Table Of ContentJ.HYM.RES.
Vol.5,1996,pp.240-248
Nesting Behaviorand Nest Distributions ofAmmophilagracilis
Lepeletier (Hymenoptera: Sphecidae) in Brazil
S.D.GaimariandR.P.Martins
(SDG)DepartmentofEntomology,320MorrillHall,505SouthGoodwinAvenue,Universityof
Illinois,Urbana,Illinois61801,U.S.A.;(RPM)Laborat6riodeEcologiaeComportamentode
Insetos,DepartamentodeBiologiaGeral,CaixaPostal486,ICB/UniversidadeFederalde
MinasGerais,30161-970BeloHorizonte,MinasGerais,Brazil.
—
Abstract. Ammophilagracilisisamassprovisioner,supplyinganeggwith1-2geometridcat-
erpillarsover1-2daysbeforefinalnestclosure.Nestingofmarkedwaspswasobservedattwo
sitesinBeloHorizonte,MinasGerais,Brazil.Nestsatthemorehomogeneoussite(n=54),anopen
dirtroad,hadaclumpeddistribution,comparedtothoseattheothersite(n=30)whichconsisted
ofaseriesofsmallpatchyclearings.Adultwaspslivedupto84days.Developmentaveraged
56±10days.Sevennestsweredestroyedbymiltogrammineflies{Metopian.sp.nr.sinipmlpis).Ant
predationwassuspectedasthemajorcauseofmortalityfor59neststhatdidnotyieldadult
waspsorparasites.Adistinctive"crouching"behaviordisplayedbynestingfemaleswhenmil-
togramminefliesweredetectedisdescribedforthefirsttime.
INTRODUCTION MembersofAmmophila display a wide
range of nesting tactics (discussed by
SphecidsinthegenusAmmophilaKirby Evans1959;Powell1964;BohartandMen-
eoapsMrrripepeeosinynla,k(lleleaEaervgscsaprh1neao9scnnu7ie6nda1s)dl9t.-sl5ny9wye;HhmnsoepPtawrhoikeeywnevetgdaelawrlnsla,ie1snlp9gapi6llrsd4aevo;ratvplehBata,aloretvhtraoaowcrueapdtseprevtovaciuveanlriltdse--- eppktirenreerporwi1vilhe9iol7igsda6igcr;osohnfePiatcarn1hlreg-eok,2seplirauidosnntate;ytwpsian;rhal.teinodyc1de9hali8pt1asor)eyi,nomengedgirilnseetpcsoprlsnrouieavodvsivitefsiniedgoopwevnrmdeioacrnvaasgitfsa,---
Chaavmeerboenen(Erveacnosrd1e9d65)a.sTphreeycofmorplAe.xiatzytecoaf nseisotnsintgo,pirnowvihdiechfotohdeythcroonutgihnumeutcohroefoplaern-
nestingbehaviorsisnoteworthyinthisge- val life. The progressive-provisioning
nus(EvansandWest-Eberhard1970;Tin- members,suchasA. harti(Femald),may
bergen1974;Field1989),andisamongthe maintain several nests indifferentdevel-
mostdiverseintheSphecidae.Theimpor- opmental stages at one time (Baerends
tanceofethologicalstudiestothesystem- 1941; Evans 1965; Tsuneki 1968; Hager
atics of the group was demonstrated by and Kurczewski 1986). Also, as Weaving
Baerends (1941) and Adriaanse (1947), (1989)pointedout,manymassprovision-
whodiscussedtheinter-andintra-specific ing species can be facultatively delayed
nesting behavior, and Rosenheim (1987), provisioners due to inclement weather.
