A note on facultative kleptoparasitism in Prionyx kirbii (Hymenoptera: Sphecidae) as a consequence of multi-specific shared nesting site, with description of its prepupa

2006·0.36 MB·

by  Carlo Polidori

©EntomologicaFennica.8December2006
Anote on facultative kleptoparasitism in Prionyx kirbii
(Hymenoptera: Sphecidae) as a consequence of multi-specific
shared nesting site, with description of its prepupa
CarloPolidori,JoséTormos,JosepD.Asís,PauMendiola&FrancescoAndrietti
Polidori,C.,Tormos,J.,Asís,J.D.,Mendiola,P.&Andrietti,F.2006:Anoteon
facultative kleptoparasitism in Prionyx kirbii (Hymenoptera: Sphecidae) as a
consequenceofmulti-specificsharednestingsite,withdescriptionofitsprepupa.
—Entomol.Fennica17:405–413.
ObservationsofafemaleofPrionyxkirbiiatanaggregationofStizuscontinuus
revealedanalternativenestingbehaviourofthefirstwaspduetothepresenceof
thesecond one. Theoverlap ofsomeresources (prey, kind ofsoil) allowed P.
kirbiitoactasakleptoparasiteofS.continuus.Theobservedfemaleoftenre-used
asnestspre-existingS.continuusemergenceholes,butnoS.continuusnests.The
prey wereobtainedsometimesby stealingthegrasshoppersfromS.continuus
nestsand/orbyattackingthefemaleswhilecarryingtheprey.Fromareviewof
interspecific kleptoparasitism among sphecoid wasps it appears that family
Sphecidaeincludesthehighestnumberoffacultativeparasitismcasesandthis
finding is discussed. In addition, the description of the prepupa of P. kirbii is
givenandisshowntobeinagreementwiththedescriptionofthematurelarvaeof
theotherspeciesofthegenus.
C.Polidori,DipartimentodiBiologia,SezionediZoologiaeCitologia,Univer-
sità degli Studi di Milano – Via Celoria, 26, 20133, Milan, Italy. Tel.
[phone];E-mail:[email protected]
J. Tormos, Unidad de Zoología, Facultad de Biología, Universidad de Sala-
manca–37071,Salamanca,Spain
J. D. Asís, Unidad de Zoología, Facultad de Biología, Universidad de Sala-
manca–37071,Salamanca,Spain
P.Mendiola,DepartamentdeZoologia,UniversitatdeValència–c/Dr.Moliner,
50,46100,Burjassot(València),Spain
F. Andrietti, Dipartimento di Biologia, Sezione di Zoologia e Citologia,
UniversitàdegliStudidiMilano–ViaCeloria,26,20133,Milan,Italy
Received3July2004,accepted13December2005
1.Introduction pactness,soilmoistureandamountofvegetation
amongothers),aresimilarinmanycasesfordif-
Sphecoidwasps(SphecidaeandCrabronidae)of- ferentspecies,nestingsitesareoftensharedbya
tennestinlargeaggregations(e.g.Evans1966& number of species. When this occurs, inter-
1974,Evans&Hook1986,Rosenheim1990).As specific interactions may become frequent and
thephysicalnestingrequirements(likesoilcom- lead to nest usurpation (Matthews 1983, Hook
406 Polidorietal. (cid:127) ENTOMOL.FENNICAVol.17
1987),kleptoparasitism(Kurczewski&Spofford soryburrows(falsenests2–3cmlong,seeEvans
1998,Casiraghietal.2003)or,morerarely,nest 1966)aredugclosetothenestentrance(Asìset
sharing(McCorquodale&Thomson1988).Fac- al.1988).
ultative kleptoparasitism and brood parasitism Our aims are twofold. Firstly, we report ob-
(i.e.usebyawaspofanotherwasp’spreytofeed servationsonanunusualstrategy adoptedby P.
herownbrood)couldhavebeenanintermediate kirbii to obtain prey and nest; secondly, we de-
step in the evolution of the obligate klepto- scribethematurelarvaofP.kirbii,inordertopro-
parasitismandbroodparasitismtypicalofgenera videtraitstoseparatethisspeciesfromothersof
such as Nysson and Stizoides (Evans 1966, thegenus.
