Phylogenetic relationships of the thynnine wasp tribe Rhagigasterini (Hymenoptera: Tiphiidae)

20 Pages·1996·6.4 MB·English

by  L S Kimsey

J.HYM.RES.
Vol.5,1996,pp.80-99
Phylogenetic Relationships oftheThynnineWaspTribe
Rhagigasterini (Hymenoptera:Tiphiidae)
LynnS.Kimsey
DepartmentofEntomology,UniversityofCalifornia,Davis95616
—
Abstract. ThethyrminetribeRhagigasteriniisreviewed,withanextensivereevaluationofge-
nericgroupings.PhylogeneticanalysesofsevenRhngignsterspecies,twospeciesofDimorphoth-
aysnnouus,tgtrowuopsA,elurervuesalsptechieesr,elaantidonfsohuirpts:eeAnnstpheocbioesscoifnaEiero+ue,(TwhiytnhniAnnith+ob[o{sAceilnurauesa+ndEiTrhoynne)ni+ni(sD.is-.
morphothi/twus + Rhagigaster)]).Aconsensustreegeneratedfromall27taxayieldedaciof59.
Analysisofonlyspeciesrepresentinggenera(generictypespecies)resultedinacladogramwith
thesametopographyandaciof73.Rhagigasterspecieswerefoundtoconstituteamonophyletic
EuinriotnedeaspnidteAesluugrguesstiisopnrsobtlheamtatthiics.gAeenluursusm,atyhehaovrUeybSeoeunthpolAympehryilectainc.geTnhues,reelnadtsionusphiepmbbeetdwdeeedn
inEironeinalloftheseanalyses,andmayindeedindicatethatEironeisaparaphyleticgenus.
However,derivedfeaturesofthemaleterminaliaineachgroupclearlyseparatethem,andsince
hcfeheremeeaislnme.asnFahieav,evsenchebleiwnegnseprseiceaieennsdoonsfpleyEciirfooosrnuesaafrfreeowdmessPpcaerpciiubeeasd:NineaqbwuoitlGohuniignueesnaea.rnad,vAaelloukrauesnsiwsilflrnoomtNbeewsyBnriotnayinm,iaznedd
INTRODUCTION characteristics or phylogenetic analyses,
vcTsTagareuuehnnebrTrnyddtfshenneaerenymTcairnfishlannluay(maiibadn1ieefinnl9ns.saay1di,tm,0enSiibaucDsw)liebpiysh,estaaicrenimtcienqhatmehaeulhistteyoenhhlnsdueaewategereTtshgd,hsireiitpess(uvhtRtdBiihghipirdearaedloegonsaatdimudech(ga.peeiPajdnrmasoIatstonrtsoeiiesitatnr1t1ityia99hdln74rtidla57ehoisy),ee--e,f aoi1mrt1rr8on8neia3n4cTdmlg9e2lhai,.buisnedsWieaADealergenislels.myuonutotfberThrdwrfuhepteaosiahhslsiomoclsgitidrsyeglglinrntrbuye/h1oeonren8uduaaRn4ppcohu4b,cnsayeiRlagspahlyTitTlannugeugdStoradirhonsg.wneuaAetetserrerthtelrrseui(1etrAra1n9umat91asree10Geedr0bub,iKAewa)clrusaEauistisgnn-ao-
Brothers&Carpenter1993, Kimsey1991) tribe in the subfamily Thynninae (Given
cwtloaesatrhlbeyasdfeeadmmioolnnystaTriasptuhieitietdhaoaeft.cthhTayhrinasnctiencreoisnsctblieculssoiinongn- n19u5Mm4eb,meSbraeltorefsrt1ho9ef54dt,ehreKiivRmehdsaegfyiegaa1t9su9tr1ee)rs.icnhiarlaacctkera-
cluding the presence of mesopleural la- istic of other Thynninae, having instead
mellae, a quadrate pronotum, the unci- theprimitivecharacterstatesasfollow:1.
