Ethology of three Southern African ground nesting Masarinae, two Celonites species and a silk-spinning Quartinia species, with a discussion of nesting by the subfamily as a whole (Hymenoptera: Vespidae)

11 Pages·1992·4.6 MB·English

by  F W Gess

J.HYM.RES.
1(1),1992 pp.145-155
EthologyofThreeSouthernAfricanGroundNestingMasarinae,
TwoCelonitesSpeciesandaSilk-spinningQuartiniaSpecies,
withaDiscussionofNestingbytheSubfamilyasaWhole
(Hymenoptera:Vespidae)
F.W.GessandS.K.Gess1
AlbanyMuseum,Grahamstown,SouthAfrica
—
Abstract. AccountsaregivenofsomeaspectsofthenestingofCeloniteslatitarsisGess,C.wahlenbergiaeGessandQuartinia
vagepunctataSchulthess.Celoniteslatitarsisnestsinsandysoil,excavatingaslopingburrowterminatinginacellinwhichit
constructsanearthencell.C.wahlenbergiaeusesapre-existingburrowinwhichitconstructslinearlyarrangedearthencells.Both
speciescollectsoilforcellconstructionfromaquarryatsomedistancefromthenestandcircumstantialevidencesuggeststhat
bothusenectarasthebondingagent.Quartiniavagepunctataexcavatesaverticalburrowsurmountedbyaturretandterminating
inacell,thewholebeingstabilizedbytheuseofselfgeneratedsilkasthebondingagent.Inorderthatthenestingaccounts
shouldbeputintocontext,nestingbytheMasarinae(sensuCarpenter1982)asawholeisoutlinedanddiscussed.
CelonitesLatreilleandQuartiniaEd.Andreare wardsintoAsiaticRussiaandIndia,andinthe
twoOldWorldmasarinegenera.Celonitesoccurs AfrotropicalRegioninsouthernAfrica(Richards
inthePalaearcticRegioninthecountriesbordering 1962).InsouthernAfricathegenushasalargely
theMediterraneanSea,northwardstoSwitzerland southern and western distribution with, like
and southern Germany and eastwards to Celonites thegreatest numberofspecies having
Transcaspia and southwestern Iran, and in the beenrecordedfromthewesternareas(Richards
AfrotropicalRegioninnortheastAfricaandsouth- 1962,GessandGess1989,labeldataAlbanyMu-
ernAfrica(Richards1962).InsouthernAfricathe seum).
genus has a southern and western distribution AroundfiftyspeciesofQuartiniaareknown,
withthegreatestnumberofspecieshavingbeen morethanhalffromsouthernAfrica.Concerning
recordedfromthewesternareas(Richards1962, thenestingbehaviourofthisgenusthereseemsto
GessandGess1989,labeldataAlbanyMuseum). beonlyonecasualobservationlistedinthediscus-
AroundthirtyspeciesofCelonitesareknown, sion. The present account for Q. vagepunctata
nearlyhalffromsouthernAfrica.Littlehasbeen Schulthess,whichusessilkforstabilizingitsturret,
recordedconcerningtheirnestingbehaviourdue burrowand cell walls is thereforeofparticular
undoubtedlytothecrypticnatureofthenests.Brief interest.
noteshavebeenpublishedonthenests,allaerial,of Theinvestigationspresentedinthepresentpa-
sixspeciesaslistedinthediscussion.Itseemed perwereundertakenduringthecourseoftwofield
likelythatCelonitesasagenuswouldbefoundto tripstothesouthwesternCapeinearlysummer,
construct aerial nests. The present accounts of September/October,of1989and1990.
ground nesting by C. latitarsis Gess and C. Voucherspecimensfromthesestudiesarede-
zvahlenbergiaeGesstherefore,althoughbased on positedintheAlbanyMuseum.
onlyonenesteach,addconsiderablytotheknowl-
edgeofthenestingbehaviourofthegenus. ETHOLOGICALACCOUNTS
QuartiniaoccursinthePalaearcticRegionbor- CeloniteslatitarsisGess
deringtheMediterraneanSeaandextendseast-
Ge—ographicdistributionanddescriptionofnesting
area. Celoniteslatitarsishasasyetonlybeenre-
s'Thhoeulodrdbeeruonfdnearmsetosoids.alphabeticalandequaljointauthorship cCloarndweidllfiraommonthtehetyrpoeadltoocaGlirtaya,fw1a1tekrm(Fwige.st1)oinf
JournalofHymenopteraResearch
Figs.1-3.1,NestingareaofCeloniteslatitarsis11kmwestof
Clanwilliam on the road to Graafwater. 2, Flowerof
Wahlenbergiapsammophila(Campanulaceae).(approx.x5).3,
EarthencellofCeloniteslatitarsisshowingtheroughouter
surfacewithadistinct"fish-scale" pattern,(x6)
thehillyareabetweentheOlifant'sRiverValley sandsuitablefortheexcavationofburrowsand
andthesandycoastalplain. used for this purpose by Scrapter (Colletidae),
Thevegetationischaracterizedbythepresence BelomicrusandBembecinus(bothSphecidae).The
ofRestionaceae,shrubbyProteaceaeandscattered nestofC.latitarsiswassitedinthegentlysloping
Aspalathus spinescens Thunb. (Fabaceae) with a sideofsucha"hillock"whichwasalmostentirely
sparse general ground cover of predominantly coveredwithloosesandandwassitedatthebase
WahlenbergiapsammophilaSchltr.(Campanulaceae) ofasmall,dry,de—adshrub.
andHelichrysumcf.hebelepisDC.(Asteraceae),and Plant visiting. All plants in flower were
a moist ground cover of Monopsis debilis (L.f.) sampledforflowervisitors.Femalesandmalesof
Presl.(Lobeliaceae).Itisbestcategorizedas"Dry C.latitarsiswerevisitingflowersofWahlenbergia
Mountain Fynbos", as described by Moll et al. psammophila (Fig. 2) in company with Celonites
(1984),withanintrusionofsandveldelementson wahlenbergiaeGess,CelonitesbergenumhliaeGessand
disturbedground. MasarinamixtaRichards(alsoMasarinae). Apart
Thesoilissandy,relativelycoarseandlooseon fromW.psammophilatheonlyplantspeciesfrom
thesurfacebutfinerandmorecompactbeneath. whichC.latitarsiswasrecordedwasCoelanthitm
ThefinersandisbroughttothesurfacebytheCape grandiflorumE.MeyexFenzl(Aizoaceae)(1male).