Wcislo1987).
In this paper we report a few observations
takenonasinglefemaleofPrionyxkirbii(Vander 2.Materialandmethods
Linden) (Hymenoptera: Sphecidae), a common
speciesoftheoldworld,nestinginthesamearea ObservationsweremadeonasinglefemaleofP.
where a large aggregation of Stizus continuus kirbiibetween24.VI.2003and10.VII.2003,the
Klug(Hymenoptera:Crabronidae)hadbeenes- datewhenthewaspwascollectedfordetermina-
tablished for many years (Asìs et al. 1988 & tion.P.kirbiiwasobservedonanaggregationof
2006).Thebehaviouralconsequencesoftheco- thesolitary waspStizuscontinuus,inanareaof
existenceofthetwowaspsareanalyzed. salt-marsh(ca.9m×15m),characterizedby a
Prionyxfemalesusuallyestablishtheirnests bare,compactsoil,covered(30%ofthesurface
onbare,compactsoil.Theunicellularor,infew approximately) by Arthrocnemum fruticosum
cases, multicellular nests are filled with Ortho- (L.)andJuncusmaritimusLam.andboundedby
pthera Acrididae to feed the larvae. One grass- Pinus pinaster Aiton and Phragmites australis
hopperisusuallyprovidedtoeachlarva(Evans (Cav.). The female was marked with non-toxic
1958, Benz 1959, Bohart & Menke 1976). De- paintstopermitidentificationonfield.Nestsdug
pendingonthespecies,nestsareconstructedbe- andprovisionedbythewaspweremarkedwitha
foreorafterhuntingtheprey.Somespeciesclose coloured stick placed in theground closeto the
thenesttemporarilybeforehunting(Evans1958, entrances.
Genise1980,Grissell1981).P.kirbiibelongsto ObservationsonP.kirbiiweremadeonadis-
thegroupofspeciesthatdigthenestbeforehunt- continuousbase,sincetheywereintermixedwith
ing and make a temporary closure before the alongtermplannedresearchonS.continuus.
provisioning trip (Ferton 1902, Fabre, 1914, p. In the following text, we will use the term
202,Benz1959,Bohart&Menke1976).P.kirbii “kleptoparasitism”toindicatewhenawaspstole
females usually hunt one large prey, sometimes thehostprovisionsfrominsidethenestandthen
two,iftheyaretoosmall(Benz1959).Thereis carriedthemtoherownnestorwhenshestolethe
only one report of Ferton (1902) of a nest with hostprovisionsattackingdirectlythehostoutside
twocells,onewithtwo-prey,theotherwithonly thenest;wewillusetheterm“broodparasitism”
one,buthisfindingisdubious.Thenestsstruc- ifthewaspenteredthehostnestandovipositson
tureconsistsofaverticalburrowfrom3to4cm the host provisions, eventually eating the host
deepfollowedbyanhorizontalsectionofabout4 egg.
cm(Benz1959). The postdefecated larva (prepupa) was ob-
S.continuusfemalesdig(multicellular)nests tainedatSaler,Valencia(6.IX.1992).Themethod
before hunting on bare and compact soil, pro- employedtopreparethespecimen,aswellasthe
visioning them massively with grasshoppers terminology of larval morphology and format
(several for each cell). The nest remains closed usedinthedescriptionfollowfundamentallyEv-
duringhuntingflights(Asìsetal.1988).Thenest ans (1987). The following abbreviations have
of S. continuus is composed of a main burrow been used in the description: d = diameter; h =
(oblique to the soil surface) about 50 cm long height;l=lengthandw=width.Voucherspeci-
fromwhich3–8secondarytunnelsdepart,eachof mensaredepositedatthe“Torres-Sala”Entomo-
themendinginabroodcell.Generally1–2acces- logicalFoundation(Valencia,Spain).