who also discussed the importance of Krombein (1984) discusseci the general
prey-nest sequences, though Weaving provisioningtacticsforseveralspecies,in-
(1989)stressedthatprey-nestsequencesin cludingA. laevigataSmith (a mass provi-
Ammophila do not always reflect thesys- sionerofseveralpreyitemspernest)and
tematic relationships supported by mor- A.atripesSmith(amassprovisionerofone
phology. large prey item per nest). Ammophila are
Volume5,19% 241
alsonotedfortooluse,especiallyforusing uppersurfaceofthethoraxwereexposed
apebble in the mandibles to push down (seeFig. 9ofMartinsand Pimenta 1993).
and pack soil into their completed nests Head and thorax widthsweremeasured,
(PeckhamandPeckham1898;Evans1959; andeachwaspwasmarkedonthemeso-
Powell 1964; Tsuneki 1968). A classicac- thoraxwiththreedotsofacrylic,fast-dry-
countofnestbuildingandprovisioningin ingpaint,inauniquecolorcombination.
Aniuipphilaandotherwasps,includingnu- It was carefully noted where each was
merous outstanding, informative photo- originally captured, and any subsequent
graphs, is provided by Olberg (1959). sightings were noted astoexactlocation
Many of the behaviors discussed in the anddate.Fifty-fourwaspswerecaptured
presentworkarealsoillustratedwithpho- andmarked.
tographs for other Ammophila species in Wecarefullyrecordedfemaleactivities,
Olberg(1959). including searching and nest building,
Weobservedthenestingbehavioroffe- preyhandlingandnestprovisioning,nest
maleA.gracilisLepeletier,gatheringinfor- plugging and camouflaging, breaks for
mation about their general habits of takingnectarfromflowers,andanyinter-
searching and nest construction, provi- actionswithotherinsects.Inaddition,we
sioningandnestclosure,interactionswith marked 84nests(54atPrefeitura,and30
otherinsects,andnestdistributionsattwo atEstaqaoEcologica) toidentifytheindi-
sitesonthePampullacampusoftheUniv- vidualwaspandthedateofhernestcom-
ersidade Federal de Minas Gerais pletion.
(UFMG), Belo Horizonte, Minas Gerais, Uponseeingawaspwithprey,wefol-
Brazil. lowed to her nest. After the nestwas fi-
MATERIALSANDMETHODS nallyprovisionedandplugged,weplaced
aglasschipovertheentrance(afterWeav-
feiTthurea,f"irswtassitea, w15h0icmh wleongwilalndcal6l-1"0Prem- minogve1d989f)o.roAvfetrerawteheek,chiinpdicraetmianigntehdatusnh-e
wide, homogeneous, compact sand and probablyhasnotreturnedandreexcavat-
dirtroad(Fig. lA)withina3hectareplot ed thenest,wesecured a markedplastic
oflandcontainingvegetationinsecondary cup over the nest to capture whatever
succession.Thesecondstudyarea,which emerged.
we will call "Estaqao Ecologica," was a During ongoing studies of various
long trail with a series of small, patchy ground-nestingwaspsandbeesatthetwo
clearings(Fig.IB),allwithina156hectare sites(e.g.MartinsandAlmeida1994;Mar-
researchfacilityof2ndgrowthvegetation. tins and Antonini 1994), A. gracilis has
DetailsofthevegetationofEsta^aoEcol- beenactiveduringtheperiodofFebruary
ogica are discussed by Martins and Al- through September. However, they have
meida (1994) and Martins and Antonini been noticeably absent during the rainy
(1994). Both sites had dense, grassy and season of October through January,
shrubby vegetationalong theedges.Am- though the nature of this apparent dor-
mophila gracilis was commonly encoun- mancyremainsamystery.
tered atboth of these sites, atwhich we All values presented are expressed as
logged over 100hoursofobservationsat the mean ± standard deviation. Voucher
eacWhefrsopmenAtprtihletfoirDstecfeemwbewree1k9s93.making sinpetchieme"nLsaboofraAt.ogrriaocildieshEacvoelogbieaenepClaocme-d
preliminary observations and marking portamento de Insetos" at UFMG, Belo
and measuring female A. gracilis. They Horizonte,MinasGerais,Brazil,andinthe
were hand netted and worked into a Illinois Natural History Survey, Urbana,
matchbox such that only their head and Illinois, USA. Voucher specimens of the
242 JournalofHymenopteraResearch
Fig.1. A-B.Nestingsites,A."Prefeitura,"showingthehomogeneousnatureofthearea,B."Esta^aoEcol-
6gica,"showingtheheterogeneous,patchynatureofthearea.
