniisForel1920newsyn.];AztecasapiiForel1912,newstat.
INTRODUCTION interpretationoffieldresultsand/oranun-
ctitdsnc,inohoopoeebn^Smie'ylps^mc1c1eep\uu9o9hcrin8ol.8,iac59.iouav&uagtsle,o,.yiiDuzWe,baasae1isnevdn9tl,ei9do-sfrn,1dafpeoanesalan,tivtoan-uofnn,tp1Wrrlt,el9e,eau1r9a&q.aha0rnuo)csotde.rtsMrnnioaacntMc,1strgK1io9hsysae9esousu1tc,pbytb,ieii.Lit^.oecaoro1nciFoh,lrf.9tseipoo9asingtoi3lcisyrh,as(ne,L-/(vsaoHeDBotrtcnoejauleoglandvnailtld:yea----J. dsh,"p^'^N"^"oelea,^'^craovT^"inet.'eh(^ats^.r1'te".to9'i^r^ei4^poge2emni°e)c^snac^",^e.nte's-i•'uir,'w,v„T^^oasheeanw"disjnC'C^oec'oodo^fl,rhra->iadt,ts'd.dnit,c^wsi.miltao'^oeov-nceop^atiasr.lc',anatps,a^cit.dtein^juni.cett°r1osin.yT's-netiray-sasiiwst,ron^ldswwhna^oCaerra-*.cinao^siist'lrunn.at,bovgJhp-hdoyhTololoinhasnvo,Wusne.afehtt„ieltatltt,Uhaaena..hhiasuxsd-ee-.t-t
1991, Davidson et al. 1991, Davidson & C. alliodora, are specialized ant-plants
Fisher1991,McKey1991).Studiesofthese (Wheeler 1942). Both species have pyn-
communitiesarehamperedbyalackofba- form cauline swellings at nodes where
sictaxonomyandnahiralhistoryoftheor- whorls of branches arise. These domatia
ganismsinvolved.Lackofnamesand/ora are hollow and are usually inhabited by
confusedstateofnamesimpedeseffective ants.CordianodosaisSouthAmerican.Cor-
communication ofresults. An inability to diaalliodoraiswidespreadinSouthAmer-
distinguishamongspeciescanleadtomis- ica and also extends through Central
132 JournalofHymenopteraResearch
America to southern Mexico. In Costa throughout the Neotropics, all of which
Rica,C.alliodoraistheonlymyrmecophy- have hollow stems that are inhabited by
ticCordia. Itisaverycommontreeinan- ants(Brandbyge 1986,Wheeler1942).The
thropogenic habitats, occurring along hollow stems are much like Cecropia or
roadsidesandinpastures.OnthedryPa- bamboo, withshortcylindricalintemodes
cificside, thetreesareusuallysmalland separatedbysolidsepta.Inmanypartsof
bushy, with crooked trunks. On the wet SouthAmerica,Triplaristreesaredominat-
Atlantic side, the trees are tall and ed by the Psendomyrtnex triplarinus com-
straight.Ithasthecommonname"laurel" plex,asetofatleastfourspeciesofobligate
and is considered a valuable timber tree Triplarisants(Ward1991).Triplarismelaen-
(Opler&Janzen1983). odendron subsp. melaenodendron {sensu
Wheeler(1942)consideredthefollowing Brandbyge 1986; often identified as T.
to be obligate inhabitants of Cordia allio- americanainearlierliterature;referredtoas
dora: "Azteca longiceps anditssubspecies, T. melaenodendron in this paper) occurs
A. pjittieri and its var. emarginatisquamis, fromMexicotosouthernCostaRica,andis
Pseiidoim/rma sericea and its varieties ita theonlyTriplarisspeciesinCostaRica.Tri-
and cordiae, and Ps. alliodorae." Wheeler plaris melaenodendron is moderately abun-
misidentified thematerialhecalled Pseii- dant on the Pacific side of Costa Rica,
domyrmex ita (Forel). True ita is a gener- where itmostoften occursalongstreams
alist inhabitant ofdead twigs, and isoc- indryforestareas(pers.obs.).
casionally found in Acacia thorns (Ward Wheeler (1942) examined numerous
1993). Thematerial Wheeleridentifiedas specimensofwhathecalledTriplarisamer-
"ita" isclosetoorthesameasP. cordiae icanainPanama,whichIassumetobeT.