Dune Molerat, Bathyergus suillus (Schreber) On sunny days activity on W. psammophila
(Bathyergidae)whichiscommoninthearea.The flowerswasfrommidmorning,whentheflowers
molehillsstabilizeforming"hillocks"ofcompacted opened, until late afternoon, when the flowers
Volume1,Number1,1992 147
closed. The exact times of flower opening and
closingandmasarineactivityvariedaccordingto
theweather. D
FemalesofC.latitarsisvisitingW.psammophila
flowersentereduptoeightflowersinsuccession,
usuallyondifferentplants.Theyalwaysalighted
ontheoutwardlycurvedfreetipofacorollalobe
beforeenteri—ngtheflower.
Provision. Pollen from the provision taken
fromthenestandexaminedmicroscopicallywas
allofonetypeandma—tchedthatofIV.psammophila.
Descriptionofnest. OnlyonenestofC.latitarsis
waslocated. Itconsisted ofanarched entrance
elmxemcadaivtneagrtmetidoncaaetlsilhnoogfratdtisaaldmoepepitnteghrob5fum3rm5r.mowmWiofitnhdaiinhaomtrehitezeoernxtc4aa,l-5 aFingd.t4.uDmiualugsrafmrsomofanbeosvte;ofb.Cevlerotniictaelsplaltaint.ars(iSsc:aal.enbeasrt=e5n0trmamn)c.e
vated cell wasaconstructedearthencellofthe
samediameterastheexcavatedcell(Figs3and4). seconds(n=36),alternatedregularlywithperiods
dfmliamarTmeheditenleeirlpaearenttgxthttheheen,nwcditeilhdnleegissltbirpepooyubiogenhnitldniygst5ohuvmenoenimevdceawknn.i.dtIhtaTtahitseshl1ei0ognuh-ettceflklry cftieinhlveltebhluteaoisnltesedseditvn.aegnEnmaaacsvtuheecrrcpiaeeagsrlesimioovudfes4it8vnihsstaihetvecseonntbeodsestetnh(denuar=dqidu3nae7g)rd.rwtyAhofitttcehhhere
4mm.Thecellwallsareextremelyhard,thesand waspvisitedasuccessionofW.psammophilaflowers
grainsbeingveryfirmlycementedtogether.The andwasthenlosttosightfor10-20minutes.Onher
outersurfaceisroughandhasavisible"fishscale" returnshealightedateitherthenestorthequarry
pattern.Theinnersurfaceissmoothed.Thecellis site.Itispresumedthatshewascollectingliquidto
pstriolvlMioesptieohnno,idnpgor.fo—cvoinsTsihtorenuricentgiaonrnoetofthtawhvoeinndegisstb,teionevcnitpcoopsmihtpaisloeenstaeniddn. cdmouinrxsatbwrliuetchttithohanen.dAtrhsyetyshawenocdeulltlodwmablaelksheaaridtemhwaaalrtldeeerarbblaeenednfomruoscreeledl
nestconstruction;burrowexcavationandcellcon- andasC.latitarsishasneverbeenobservedatwater
struction.Sandremovedduringburrowexcavation itseemsprobablethattheliquidcollectedwasW.
is not used for cell construction. Sand for this psammophilanectar.
purposeisminedatsomedistanceandcarriedinto At 14h00afterapproximately36additionsto
theburrow.Theburrowentranceisleftopenwhile thecell,thewaspflewofftovisitasuccessionofW.
thewaspisawayfromthenest. psammophilaflowers.Afteranabsenceof49min-
Atllh00on3.x.l990afemaleC./flr/fars/swasseen utesshereappeared.Insteadofgoingtothequarry
tobeinitiatingaburrow.Sandexcavatedfromthe shereturneddirectlytothenest.Afteraperiodof
burrowwasdrawnoutbythewaspasshereversed 11minutes45secondsinthenest,duringwhichit
moumtodfothwenbuslrorpoewftrooamdtihsetabnucreroofwapepnrtroaxnicmea,twehlyer2e0 tisoptrhoebafblloewetrhsa.toPvriopvoissiitoinoinntgoohkadplaccoem,msehnecfeldew.ofAf
itaccumulatedformingatumulus.Fromtimeto regularpatternofflowervisitingalternatingwith
timeacertainamountofrakingofthe"path"tothe aperiodinthenestcontinueduntil16h00whenthe
burrowtookplace.Alsofromtimetotimethewasp waspenteredthenestanddidnotreappear.
flewup,circledandreturned. At17h00itwasdecidedthatthewasp'swork
At11h52burrowexcavationhadbeencompleted forthedaywasover.Thenestwastheninvesti-
and cell construction commenced. During cell gatedandthefemalewhichwasshelteringinthe
pcqorunoasxrtirrmyuacsttieitleoynon2t,h5aemswtaabfsirploimzmeadtdhmeeolnreeersgtau.tlaW"rhhivelilnsoictksa"ttaotph-ea vceilcliMnwaialteysocbfoelhtlahevecitoneuedrs..t.—Nomaleswereseeninthe
quarrysite,thewaspvibratedupanddownvig-
orously whilstscrapingupa load ofsand. The
visitstothequarry,eachtakinganaverageof29
148 JournalofHymenopteraResearch
CeloniteswahlenbergiaeGess
Ge—ographicdistributionanddescriptionofnesting
area. Celonites wahlenbergiaeGess hasbeen re-
cordedfromthreesitesintheClanwilliamDistrict,
oneattheClanwilliamDamandtwoontheroadto
Graafwaterat5kmand11kmwestofClanwilliam.