Metopia species (Diptera: Sarcophagidae: sionallines.Totestwhetherourdistribu-
Miltogramminae)havebeenplacedinthe tionsweredifferentfromrandom,wecal-
Swedish Museum of Natural History, culated a ;-valuebased on theS-statistic
Stockholm,Sweden.Thenestdistributions suggestedbyDurbin(1965)andcompared
underwent nearest neighbor analyses in itwiththenormal.
one dimension (after Boots and Getis RESULTSANDDISCUSSION
1988), usingeachnestasa pointalonga
line. The nesting sites were linear in na- Themeanheadwidthandthoraxwidth
ntuarlel,yasnodassotowebreerceodmucperdestsoedonleondgiitmuedni-- omfmtheanmdar2k.e6d8±A0..2j^5racmilmis,warses3p.e3c3t±i0v.e2l9y
Volume5,1996 243
01
1/3
«
9
V
S
e
u
Ol
ZS
1-10 11-20 21-30 31-40 41-50 51-60 61-70 71-80 81-90
NumberofDaysBetweenMarkingandLastSighting
Fig.2. Histogramrepresentingtheminimumlongevitiesoftheindividual,markedAtmnoplulagracilisfe-
males.
(n=53), with no differences between the wasps were typically active only during
sites. sunnyperiodsoftheday,withlittleorno
Ammophiliigracilisisamassprovisioner, activityoncloudydays.
alwayssupplying 1 or2geometridcater- TheadultlifespanofA.gracilisiscon-
pillars (Lepidoptera: Geometridae) to a siderably longer than for any otherAm-
nestovera 1-2dayperiodbeforeperma- mophila species recorded. The minimum
nentclosure. Otherspeciesareknownto longevityhistogram(Fig.2)representsthe
useavarietyofprey,suchasA.ahertiHal- longestperiodoftimebetweenourinitial
deman,whichpreyonmembersof14gen- markingandlastsightingofanindividu-
erainfivefamiliesofLepidoptera(Parker al,with84daysbeingthelongestinterval.
ctal.1980),andA.harii,whichusesatleast Eighteenindividualswereneverseenafter
16 genera in seven families (Hager and theinitialmarking(notshowninFig.2).
Kurczewski1986).
Marked indi\'iduals were never ob- NestDigging
served farfromwheretheywereinitially Thegeneralnestingbehaviorwassimi-
coanpltyurseede.nAwtitPhrienfeaitbuoruat, 1in0d-i2v0idmualosfwtheerier wlahrilbeetsewaerecnhintgh,ethsetufdeymaslietesw.asTpypiwcoalullyd,
originalmarkingsite.AtEsta^aoEcologi- notactaggressivelytowardsotherinsects.