(Forel) (Ward,pers.comm.),andisprob- cumingiana based on Brandbyge's (1986)
ablyanobligateCordiaant.Pseudoinyrmex revision. Wheeler found two ant species
alliodorae, describedbyWheelerfrom his heconsideredtobeobligates:Aztecamen-
studyofCordia,isajuniorsynonymofP. ceps and "Pseudomyrma loeivensohni". Az-
elougatits (Mayr),acommonandgeneral- tecametKepsisnotknownfromCostaRica
ized inhabitant of plant cavities (Ward (pers. obs.). Thematerial Wheeleridenti-
1989). In the Canal Zone of Panama, fied as "loeivensohni" (an unavailable
Wheelerencounteredwhatheinterpreted name; Ward 1989) issymbioticus, anobli-
tobetwospeciesofAztecathatwerespe- gateTriplarisantintheP.triplarinuscom-
cialized inhabitants of Cordia. He identi- plex,andknownfromPanamaandnorth-
fiedthemasA.longicepsandA.pittieri.Az- ernSouthAmerica(Ward,pers.com.).As
tecalongicepswasthemostabundant,oc- reported below, Costa Rican T. melaeno-
curringin85%ofthedomatia.Thespecies dendron is inhabited by a somewhat less
waspreviouslyknownonlyfromthetype specialized community of ants, without
queen,collectedinCostaRicawithnobi- close affinities to the specialist Triplaris
ologicaldata,andheredescribeditbased antsfromSouthAmerica.
on abundant material from C. alliodora. AntsinthedolichoderinegenusAzteca
His results have influenced subsequent are major elements of neotropical forest
identificationsofCordiaantsasA.longiceps antcommunities (Forel 1899). All arear-
(e.g. Opler & Janzen 1993). As reported boreal.Thenumerousspeciesexhibitava-
below, true A. longiceps is a Triplaris me- rietyofnestinghabits,inhabitingexternal
sliateensodienndCroosntaspReicciaa,liasntdkWnhoewelnerf'rsotmwotwCo. claivritnogntnreesetss,,adnedadlibvreabnrcahnecsh,esd.eaAdncourmebseorf
alliodora ants should be interpreted as ofspeciesareobligateinhabitantsofspe-
meTmrbipelrarsisofctohnetaAi.nspitatiterilecaosmtpl1e7x.species &cialMiczeKdeyant19p9l3a)n.tsFo(rreelvi(e1w8e7d8)inesDtaavbliidsshoend
Volume5,1996 133
the genus, and hisdefinitionremainses- of phenetically similar organisms for
sentially unchanged. Shattuck (1992) rec- which 1) therangeofcharactervariation
ognized thegenusasamonophyleticlin- is greater than that typicallyobserved in
eagewithintheTapinomini.Emery(1893) singlespecies,2) charactervariationisat
providedthefirstandasyetonlyrevision least partially discontinuous, suggesting
ofAzteca,recognizing25validnames,but multiplespecies,and3)eitherthediscon-
there are now over 150 available names tinuityisgeographicallyunstable(e.g.Az-
(Shattuck1994)duetosubsequentdiscon- tecapittiericomplex)orthereisinsufficient
nected descriptions. Longino (1989b, material to evaluate its stability (e.g. Az-
1991b) has recentlyreviewed thetaxono- tecanigricanscomplex).
myofthespeciesthatareobligateinhab- A provisional taxonomy is provided,
itantsofCecropiatrees. alongwithkeystoqueensandworkers.The
MystudiesoftheCostaRicanantfauna taxonomicresultsarebasedonqueensbe-
haverevealedatleast20speciesofAzteca causetheyshowgreaterdifferentiationbe-
inthecountry.Sevenofthesespeciesnest tweenspeciesthanworkersormales(Lon-
inliveplantstemsandhavequeenswith gino 1991b, Wheeler and Bequaert 1929).
