J.HYM.RES. Vol.6,1997,pp.75-91 Update on the FlowerAssociations of Southern African Masarinae with Notes on the Nesting ofMasarina stntcki Gess and Celonites gariepensis Gess (Hymenoptera: Vespidae: Masarinae) SarahK.Gess,FriedrichW.Gess,andRobertW.Gess AlbanyMuseum,Grahamstown,SouthAfrica — Abstract. FlowervisitingrecordsarepresentedforsevennewlydescribedspeciesofMasarinae fromtheRichtersveldandforMasarinastruckiGess.Thepotentialofthesewaspsaspollinators isdiscussed.CelonitesgariepensisGessandMasarinapeliostoiniGessareassociatedwithPeliostomum andtheformer,atleast,withAptosimum(Scrophulariaceae),apreferencesharedwithfourother speciesofCelonites.Masarinati/lecodoniGessappearstoberestrictedtoTylecodonhallii(Tolken) Tolken(Crassulaceae)and isapparentlythefirstrecordedinsectvisitortoaTi/lecodon species. JugurtiacodoniGessisanabundantvisitortoCodonroyeniiL.ofthefamilyHydrophyllaceae,one ofthepreferredforageplantfamiliesoftheNorthAmericanmasarinesbutotherwisenotknown tobevisitedbymasarines.RecordsforMasarinastruckifromawiderangeoflocalitiesconfirm its,foramasarine,unusualassociationwithHernianniaspp.(Sterculiaceae).IntheRichtersveld Zygophyllumspp.(Zygophyllaceae)werebeingvisited,thoughnotapparentlyfavoured,byCer- aniiusbrevitarsisGess,twospeciesoiJugurtia,J.codoniGessand/.koeroegabensisGess,andMasarina tni.xtoidesGess.TheserecordsareofinterestasZygophyllumhasonlyotherwisebeenshownto attractoneotherspeciesofMasarinae,aQuartimoidesspecies,intheCentralNamibDesert.Some aspectsofthenestingofC.gariepensis,whichconstructsaerialmudcells,andofM.strucki,which nestsintheground,arepresentedanddiscussed. Gess(S.K.1996)presentedasynthesisof /. braunsiella(Schulthess),/.calcarataRich- theavailabledataonthenestingandflow- ards,and/.duplicataRichards,twospecies er visiting of the Masarinae in southern of Celonites, C. ch/peatus Richards and C. Africa. Recent fieldwork insouthwestern pcliostomiGess.Flowervisitingrecordsfor southern Africa, mostnotablyinthepre- thesespeciesofJugurtiaandCelonitescon- viously under collected Richtersveld Na- firm previously recorded flower family tional Park in the extreme north of Na- choices(Gess,S.K. 1996). m18a.qixu.a1l99a6n),dh(aFisg.re1s)ul(t1e6d-24i.nixt.hle99d5isacnodve5r-y CelonitesLatreille ofnewspecies,additionalflowerassocia- Celonit—esgariepensisGess tionsofparticularinterest,andanesteach Distribution. Celonites gariepensis Gess ofMasarinastruckiGessandCeloiitesgar- hasbeenrecordedfromonlytheRichters- iepensisGess.Descriptionsofthenewspe- veldNationalParkinnorthernNamaqua- ciesaregiveninGess(F.W. 1997)andthe land(Gess,F.W.—1997). flower visiting and nesting data are pre- Floioervisiting. OntheKoeroegabvlak- sentedanddiscussedinthepresentpaper. te(28.11S,17.03E)(Figs.2-5)C.gariepensis Some other masarines collected in the femalesandmaleswereforagingonlyon Richtersveld National Park, allextending flowers of a species of Peliostomum the known distributions for the species, (Scrophulariaceae),togetherwithCelonites are: Ceramius cerceriformis Saussure, four ch/peatus Brauns, Celonites pcliostomiGess speciesofJugurtia,j. braunsi(Schulthess), (Fig.6)andMasarinapcliostomiGess(Fig. 76 JournalofHymenopteraResearch 27 29 31 33 Volume6,1997 77 W^^v 78 JournalofHymenopteraResearch