ENTOMOL.FENNICAVol.17 (cid:127) KleptoparasiticstrategyofPrionyxkirbii 407
Table1.NumbersofobservationsmadeonthefemaleofPrionyxkirbiibetween24.VI.and10.VII.2003.
Date Patrollingthearea: Patrollingthearea: Attacking Carrying Closing Stealinga
stoppingonStizus “touching”S.con- S.continuus apreyto anest preyfrom
continuusnests tinuusfemales femaleson anest definitively S.continuus
ontheirnests theirnests females
24.VI. 3 1
25.VI. 4 1 1
26.VI. 2 1 1** 1 1**
27.VI. 1***
28.VI. 4 1 1
29.VI. 2 1****
30.VI. 2 1*** 1*
01.VII. 1
02.VII. 1 1*****
03.VII. 4 1 1
04.VII. 3 2
05.VII. 5
06.VII. 2
07.VII. 2
08.VII. 3 1
09.VII. 4
10.VII. 1
*=preycarrieddeepinagrassarea;nestnotidentified;**=preystolenfromthewaspafteranattack;***=theS.continuusfemaleflownawayafter
the“touch”;****=thisnestdidnotcorrespondtoanemergingholeofS.continuusastheotherones;*****=preystolenfrominsidethenest.
3.Results ofP.kirbiitooktheprey,stungitandtransported
it away, stopping sometimes to sting again the
3.1.BehaviourofP.kirbii grasshopper.TheothertwotimesthefemaleofS.
continuusdidnotabandontheprey,andP.kirbii
Table 1 summarizes the results of the observa- flewaway.Onanotheroccasion,P.kirbiiwaited
tions.ThefemaleofP.kirbiiwasobservedforthe on a closed nest the return of the owner with a
firsttimeonthe24.VI.2003,whilepatrollingthe preyandfollowedherinsidethenest,comingout
nesting site of S. continuus, and she frequented afewsecondslaterwiththegrasshopper.Evenin
theareaeverydayoftheobservationperiod.The this case the wasp re-stung the prey during the
patrollingbehaviourincludedalwaysanumberof transport. Neither females nor males of S.
stops on some nests of S. continuus (open or continuusshowedany aggressivenesstowardP.
closed). The stops lasted 5–30 seconds and, on kirbii.
average, the wasp stopped on a nest 0.8 times/ Atotalof5nestsofP.kirbiiwerelocatedin
minute(range:0–5min–1,SD=1.31,n=19ob- the nesting area of S. continuus, and 4 of them
servationsthatlastedmorethan1minute). were identified as re-used emerging holes of S.
Onsomeoccasions(6outof42contactswith continuus, but no nest of S. continuus was ob-
thewasp)thePrionixfemale,whileoverflyinga servedtobeactivelyusurpedbyP.kirbii.Atotal
S.continuusnestwhentheownerwaspresentat of 4 prey-carryings by P. kirbii were observed,
the entrance, flew rapidly down and “touched” but no more than one to each nest. Afifth prey
theotherwasp,disturbingheractivity(in2cases wascarriedinaperipheralareawithdensevege-
outof6,thefemaleofS.continuusflewawayfor tation,butthewaspdisappearedinthegrassand
awhileafterthe“touch”).Threetimesthefemale thenestwasnotfound.Consideringthattheob-
ofP.kirbiiattackedmoreviolentlyaS.continuus served 5 nests were closed definitively by the
femaleasshewasopeninghernesttoenterwitha wasp within one-two days since they were first
prey.Onetimeshecausedtheabandonmentofthe recorded,andthatthefemalewasobservedfora
preybytheattackedwasp:atthispointthefemale period of 17 days, probably many other unde-
408 Polidorietal. (cid:127) ENTOMOL.FENNICAVol.17
Fig.1.PrepupaofPrionyxkirbii(VanderLinden).–a.Generalaspect(lateralview).–b.Portionofintegument.