relativelylong,narrowheads(headlength Distinguishingspeciesfromworkersalone
greater than or equal to 1.3 times head isproblematic,becauseworkersexhibitcon-
width). This report addresses the taxono- tinuoussizepolymorphism,andcoloniesof
myandnaturalhistoryofthesesevenspe- thesamespeciesvarygreatlyinthesizeof
cies. The species treated here include the the largest workers. Following the species
obligate inhabitants of C. alliodora and T. accountsareasummaryofCordiaandTri-
nwlaenodeiidwn, as well as other less spe- plaristintcommunitycompositioninCosta
cialized inhabitants of a variety of plant Rica,anddiscussionsof1)theadaptivesig-
species.TheremainingknownCostaRican nificanceofqueencharacters,2)problemsof
Azteca,includingthefiveobligateCecropia speciesdefinitionsasrevealedbythisstudy,
ants (Longino 1989b, 1991b), havequeens 3)possiblemechanismsgeneratingcomplex
withrelativelybroaderheads(headlength characterdistributionsintheA.pittiericom-
lessthan1.3timesheadwidth). plex,and4)thecontrastingrolesofregional
This is an intentionally regional work. faunasandglobalrevisions.
Differentiating species within Costa Rica LIVE-STEMNESTINGAZTECA
hasprovendifficult.Distinguishingcontin-
uousgeographicvariabilityfromdiscontin- Allmembersofthegroupdescribedhere
uouscharacterchange is difficulteven in nestinlivestems.WheninCordia,Triplaris,
an area the size ofCosta Rica, and chal- orCecropiatheyinhabitpre-formedcavities
lengesspeciesconcepts. Thus, formaltax- inthestems.Wheninplantspecieswithout
onomicchangesarerestricted to"nomen- pre-formedcavities,theyoccupyirregular
claturalhousecleaning" fora fewobvious chambersthroughoutthestems,apparent-
cases. Species that are insufficiently well- ly excavated by the workers themselves.
known, primarily due to inadequate Only young stems are occupied; older
knowledge of character variation within partsoftheplantaregraduallyabandoned.
andoutsideofCostaRica,aregiventaxo- Coloniesareusuallypolydomous,withall
nomically unavailable code names. These or large parts ofa plant crown being in-
codenames are developedby the author habited. Brood, often including sexual
andareuniquewithinthegenusAzteca.In broodoralateadults,isdispersedthrough-
thefuture,ifnumberedtaxaarenamedor outthecolonyspace.Nocasesofpolygyny
associated with existing available names, are known, and workers must transport
thenumberswillberetiredandnotreused. broodamongnestsinapolydomouscolo-
Theterm"complex"isusedforclusters ny. Carton construction is common, usu-
134 JournalofHymenopteraResearch
allyintheformofsmallplatformsandbaf- weremainlyfrommyresearchcollection.
fles inside the stems, but sometimes ex- Thesespecimenswillbedeposited inre-
tendingoutsideofthestemstoformrun- search museums (primarily LACM and
waysalongthestemsurfaces.Theinsides IBCR).
ofthestemsareusuallypackedwithHo- CHARACTERS
moptera (Coccoidea: mealy bugs and
scales), and when carton galleries extend Speciesdefinitionsarebasedonthefol-
outside of the chambers, abundant Ho- lowingcharacterset:
moptera are often found underneath.
Workersdonotappeartoforageofftheir Queens
host plant, and conspicuous patrollingor • Headshape(Fig. 1).
foragingoutsideofthestemsisrarelyob- • Plot of head width vs. head length
served.Thus,coloniesofthesespeciesare (Fig. 2, 3; measurement definitions
relativelyinconspicuous.Workersareoften providedinfigurecaption).
somewhattimid,emergingandbitingonly • Mandiblesculptureandpilosity(Fig.
whenthenestspaceisviolentlydisturbed 4).
oractuallybrokenopen.Acompletelydif- • Pilosityofthepropodeum(Fig.5).
ferentantfaunamayoccupytheoutersur- • Shape of the petiole in lateral view
facesofthehostplant. (Fig.6).
• Color
METHODS
• Plot of head width vs. scape length
Measurements were taken at 50X mag- (Fig.7).
nwiiftihcatainon,orutshionggonaalNipakiornomfiBcoreocmkelteerrrsottaagrey Workers
micrometers, wired to a dual-axis digital • Head shape of largest workers (Fig.
readout. The outputofthe measuringde- 8).
vicewasin0.0001mmincrements,andthe • Plot of head width vs. head length
rawdatawererecordedassuch,but10rep- (Fig.9).
licatemeasurementsofheadlengthofone • Plot of head width vs. scape length
specimen had a standard deviation of (Fig. 10).