gfeonremram,aalnedsuwbignegneitdalfoprlmastehafvoiunngdtihnempoosst- omrettarsutnceartniuomnwinitbhootuhtsseixmeps,le2.memdailaelarpiidcgael
teriorangleofthepronotumreachtheteg- sternumapicallyroundedorwithalinear
ula. Argaman and Ozbek (1992) made a uncus,3.aedeagussimpleandlinear(ex-
retrogradeproposal to restrictthefamily ceptDimorphathyiunts),4.femaletergumI
Tiphiidaetoincludeonlymembersofthe and II simple, without carinae or rugae
Tiphiinae.However,thisproposaliscom- (except Diniorphotln/unus), and 5. female
pletely unsupported by any apomorphic withoutdiscretepygidialplate(exceptDi-
Volume5,19% 81
morpholhynnus).Thereforetheyaretreated England, M. C. Day; U. S. National Mu-
asabasal lineageofthesubfamily(Kim- seum,Washington,D.C,K.V.Krombein
sey1991).Derivedfeaturesofrhagigaster- and A. S. Menke; ZoologischesMuseum,
inesare: 1. maleparamereswithadense Humboldt-Universitat, Berlin, F. Koch;
apicalrowofelongate,oftenflattenedse- Zoologisk Museum, Copenhagen, Den-
tae,and2.femalemesopleuronwithadis- mark, B. Petersen. The species examined
cretedorsalsurface. for this study are indicated below by an
Afterexaminationofalloftherhagigas- asterisk(*).
ugtreeranilenreHaitsEytiporerosyn)einiatntbdheecAeBarlmiuetniisashppwMaeurresenetvuetmrhya(tsNiatmth--e a=meTtfhleearg,feolPllDloomew=rienp,gunaMcbtbOurrDeevida=itaimoneintisedrao,rceeSlul=suesdg:adsiF--
ilar,andhaveverysimilarfemales,where tralsternum,andT= gastraltergum.
known. There has also been the sugges- Parsimonyanalysesofthegenericrela-
tionthatthegenusRhagigasterispolyphy- tionshipswereperformedusingtheHen-
leticandshouldbefurthersubdivided(G. nig86software(Farris1988).Thedatama-
Brown,personalcommunication).Finally, trixgeneratedbydetailedexaminationof
Eironeisastructurallydiversegenuswith thespecieslistedinTable1wasanalyzed
somespeciesgroupsapparentlydivergent using explicit enumeration with branch
enoughtojustifytheirseparationintodis- swapping.Allcharacteristicsweretreated
crete genera. Therefore, previous treat- as unweighted,and multistatecharacters
mentsoftherelationshipsamongtheRha- asadditive.Thesespecieswerechosenbe-
gigasterini areinadequate,and theentire causetheyrepresentverydifferentgroups
groupneededanobjectivereexamination. within each genus or are types of pub-
MATERIALS lished generic names. They were specifi-
callychosentotestthenotionthatthecur-
Specimens were borrowed for this rent generic categories were either para-
studyorstudiedinsitufromthefollowing orpolyphyletic.Characterstateswerepo-
institutions and individuals: Australian larizedusingtheThynniniandAnthobos-
National Insect Collection, CSIRO, Can- cinae as outgroups. Character polarities
berra, ACT; California Academy of Sci- arediscussedbelow,andaregiveninthe
ences, San Francisco, USA, W. Pulawski; datamatrixprovidedinTable1.
California DepartmentofFoodandAgri-
culture, Sacramento, USA, M. S. Was- RESULTS
bauer; Canadian National Insect Collec- TribalCharacters
tion,Ottawa,Ontario,L.Masner;Carnegie —
Museum, Pittsburgh, Pennsylvania,USA, Head. In the Thynninae a transverse
J. E. Rawlins; Cornell University, Ithaca, carinaextendingacrossthefronsisfound
N. Y., USA, J. K. Liebherr; Florida State only in males of Rhagigaster and Diinor-
Collection of Arthropods, Gainesville, phothynmis. The majority of Rhagigaster
USA; L. Stange; Charles Porter, personal haveatleastatraceofatransversecarina.
collection;Gainesville,Florida, USA;Mu- In Dimorphothynmis thecarina iswell-de-
seo ed Istituto di Zoologia Sistematica, veloped and extends from eye to eye
Universita diTorino, Italy, P. d'Entreves; acrossthefrons,withadorsalbranchthat
Museum of Comparative Zoology, Har- forms an almost heart-shaped enclosure
vard University, Cambridge, Massachu- (Fig.8).Thisfeatureisclearlyauniquely
setts,USA,J.M.Carpenter,andD.Furth; derivedch—aracteristicofthesegenera.