ca,theywereneverobservedtomovebe- She usually concentrated her search in
tweenclearings,and theywereonlyseen sandy patches and along cracks of more
inthesamegeneralareaastheywereorig- compact ground, and would often start
inallymarkedinagivenclearing.AsinA. digginginseveraldifferentspotsbeforefi-
Iiarti (Hager and Kurczewski 1986), nallysettlinginonone,similartothatof
markedA.gracilisfemaleseachconstruct- its close relative (another Ammophilini),
edtheirownnestswithinthissamearea. Podaloniarobusla(Cresson)(Kurczewskiet
However, each individual wasp was not al. 1992). On most occasions she would
seeneveryday,andtheiractivityonthose use pebbles and sand to rebury unsatis-
days is unknown, but of interest. The factory holes, and would occasionally
244 JournalofHymenopteraResearch
abandon nearly completed nests, as was left.Shewouldoftencarrytheantintothe
also seen in A. sabulosa (L.) (Field 1989). air, dropping it a few cm away, as was
Interestingly,ononeoccasion,anA.grac- also observed forA. dysmica (Rosenheim
ilisfemalewasobserveddiggingtwonests 1987). When she encountered a female
simultaneously, each a few centimeters conspecific, or another ground-nesting
apart. This was probably reminiscent of wasp, which were common to the area,
the false burrows discussed by Evans she would attack it aggressively, driving
(1966a), where more than oneburrow is itawayinasimilarfashionaswithants.
simultaneously constructed in a possible Thecaseofperchingsatelliteflies(Dip-
efforttomisleadparasites. tera: Sarcophagidae: Miltogramminae),
Once a suitable spot was found, she was rather interesting, and warrants fur-
would begin cutting soil with her large therinvestigation.Whenaflyorflieswere
mandibles, flying out of the hole and perching near her nest, she would often
throwingeachloadofsoilaboutone-half stopnestingactivihesandfreeze.Thishas
meterfromthehole,inalldirections.She beenaptlydescribedas "freeze-stops"in
did, however, consistently enter thehole some other sphecid wasps (Alcock 1975;
fromasingledirection.Asshedugdeeper Spofftirdetal. 1986),and was mentioned
(to approximately thorax depth), she forA.harii(HagerandKurczewski1985).
wouldstartflyingoutoftheholeinasin- The wasp would also crouch low to the
gledirection,about45°toonesideofher ground with her legs spread wide, re-
entrance direction, throwing soil farther maininginthispositionuntilthefliesleft.
fromtheholeeachtime,withamaximum Thisisdescribedhereas"crouching"be-
distanceofabout1.5-2.0m.Thisbehavior havior. Sometimes, however, she would
offlyingloadsofsoilawayfromthenest attack the parasites, temporarily driving
was also reported, and nicely illustrated themfromhernestingarea. Despitesuch
withphotographs,forA.piibescensCourtis efforts,parasitismofthenestswasatleast
and other sphecids (Olberg 195m9)m. Com- 8.37obytheseflies.
wplietthedanneesnttsraanvceeradgieadme2t1e.r6±o2f.60.4±0.8demepm, ProvisioningandNestClosure
(n=18). Oncethenestwascomplete,shewould
During the entire process she would searchforaplugtoformatemporaryclo-
regularlystopandflyintothenearbyveg- sure. In searching for a plug, shewould
etationtotakenectarfromanyofseveral pick up and manipulate numerous peb-
plantspecies,includingElephantopusmollis bles in her mandibles, often dropping
Humboldt, Bonpland, and Kunth (Aster- themwithouttryingthemintheholebe-
aceae), Vcnioinapoh/aiUlws(Sprengel)Les- fore finally finding a suitableone, which
sing (Asteraceae), Brcdcinei/crn floribunda was also observed in A. aberti (Powell
Willdenow(Apocynaceae),Mitricnrpushir- 1964).Tlienshewould plugtheholeand
tiis(L.)DeCandolle(Rubiaceaej,Wnltlierin pileseveral (mean: 6.25±1.50;n=4)more
indica L. (Sterculiaceae), and a Sida sp. smaller pebbles on top, finally shoveling
(Malvaceae).Thesesharethecharacteristic sandovertheentrance.Oncesoplugged,
ofpossessingsmallflowers. shewould fly offand disappear, usually
nottoreturnforoveranhourortwo,and
InteractionsWithOtherInsects sometimesnotuntilthenextday.Onsev-
While digging, female A. gracilis often eraloccasions,weobservedfemalesmov-
hadencounterswithotherinsects.Anant ing to tall grass and running their man-
crossing her nest building area was re- diblesupanddowntheblades,asifclean-
spondedtoaggressively,byattackingand ingthemouthparts.
hoveringabove,nippingattheantuntilit Eventually,shecouldbeseendragging,
Volume5,1996 245
weresmaller,suchthatsheneededtofind
anotherpreyitem,shewouldmakeanoth-
ertemporaryclosure,asdescribedabove.