0.0025mm.Thus,the95%confidenceinter- • Pilosity of the mesosomal dorsum
valformeasurementsspans0.003mm.Mea- (Fig.11).
csoulTrlehemecetinfotonlsdleoafwriienniugtsiaeondbs:braerveiaitnifoingsuroefcraepsteiaonrsc.h vwairtEihaatcaihodnci.hsacTruhasecstieforing(uiernxectsehpeatncdcaoplktoeiryo)nsisopfirloclhvusaitrdraeacttdeeid-r,
IBCR: Instituto Nacional de Biodiver- agnosticinformation,soitisnotrepeatedin
sidad, Santo Domingo de He- species accounts. Species accounts contain
redia,CostaRica. taxonomic changes, taxonomic comments,
LACM: LosAngelesCountyMuseumof distributiondata,andbiologicaldata.Costa
Natural History, Los Angeles, Rican placenames are used commonly in
CA,USA. thespeciesaccounts(Fig.12).
MCSN: MuseoCivicodeStoriaNahirale TAXONOMICSYNOPSIS
MCZC: "MGuisaecuommooDfoChoam,p"aGreantoiav,eIZtaoloy.l- AztcaihdtiiEmery 1893,newstat.;Honduras
MHNG: GoMgeuyns,eevCeua,mmbSrwdii'tdHzgieesr,tloaiMnrAde.,NUaStuAr.elle, t==ofsPntsaolcnliiantmFnaosr;uelbasn1pt9.-1p2Iln,catnintieWgwheneseeyrlnaei.rist1942,newsyn.
AztecncordiiicolciForel1920,newstat.,Boli\'ia;
The specimens examined in this work Cortiinspecialist?
Volume5,1996 135
AztecaJTL-003, unavailablecodename;Costa Azteca(nigrkatiscomplex)JTL-002,unavailable
AztReiccaa;JTCoLr-d0i0a7,spuencaiavlaiisltablecodename;Costa Aztceocdaepnatarmueel;isCoFsotraelRi1c9a0;8,livnee-wstesmtatg.e,neMraelxiiscto
Rica;Ocoteaspecialist? (pittiericomplex);Cordiaspecialist?
AztecajuruensisForel1904,newstat.,Brazil;in AztecapittieriForel1899;CostaRica;Cordiaspe-
Swartziastems(Fabaceae) cialist
Azteca longiceps Emery 1893; Costa Rica; Tri- = pittieri var. emarginatisquamis Forel 1920,
AztpelcaarisnisgpreicciaanlsisFtorel1899,newstat.;Panama Aztneecawsaspyini.Forel1912,newstat.,Brazil;inSap)-
Azteca{nigricanscomplex)JTL-001,unavailable iumstems(Euphorbiaceae)
codename;CostaRica;live-stemgeneralist
KEYTOQUEENS
KeytoAztecaqueensthat:1)areknowntooccurinCostaRica,and2)havesubrectangularheads,with
headlength^1.3timesheadwidth.Speciesdefinitionsinthistreatmentstronglyrelyonlengthandwidth
ofthequeenheadcapsule,andthekeyshouldbeusedinconjunctionwithFigures2and3.
lL.ab.. CCoolloorrllaarrggeellyyoorraenngtei;rehleyabdlawcki;dthhea>d1w.id2tmhm<(Fi1g..22)mm belti2i
2.a. Mandiblewithevencoveringofcoarse,piligerouspuncta(Fig.4A);mandiblesurface
appearingbristly{nigricanscomplex) 3
2.b. Mandible alwayswith row ofpiligerous puncta along masticatorymargin,butlarge
punctasparsetoabsentonmandiblesurfaceproximaltothisrow,andwithatmostfour
punctabearingsetae(Fig.4B-F) 4
3.a. PeHolarnodelowandblunt,ventrallobedeep(Fig.6D);scaperelativelyshort(Fig.7)
JTL-001
3.b. Petiolarnodesharp;ventrallobeshallow(Fig.6E);scaperelativelylong(Fig.7) ...JTL-002
4.a. H(Feiga.d1,stlronognigclepyisreacntdanJgTuLl-a0r0,3)w;ithheafdlatlesnigdetsha>nd0.2l7a5ter+al1m.a3(rhgeiandowfidvtehr)te(xabreolvaetilvienleyisnhFairgp.