AtallsitesithasbeenassociatedwithWahlenbergia.
DespiteintensivecollectingonWahlenbergiainar-
easbeyondtheClanwilliamDistrictithasnotbeen
found,suggestingarelativelyrestricteddistribu-
tion.
A nestingarea ofCeloniteswahlenbergiaewas
locatedontheeasternsideoftheClanwilliamDam
onasparselyvegetatedslopeabovethecaravan
park.Thevegetationintheimmediatevicinityof
Clanwilliam is classified as a "Mosaic of Dry
MountainFynbosandKarroidShrublands"(Moll
et al. 1984). That of the nesting area of C.
zvahlenbergiaeisbestdescribedasdepauperatedry
fynbos.ThedominantshrubisAspalathusspinescens
Thunb. (Fabaceae).Groundcoverissparse.The
dominant low growing plant is Wahlenbergia
paniculata(Thunb.)A.DC.(Campanulaceae).The
soilisofthesamenatureasthatattheC.laiitarsis
nacetsiPtvliiantnygt.sviitseitianng.d—isCesliomniiltaerslwyahsluebnjbeecrtgitaoemhoaslebereant O7rF,iongEusng.thd6h,eoo7un.lte6evf,rtiEesaiwusrrstfshoaehfcnoetwchweenilletahotrfhateChdeeilssnomtnociientolcetltsshw"eofafdihslCheei-lnnsobncneeairrltgeei"sasewpuaasrhthfltaeoecnwrbenie,nrogg(fixat6aeh)n.e.
foundcommonlyassociatedwithdeepflowered incompletecell;ontherightisshownthesealofacompleted
CAwD1W1laaaiahsmknnlp;weGminewbwlwsielaestrisshg1ati9mWfa8;oa9ofhs,uaplCnGneelcdendaibsneeswWwsria:iaghtlniwlhladieitncaWGhbomaeseh.strWlasg.Ietinan1pab9Aaae8ln.wlr9iDag)ccCiusaaa.lstaeiC5ptsnlaskaawf(mnmhalwmseowiorweWlpee.slhrtiitsalhopima.enf ctzfafevhllbllaeoou,hswwnl(eexedernrwa6e)bina.dencsr,gegpWi.saaaehIabtllwueitsnnahbdsooefaurnpiggrtnhietlaseiytaernenwsntiatu,nsltawhtvraaihitssesitAtnha.eeerdDvelfCaeaosr,rrgnwevw,eihhcsistiechtarhaeredlwalanboCdsyw.
pollenbymelittidbees.
Celonites wahlenbergiae, however, is not re-
strictedtoWahlenbergiaspeciesasitwasvisiting,in
additionbutlesscommonly,CrassuladichotomaL.
(Crassulaceae)atClanwilliamDam;andCoelanthum
grandiflorumE.MeyexFenzl(Aizoaceae),Herrea
sp. (Mesembryanthemaceae), Polycarena sp.
(Scrophulariaceae), Helichrysum cf. hebelepis DC.
(Asteraceae),anunidentifiedcompositeandPel-
argonium sp. (Geraniaceae) 11 km west of
Clanwilliam.It,however,wasnotrepresentedin
samples ofinsects from other plants in flower,
notablyAspalathusspinescenswhichiscommonly
visitedbythemasarinesCeramiusclypeatusRichards
andMasarina—familiarisRichards.
F=ig5.05.mVme)r.ticalplanofnestofCeloniteswahlenbergiae.(Scalebar cellPwroavsisoiloinve.grePerno,vivseiroynmforiosmtaanfudlldyidprnoovtiasdihoenrede
Volume1,Number1,1992 149
—
tonorwetthecellwalls.Thepollen,examined Malebehaviour. Nomaleswereseen inthe
microscopically,wasfoundtobeoftwotypes,one vicinityofthenest.
matchingonlythatfrom Wahlenbergiapaniculata
andtheotheronlythatfromaCoeknthumspecies QuartiniavagepunctataSchulthess
whichwasgrowingmixe—dwithit.
Descriptionofthenest. ThesinglenestofC. Ge—ographicdistributionanddescriptionofnesting
ivahlenbergiaelocated consisted ofthree linearly area. QuartiniavagepunctataSchulthesshasbeen
arrangedearthencellsattachedtothewallofan recordedfrom38kmwestofCeres,Calvinia,and
apparentlypre-existingburrowexcavatedinsandy DoomRiverFalls(Richards1962).Itishererecorded
swvoehirlit.cicThahlietlcybotunortariondwue,epdtohifnodafisaatpmepeertpoesxrlio5mp,ae5t.meTlmhyr,e3e0decmselcmles,natdftweedor fKRiracorhmoaortdws(o'abDsoiovteeosmothnReitvhNeeirewFueawlslotsue]dratnnvdfirltilhneegeSWkoauftietnrhsfehaoGlorlgeta1=et
sealedandthethirdinanearlystageofconstruc- Pass,15kmnorthofNieuwoudtville)andfromsix
tion,werepositionedattheupperendoftheslope sitesinNamaqualand(GoegabNatureReserveto
(Fig.5). theeastofSpringbok;Narap,Klipfonteinandthe
Thecompletedcellsareroundedattheinner Wildeperdehoek Pass, all to the south west of
eatrnnuddn,c4artoemu.gmhTlhryeeyospvaeorcietdi9vbeaultnydwii8ntmhoumttheerinsdeliaelanemgdettho,eurtaenartdet4nh,de5 SSpprrAiinngngbebosotkki);n.gaanrdeaAonfeQn.ovuagseptuonctthaetanwoarsthlowceatsetdoinf
widestpoint.Theouterdiameterattheneckof theSkuinshoogtePass(Fig.8).Thevegetationis
eachcellis4mm.Thecellwallsareextremelyhard, probablyclosesttoAcocks'VeldType28,Western
thesand grainsbeingveryfirmlycemented to- MountainKaroo(Acocks1953,1975).Thenesting
gether.Theoutersurfaceofthecellwallisrough sitewasabarepatchofsomewhatunevenlevel
andshowsadistinct"fish-scale"pattern(Fig.6). groundbetweenshrubs.Thesoilwassandyand
Theseal,constructedfromthesamematerialasthe friable.