Themeancaterpillarsize(n=9),including
npersetys,frwomamsmbo3t0h.4o±ne1-0.a8ndmmtwo-lcoantger,piallnadr
3.9±0.9 body width. Forpermanent
closure, she would set the plug deeper
intothehole,thenaddingsmallerpebbles
(mean: 12.50±1.29;n=4)beforeshoveling
in sand and packing it tight, using her
headandmandibles,orapebblegrasped
in her mandibles, to push. A typical se-
quence would be: add plug, then seven
pebbles, then shovel in some sand, then
add two pebbles and a small stick, then
shovel insomemoresand, thenadd five
pebbles,thenfinishbyshovelinginsand.
Oncefilledinwithpebblesandsand,she
would carefully camouflage the area by
Fig.3. EggofAmmophilagracilisonthefirstabdom- moving sand, pebbles, and small sticks
inalsegmentsofthepreycaterpillar. paerbobulneds atnhed setnitcrkasncuep, teov0e.n5 mreafrrroamngitnhge
nestentrance.
oprartaalkyiznegdshcoartetrphiolplapri,ngwhfilicghhtsshweithh,ada MortalityandEmergencePatterns
graspedinhermandiblesbelowitsthorax, Of thoseA. gracilis that emerged from
usually venter up, as has also been ob- both sites (n=12), the mean time period
served for several AnnnophiJa species spentundergroundafternestclosurewas
(Powell 1964; Tsuneki 1968). Then she 56.30±10.14days.Althoughtherangewas
wouldfindherpluggedhole,dropthecat- quitewide(37days),therewasnocorre-
erpillar nearby, and unplug the nest. At lation between days spent underground
both sites, ants would occasionally carry andthedate.Infact,theindividualswith
off the prey if left for more than a few the shortest (35 days) and longest (72
minutes. Rosenheim (1989)observed that days) times were initially buried within
over5%ofpreyitemsofA. dysmicawere fourdaysofeachother.Anadditionalsix
stolen by ants. After inspecting the nest, pupae were excavated from their nests
shewouldbackdownthehole,dragging priortoadultemergence.
thecaterpillardownheadfirst,andwould Ofthe remaining66nests thatdid not
remain inside for 1-3 minutesbeforeex- haveA.gracilisemergenceorpupae,seven
iting,presumablylayinganeggonthelat- nestswerefoundtohavebeensuccessful-
eral partofthefirst few segmentsofthe ly parasitized by members of an unde-
prey'sabdomen(Fig.3),oronthesecond scribed species of Metopia (Allenicia)
orthirdthoracicsegment. Townsend (nearM.sinipalpisAllen)(Dip-
Then she searched out a new plug, or tera: Sarcophagidae: Miltogramminae)(T.
occasionallyusedtheoldone,toclosethe Pape, pers. comm.). Each of these nests
nest.Ifthefirstcaterpillarwasalargeone produced from one to ten flies within
(e.g., 2x her own body size), she would 35.75±4.99 days after nest closure. After
putapermanentclosureonthenest. Ifit excavation, some prey items were ob-
246 JournalofHymenopteraResearch
servedtohaveupto15parasiteeggsclus- the mortality ofanothersphecid, Tachys-
teredoverthecaterpillar'sheadcapsuleor phextcriniuatus(Smith),duetomiltogram-
firstthoracicsegment. Nootherparasites mineflyparasitismalone,was30.6-57.9%,
wererecoveredfromnestsofA.gracilisin dependinguponnestingsite(Spoffordet
thisstudy,althoughtherewerenumerous al.1986).Only10.6%ofthemortalityoiA.