2) ^ 5
4.b. Hbreoaaddlylesrsoruencdteadngu(lFairg.,1w,itrhemsaiidneisngslisgphetcliyesc);onhveexa,dalnedngtlhate<ral0.m2a7r5gi-I-n1o.f3(vheeratdexwimdotrhe)
(belowlineinFig.2) 6
5.a. Petiolarnodelow,anteriorfaceofpetioleflat(Fig.6C);propodeumwithsparseshort
setaeconcentratedposteriortospiracle(Fig.5C);mandiblelackinglargepunctaproximal
tomasticatorymargin(Fig.4E) JTL-003
5.b. Petiolarnodehigher,anteriorfacesomewhatconcave(Fig.6B);propodeumwithsetae
sparseorabundant(Fig.5);mandiblewithabout5largepunctaproximaltomasticatory
margin(Fig.4F) longiceps
6.a. Mandiblewithabout5largepunctaproximaltomasticatorymargin,about3ofthese
bearingsetae(Fig.4C);propodeumsparselysetose(Fig.5C) JTL-007
6.b. Mandiblewithabout3largepunctaproximaltomasticatorymargin,thesenotbearing
setae(Fig.4D);propodeumdenselysetoseovermostofsurface(Pacificslope;Fig.5A)
orsparselysetose(Atlanticslope.Fig.5B) pittiericomplex
KEYTOWORKERS
Thiskeyisacorroborativedevicewhenonealsohasqueensand/orhostplantdata.
La. oPfrolamregsesotnowotrukmerwsitohftsepna,rsbeutpinlootsiatlyw(aFyisg.,I>IA1);msmet(aFeigp.re9s)entonpropodeum;headwidth 2
l.b. Pheraodmewsiodntohtoufmlwarigtehstabwuonrdkaenrstpoifltoesni,tybu(tFing.otIIaBl-wEa)y;sp,r<op1omdemum(Fiwgi.t9h)orwithoutpilosity; 3
136 JournalofHymenopteraResearch
2.a. Headandpronotumredororange,gradingtobrownposteriorly,appearingbicolored
inthefield beltii
2.b. Colormoreuniformredbrown JTL-003
3.a. Propodeumlackingorwithatmostoneortwoshorterectsetae(Fig.11D,E);mandibles
bristly(nigricanscomplex) 4
3.b. Propodeumwithmorethan5conspicuouserectsetae;mandibleswithorwithoutbristles
5
4.a. Scapesrelativelyshort(Fig.10);mesosomalpilosityrelativelylong(Fig.IIE) JTL-001
4.b. Scapesrelativelylong(Fig.10);mesosomalpilosityrelativelyshort(Fig.IIF) JTL-002
5.a. Headrelativelynarrow(Fig.9);inhabitantsofTripilarismdaenodendron longiceps
65..ba.. CHeoamdmorenlatiinvhealbyitwaindterof(CFiogr.di9a);alilnihoadboirtaantsofCordiaalliodoraorOcoteanicarapgiutetniseirsicomplex6
6.b. InhabitantofOcoteanicaraguensis,knownfromonecollectionatCarara JTL-007
SPECIESACCOUNTS Largesize,orangehead,andsparsedor-
Aztecabel(tFiiigsEm1-e1r1y)newstat. srtaoellaoptciiclvuoersliytfdyrisomtmainkcGetuiavweto.erAmkzaetlreacsatobofeltPtihaiinissasmkpane.ociweInns
AztecabicolorracebeltiiEmery,1893:142.Holo- Costa Rica it is common at Santa Rosa,
type worker, Costa Rica (Alfaro) [MCSN] Palo Verde, and along the road to Mon-
(examined). teverde.Itisoneofthemorecommonin-
AztCHeaocnlaaoltfaysZpcoieante(a,unsiBuqabursepr.osyClnaoetltyaopreWa)hdeoqeulIeeserln,a:nd1P,9a49n2:aJ2m2ua7l,.y hhaabsitalasnotsbeoefnTcroilpllaercitsedmfelraoemnoCdoernddiraonuo,dobsuat,
1924 (Wheeler #637), from a domatium of Cecropia peltata, Cochlospermum intifolium
Cordia alliodora [MCZC] (examined), new (Janzen, pers. comm.), and Pithecellobium
syn. samaii. Colonies can be large, filling the
Azteca stolli Forel, 1912:54. Syntype workers: crown of large Pitliecellobiiirii trees. The
Gamuianteedm)a,lan,ewRestyanl.uleu (StoU) [MHNG] (ex- bnersatncshpatcieps,isaenndtirweolrykweirtshianrelirvaeresltyemsseeant
maUjonrdewrorbkeeltriifrEommeroyne(c1o8l9l3e)ctdioenscarnidbetdwoa ifsormaugcinhgmooutrseidceoomfmtohensttheamns.mTuhsiseuspmeccioels-
smallworkersfromadifferentcollection. lectionsmightsuggest,becauseofitscryp-
mInajtohrepwuobrlkiecartiaosnthheedteyspiegn(actoendsitdheeresdintghlee ticAhnabiotbss.ervation ofqueen foundingbe-
holotype here), and conjectured that the haviorisdescribedbelowunderA. longi-
smallerworkersmightrepresentadistinct ceps.