–c–d.Abdominalspiracle.–e.Cranium(frontalview).–f.Antennalorbit(frontalview).–g.Labrum.–h.Epi-
pharynx.–i.Leftmandible.–j.MaxillaandLabium.
tected nests belonging to the same individual ments; atria (d = ma) goblet-shaped; walls lined
werebeenpresentinthearea. withridgesformingsmallandirregularpolygons;
opening into subatria armed with long spines;
subatriacomparativelysmall(d=64mm),with6
3.2.DescriptionoftheprepupaofP.kirbii swellings. Cranium (Fig. 1e) (w = 0.9 mm, h
[heightmeasuredtoapexofclypeus]=1.1mm)
General aspect (Fig. 1a). Body (l = 13.5 mm, higherthanwide,yellowish,withsparsesetae(l=
maximumw[3rdabdominalsegment]=4.3mm) 5mm)andsomepunctationsclosetomandibular
hymenopteriform, elongate, strongly curved at insertion;vertexdepressedmedially;coronalsu-
thelevel ofthethird thoracicsegment, with the tureandparietalbandspresent,brown;antennal
thoraxandfirstabdominalsegmentnarrowwith orbits(Fig.1f)(58×43mm)withmembraneofthe
respecttotherest,yellowish,weaklysclerotized, orbitconvex,with3sensilla.Fronswithfivede-
exceptforspiracles.Anusterminal,atransverse pressions, near epistomal suture and base of
slit. Pleural lobes convex. Integument (Fig. 1b) clypeustransverselyfurrowed;frontoclypealsu-
with large spinules (l of spinules = 30 mm, w ture unpigmented; clypeolabral suture well de-
basal=10mm)arisingfromplate-likebases,with fined.Clypeuswith12setae(l=5mm).Labrum
theexceptionofthoseofthethoracicsegments(l (Fig.1g)(maximumw=470mm;maximumh=
=5mm)whicharesmallandarrangedinsucha 150mm)bilobate;basesmooth;anteriormargin
waythattheyhavetheappearanceoftransverse weaklypigmentedwithsmallsensilla;eachlobe
grooves,andwithsparsesetae(l=3–7mm).Spi- surface with sparse setae (l = 5mm) and dense
racles (Fig. 1c–d) all in a single line, on meso- conicalsensilla(w=3mm);epipharynx(Fig.1h)
thorax, metathorax, and first 8 abdominal seg- denselyspinulose(lofspinules=25mm,basalw
ENTOMOL.FENNICAVol.17 (cid:127) KleptoparasiticstrategyofPrionyxkirbii 409
= 3 mm) centrally and laterally, with 8 sensory nesting area of several provisioning females of
pores (d = 7 mm) medially distributed en 2 epi- S.continuus(ca.30duringtheperiod).Theover-
pharyngealsensoryareas,eachwith4pores. lapoftheresourcesneededbythetwospecies,in
Mouthparts.Mandibles(Fig.1i)(l=370mm, particular bare soil and grasshoppers (Evans
maximum w = 170 mm) robust, sclerotized, 1958, Benz 1959, Asìs et al. 1988), would ac-
brown,teeth3and4darkbrown,2.2×aslongas count for the observed phenomenona. Notwith-
wide,withtwopuncturesclosetothebasalexter- standingthelackofacontinuousrecordingofthe
nalmargin,with4apicalteeth,3superiorteethon activityofP.kirbii,wehavestatedthat,atleastin
aboutthesameplane,thebasaltoothonalower part, the wasp used two resources obtained by
plane.Maxillae(Fig.1j)withsparselateralsetae S.continuusfemales:emergenceholes(asnests)
(l=11mm),mesallydenselyspinulose,withvery andprey(asfoodforthelarvae).Byaslightshift
shortspinules;maxillarypalpilongerthanwide inherbehaviour,thefemaleofP.kirbiimaybe-
(75×49mm)with5apicalsensilla;galeae(l=75 comeatruekleptoparasiteofS.continuus,atleast
mm,w=30mm)similarlyformed,with2large for what prey is concerned (strictly speaking,
apical sensilla. Labium(Fig. 1j) setose (l of the emergenceholesre-useisnotaparasiticaction).