NWontheroadfromOkaukuejotoOkon- deka,3.iv.l996,byD.W.andG.T.Gess.A sampleoftwelve—femaleswastaken. Watervisiting. C.brci'itarsiswasfound tobeabundantatatrickleofwatercross- ing the road in a rocky pass (28.10S, 17.02E)immediately to the northofKoe- roegabvlakte. Whilst imbibing water the wasps stood on wet sand or rock at the edgeofthewater.Onefurtherfemalewas observedimbibingwaterfromtheedgeof anisolatedpoolofwaterinahollowona steeply sloping rock. In this instance the surfaceonwhichthewaspstoodwasdry, indicating that water is imbibed directly from the pool and not from wetsand as has been observed for Ceramius hicolor (Thunberg)(Gess,S.K.1996)andCeramius sociusTurner(GessandGess1986).Inno casedidoneofthesewaspsalightonthe watersurface. — Flowervisiting. Allplantsin flowerin thevicinityofthewaterweresampledfor flowervisitorsthroughout theday, how- ever,onlyonefemaleandonemaleofC. brevitarsis were taken. Both werevisiting the small cream flowers of Zygcphyllum prismatocarpum E. Mey. ex Sond. (Zygo- phyllaceae).ThisisthefirstrecordofCer- amius visiting flowersofZygophyllaceae. Therecordsare,however, toofewforan Figs.6and7. Pcliosiomumsp.(Scrophulariaceae).6, associationtobesuggestedasinstancesof MCfeallsooawneriritcnasappppnreilioioxosisttcmcnavitniieGlGeyes2ss0spmremenp)tae.rriinnggtfoloewnetrer(flleonwgetrh.o7f, cpfoarrseufsaelrormveeidsishtpaienscgiooecfscafoslfioowCneearrlslaymoitbuheseren(tGherasensc,otrhSdo.esKd.e 1996). Aizoaceae: Mesembryanthema south. To the north of the Richtersveld wouldbe theexpected planttaxon tobe NationalParkthereisasurprisinghiatus favouredbyC.brevitarsis,itbeingamem- intheknowndistributionofCeramiiis,no berofGroup2A and therebeing consis- species having been recorded between tencyofflowerfamily choicewithinCer- thereandnorthernNamibiawhenceasin- amius species groups in southern Africa glespeciesfromGroup4,Ceramiusdainar- (Gess,S.K. 1996). i(i1n9i3s5)Tufrrnoemr,femwaalsesdaensdcraibmeadlebyfroTmurOnne-r JugurtiadeSaussure gandjera (1923)anda malefromKaman- Jugu—rtiacodoniGess yab (1925), part of material collected by Distribution. Jugurtia coiioni Gess has the staff of the South African Museum beenrecordedfromonlytheRichtersveld (Gess, F.W. 1965). Recently this species National Park in northern Namaqualand was recorded flying abundantly 10.3 km (Gess,F.W. 1997). Volume6,1997 79 Figs.8-12. ParadiseKloof, RichtersveldNationalPark.8,Approachtokloot.9,NestingsiteofCehnites ^mkyensisGessmarkedwitharrow.10,NestofC.gariepensisonundersideofjuttingedgeinrockcrevice (outercellbrokenduringcaptureofbuilder).11,CellofC.gariepensis.12,CellofC.gariep\'nsisopenedto showpapillateprovisionmassandpositionofegg. — weWraetceorllveiscitteidnga.ttTheweodgfeemoafleaspoofol/.ocfodwoiai-i cfrlooswseirnsgotfhCe.Krooeyrenoieigaarbevl3a5kmtem(Fhiig.gh5,).erTehcet terinParadiseKloof.Thissuggeststhatit andcampanulate(Figs. 13-15).Themany is probable that /. cocioni, like the other lobedcorollaiswhite.Thereare10-12sta- kspneocwines,uosfesJugwiaitrtei—raifnorneswthicconhstnreuscttiionng.is mroelnlsa.tTubheeafbioluamten5tsmamrefartotmachtehdetboasteheacnod- Flower visiting. Females of /. codoiii arecloselyadpressedsothattheycloseoff were found foraging abundantly on the thebaseoftheflower(Fig.15)presumably