taxon.Themajorworker,collectedbyAl- Ihaveexaminedscatteredmaterialfrom
faro and simply labeled "Costa Rica," southernSouthAmericathatiseitherthe
matches material commonly collected samespecies,acloserelative,orahighly
fromTriplarisandothertreesinthePacific convergentspecies.Theseincludethetype
lowlandsofCostaRica.Thesmallerwork- queensofEmery'sfasciataandnun/ri, for-
ers,alsoexamined,arefrom"Jimenez,"a mer syntype workers of bicolor (workers
lowland Atlanticsite,andareworkersof excluded from types and bicolor synony-
Azteca alfari. The stolli syntype workers mized withalfariin Longino 1991b), and
clearlycomefromasinglenestseries,and recentBoliviancollectionsbyP.S. Ward.
appear identical to queen associated ma- If the South American material is beltii,
terial ofbeltii from Costa Rica and Hon- then thebiologyand distributionofbeltii
duras. isstrikinglysimilarto thatofPseitdonn/r-
Volume5,19% 137
V*. S.
JH c3
^^-
»
138 JournalofHymenopteraResearch
mexvidiius (Ward 1991). Bothshowcath- PseudomyrmexvidmisandAztecamayalso
olicity with respect to which ant-plants be found. During a briefexamination of
theywillinhabit,andbotharepresentin O. nicaraguensis plants at Carara, I ob-
bothCentralAmericaandsouthernSouth served that plants in shaded understory
America. were small and all inhabited by Myrme-
AztecacordincolaForelnewstat. alalcohinsgta,stwrheialmebplaannktssiwnermeorlearsguernnayndareians-
Azteca longiceps subsp. cordincola Forel, 1920: habited by Azteca. However, only one
203.Holotype(uniquesyntype)worker:Bo- vouchercollectionoftheAztecawastaken,
livia (Bang, n.ll78) incauline swellings of fromavigorouscolonywithalatequeens
Cordin[MHNG](examined). inthestems.
0.T5h9emm)sinigslneonsdmeaslclriwpotrkaenrd I(hceaandnowtidditsh- AztecajuruensisForelnewstat.
Wtihnegeuliesrh (i1t94f2r:2o3m2)mdoesstcriAbzetdecathespqeucieeesn. AztSeycnatylponegicweoprskevrasr,.jfuermuaelnessi,s Fmoarleel(,s):190B4r:a6z9i9l.,
and redescribed the worker based on Amazonas, Jurua, Jurua Mirim, Aug 1901
Mann collections from cauline swellings (Ule),inbranchesofSclnvartzia[MHNG](ex-
ofCordiaalliodora in Ivon, Beni, and Hu- amined).
cciaoorchmlihaeagi,nbiBnaaeallmnlatbibh,daeoel"B.usogclaTrhisihvptiethtahi.iesqoWnutdheyonepeeoeensrllentoorhocnataatlltiashtWpoeyhpletefioyaesrptrleeecdiorsn"rpdCdetioehcn--e-- hp1al.beA2ixh4.missmspy,enjcutihreyeuspae,endsiiwnsqiculldeouteodhknisn0g.hli7atk7shememah.sepaiatImtdnieerglieleancnteoegrrmtaa-hll
bascunrtdibMseaodnmnha'asyscbhoeellapedacrttlioeonnfsgttchhaen2pnixotttihebreeiacdaosmwspiuldmetexhd,, epterrroaifbailll.yeIstomfiasltlvheeerryptethsiiaomnliela.anrTyhaeCnodssitpzaoesRsiiisbclcayonncsmoiand---
conspecificwiththetype. dspiesctiifnigcuiwsihtthhesatpwioi,(bsueetIbedleofewr).sy1ncoannynmoyt
AztecaJTL-003 for lackofdata oncharactervariationin
(Figs1-11) Amazonianstem-nestingAzteca.