setae=12mm);labialpalpilongerthanwide(75 Thisstrategyshouldallowreducedtheexpendi-
×49),with4apicalsensilla(d=7.5mm);salivary tureofenergy indigginganest,butprobably it
orifice(l=450mm)transverse. does not reduce the time necessary to find the
prey.Infact,weobservednomorethanonenest
provisionedandcompletedperday,thecommon-
4.Discussion est value found in many other Prionyx species
studied (Evans 1958, Genise 1980, Grissell
ThesearethefirstobservationsmadeonP.kirbii 1981).
(aswellasonthegenusPrionyx)behavingasa Since re-using a pre-existing burrow (the
kleptoparasiteofadifferentspeciesofsphecoid emergencetunnelsofS.continuus)evidentlyre-
wasp,andprovidenewdataontheseinterspecific ducesthetimespenttoobtainanest,prey-finding
relationships phenomena, which are probably shouldbetheactivitythatmainlydeterminesthe
more widespread than previously supposed timenecessarytocompleteanest.Infact,attack-
amongfossorialHymenoptera.Theobservations ing a S. continuus female to steal her prey is
arebasedonasinglefemale.However,thebehav- greatly more dangerous and time-consuming
iourisprobablyrare:innoneofthe14published than re-using an emergence hole, which is no
works on the nesting habits of Prionyx spp. moreusedbyanyS.continuusfemalesandonly
kleptoparasiticbehaviourwasdetected(mostim- attracts the attention of the males (Asìs et al.
portantreferencesquotedinthetext;thecomplete 2006). Moreover, although territorial S. conti-
listofworksonPrionyxspp. couldbefoundin nuusmalesattackconspecificmalesandmalesof
Pulawski (1999)). For the much more investi- themutillidparasiteNemkaviduata(Pallas),they
gated genus Ammophila, only in 4 out of more wereneverobservedtointerferewiththeactivity
than50studiessuchbehaviourwasrecorded(Ta- ofP.kirbii.Inconclusion,wemaysupposethat
ble 2 and Pulawski 1999). This means that this thefemalesofP.kirbiiprefertogaintimeanden-
behaviourisdifficulttoobserverepeatedly.Ac- ergy colonizing an emerging tunnel of S. conti-
cordingly,weconsiderthisphenomenonworthof nuus;subsequently,theyfirsttrytofeedtheirlar-
reportinginspiteoftherestricteddata. vae at the expense of S. continuus (prey). Any-
way,usuallyafterhavinglosttimebypatrolling
theS.continuusnestingareainsearchofapreyin
4.1.ThebehaviourofP.kirbii an “easier” way, they resume their “normal”
onthenestingsiteofS.continuus huntingactivity.Re-usingS.continuusnestsdoes
notseemtoberelatedtosimilarityinnestarchi-
Thenestingbehaviourexhibitedbytheobserved tectureofthetwospecies.InfactP.kirbiifemales
female of P. kirbii showed characteristics that havebeenobservedtodigtheirnestseveninside
seem to be related to the presence in the same already pre-existing nests of bees of the genus
410 Polidorietal. (cid:127) ENTOMOL.FENNICAVol.17
Table2.Observedfacultativeinterspecificparasitismamongdifferentspeciesofsphecoidwasps(Sphecidae
andCrabronidae).Thekindofbehaviourofpreystealingisreportedonlyforfacultativekleptoparasitism.