flowersofthespiny, metrehighherbCo- protectingthenectarfromevaporation,ju- don royeniiL. (Hydrophyllaceae)growing giirtia codoni, when visiting a flower, inthegravellybed ofa drywatercourse, alights on the outwardly curved corolla where it emerged from the hills before lobesand,whenpreparingtoimbibenee- 80 JournalofHymenopteraResearch Steud. (Amaranthaceae). Atothersitesin theParkitwasfounduncommonlyvisit- ing flowers of Zi/gopln/Uiim meyeri Sond. (Zygophyllaceae),SeuecioarenariusThunb. (Asteraceae) and Pelargonium klitighard- tenseKnuth(Geraniaceae). Duetothesmallnessinsizeof/.codoni comparedwithCodonroyeniiitispossible for/.codoniwhenobtainingnectartoenter andleavetheflowerswithoutcominginto contactwitheithertheanthersorthestig- masandtocollectpollenfromtheanthers withoutcomingintocontactwiththestig- mas.Itisthereforeprobablethat,whereas C. royenii is clearly an important forage plantfor/.codoni,/.codoniisnotapoten- tialpollinatorofC. royeniibutisrathera pollenandnectarthief.Amorelikelypo- tentialpollinatoristherelativelylargebee XylocopaliiguhrisGerstaecker(Apidae:Xy- locopinae) which, although it is a less commonvisitor and not restricted toCo- don, inbehaviourand fitisideallysuited toactasapollinator. Jugurtiak—oeroegabensisGess Distribution. Jugurtia koeroegabensis Gess has been recorded from only the Richtersveld National Park in northern Namaqualand(G—ess,F.W. 1997). Floioervisiting. Only seven femalesof /e.rsk.oeOronegatbhoeisbiasnwkesreofcoallderctyedwaftreormcofulroswe- cF13oi,glsl.Jeucg1tu3irntaginapdocl1ol4de.on.niCGodeosisiiromi/bciiibiiinLg.(neHcytdarro.ph14y,ll/.acceoadeo)u.i cf17rr.oo0sm3sEit)nheg(Fsitmghsa.ell2K-c5or)eertaowmeogffaleobmwvaelrlaseksotfweeZry(ge2o8t.p1ahkyIelSn-, luniprismatocarpuinE.Mey.exSond.(Zy- gophyllaceae) and one female was taken tar, walks down the side of the flower from a yellow rayed capitulum ofOsteo- (Fig. 13) and inserts its long tongue be- spennumsp.(Asteraceae).AtPootjiespram tweenthefilaments.Whencollectingpol- (20.OSS,16.57E),inabroadsandydrainage len,thewaspeitherreachesoverfromthe area, four females were taken visiting corolla or moves across onto a filament flowers, one from the yellow petalled and ingests the pollen directly from the flowers of Cleoine paxii (Schinz) Gilg. & anther(Fig.14).Althoughapparentlypre- Benth. (Capparaceae),onefromthe flow- ferringC. royenii,/. codoniwasfoundless ersofPeliostomum leucorrhizum (Scrophu- commonly visiting the flowers of P. leii- lariaceae) and two from the green and corrhizum (Scrophulariaceae) and of brown flowers of Ferraria cf divaricata Hermbstaedtia glauca (Wendl.) Reichb. ex Sweet (Iridaceae). These records are too Volume6,1997 81 shrubbyZygophyllumprismatocarpum (Zy- gophyllaceae) (Fig. 4), and four from the largeryellowflowersofcompact,shrubby Zygophyllum meyeri. In addition one fe- malewascollected fromtheflowersofa species of Wahlenhergia (Campanulaceae) andonefromayellowrayedcapitulumof Asteraceae. Masarin—apeliostomiGess 20mm Distribution. Masarina peliostomi has beenrecordedfromonlytheRichtersveld NationalPark(G—ess,F.W. 1997). Flowervisiting. Bothfemalesandmales ofM.peliostomiwereforagingabundantly ontheflowersofPeliostomumleucorrhizum (Scrophulariaceae)(Fig. 7)growingalong the banks of the upper