Rijksmuseum van Natuurlijke Historie, Thorax. The dorsally bulging meso-
Leiden, Netherlands, C. van Achterberg; pleuron in female Rhagigasterini isdiag-
TheMuseumofNaturalHistory,London, nostic for this tribe. A hindcoxal carina.
82 JournalofHymenopteraResearch
extending along the dorsum of the coxa analyze phylogenetic relationships. The
fromthebasenearlytotheapex,ispresent resultingmatrixisgiveninTable1.Polar-
inallmalethynnines,andisgenerallyun- ity is indicated in parentheses, (0) is the
developed or is elevated and broadly primitivestate,and(1)or(2)arederived.
roundedbasallyinrhagigasterines.How-
ever, in Rhagigaster and Dimorphotliynnus 1. Male transverse frontal carina: Ab-
itiselevated and toothlike (Fig. 11). Un- sent (0), present (1) [Rhagigaster, Di-
like most other thynnines, female Eiwne inorphotln/ivnis], joined by U-shaped
have greatly reduced mesopleural lamel- dorsalcarina formingafrontalenclo-
lae. sure(2)(Fig.8)[Dimorphothynnus].
In the majority of tiphiid subfamilies, 2. Female maxillary palpus: Four-seg-
includingMyzininae,Anthoboscinae,and mented (0), three or two segmented
Tiphiinae,themetapostnotumisobscured (1). [Aelurus and Eirone]. Female Ae-
mediallyby the scutellum. In the Rhagi- lunis and Eirone have reduced palpi.
igbalseteraisniathbeamnedtadporossatlnlyo.tuTmhiissclfeeaartluyrevisi-s sReedgumcetnitosniisnctohemmnounmbeinr aocfulpeaaltpeasl;
foundinnootherTiphiidae,andthepat- however,itappearstobeaconsistent
ternseenintherestoftheAculeatasug- genericfeatureintheTiphiidae.
gests thatthepresenceofa visiblemeta- 3. Female labial palpus: Five or 6-seg-
postnotumintheformseeninthesegen- mented (0), four segmented (1). [Ae-
eraisder—ived. lurusand Eirone].
Gaster. Sculpturing of the female T-11 4. Female mesopleural development:
isaprominentfeatureinmembersofthe Tapering evenly toward scutvim (0),
ThynniniandScotaenini(Kimsey1992).In stronglybulgingtowardscutumwith
RhagigasterinionlyDimorphothynnushave distinctdorsalsurface(Fig.4)(1)[Rha-
sculpturing on this or any other gastral gigasterini].
terga. In Anthoboscinae the subgenital 5. Femalemesopleurallamella:Wellde-
plate is simple and evenly rounded, al- veloped and apically rounded (0),
thoughinafewspeciessuchasAnthobosca strongly reduced to small point-like
chilensis Guerin, the apex may be spine- process(1)[Eirone].
rimmed and thickened medially (Fig. 27) 6. Male hindcoxal carina: Present but
asisseeninsomespeciesofEiwneandin evenly curved from apex tobase (0).
Aelunis (Fig. 31). Thissimpleandevenly Abruptlyexpanded and angulateba-
roundedapicalsternumisassumedtobe sally (Fig. 11) (1) [Dimorphothynnus
the primitive form in the Rhagigasterini. andRhagigaster].