cellwalls,ispositionedwithintheneckofthecell Thesiteapproximatelyonesquaremetre,was
(Fig.7).Thebaseofasucceedingcellisattachedto locatedinOctober1989andrevisitedinSeptember
thesealofaprecedingcellsothatitispositioned 1990whenitwasagainbeingactivelyused for
withintheopeningofthatcell.Theinnersurfaceof nesting. —
thecellsissmooth(Fig.7). — Plantvisiting. Quartiniavagepunctatawasfor-
Methodofconstructionofthenest. Nestinitia- agingalmostexclusivelyonAsteraceae:extremely
tionwasnotobserved.Apre-existingburrowwas commonlyonflowersofRelhaniapumilaThunberg
probablyusedasthecellsinthenestinvestigated (Fig. 9) in the Skuinshoogte Pass and Leysera
werepositionedontheburrowwallwellabovethe gnaphalodes(L.)L.intheGoegabNatureReserve,
innerendoftheburrowand astheirdiameters andatNarapandAnenous;andlesscommonlyon
werelessthanwasthatoftheshaft. flowers of Leysera gnaphalodes above the
Sand forconstruction ofthecells wasbeing NieuwoudtvilleWaterfall,Cotulacf.leptaleaDC,
q"uhialrlroicekd"fsirtoumattehde3smurffracoemotfheabsutrarboiwli.zAedtmmoildedraayt SHeuntecchi,oepxrMobe.rxnimv.euasndLeOssst.,eoPsepnetrzniuaansucfff.rouptpiocsoistaifo(lLi.a)
on19.x.1989thewaspwasseenvisitingaCrassula intheSkuinshoogtePass,Pentziasuffruticosainthe
dichotomaflowerafterwhichsheflewtothequarry. WildeperdehoekPass,IHelichrysumsp.andCotula
Atthequarrysiteshevibratedvigorouslyappar- sp.atAnenous.
entlylooseningsandwithhermandibles.Having Theexceptionswereafemalecollectedonalow
gatheredaloadsheflewwithittotheburrow.Sand growingGaleniasp.(Aizoaceae)growingamongst
gatheringwasrepeatedseveraltimes.Thewasp LeyseragnaphalodesatAnenousandamalecollected
wasthencapturedandthenestinvestigated. onaflowerofLebeckiacf.sericeaThunb.(Fabaceae)
Asthecellwallsareharderandmoredurable at Klipfontein. It is of interest that these were
thanwouldbeexpectedhadwaterbeenthebond- visiting flowers of families favoured by other
ing agent it is probable that nectar was used. masarines.
Celonitesivahlenbergiaehas,notably,neverbeenseen Theflowersofallbuttwoofthesespecieswere
atwater.ItseemslikelythatthevisittoCrassula yellow.Senecioprob.niveushaswhitishflowersand
dichotomawasforthepurposeofcollectingnectar. theGaleniasp.pinkflowers.
150 JournalofHymenopteraResearch
Figs.8-11.8,NestingareaofQuartiniavagepunctata,Skuinshoogte Pass,15kmnorthofNieuwoudtvilleontheroadto
Loeriesfontein.9,QuartiniavagepunctataonflowerofRelhaniapumila(Asteraceae).(approx.x3).10,NestingsiteofQuartinia
vagepunctata,arrowindicatingsandandsilknestentranceturret,(approx.x1). 11,Dorsalviewofsandandsilknestentrance
turretofQuartiniavagepunctata.(x14,4).
Provision.— Provision from four nests of size:0.16mm- 1,2mm) held togetherbyasilk
Quartinia vagepunctata investigated in the lining.Theturretisbag-like,approximatelycircular
SkuinshoogtePasswasintheformofarelatively incross-sectionwithitsdiametergreatestatits
moistbrightyellownectarandpollenmassalmost outerendandsmallestatitsinnerend.Theopen-
entirelyfillingthecell,adheringtothecellwalls ingtotheburrowentranceisatsomelittledistance
andthereforenotformingadiscretepollenloaf. fromtheclosedinnerendofthebag.
Tfhroempoonlleenneswtaswaesxoafmionneedtympiecroonslcyopaincadllmya.tcThheadt ofaTshuebvbeurrtircoawlsihnatfth,e1n,e5s-t2sminmveisntidgiaatmeedtecro,nstiesrtmeid-
thatofCotulaandthatfromtheotherthreenests natinginasealedroughlyovoidcellatdepthsof
wasoftwotypesmixedandmatchedthatofRelhania 25-30mm.Thecellwallswereconstructedofsand
andCotula. — bondedtogetherwithsilkandwellcementedwith
Descriptioti of the nest. Seven nests of Q. anunidentifiedsubstancesomewhatresinousin
vagepunctata wereinvestigated,fouron 7.x.1989 appearance. Inoneofthe nests thefemalewas
andthreeon27.ix.1990.Eachhaditsentranceto foundshelteringinalateralshaft,suggestingthat
onesideofanearthclodorstone(Fig.10). morethanonecellpernestisprobably—constructed.