reaches ofa dry Fig. 15. Coihii nn/ciiii L. (Hydrophyllaceae), Dia- watercoursecrossingtheKoeroegabvlakte gtlyerda)a.mnmdatoifclurgeiipirteisaecnotdactiinionGeofssfl(owwienrgscutanldonlgeigtsudoimniatl-- aa(Fsnisdgosc.iaa2t-si5po)en.cibTeehstiwsoefiesnPetlahiesopsfeticrosimteusmr.eocfoHMrodawseaovfreiran,na few to establish a particular preference, afisvseocsipaetceideswiotfhCeplloannittsesofartehiksngoewnnusto(sbeee nhootwerveestrr,icittedistcolveiasrittihnagtf/.lokwoeerwsegoafbaenssiinsglies asibzoevea)n.dMabseahraivniaouprelitoosttohmeiCbeelionnigteosfsspiemciileasr plantfamily. should, likethem,beconsideredapoten- MasarinaRichards tial pollinatorofPeliostomum intheRich- tersveld. Although not collected from Masari—namixtoidesGess flowers of Aptosimum spinescens, which Distribution. Masarina mixtoides Gess was growing further down the water- hasbeenrecordedfromonlytheRichters- course, it is probable that M. peliostomi veldNationalParkinnorthernNamaqua- wouldvisittheseflowers. lanFdlo(wGeerss,visFi.tWin.g.1—99M7a).sarina mixtoides is Masa—rinastruckiGess notrestrictedtoasinglefamilyofplants. Distribution. Masarinastruckihasbeen At a site 1.5km from theHelskloofGate recorded from Namaqualand (Goegap (28.18S, 16.57E),theabundantwhiteflow- NatureReserve,Springbokand thewest- ersofanisolatedplantofPelargoniumkling- ern side ofthe Kamiesberg), theOlifants liardtenseKnuth(Geraniaceae)growingon RiverValleytothesouth(eastbankofthe theedgeofadry,stoneydrainagechamiel Clanwilliam Dam), immediately east of onahillsidewereclearlyattractivetothis the Western Escarpment (Skuinshoogte wasp, a sample of 15 females and one Pass between Nieuwoudtville and Lour- malehavingbeentakeninunderanhour. iesfonteinandBiedouwValleyintheCed- However,ontheKoeroegabvlakte,where erberg) and in the western Little Karoo flowers associated with a dry, gravelly (Ladismith)(Gess—,F.W. 1997). drainage chamiel crossing the flats (Figs. Flowervisiting. Thefirstrecordwasfor 2-5) were sampled, three females were a single female collected by Michael takenfromthesmallcreamflowersoflax. Struck, 20.viii.l985, visiting the yellow 82 JournalofHymenopteraResearch flowers of Hermannia disermifolia Jacq. of andS.K. Gess)visitingyellow flowersof thesubgenusHermannia(Sterculiaceae)in HermanniavestitaThunb. ofthesubgenus Goegap NatureReserve (formerlyHester Hermanniagrowinginlevelgroundonthe Malan Nature Reserve) in theSpringbok roadsidehalfwayuparockyslope. district, Namaqualand. An intensive It seems probable thatM. strucki is al- search by the Gesses for this masarine ways associated with Hermannia spp. as duringsixsubsequentspringvisitstoNa- careful collecting by the Gesses from a maqualand yielded only one further fe- widerangeoffloweringplantsbothinthe male,fromthesamelocality(10-12.x.1988, vicinity ofits collection sites and further F.W. and S.K. Gess), until spring 1994 afieldhasbeenundertaken. Italsoseems v^'hentheyfoundfemalesandmalesrela- to be of note that the Hermannia plants tivelyabundantinassociationwithH.dis- with which it has been found have all ermifolia in the Goegap Nature Reserve beenassociatedwithrockyhillslopes. (primarily at 29.37S, 17.59E) (3-8.X.1994, The flowers of the Hermannia spp. re- F(3.0W..13aS,nd1S8..K0.3EG)essi)nantdheatKaBmakileesibkrearagl