BasedontheconditionseeninAnthobos- 7. Male metapostnotal development:
cinae and American Thynnini and Sco- Highlyreducedandsunkenmedially
taenini, thelackofpenisvalvesisprimi- (0). Broadly exposed dorsomedially
tive for Rhagigasterini. All Thynninae (1) [EironeandAelurus].
haveahighlyderivedanddistinctiveae- 8. Female metasternum (first variable):
deagus. Furthermodificationcanbeseen With low medial ridge (0). Medial
in the Scotaenini where the aedeagus is ridge modified intobilobate, apically
relativelyshortwith membranous lateral flattenedprojectionbetweenmidcoxae
lobes and no apical loop, an autapomor- (1) [Rhagigaster].
phyforthatgroup(Kimsey1992).Within 9. Female metasternum (second vari-
theRhagigasterini,themajorityofgenera able):Flatorcarinate(0).Stronglyex-
havean aedeagus with a basal bulb and panded ventrally, appearing triangu-
slenderapicalneck(asinFigs.43-53). larinprofile,withventralapex(which
The following characters were used to maybebilobate)locatedconsiderably
Volume5,1996 83
Table 1. CharactermatrixforthegeneraofRhagigastermi(Tiphiidae).Anthoboscinae(representedby
Anthoboscacliilensis(Saussure)andA.crosiicorniiSmithandThynnini(representedbyThynnusdentatusFa-
briciusandZaspilothynnnsinlerruptufWestwood)aretheoutgroupsusedinthisanalysis
T<i\on Characters
Anthoboscinae 00000
Thynnini
Aelurusiiasutus
Aelurussep'ti'iitriciwUs
Diniorphotln/inuisapicalis
Diinoiplwtln/uiiusmorio
Eiroticaliciiw
Eironeaquikniiiis
Eironecheesrnanac
Eironeemarginatus
Eironeferrugineicornis
Eironemajor
Eironemutabilis
Eironeparca
Eironerufopictus
Eironeichizorhina
Eironexhlingeri
Eironespeciosiis
Eironevalokaensis
Rhagigasteraculeatus
Rhagigasterephippiger
Rliagigastercinerelhis
Rhagigasterlaeingatus
Rliagigasterlatisulcatus
Rhagigasterlyelli
Rhagigasterunicolor
84 JournalofHymenopteraResearch
15. Female tergum III sculpturing: Dor- meresimpleandevenlyroundedapi-
sally smooth, without cross-ridging cally (0). Apically with awl-like lobe
(0). Extensively cross-ridged (Fig. 4) (Fig.48)(1) [severalEironcspecies]
(1)[Diniorphotln/)iiuis]. 25. Paramere shape (second variable):
16. Male epipygial plate: Absent (0). Parameresimpleandevenlyrounded
Present and carinate (1) [Dimorpliotli- apically(0).Submediallywiththumb-
ynnus].Theabsenceofapygidialplate like lobe (Fig. 37) (1) [several Rhagi-
is assumed to be the primitive state gasterspecies]
sincetheAnthoboscinaelackone. 26. Digitus shape: Digitus forminga se-
17. MalegastraltergumVIIlobate:Gas- tose, often small, rounded lobe (0).
tral tergum Vll evenly rounded (0). DigitussetoseandC-shaped(1)[some
Tergum withapicomedial lobe (as in Rliagigasterspecies]
Fig.26)(1)[someRhagignsterspecies] 27. Penisvalves:Absent(0).Present,sim-
18. Male gastral tergum VII carinate: pleandfoliaceous(asinFigs.45,46)
TergumVllevenlyrounded (0). Sub- (1) [Rhagigaster]. Elongate, spoon-
laterallycarinate(1)[someRhagigaster shaped (2) [Thynnini s.s]. Extremely
species] bilobate, with elongate dorsal and
19. MalegastralsternumI(firstvariable): ventrallobes(3) [Einvws.s.].Bilobate
Evenlyroundedwithasinglebasalca- withventrallobesecondarilybifid(4)
rina or ridge (0). With medial hook- [some Eirone species]. Multilobate,
likeridge(Figs.20,21,23)(1)[Dimor- without one or two unusually elon-
phothi/iiiiiis, some Rhagigaster species, gatelobes(5) [Aehirus].
someEiwnespecies]. 28. Aedeagal form: Aedeagus consisting
20. Malegastral sternum I (second vari- oftwoseparate,simple,elongatelobes
able): Sternum 1 simple basally (0). (0), Lobes separate, forming robust,
Withshelf-liketransversecarina,often basally lobed or angulate structure
connectedtoshortbutprominentme- (Figs. 41, 42) (1) [Dimorpiiothynuus].