Thenestconsistsofasubterraneansilk-lined Methodofconstructionofthenest. Thesoilin
burrowsurmountedbyahorizontalturret(Figs11 whichthenestisexcavatedisfriable.Waterisnot
and12),theoutersurfaceofwhichisofsand(grain requiredfornestexcavationandisnotusedasa
Volume1,Number1,1992 151
DISCUSSION
Richards(1962)sawthefamilyMasaridaeas
constitutedofthreesub-families,theEuparagiinae,
GayellinaeandMasarinae,andtobea"sistergroup"
ofthefamiliesEumenidaeandVespidaeinasu-
perfamilytheVespoidea.Carpenter(1982)assessed
thephylogeneticrelationshipsofthecomponents
oftheVespoidea(sensuRichards)usingcladistic
methods.HetreatedRichards'Vespoideaasasingle
familyVespidaeinwhichherecognizedsixsub-
families, Euparagiinae, Masarinae, Eumeninae,
Stenogastrinae,PolistinaeandVespinae.Hethereby
disassociated theEuparagiinaewhichprovision
withbeetlelarvaefromRichards'Gayellinaeand
Masarinaewhichprovisionwithpollenandnec-
tar.Atthesametimeheassociatedmorecloselythe
Gayellinae (sensu Giordani Soika 1974) and
Masarinae{sensuRichards1962)byplacingthem
togetherastribes(GayelliniandMasarini)inhis
subfamilyMasarinae.Followingthisgroupingthe
presentauthorshaveexcludedtheEuparagiinae
fromtheirdiscussion.Ceramius,themoststudied
genus, isdivided intospeciesgroups following
Richards(1962)amendedbyGessandGess(1986,
Fig.12.PlansofturretofQuartiniavagepunctata.a.fromabove; 1990).
b.verticalsection;c.frombelow.(Scalebar=5mm). InorderthatthenestingaccountsforCelonites
latitarsis, Celonites wahlenbergiae and Quartinia
bondingagent.ItisthereforenotsurprisingthatQ. vagepunctatashouldbeputintocontext,nestingby
vagepunctuta,thoughcollectedcommonlyatflow- theMasarinae(sensuCarpenter1982)asawholeis
ers,hasneverbeencollectedatwater. outlinedanddiscus—sed.
Thesilkusedinnestconstructionisspunbythe Basicnesttypes. Fromareviewofthepub-
nestbuilder.Oneindividualwasobservedwhilst lishedandpresentaccountsofthenestingofthe
itwasjoiningtogethersandgrainswithsilk.Itwas Masarinaeitispossibletorecognizesevenbasic
rotatingitsheadandthesilkwasapparentlyissuing nesttypes:
from its mouthsuggestingthatthesilkmaybe Nesttype1:amulticellularsub-verticalburrow
producedbymandi—bularglands. inhorizontaltosub-horizontalgroundexcavated
werMealceombemhoavnioounrt.hegMraoluensd,iunnltihkeevitchieniftyemoafletsh,e bfyrotmheeanretshteer,xtwriatctheadnfernotmrawnictehitnurtrhetecbounrsrtoruwcbtuedt
forageplants.Theywereobservedtoriseupand withtheexcavatedcellsnotcontainingconstructed
mountfemalesvisitingtheseplants. cells:
Maleswerealsopre—sentatthenestingsite. FourspeciesofCeramius:allspeciesofGroup8
AlloAcsoseolciaiatmeodscainrsyeicts(Br.aunSse)ve(rCahlryisniddiivdiadeu)alwseroef -GeCs.sca1p9i8c0o)la,BC.rabuincosloarn(dTCh.unlbineeragri)s(KGleusgs(aGnedssGaensds
present in close proximity to the nests. As this 1986)andC.sociusTurner(GessandGess1988b).
chrysididhasinadditionbeenfoundbytheau- Nesttype2:amulticellularsub-horizontalbur-
thorsinassociationwiththreeQuartinioidesspecies rowinverticaltosub-verticalgroundexcavatedby
andasallknownAllocoeliaassociationsarewith the nester, with an entrance turret constructed
masarines(KimseyandBohart1990)itissuggested fromearthextractedfromwithintheburrow,and
thatthiswaspismostprobablyanestparasiteof withthewallsofeachexcavatedcelllinedwith
QuartiniaandQuarUnhides. cementedearthexcavatedwithintheburrow:
152 JournalofHymenopteraResearch
OnespeciesofMasarina:M.familiarisRichards Nesttype5:constructedearthencellslocatedin
(GessandGess1988a). apre-existingcavity;soilforcellconstructioncol-
Nesttype3:amulticellularsub-verticalburrow lectedfromaquarrysiteatsomedistancefromthe
inhorizontaltosub-horizontalgroundexcavated nest:
bythenester,withorwithoutanentranceturret OnespeciesofCelonites:C.ivahlenbergiae(present
constructedfromearthextractedfromwithinthe paper).
burrow,andwitheachexcavatedcellcontaininga Nesttype6:aself-excavatedslopingburrowin
constructedcellformedfromearthexcavatedwithin friablesoilwithanexcavatedcellinwhichisan
theburrow: earthencellconstructedfromsoilcollectedfroma
Three species ofParagia: P. (Paragia) tricolor quarrysiteatsomedistancefromthenest:
Smith (Houston 1984); P. (Paragia) decipiens OnespeciesofCelonites: C. latitarsis (present
Shuckard (Naumannand Cardale 1987);and P. paper).