diggerwaspandbeeparasites(especially graciliscouldbeexplainedbymiltogram-
Diptera: Bombyliidae, and Hymenoptera: mLne fly parasitism, although the actual
Chrysididae, Mutillidae, and Leucospi- rateofparasitismisprobablyconsiderably
dae) present at each site. We can only higher ifantpredationofclosed nests is
speculate about the remaining mortality great.
nfaocnt-oresm,erwgheinccheowfer5e9orfestphoens8i4bltoetalfonrestthse. NestDistributions
We suspect that there is extremely high For our nearest neighbor analyses, us-
ant predation, as all of these nests were ingDurbin'sS-statistic(Durbin 1965),we
excavatedtoyieldnoremainswhatsoever. concludedthatthedistributionofA.grac-
Rosenheim (1987) observed that ants ilis nestswas clumped at Prefeitura. The
wouldalsopreyonnestcontentsafterfi- calculationoftheS-statisticforthePrefei-
nal nestclosure inA. di/sniica. Therefore, turanestsyieldeda2-valueof—2.406.Be-
we do not know the true rate of milto- causethiscalculatedvalueofzisnegative
grammine fly parasitism, as these nests andthevalueobtainedfromthetablesof
could also have been removed by ants. the normal distribution is smaller than
Assumingnestswithfliesweredestroyed 0.05 (P=0.016), the H., (that the distribu-
by ants at the same rate as those with tionofnestsisrandom)isrejectedinfavor
wasps, flyparasitismcouldhavebeenas ofone indicating a clumped distribution
highas28%. ofpointsalongtheline.Regardingthedis-
On only one occasion did we observe tributionofnestsatEstaqaoEcologica(us-
theresults(butnottheeventitself,unfor- ingonlythemostheavilynestedclearing,
tunately) of physical removal of a prey at the beginning of the series of patchy
itemafternestclosure.Withinonedayaf- clearings),wefoundthatwecouldnotre-
ter an observed final nest closure, we ject H„. The calculation of the S-statistic
found thenestunpluggedwiththepara- yieldeda::-valueof-1.027.Thevalueob-
lyzedcaterpillarbesidetheentrance,with tained from the tablesofthenormal dis-
noA. gracilis egg attached. This removal tributionislargerthan0.05(P=0.306),in-
ofprey may havebeenbya conspecific, dicatingthedistributioncannotbeconsid-
ashasbeen observed, forexample, inA. ered different from random. This could
sabulosa (Field 1989), A. di/suiica (Rosen- possiblybeexplainedbythesmallersam-
heim 1987), and A. aberti (Parker ct nl. plesize,oritmaybea realdifferencein
1980). the distributional patterns between the
Interestingly, the total mortality forA. twosites.
graciliswasquitehighcomparedtoother If the differences in nest distributions
published accountsofAtuuiophilaspecies. betweenthesitesarereal,theycanbeac-
The total mortality for both study sites counted for. It is possible that there isa
was78.6%(66of84nests),withPrefeihira differential parasite and predator pres-
mortalityat72.3%,andEstaqaoEcologica sure, causing more clumping and aggre-
mortalityat907o.Mortalityratesforother gationatthePrefeiturasite,butmoredata
speciesinclude:52.5%forA.lii/smica(Ro- is needed to support this. If that is the
senheim 1987); 51.7% forA. liarti (Hager case, there may be less pressure on A.