diallongitudinalcarinawhen thelat- Aedeagal lobes fused intoa structure
ter is present (Figs. 20, 21) (1) [most with a distinct basal bulb and short
Rhagigasterspecies,Dimorphothyimiis]. apicalneck(Figs.45^8)(2)[someEi-
21. Malesubgenitalplate(firstvariable): ronespecies].Apicalneckgreatlyelon-
Simple and evenly curved, or some- gateandlinear(Figs.49-53)(3)[some
what thickened apically and spinose Eironespecies,Aelurus].Neckflattened
(0). With discrete, flat, spine-rimmed and coiled (4) [onespeciesofEirone\.
apicomedialplate(1) [Aeliirus]. Aedeaguswithbasalcolumnandneck
22. Male subgenital plate (second vari- (apical loop) flattened and coiled (5)
able): Simple, broadly and evenly [onespeciesofRhagigaster].
curved (0). With long curved ventral
spine(uncus) (Figs. 32-36) (1) [Rhagi- PhylogeneticRelationships
gaster, Dimorphothyniiiis]. Uncus pro- The tiphiid subfamily Anthoboscinae
truding below broad flat shelf-like andthynninetribeThynnini,areincluded
rim, fitting broad apical tergum in in thisanalysisasoutgroups. The result-
outline (Figs. 32, 33) (2) [Dimorpihoth- ing14treesfoundbyimplicitenumeration
ynnus]. hadaciof59. ANelsonConsensusTree,
23. Paramere setation: Apical setae un- shown in Fig. 1, produced the following
modified(0).Apicalsetaedense,elon- relationships: Anthoboscinae + (Thynnini
gateandmostoftenflattened(Figs.48, + [{Aelurus/Eirone)+{Dimorphothynnus+
52,53)(1)[Rhagigasterini]. Rhagigaster)]). Theci wassolowbecause
24. Paramereshape (firstvariable): Para- therewasrelativelylittleresolutionofre-
Volume5,1996 85
Anthoboscinae
r'
h37r r;=2 Thynnini
i!=3& i=26 Rhagigaster unicolor
=25 Rhagigaster lyelli
=24 Rhagigaster latisulcatus
=23 Rhagigaster laevigatas
22 Rhagigaster cinerellus
lr=32 E- Rhagigaster blnodatus
lt=2 Rhagigaster aculeatus
n=34:^ [p=5 Dimorphothynnus morio
1!=31:!!=6 Dimorphothynnus aplcalls
l!=35^ [r=10 Eirone emarglnatus
=33: tr=13 Eirone mutahills
L3oiL=i6 Eirone schlzorhlnus
9 Eirone cheesmanae
r=7 Eirone allciae
l!=28 =8 Eirone aqullonius
=11 Eirone ferrugineicornis
=12 Eirone major
=14 Eirone parca
=15 Eirone rufoplctus
=17 Eirone schllngerl
=18 Eirone speclosus
r=19 Eirone valokaensis
Aelurus septentrionalis
[|=3
^27l=4 Aelurus nasutus
Fig. 1. NelsonConsensusTreeshowingphylogeneticrelationshipsofrhagigasterinespeciesandgeneric
groupings.
lationships among species of Eirone and male palpal reduction, triangular female
Rhagigaster. However, implicit resolution metastemum,andabroadlyexposedme-
ofrelationshipsamong thesespecieswas tapostnotum and projecting lobate meta-
less important than determiningwhether stemuminmales.InthisanalysisAelurus
ornottheyactuallyrepresentedsufficient- s.s.fitsreadilywithinanassemblageofEi-
ly divergent groupings toconstitutesep- ronespecies.ItispossiblethatAeluruss.s.
arategenera. mayrepresentaspeciesgroupinthislarg-
Byeliminatingallspeciesbutthosewith er taxon. The femalesofboth Eirones. s.
uniquecombinations ofcharacterstates a andAeluruss.s.arethusfarindistinguish-
singlecladogram was generated with the able. Inbothgroupstheapicalmarginof
same length and ci as the one using all the male subgenital plate varies from a
taxa.Thistreealsohadthesameunderly- thin,evenlycurved,spinelessrimtoadis-
ingtopologyastreesgeneratedusingall27 cretespine-rimmedapicalplatform.These
taxa. modifications of the subgenital plate do
Asdiscussedabove,EironeandAelurus not appear to correlate with any other
areverycloselyrelatedgroupssharingfe- modifications of the head or genitalia.