calolr"dbeedlla-sshbaepiendg"vaisnidte7d-1b0ymM.msitnrulcekingathr.e a(9n-d11.1X7)..19T94h,eFf.oWl.loawnidngS.sKp.riGnegs,s)19(9F5i,gst.he1y6 Maansaarvienraagsteruclkein,gotfhwohfic7h.4thmemfemaalnedsatrhee againfoundfemalesandmalesattheKa- males 6.6 mm, alights on theside ofthe miesberg site (28.ix.1995, F.W., S.K. and downwardly hanging flowers, moves R.W. Gess) but none were found at the down, and curves around the lip to face Goegapsite.ThesitesintheGoegapNa- upwardsintotheflower(Fig.22h).When tureReserveandtheKamiesbergarelevel remaininginthispositionanindividualis sandyareasatthebaseofrockyoutcrops, apparentlyingestingpollen.Whenthein- in which H. disermifolia is the dominant dividual enters into the flower leaving plant. only its abdomen exposed it isundoubt- IntheOlifantsRiverValleyontheeast edly imbibing nectar. The nectarheld in bank of the Clanwilliam Dam, 19.2 km the inrolled bases of the petals will be s7fEoo)ruatMghi.nogfstotrhuncekaicnawroaarsvaanfngoepuafnlrdokwbe(y3r2eR.d1o7bsSep,retc1i8eG.se56so/sf raaennadacvh4e.e0rdamgbeymlteihnnegttwhhaesofmpa'4ls.e2.tmonmguienwthheicfhemhaalse HermanniasubgenusMahernia (3 females, AsaflowervisitorM.struckiappearsto 5.X.1995,F.W.,S.K.andR.W.Gess)grow- berestrictedtoHermanniaandthereforeto ing in sandy soil at the foot of a rocky bedependantuponthisgenusforpollen slope. andnectar.Itsdistributionisfarmorelim- At the Skuinshoogte Pass site (31.16S, itedthanthatofHermanniaandwithinthe 19.08E) only two females were collected areaswhere it doesoccur it seems tobe (23-30.ix.l994,F.W.andS.K.Gess),oneon restricted to rocky slopes and on these thegroundinadryriverbedandtheoth- slopestoalimitedrangeofHermanniaspe- eronthegroundbeneathashrubletofa cies.Itis,however,notthesolevisitorto yellowfloweredspeciesofHermanniasub- these species ofHermannia (Table 1). As- genusHermannia. semblagesofvisitorsto the threespecies InthewesternLittleKarootwosamples of Hermannia subgenus Hermannia with havebeentakenintheLadismithdistrict, which it has been associatetl are charac- one at Buffelspoort (3320BD) (3 females, terizedbythepresenceof,inaddition,one 14.viii.l995, V.B. Whitehead)onayellow or more species ofAnthophora {Py^antho- flowered shrubby Her)nannia sp. and the phora) (Apidae: Apinae: Anthophorini) other6kmfromLadismithontheroadto and—one or more species of Megachili- Barrydale (3 females, 21.viii.l995, F.W. dae oneormorespeciesofPlesianthidium Volume6,1997 83 Figs. 16-18. lb.NestingareaotMasaiinastnickiGess, Kamiesbcrg,bushesmtheforeground,Hcrnmiuim disermifolinJacq.(Sterculiaceae).17,FlowersofH.liiscniiifclin(lengthofflowerapproximately9mm).18,Cell ofM.ifruckiopenedtoshowpapillateprovisionmassandpositionofegg. (Spinaiithidium) (Megachilidae: Megachili- hairy faces. Such pollen would be trans- nae: Anthidiini) and/or a species of La- ferred to receptive stigmasby the wasps sioglossum(Halictidae).Thoughthesebees andthebeesvisitingflowersintherecep- arethereforealsoexpectedvisitorstothe tivephase.Allcanthereforebeconsidered flowerstheyarelessconstantastheyare tobepotentialpollinators,butwherethey allknown tovisitinadditionflowerspe- arepresentthemasarineswillbethemore cies of other genera belonging to other dependable. plantfamiliesoccurringincloseproximity