(Cygnea)vespiformisSmith(Houston1986). — Nesttype7: asub-verticalburrowinfriable
Eleven species of Ceramius: Group 2a C. soil,surmounted bya sand and silkturretand
cerc—eriformisSaussure(GessandGess1988b);Group havinganexcavatedcellinwhichisaconstructed
2b C.clypeatnsRichards(G—essandGess1990); sandandsilkcell:
Groupuncertain,probably2b C.micheneri(—Gess OnespeciesofQuartinia.Q.vagepunctata(present
and Gess 1990); all species of Group 3 C. paper).
nigripennisSaussure(GessandGess1986),C.jacoti Groundnestinghasbeenrecordedforafurther
Richards(GessandGess1988b),C.braunsiTurner elevenspecies,however,theobservationsaretoo
andC.torigerSchulthess(—GessandGess1990);the incompleteforthesespeciestobeattributedtothe
singlespeciesofGroup5 C.li—chtensteinii(Klug) nesttypesassetoutabove:(Paragia(Paragia)smithii
(GessandGess1980);Group6 C.rexSaussure Saussure(Wilson1869);Rolandiamacidata(Meade-
(GessandGess1988b)andC.metanotalisRichards Waldo) and an undescribed species of Riekia
(—Gess and Gessunpublished fieldnotes); Group (Houston1984);CeramiopsisparagnayensisBertoni
7 C. tuberculiferSaussure(Giraud 1871,Ferton (almostcertainlyasynonymofC.gestroiZavattari
1901). (Richards 1962)) (Bertoni 1922 as reported in
TwospeciesofJugurtia:J.confnsaRichards(Gess Richards1962);threespeciesofCeramius,Group1
andGess1980)and/.brannsi(Schulthess)(Gessand -C.fonscolombeiLatreille(Fonscolombe1835),Group
Gessunpublishedfieldnotes). 7-C.bischoffiRichards(Richards1963a),andGroup
Nesttype4:agroupofconstructedearthencells 4-C.beyeriBrauns(Brauns1910,GessandGess
attachedtoplantstemsorstones: 1988b);twospeciesofTrirrteria,T.howardiBertoni
SixspeciesofCelonites:C.abbreviatus(Villers) (Zucchietal. 1976asreportedinHouston1984)
(Lichtenstein1869(asC.apiformisFabricius),Ferton andT.buyssoniBrethes(NeffandSimpson1985);
1901, 1910, Bellmann 1984); C. fischeri Spinola Quartiniasp.(JacotGuillarmodpersonalcommu-
(Bingham1898asreportedinRichards1962);C. nication), and Quartinioides sp. (Gess and Gess
mayetiRichards(Lichtenstein1875,Ed.Andre1884 1988a,1989).Conflictingaccountshavebeengiven
asreportedinRichards 1962);C.jousseaumeidu for Masaris vespiformis Fabricius. Morice (1900)
Buysson (Richards 1962); and C. andrei Brauns suggestedthatthisspeciesisgroundnestingand
(Brauns1913);andinadditionaputativenestofC. Ferton(1920)thatitmakesaerialmudcells.
promontorii(Brauns)(GessandGess1989). DorrandNeff(1982)describedanestinabeetle
RohEwigehrt(sRpiecchiaersdsof19P6s3eb)u;doPm.aseadrwiasr:dsPi.i(coCcrjeuislsloetnt)i buonrliinnge.dTcheellnsesstepcaornastiesdtedbyofmauldinepaarrsteirtiieosnso.fTfhoiusr
(Torchio1970);P.maculifrons(Fox)(Parker1967); theyallegedtobeanestofPseudomasarismarginalis
P.occidentalis(Cresson)(Hungerford 1937asre- (Cresson),howeve—r,confirmationisrequired.
portedinTorchio1970);P.phaceliaeRohwer(Parker Bondingagent. Threebondingagents,water,
1967,Torchio1970);P.texanus(Cresson)(Bequaert nectar,andsilk,areknowntobeusedbymasarines
1940 as reported in Torchio 1970); P. vespoides innestconstruction.
(Cresson)(Torchio1970);andP.zonalis(Cresson) Useofwaterinexcavationandasthebonding
(Parker1967). agent is either stated or implied in all nesting
OnespeciesofCayella:G.eumenoidesSpinola accountsofNesttypes1,2and3.Inadditionthe
(Claude-Joseph1930asreportedinRichards1962). innersurfacesofthecellsofParagia(P.)tricolorare
Volume1,Number1,1992 153
polishedandwaterproofedwithanunidentified masarinesbutforVespidaeasawhole.Further-
substance(Houston1984). moreitseems thatnesting in friablesoil in the
Nectaristheprovenbondingagentemployed Vespidaeisprobablyderivedratherthanprimitive
byPseudomasarisedwardsiiofNesttype4(Torchio asinthePompilidaeandSphecidae.ApartfromC.
1970). Circumstantial evidence, available to the latitarsisnoneofthosespeciesknowntoexcavate
presentauthors,furthermoresuggeststhatnectar nestsinfriablesoilhasforetarsalsandrakesas
isusedbyCelonitesofNesttypes4,5and6. possessedbymanygroundnestingSphecidaeand
Theuseofself-generatedsilksetsNesttype7as Pompilidae.Soilremovaliseffectedbytheman-
exemplifiedbyQ.vagepunctataapartfromallthe diblesasinthosespeciesexcavatinginnon-friable
others.Theuseofsilkinnestbuildingbywasps soil.ForexamplePseudepiponaherrichii(Saussure),
seems to be altogether uncommon. It has been aeumeninenestinginaverticalburrowinfriable
notedfortwoeumenines,onegroundnesting(Gess ground,removessandparticleswiththemandibles
andGessunpublishedfieldnotes)andonenesting oneatatime(Spooner1934asinSpradbery1973).
inpre-existingcavities (Weavingpersonalcom- Theonlyrecordedmorphologicalmodificationfor
munication),andhasbeenrecordedfortwosocial sand removal is that of the mouthparts of
sphecids,oneconstructingaerialnests,Microstignms Pterocheilus (Bohart 1940) for which nesting in
comesKrombein(Myers1934,MatthewsandStarr verticalburrowsinfriablesoilbytwospecieshas
A1r9p84a)ctoapnhdiluosneminemsitiNngauimnapnren-ex(iMsattitnghecwasvitaiensd, btheeenmarseacroirndeesd(tIusrerleytl1e9s1s4,inEcvlainnesd19b5u6r)r.oAwmsonexgcsat-
Naumann1988).Inbothsphecidspeciestheadult vatedinsandygroundhavebeenrecordedforan
waspssecretethesilkfromglandsnearthetipof undescribed species of Riekia and for Rolandia
the metasoma. Adult Q. vagepunctata observed maculata(Houston1984).Unfortunatelythemethod
appearedtoproducesilkfromtheirmouthsandit ofexcavationwasnotnotedandthenestswere
issuggestedthereforethatsilkismostprobably incomplete.