86 JournalofHymenopteraResearch
However,EironeandAelurusmalescanbe many of the thynnine genera, including
consistentlydistinguishedbyuniqueelab- the transversely ridged female tergum,
oration of the penis valves in each. The and a delineated pygidial plate. Synapo-
valvesin Eironearestronglybilobedwith morphiesforRliagigasterinclude:theflat-
onelobeextendingventrallyandtheother tenedandbilobatestructureofthefemale
dorsally,withsecondarysubdivisionofthe metasternum,foliaceouspenisvalves,and
dorsallobeintoapincher-likestructurein maleapicaltergumapicallynarrowedand
afewspecies.Aelurusmaleshavethepenis carinateorwiththumblikelobe.
valveselaboratelylobed,withdorsal,ven-
tral, and even lateral lobes, which may Biogeography
themselves be secondarily lobate. Addi- Thynninae exhibit a typical trans-Ant-
tionally,althoughAelimisspeciesbasedon arcticdistribution(Figs.2,3),withspecies
malesarelargelydescribed(Kimsey1992), occurring inSouthAmerica and Austral-
many Eirone species remain undescribed asia.Althoughtherearecurrentlynogen-
from either sex, and so few females are era shared between these continental
knownforeithergenus,thatfurtherstudy regions, Aelurus and Eirone show close
is essentialbefore making thedecision to phylogenetic links between the South
synonymizeonewiththeother. American and Australasianfaunas. Aelu-
Eirones.s. isa largestructurallydiverse russ.s.isrestrictedtotheNeotropicalRe-
groupofspecies,butduringthecourseof gion,whileEirones.s. isAustralasian. Ei-
thisstudytheonlygroupthatmightjustify rone is the most widespread member of
genericstatuswasabasalcladeconsisting theAustralianrhagigasterines,andisone
of Eirone miitabilis and schizorhinus, and ofthe fewgenera withspeciesoccurring
severalasyetundescribedspecies.Thiscla- outside of Australia, on New Guinea,
deischaracterizedbyhavingtheparamere NewCaledoniaandNewBritain.Thisge-
terminate in aprongand thepenisvalve nus is also the memberofthe Rhagigas-
secondarilybifid.However,nofemalesare teriniwiththelargestnumberofprimitive
knownforthesespecies,andeachofthese features resembling thoseofAnthobosci-
apomorphiccharacteristicsturnsupinoth- nae.Thedistributionandphylogeneticre-
erspeciesofEirone,forexample,thepenis lationships of these genera suggest that
valve structure also occurs in cheesnimme. theThynninaeevolved afterthebreakup
Asshownintheconsensustreethisclade of Gondwanaland, during the period
endsupembeddedinEirone,asitdoesin whenAustralia and SouthAmericawere
themajorityoftheiterationsresultingfrom connected to one antither via Antarctica,
this analysis. Thereforethere is nojustifi- between70and30mya(Fig.3).Members
cationfortreatingthisgroupofspeciesas of other thynnine tribes show no close
adiscretegenus. phylogenetic relationships between the
DimorphothynmisandRliagigasteraresis- two regions, and in fact are divergent
tergroupsbasedonthepresenceofafron- enough to suggest that the Australasian
talcarinaontheface,unciformsubgenital genera belong to one or more tribes and
plate, and toothlike hindcoxal carina. the South American genera in twoother
However,Dimorphothynnusishighlymod- tribes (Kimsey 1991), with none ofthese
ifiedandsharesanumberoffeatureswith occurringinbothregions.