The species of Hermannia subgenus totheHermanniaplants. Makernia was sampled on one day only. Masarinastriickiand thebeesall visita Onthisdayitwasonlyvisitedbytwoma- considerablenumberofflowersinsucces- sarines, M. strucki and Masarina mixta sion,and allwhilevisitingflowersinthe Richards.Bothcanbeconsideredpotential pollen-presenting stage receivea dusting pollinators, however, M. strucki is more ofpollenwhichadherestothem.Masarina dependablethanM. mixtawhichvisitsin struckiinenteringtheflowersreceivesthe additionflowersofplantsofseveralother pollendorsallyandthebeeswhichareall familieswhichgrowincloseproximityto too large to enter receivepollenon their theHermantimplants. 84 JournalofHymenopteraResearch Table 1. FlowervisitorstothosespeciesofHer- Table1. Continued. nujuiiiaL.onwhichMnsnrimistnickiGessisknown toforage.Numberinbracketsindicatesthenumber Apidae:Apinae:Anthophorini ofplantfamiliesinadditiontoSterculiaceaewhich Ainlit>ph(}rciiPygiinllutphiirti)kiugeriEardley,Ladi- areknowntobevisited. smith(1) Nemestrinidae sHiurbmgaenniuUsLiHcntiluiisieirimiiufioliaJacq.andHcnucinniuct.(/;.v- Prosoecasp. ennifolia subgenusMahernia Vespidae:Masarinae Hcrmanniasp, MiisariiwsiruckiGess.Goegap,Kamiesberg.(Skuins- Vespidae:Masarinae hoogte)(0) MasariunmixtaRichards,Clanwilliam(3) Vespidae:Eumeninacapparentlyhunting MflSfln'm!struckiGess,Clanwilliam(0) Megachilidae:Megachilinae:Anthidiini Vespidae:Eumeninaeapparentlyhunting Plesiiinlhidium{Spiiuiiuhidiiim)callcscens(Cocker- ell).Skuinshoogle(1) Plesianthidium{.Spinanlhiiliiint)iniclnisijorme — (Friese),Goegap(1) Nesting area. Unfortunately despite Apidae:Apinae:Anthophorini carefulsearchonlyonenesthasbeenlo- Antluiphorci[Pygmilhophnni)ubrochiuEardlcy.Goe- cated. This was at the Bakleikraal site in Aiiglahpo,plKuiarmaieisPbyegirignt(h1o)phoni)di\'cisipfsFriese,Ka- tthreancKeamwiaessbbeertgwee(Fnig.two16)s.toTnhees innesgtentelny- Antgmhaioepps(bh1eor)ngi((3.)Pygiinthophoni)krngeriEardley.Goe- Hsleorpmianngnigarobuunshdesin.aThbearseoialrewaasbestawnedeynatnhde Anlhophoni{Pygantliophont)scliiill:i'iFrcise.Skuins- friable. — hoogle(2) Provision. The provisionfrom thesin- APmaec(bg3eh)iryglm.lealSukiusMiicpnnesirhiioinoigegigilieel\li(u1)(6F)nriiveesael)aGo(Fergiaespe.),KKaammiieessb-erg Tglilhkeee,clelpolanlgoeibttuyadeiilnnleaodlwswuoarcsfharvceee,rwyaanmsdoirsetgtruablnausrltuljcyeelpnlaty-.- Apidae:Apinae:Apini pillate, each papilla ending in a nipple ApismellifeniL.(verymany) (Figs.18and19b).Attheinnerendofthe Scarabaeidae:Mclalonthinae:Hopliini provision was a larger central papilla Asinglespecies which supported theegg. The outerend Hcrniniiiiinct.cuncifolinJacq. Vespidae:Masarinae MasarinastnickiGess,Goegap,(Skuinshoogle)(0) Vespidae:Eumeninaeapparentlyhunting Halictidae Lasioglossumsp.,Skuinshoogte(?) Megachilidae:Megachilinae:Anthidiini Plcsiiiiitliidiiiiii{Spiiiiiiilludiiiiii)ciilli'scciif(Cockerel!), Skuinshoogte(I) Apidae:Apinae:Anthophorini Anthophora{P\/ganthoplwrn)ichuUzctFreise,Skuin- shoogte(2) HcnihviiiioI'cstitaThunb. Vespidae:Masarinae MasarinastruckiGess,Ladismith(0) Halictidae Fig.19. Diagrammaticrepresentationsoflongitudi- Lasioglcssiinisp.,Ladismith(?) ananldseegctgioonfoMfasnaersitnaansdtruceclkli,Gpeaspsi.llateprovisionmass