producedbythemandibularglands. Notonlyis thesubstrateand themethod of
Usingnectarorsilkasabondingagentfreesthe excavation of theburrows of Nest type 6 very
userfromdependenceonwater,anoftenephem- differentinnaturefromthatofNesttype3but,as
eralresourceinaridareas.Theuseofsilkfurther- importantly, so isthenature ofand method of
moremakesitpossiblefortheuserstoconstruct constructionofthecells.Whereastheearthencells
nestsinandwithfriablesoilwhichotherwisebe- ofNesttype3areconstructedfromsoilquarried
comesreadilyunstableunderdryconditions. — withintheburrowandbondedwithwaterthoseof
Methodofnestconstructionbygroundnesters. Nesttype6areconstructedfromsoilquarriedat
Inthefirstthreenesttypeswateriscarriedfroma somedistancefromtheburrowandbondednot
watersourceinthecrop.Onarrivalatthenestitis with waterbutmostprobablywith nectar. The
regurgitated and worked into thesoil with the methodofconstructionandnatureoftheearthen
mandiblestoformmud.Thespoilsofexcavation cellsofNesttype6asexemplifiedbyC.latitarsisin
areremovedwiththemandiblesintheformof nowaydiffersfromthatofNesttype5asexempli-
mudpelletswhichareeitherdiscarded,usedfor fied by C. wahlenbergiae nesting in pre-existing
theconstructionofaturretorfortheconstruction burrowsandthatofNesttype4asexemplifiedby
ofcells. theaerialnestingCelonite—sspecies.
InthesixthnesttypeexemplifiedbyC.latitarsis, Evolutionarysequence. Apossiblesequenceis
inwhichtheburrowisexcavatedinfriablesoil, discernablewithintheMasarinaefromexcavated
waterisnotusedandthespoilsofexcavationare burrowswithexcavatedcellsonly(Nesttype1)
rakedoutandaccumulatetoformatumulus.This through excavated burrows with constructed
isofparticularnotewhenthestructureofthefore earthencellswithinexcavatedcellswithearthfor
tarsiofC.latitarsisiscomparedwiththatoften constructionbeingderivedfromwithinthebur-
otherAfrotropicalspeciesofCelonites(Gess1992). row(Nesttype3)tothepresumablymoreadvanced
Ofthosespeciescompared,onlyC.latitarsishas constructionofaerialearthencells(Nesttype4)
widelyexpanded tarsomeressuitableforraking (discussedinGessandGess1980).
soilwhichsuggeststhatitsnesttypemaybeunusual Afurtherpossiblesequencewithinthegenus
forCelonites. Celonitesisheresuggested,thatisareturntothe
Sandrakingseemstobeunusualnotonlyfor ground from aerial earthen cells (Nest type 4)
154 JournalofHymenopteraResearch
throughconstructedearthencellsinpre-existing Bohart,R.M.1940.ArevisionoftheNorthAmericanspecies
bccaauvvriarttioeewsdsincwetilhtleshgcwroiontushntrdeua(crNtteehsdtfetoayrrptecho5e)nnsttcoersluelclsftweiioxtnchaivbnaetieenxdg- Brau3oV3fne(ssP1p,t)ie:rdo1Hac6e.l2)w-.i21l09Au81sn.0na.anldsBnioooftletoshgeoinEsncrtheoelmasotleoidgiigbceearnlersSaioci(idHeatyfymreoinfkoaApnmtieesrrciach,ae
minedoutsidetheburrow(Nesttype6). Hymenopteren. Zeitschrift fiir wissenschaftliche
Thissecondsequenceissuggestedbythemethod Insektenbiologie6:384-387,445-447.
oomrffaaskcsaioarnnnigsdntb-erersuha,ckatsevoisiiolonaufsororfywceeNitltelhsnctoottnhtseyrtpcereoucnco6tsr,iedonqenoudtebanefbotilrnypgaootnshbsyeteasosistanihneoeddnr CBarrapun8(en)aH:nstyt,u1mer9Hr.a3.,)l-21Jv0c.99lo1.anM3s..sSiDif1rii9dic8ata2ftt.reiirokTnBah.eoeifEtnrpttahhogyemlzooVulgeroesgnKpieeostnciinhdcetenaMriies(lltHadetyeiimrloeunMnnsaoghspeiantprse2irada(ne)7dn./
fromasiteatsomedistancefromthenestnotfrom SystematicEntomology7:11-38.
awNsietsuNhtseietsndyttpihteney3Nnp.eeesst7t,atisynpddiets4thieanbncotdna5dnindnotgiswaagptoesenstribabesliyinsdguensreeicdvteaidrn CDlorarusiZ,dnoueorLl-.loJJlog.egossiasieepnnprd(Che1oN0,dle)a1fFot.3fre:,(auJJ2rd.a3sonL5.v(-di2H1ue59yr48Cm,2.he.iHnl.Poi)s.pe1tAu9end3rno0aam.:laResMesaacrsdiheasesrrmiscadchraigeeeisn)nc.aebslPiiaonsnla-otnguePrisaeqctliuilfeneisgsc
fromaverticalburrowexcavatedinstablefriable Entomologist58(2):81-91.
soilwithouttheuseofabondingagent. Evans,H.E.1956.Notesonthebiologyoffourspeciesof
ground-nestingVespidae(Hymenoptera).Proceedingsof
ACKNOWLEDGMENTS theEntomolocicalSocietyofWashington58(5):265-270.