KEYTOTHEGENERAOFRHAGIGASTERINAE
1. Winged,with7gastralsegments,andelongateslenderantennae;males 2
- Wingless,andant-like,with6gastralsegments,andshortcoiledantennae(asinFig.4);
females 5
Volume5,19% 87
Fig.2. WorlddistributionofthetiphiidtribeRhagigasterini.
2. Apicalabdominalsternumunciform(Figs.32-36),apicaltergumbroadandshovel-likeor
narrowedandsublaterallycarinate(Figs.25-26) 3
- Apicalabdominalsternumevenlyrounded,unmodifiedorwithanarrowplatformmar-
ginedwithshortspine(Fig.31);apicaltergumevenlyrounded,orslightlyindentedand
otherwiseunmodified(Figs.29,30) 4
3. Apicalabdominaltergumbroadlyroundedapicallyandshovel-like(Fig.25),apicalster-
numwithbroaddorsalplatformaboveelongatecurvedapicalspine(uncus)(Figs.28,32,
33) DimorphothynnusTurner
- Apicalabdominaltergumstronglynarrowedoralmosttrilobateapically,oftenwithac-
companyingsublateralcarinae(asinFig.26);apicalsternumwithdorsaltoothornarrow
rimaboveuncus(Figs.34-36) RhagigasterGu^rin
4. Neotropical;apicalabdominalsternumapicallythickenedwithadiscrete,narrow,spine-
rimmedplatform(Fig.31),andpenisvalveselaboratelyprongedwithoutanelongatedor-
sallobe(Figs.52,53) AclurusKlug
- Australasian;apicalabdominalsternumusuallywithoutadiscreteapical,spine-rimmed
appliactaflorlmob(easanindFsiignsg.le29,or30b)i,loabneddpdeonrissalvaolnvees(Fsitgrso.n4g5l-y5b1i)-ortrilobedwithwelEli-rodneeveWleosptedwood
5. TergumIIIwithnumeroustransversecarinae(Fig.4);tergumVIwithdiscrete,carinate
pygidialplate;genalbridgestronglybulginginprofile(Fig.6);sternumIventrallysimple
DimorplmthynnusTurner
- TergumIIIwithoutcarinae;tergumVIsmooth,withoutcarinaeordiscretepygidialplate;
genalbridgenotbulginginprofile;sternum1ventrallydentate 6
6. Hindtarsalclawsdentate;metastemummediallyprojecting,withanapicallyflattenedand
stronglybilobatestructure ; RhagigasterGu^rin
88 JournalofHymenopteraResearch
Fig.3. Projectionofcontinentsinsouthernpolarview60millionyearsago,withmodemdistributionof
Rhagigasteriniindicatedbyshading.
Hindtarsalclawsedentate;metastemumexpandedandtriangularinprofile,apexofthe
trianglesharpapically(Fig.5) 7
Mesopleural lamellawelldeveloped,apicallyrounded;neotropical;propodeumusually
withoneortwoovoiddepressionsmedially(Fig. 10);fourmaxillaryandthreeorfour
labialpalpalsegments AelurusKlug
Mesopleurallamellareducedtosmallpointedprocess;Australasian;fourorfewermaxil-
glajrQy^gandthreeorfewerlabialpalpalsegments;propodeumwithoutovoidmeEdiiraolncdeWpersets-wood
AelurusKlug rarely tridentate; labrum broadly ovoid
(Figs.10, 15,31,52,53) with narrowed base; maxillary palpus
with6articles,labialpalpuswith4;occip-
Aelurus Klug 1842:42. Type: Adurus nasutus ital carina faint dorsally; frons smooth
CopKphlloautcgheym1n8en4nu2t:s4n3T.aumrOerniegfr.or1d9Ae0es8li:gu7.r9u.sUKmliuegce1s8s42a.ryre- wmdieedtuihmaolulteyverpnirldoygjeesrcotuoinrndgecdaar,nidneaceba;irlionmbaeatttee;a;satpberdroonpmuo-m-
Mfl/e.—Mandible simple, bidentate, or inal segments weakly punctate, without
Volume5,1^% 89
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subapicalconstriction,withanteriorzone Includedspecies. AelurusalbofaciesKim-
ofcoarsepunctation;apicalsternumthick- sey*,ater Duran, brasilianus Kimsey*,cly-
enedapically,withmarginalrowofspines peatus Klug, concazm Kimsey*,enigmaticus
alongapex (Fig. 31);parameresgenerally Kimsey*, gayi (Spinola), grande Kimsey*,
broad,withrowoflongflattenedsetaeon nasutusKlug*,nigrofasciatus(Smith)*,pen-
apicalmargin(asinFig.53);volsellausu- alKimsey*,septentrionalisKimsey*,tridens
ally apically bilobate; penis valves long, (Spinola),unciferTurner.