andKurczewski1986);and33%forA.sa- gracilisintheverydiverse,patchyareasof
bulosa(Field 1989).OutsideofAmmopiiila, Esta^ao EcokSgica, where they could be
Volume5,1996 247
more difficult to find by searching, gen- gleichendenVerhaltensforschung.Beliariour1:1-
nelaruramgleei,zreoodupespna,roatashinetders.hgormPorouegnfedeni-tneueorsuatsinissgitea,wawvsieptrshy Alcoc3diki4an.,ei)l.]l.iAnmu1Ke9i7ri5ih.jctauTnshMCeiadmnleeasrmtoiinnNgn(tbiHeiyhmmateivnsitoopr9t3:eofr2a2P:2h-iS2lp2nhn6et.chius-
continuously present. This, coupled with Baerends, G.P. 1941. Fortpflanzungsverhaltenund
thenumerousparasitescouldpressurethe orientierungderGrabwespeAmtnophilacampei-
swntupahmosebnpmessreasti,n(tlHoaeaassmstmiaalslttloyopmnaeegg1orp9fer7go1"at;steeiclWotfcinissoih,nslhaboefyrfod1sr9"hd8e4ir)ene,rg- BBooohtasrtft,trhoi,erfB.n.WRiNToa.i.rMjlPd.ardsne:c&shdsrAi,AfA.tG.BSeeGr.nreoektorMeirilsec.enEynkR.1tee9ov6.8im98so.5il1oo9npPg7.poi6i.e.Unnt8iS4Pvp:aeht6ret8csei-ird2tn7yA5Wn.oaafslpyCssailsio.-f
where the probability of nest parasitism ScientificGeographySeries,Vol.8.SagePubli-
mpdHaeaorctwarieseoivantesereos,fprneiwetsisshttuahirsnegiamnalcsgarogyerabesaegciaetntngiaogpnnarseio,snptsaotnsdtdeehdneisnitfthofyara.t-- EDvuarnbsihcRa,noaty,viaiHolo.Jn.ErsS.,to1a9fNt61ied59stw.5ii9gbc.gauDleirrOsSybocwcsuiaPeseasrstrypvik,aso.tnsio9efor2oinntepshspeP.oByngk2ee7n.t:huesj4o3nuA7rem-ns4mat3oli8pn.hogiflbtaeh.-e
vorofdelayed nestprovisioning (Rosen- AmericanMidlandNaturalist62:449-473.
heim 1989) or progressive provisioning Evans,H.E. 1965. Simultaneouscareofmorethan
(Evans 1966b; Hager and Kurczewski onenestbyAmmophilaaztecaCameron(Hyme-
1985). Evansn,opHt.eEr.a:1S9p6h6eac.idTahe)e.aPcscyecshseor7y2:bu8-r2r3o.wsofdigger
ACKNOWLEDGMENTS wasps.Science15:465-471.
Evans,H.E. 1966b.TheComparativeEthologyandEvo-
Theauthorswishtothankthefollowingpeople lution ofthe Sand Wasps. Harvard University
fromUFMG;G.WilsonFemandes,whowaslargely Press,Cambridge.526pp.
responsibleforourcollaboration,AlexandreGodinho Evans, H.E. and M.J. West-Eberhard. 1970. The
andIvanaReisLamas,fortheirhospitalityandmore, Wasps.UniversityofMichiganPress,AnnArbor.
HelcioRibeiroPimentaandLourdesAragaoScares 265pp.
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da<;aoBiodiversitas,inBeloHorizonte,whofurther Hager,B.J.&F.E.Kurczewski. 1985. Cleptoparasi-
aidedthisresearchinmanyways,andHelenG.Gai- dsmoiAmmophilaharti(Femald)(Hymenoptera:
imanrgi,thfeormahneulspcriniptth,ewfieeldthaanndklDabo.ugFloarskiAn.dlYyanreevgiaewo-f Sseprh\e'caitidoanes)obnyPShernoostianeilnliaaavuirgiiflaacnisesAlDloenw,newsith(Doibp--
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eAdlistoo,rwoefatrheegJroeuartnlayl,inadnedbtaendatonoBneytimnoausAternevioefwetrh.e Hagerio,rBo.Jf.A&mmFo.pE.hiKluarchzaerwtisk(iF.er1n9a8l6d.)N(eHsytmienngopbteehraav:-
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