Ferton, C. 1901. Notes detachees sur l'instinct des
Hymenopteresmelliferesetravisseursavecladescription
Thefollowingarethankedwithappreciation:MrD.W. dequelquesespeces.AnnalesdelaSocie'teentomologiquede
Gessforassistanceinthefield,inparticularforhisdiscovery France70:83-148.
ooQffuatrththeienneiMsatuisvnaegguseipmtuenoocfftaQtuCaao;rmtpiMansiraatJv.aigvBeeepyuneZcrotsoaltoaof;gyDtrhfeJo.rMS.tiedClealnretpniebfnoytsiecnhrg FertdHoeynml,aeSnCooc.ipett1ee'9r1ee0ns.tommNeolollotigefiesqrueedsedetetarFacrvhainesceseesu7r9ss:.u1r(4b5el-S'1ei7rn8is.et)i.nAcntnadleess
HerbariumforidentifyingWahlenbergiacostata,Wahlenbergia Ferton, C. 1920. Notes detachees sur l'instinct des
panicitlata, Wahlenbergia psammophila and Coelanthus Hymenopteresmelliferesetravisseursavecladescription
grandiflorum;MrA.J.S.WeavingoftheAlbanyMuseumfor deuxespecesnouvelles.(9eSerie).AnnalesdelaSociete
assistancewithtakingthephotographsreproducedasFigs4, entomologiquedeFrance89:329-375.
6,7and12,andforproducingblackandwhitenegativesfrom Fonscolombe,H. Boyerde1835.DescriptionduCeramius
theauthors'colourtransparenciesforFigs1,2and8-10;and fonscolombii(Latr.)AnnalesdelaSocie'teentomologiquede
the Foundation for Research Development for running France4:421-427.
expensesgrantsforfieldworkduringthecourseofwhichthe Gess,F.W.1989.NewspeciesofthegenusCelonitesLatreille
presentinvestigationswereundertaken. (Hymenoptera:Masaridae)fromSouthAfrica.Annalsof
theCapeProvincialMuseums(NaturalHistory)18(4):83-94.
LITERATURECITED Gess,F.W.1992.AnewsouthernAfricanspeciesofthegenus
CelonitesLatreille(Hymenoptera:Vespidae,Masarinae)
AAABBBBBcceneeilrooqdlnutucclrgtCtvRZtAaoA((EoakkmeohhefeohhknnnHHdelsseeal,voitaga.tdyry,e,oinl,ohdemBBtmnsmo2Ennemooe]Jn,,.iteogd,..eottAmta,Bibnn.laaBn].CePPiso.yoonnHde...soce1giimtp.eaC1Ph8pdcciHHhTbt.u9oea8daalce.te..a4ler1nb4aellae0.9by.rer1l111ASf.W8rrSS:8aY999noo4eSuuN9.aSW,57e6Ncz.opva8rroAi35,ri1eeoid.tcvvs.n.Debc.iaB1tiuseeididTutg9eereeVpVVyyepautre2rhanloseeseDldys2ryoosleeabllosi.ff,cdnoc3pdad2pHeehiaaVt5SnS1tRciiesaic:sNnoo2TTen.shdflu:utuortdFyyH3Atltatloihttav7yopepa.eef3efueh)hA-rrmniee,ea2l)n.4sncyea1tsMddsAAt.5agdzn-eifnfaAa.Seeru(3odroosrdbsodHnneie2ffitpBHhcalCecc8ryuBw.tiyarSSaaa.iezlseemPenmioorpbltrdau42eeslduuutye09leriHnnmpti.ttt::isna1iioinohhei2Ngomniipapd:Bnn(u-u-neAAte'riKtgiraffn1vvEeloueoesyrr,,0oMafuaftrniier1spri1vn1acc1,-saPto-:-dtetaad:e1s11peeh.I.rt1M1pe92r.eerhf4e1GaS28urMMioa-..a.Di.lesdoeell1CrBteyeoacmm.2otruemACur.uooegjotllahririinoiogslseeuodrreckciamrrvdpasoslhnrinioeasyeiaeaeaoon.)snnse,.ltff GGGGeeeessssssssSacaCbiC(Mt3kt1k(,,,,inoh64hasaaaCnHnnce4n3FeFFsuppsadytoooF.....faseeCmoh.lmwwuEWWWroesCapcesallPPr.i...upaWianWsrreenerna.otoeol(aaaRddaeutvivepsTPinnnggciilertrhtacaRdhddneenandouehetnnicccveGnaGGorriicddCoineerbfeeaaanhfnsaasdllessCt:acwtGCepss.ssretirhiMMe,,i),,agehaedsMstnuu.l)epoSsCSSaisehss,.cP..ieeMs,jttieer(rKKaKouhu(aCPoH.uu.a.ursSoKs.rvxyrt.mmmill111Teoinohmidssu9oc99uvanueKune8aglg88lirc.ms0nyo)((e6y8nen.poNN.so)bmccefoe1(aa.pfayEoEefar9tt(HHiptlftttA8uunNiyyotieohu(a8rrcafShmmosorwtoHaaanfteoeleSllh.alleuuyduonoanoRgerr:rmgtgHHoiouetaABseeisiiihcphltpnrMtncsscohethtuaahattHaouAarceeussooliolfprAtoorncarrrslrfdthfnsCanyyostriaroseteteo))on,ici:g,rurRrrrtatdcydCie11nya.earaaWM.6sCib3)ms.i:e(aao.eeuAaibl)7Afao(1shstnniMuu)6.rn6nanlinn):tdasocteni:oreaifsA(hhaCgan1iLn1lo.6rwneajerl6a4disnb3neruir)t1ss1ta-gegpa:sira:-to8oeueKeoreltd1omf331rn)filngs7eai.5tt4tnltt8uuireu1hh1iilihho.gnsa)s-eee-asneef
GeologicundBiologic)No.2:1-251.