welldeveloped;aedeagusbasallybulbous
and apically long and slender (Figs. 52, DimorphothynnusTurner
53). — (Figs.4,6,8, 11,17,20,24,25,28,32,33,
Female. Headaslongasbroadorlon- 39,41,42)
wgmeiortu;hth4epyaoerrstfseslwiregerhdtaulrycteicdll,aersgmeaarnxdiltllhaaabnriyalppepadaillcppeuulss; DimgaosrtpchrohtaheymnonrruhsoiTdualrinserGu1e9r1i0nb:158.42T:y2p.eO:riRgh.adgei--
with 3 or fewer; pronotum longer than sig. —
broad; mesopleuron with clearly devel- Male. Mandible slender and apically
oped dorsal surface; propodeum with bidentate; labrum small and linear; max-
longflatslopingdorsalsurface,oftenwith illarypalpuswith5articles,labialpalpus
1 or2 medial depressions(Fig. 10); terga with 4; occipital carina dorsally obsoles-
relatively smooth without differentiated cent; fronswith transversecarinajoining
areas,carinaeorrugosihes;apicaltergum carinaeextendingdorsallyfromthefron-
smoothwithoutcarinaeordefinedpygid- tal lobes, forming a bell or heart-shaped
ium;apicalsternumwithU-shapedapical enclosure (Fig. 8); regionbetweenanten-
liphaving2in—foldedflaps. nal sockets strongly protruding; genal
Distribution. Aeliiriis species have a bridgeprotrudinginlateralview;prono-
patchydistributionintheNeotropicalRe- tum strongly angled laterally, transverse
gion, occurring in Costa Rica, Panama, anteriorcarinastronglyflared;mesopleu-
Colombia, Ecuador, Peru, Argentina, ron sharply declivous anteriorly, with
ChileandBraz—il(Kimsey1991). scrobeobsolescent;metasternumstrongly
Discussion. Aclurus and Eirone are ventrally bilobate; propodeum strongly
closelyrelatedgenera,somuchsothatthe angulate laterally, with transversecarina
femalesarevirtuallyindistinguishable,ex- separatingdorsalfromposteriorsurfaces;
cept for the development of the meso- tarsal claws dentate; hindcoxal carina
pleural lamellae. There are other subtle stronglyangulate(Fig.11);abdominalseg-
differences,butwhetherthesewouldsep- mentsoftencoarselypunctate,andsome-
arate all species in both genera is uncer- what constricted subapically; sternum II
tain.Aelurusfemaleshave5maxillaryand with Y-shaped basal carina, forming a
4labialpalpalsegments,themesopleuron largetransverseridgesubbasallyconnect-
isstronglyconvexmedially,and thepro- ed to a medial carina extending posteri-
podeumisusuallydorsallyflattenedwith orly (as in Fig. 20); apical sternum with
oneortwomedial depressions. In Eirone slender curved unciform prong below a
femalesthereareusually4orfewermax- flat, often greatly expanded dorsal lip or
illary and 3 or fewer labial palpal seg- rimthatmatchesthemarginoftheapical
ments, and the propodeum is dorsally tergum (Fig. 32); apical tergum apically
gently convex without depressions. Rlia- broadlyroundedandsometimesmedially
gigaster females are also similarbut they weaklyindentedaswell,withshortlateral
canbeimmediatelyseparated fromthose carina(Fig.25);paramereslongandslen-
ofAelurusandEironebythedentatehind- der with elongate flattened apical setae
tarsalclaws. (Fig. 39); volsella elaborately foliaceous.