Facultativekleptoparasitism Preyof Kindof References
thehost parasitic
Parasite Host behaviour
Ammophila Ammophilaurnaria Lepidoptera Theftof Kurczewskietal.
kennedyi*(Murray) Dahlbom[S] larvae provisions 1992
[S] frominside
nests
Ammophilaurnaria* Ammophilakennedyi Lepidoptera Theftof Kurczewskietal.
[S] [S],Podalonia larvae, provisions 1992;
robusta(Cresson)[S] Lepidoptera frominside Kurczewski&
larvae nests Spofford1998
Microbembex Tachysphex Orthoptera, Preytheft Kurczewski,
monodonta**(Say) terminatus(F.Smith) Diptera, outsidethenest personal
[B] [C],Lyrodasubita Diptera, communication,
Say[C],Bembix Lepidoptera quotedbyEvans
pruinosaFox[B], larvae 1966,p.378;
Ammophilaharti Evans1966,p.
(Fernald)[S] 376,378
Passaloecuscorniger Passaloecus Homoptera Theftof Danks1970
Shuckard[P] singularisDahlbom provisions
[P] frominside
nests
Passaloecuscorniger Passaloecusgracilis Homoptera Theftof Corbet&
[P] (Curtis)[P] provisions Backhouse1975
frominside
nests
Passaloecuscorniger Passaloecusinsignis Homoptera Theftof Corbet&
[P] (VanderLinden)[P] provisions Backhouse1975
frominside
nests
Passaloecuscorniger Psenulusatratus(F.) Homoptera Theftof Chevalier1923,
[P] [P] provisions quotedbyEvans
frominside &West-Eberhard
nests 1970,p.212
Podaloniaaffinis* Ammophilasabulosa Lepidoptera Theftof Field1989,1992;
(Kirby)[S] L.[S] larvae provisions Casiraghietal.
frominside 2003
nests
Ammophilakennedyi Podaloniarobusta Lepidoptera Theftof Kurczewskietal.
[S] [S] larvae provisions 1992
frominside
nests
Prionyxkirbii Stizuscontinuus Orthoptera Theftof Presentstudy
VanderLinden[S] (Klug)[B] provisions
frominside
nests;Prey
theftoutside
thenest
Stictiasignata(L.) Stictiaheros Diptera Preytheft Sheehan1984
[B] (Fabricius)[B] outsidethenest
ENTOMOL.FENNICAVol.17 (cid:127) KleptoparasiticstrategyofPrionyxkirbii 411
Table2continued.
Facultativebroodparasitism Preyof References
thehost
Parasite Host
Ammophila Ammophilaurnaria Lepidoptera Kurczewskietal.
kennedyi*[S] [S] larvae 1992
Ammophilaurnaria* Ammophilakennedyi Lepidoptera Kurczewskietal.
[S] [S] larvae 1992
Podaloniaaffinis* Ammophilasabulosa Lepidoptera Field1989,1992
[S] [S] larvae
Lirisniger Sphexmaxillosus Orthoptera Fabre1914,p.102
(Fabricius)[C] (Fabricius)[S]
S=Sphecidae;Crabronidae:B=Bembicinae,P=Pemphredoninae,C=Crabroninae;*speciesthatappearbothasfacultativekleptoparasitesand
broodparasites;**thisspeciesisastrictscavenger,accordingtoEvans1966,p.376.
Halictus (Benz 1959). However, burrowing a iour.Thisseemstobetrueinparticularfornest
new nest is very time-consuming activity: 22 usurpation(withoututilizationofanyotherhost
minutes/cm, according to Benz (1959). Even if resources).However,intherestrictedfieldoffac-
some amount of labour should be necessary to ultativekleptoparasitism(andbroodparasitism),
adaptaforeignnest,itcouldbeworthdoingit. sharingthesamepreyisnotasufficientcondition
andparasitesandhostshavetoberelatedspecies.