This species is known only from be-
tween 400-500m on the road to Monte- AztecalongicepsEmery1893
verde, from six different Cordia alliodora (Figs1-11)
sctrlyeumespst.aetrIsrtihcoafswibtrteeheensA.caollbelolentcigit,etdAh.ienlrtoonwagoidc,deipsfa,fnedraennidts Azt((eAuclnafiaqrulooe)ngs[iycMneCtpySspNe]E)m(qeeurxeyae,mni:n1Ce8od9)s3.t:a34R4i.ca,HAollaojtuyeplea
twoformsoftheA.pittiericomplex. Thespeciesisnowknownfromthetype
AztecaJTL-007 queen, collected in Alajuela before the
(Figs1-11) turnofthecentury,andsevencollections,
Thisspeciesis known fromonecollec- allfrombetween700and900melevation
tionfromCarara Biological Reserve.Oco- in theGuacimal rivervalleybelow Mon-
tea )iicaraguensis is an understory laura- teverde. The typehas nobiological data.
ceoustreeatCarara. Itispartofagroup Thesevennewcollectionsareallfromlive
ofunderstoryLauraceaewhosestemsare stems of Triplaris nwlaenodendron. Some
always occupied by ants (Stout 1979, collectionsare from maturecolonies,and
Hammel 1986, Burger & van der Werff others are founding queens from stump
1990).Theantsareusuallyobligateinhab- sprouts. Extensive collections in the area
itants in the genus MyniielacJiista, but havenotrevealedlongicepsusinganyoth-
139
0.8 0.9 1.0 1.1 1.2 1.3 1.4
HeadWidth(mm)
Fig.2. Queenheadwidthvsheadlength.Headwidthisthegreatestwidthoftheheadinfull-faceview.
Headlengthismeasuredalongthemedianaxis,fromtheanteriorborderoftheclypeustoalinetangentto
ttyhpee;po"s3t"eri=orJmToLs-t00e3x;te"n7t"o=ftJhTeL-v0e0r7t.exlobes.Line:HeadLength=0.275+1.3(HeadWidth),"n"=nigricans
erplantspecies,and solongicef^sisprob- lyafterexaminationofthetype,theMon-
ablyahostspecialistinT.melaenodendron. teverde population of longiceps was dis-
ThediscoveryoflongicepsasaTriplaris covered in Triplaris trees. As interpreted
ant was unexpected. Subsequent to the here,longicepjsisanarrowlycircumscribed
namingoflongiceps, Forelnameda num- Triplarisant,withthequeenheadrelative-
beroflongiceps subspeciesbased on ants ly narrower than most Cordia ants. The
from Cordia and other plants (cordincola, Cordia antsexamined by Wheelerare in-
patruelis, juruensis, and sapii). Wheeler terpreted hereas moreclosely related to
(1942) encountered two species ofAzteca pittieri. Inordertodissociatetheinfraspe-
regularly inhabiting Cordia alliodora in cifictaxacordincola,patruelis,juruensis,and
Panama (Canal Zone). He identified the sapii from longiceps, all are raised tospe-
morecommononeaslongiceps, and thor- cieselsewhereinthispaper.
oughlydescribed theworker,queen,and The following observations, derived
male based on his new material. Myself fromfieldnotes,describethenestinghab-
and other contemporary workers have itsofA. hvigiceps:
continued to identify thecommon Cordia 5July 1991,Longino#2956:Iclimbeda
antsaslongicepsorcf.longiceps. Examina- Triplaristreeandcutout3smallbranches
tionofthetyperevealedthatlongicepswas that all contained parts of a colony. No
notoneofthecommonCordiaants.Short- workers appeared as I climbed the tree.