In fact, there exists a considerable overlapping
4.2.Interspecificfacultativeparasitism between nesting habits and phylogeny in sphe-
insphecoidwasps:distribution coidwasps:allthemembersofthesamegenusof-
andoccurrence tenusesimilarresources(Evans1966,Bohart&
Menke1976).Thingsseemtoworkdifferentlyin
Despite of the many studies devoted to thecaseofobligatebroodparasitism,forwhich
intraspecific parasitism among solitary wasps onlytherecognitionofthehostnestisrequired.
(see Field 1992, for a review on its occurrence Parasitismwillbeperformedlater,intheabsence
andperformanceinsolitaryHymenoptera),very ofthehost,whenitsnestwillbereopenedandits
little information exists in literature about eggdestroyed(e.g.Stizoidesspp.,Nyssonspp.).
interspecificrelationships(seeField1992,fora Onthecontrary,facultativebroodparasitismand
briefreview).Table2presentsasummaryofthe facultative kleptoparasitism have probably fol-
actualknowledgeontheoccurrenceofthisphe- lowed apathway different fromthat ofobligate
nomenon,restrictedtothesphecidwasps(sensu brood parasitism. Supporting this view, faculta-
Bohart & Menke 1976; now the family is split tivekleptoparasitismhasbeenobservedmorefre-
into Crabronidae, Sphecidae and Ampulicidae) quently in behaviourally related genera like
andconsideringonlyfacultativeparasitism.Ac- Ammophila and Podalonia, whose members
cordingtoField(1992),onemayclassifythedif- sharesimilarresources(Table2).Inassociations
ferentpatternsofinterspecifickleptoparasiticbe- betweenrelatedspecies,parasiticfemalesknow
haviour as in the intraspecific case (see Table 2 thecharacteristicsofthehost’snestandmayrec-
andField1992).Infact,thetacticdoesnotchange ognizeitmoreeasily.
ifthehostisthesameorarelatedspeciesshowing Asecondpointconcernstheoverlapoffood
similarnestinghabits.Field(1992)notesthatfac- resources. For example, species of the genus
ultativeparasitismmayoccurevenbetweendis- Cerceris often nest in sympatry (Evans 1971,
tant species (up to bees vs. wasps), because the Hook1987,Evans&Hook1986),butnorecords
parasiteonlyneedsaresourcesimilartothatem- exist of interspecific kleptoparasitism or brood
ployed in its normal and non-parasitical behav- parasitism, even if interspecific nest usurpation
412 Polidorietal. (cid:127) ENTOMOL.FENNICAVol.17
was observed in some cases (Evans & Hook Acknowledgements.ThanksareduetothestaffofOficina
1986, Hook 1987). This is probably due to the Técnica, Viveros Municipales, Ayuntamiento de Valencia,
forthefieldworkpermit,toJesúsSelfa(UniversitatdeValen-
largevarietyofpreyhuntedbythedifferentspe-
cia)forthekindattentiontothestudyandtoS.F.Gayubo
cies, which rarely overlap. On the other hand,
(UniversityofSalamanca)fortheidentificationofP.kirbii.
genera that greatly overlap their prey spectrum ThanksareduetoHenryDisney(UniversityofCambridge)
andnestingareas,asOxybelusandBembix(Bo- forreadingandcorrectionofthetext.CPandFAwereinpart
hart&Menke1976),donotshowanykindofre- supported by a three-years grant FIRB (Fondo per gli
Investimenti per la Ricerca di Base) RBAU019H94-001
ciprocal parasitism, probably because of the
(2001)and(CP)byaone-yearfellowshipfromtheUniversity
muchdifferenthabitsperformedbythesegenera,
ofMilan.JTandJDAweresupportedbyagrantfromthe
which could prevent any efficient kleptopara- Spanish Ministerio de Ciencia y Tecnología (BOS 2001-
sitism. 0864).
Unfortunately,dataaretooscarcetosubstan-
tiateanyhypothesisabouttheevolutionofpara-
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