140 JournalofHymenopteraResearch
nor after I cut branches. A few workers cupied apical shoot area, were the most
emergedfromcutbranchbases.OnlyasI recentlyentered.
begantosplitstemsdidlargenumbersof 5July1991,Longino#2972:1climbeda
workers swarm out. Abundant Homop- 4m tall Triplaris tree. Itcontained a pop-
tera were inside stems, and a few males ulouscolony, and workersemergedonto
andafewalatequeens.Therewasabun- trunk when I climbed tree. The largest
dantworkerbroodthroughout. branchsegments1examinedfromthistree
Iexamined a 24cm long section inde- were3cmdia.,andstillcontainedhollow
tail. The intemodes contained "knollen," intemodeswithants.Alargebasalsection
discretemoundsofstickybran-likemate- containedrelativelyfewworkersandscat-
rial filled with nematodes, tiny dipteran teredpseudococcids,withnococcids.Exit
larvae,andwhatappearedtobeabundant holes were still maintained through 1cm
stylets of Homoptera. (Knollen are also of wood. I dissected 180cm of occupied
found in nests of Cecropia ants (Miiller branch. There were abundant brood,
a1b8l8y0-c18o8m1m,oLnontgoimnoan1y99o1ar)a,llanstdema-rneepsrtoibn-g wmourckhersl,ikcear#t2o9n56p.artTihtieornes,waenrdeexsictathtoelrees,d
Aczutpeicea.d)iTnhteermeodweesr:e5p,i7n,k3c2o,c3c1i,ds8,in7,t4hecoocc-- a#l2a9t5e6,quteheenrse,waansdnaotlteraascteoonfepmianlke.coUcncliidkse,
wciadss ainsitnhgele7poscecuudpoiceodcciindteimnodtehse.seT7heirne- canodmmposne.udococcidswerewidespreadand
qotTtsisrcrieueehnehzipesmetTmeegtieoerhowianndmewen.deo,eoairaduwlsdStel-slbee.hooyuwsrsmutcre,aelMteapeasratshrpoitawigeantoonneerhdnnwytofre,eeeeuorebnmrsrsouweooeaffnchedtuwoaopniaegertiecdlcthrhrmcrhoefedupsnetoaep.odrrpfepiiauttxnaSaoetcaiircrodet.tettmdcimihd,ioeaoeehnlndmootslmtltmooyieehfmaodsemrwcadttrdooliadhslodwfreteiesneekceioireitnooniaera.g-ffnl bj2oaqLwuafrlu0oacpaa5ew^seAetnenez0erJcnttfrucahsemolsmcarycyi,ffataanmorvtt1milrote9etetomh9mdrhintee1geer,risssaicl.leebceeLyTpprpaaro'ataTfliwsnrnuyhp,egaclmearathpi,anraee,nsirdcdbostca2u,tvobttt#iehnhrfw2lteoteat9oiaeiiu6eu.istp9nsgnwl-uoThdosuuochoi:.ngucfebnosuugTIocipttfhtclciehqthueuepreuprtadmefiropittoereulnaiwnndoguadce.osmlhg--
themiddleofthebranch,withsolidsepta thebeltiiside(Fig. 13).
on both sides. The sclerenchyma was
thickerontheoccupiedsidethantheun- Aztecanigricanscomplex
sooitofiofclccnnanclctaokeuueenycrpprmtciierodupeenediprddcdeaiehmssebsyeiedirmtnnwdaoecteierewenrw.sonmteefoeoTrmfdmthbaeelhoetuaesdenacwreoksiasw.scealpeecllwtTc.rusaeohep,Ironeinfedaecnasdawoegrnarvrtiylteehndl-raotrsioteeeuacedmsmcogrofpeuhwodpttiunieehttnesrsah-dhesne, bm0lfRri0iaricg1Tnosaethdm,rolieoycb.utaclshuqnAeerudzpssetuteenJwiycncTnipastLetta-hhn,0oqei0afgus2Prneoaiteccehotianivchfnsaceistfucncrrstlo.scoshom.omewpviilnmelsParaeanktnxnodhndafesiohmblowaAaalftvn,releCgsaeooJnsnTattptlLhriiay-ee-c
btuhetftohremserc,leargeanicnhysumgagelasyteirngwathsatthtihcekesrclien- hleorweladnidfsf.erTihnequtehreeneh"esapdecsiiezse"arnedcorgelnaitzievde
ranetns.chyma layer was a response to the asbclaepefrloenmgtfhe.wHloocwaelivteires,,saanmdpklensowalreedagveaiol-f
Theantentranceholeswereirregularly geographicvariationinthesecharactersis
scattered,notinanypredictablelocation. inadequatetoconfidentlyestablishspecies
The terminal intemodes, near the unoc- boundaries.