ee )=3—)lsC ne Seok Vl, a9 1889 seetactepte l Enems @seets nek cw RRP ar talon of usonoran of Kaeser) (aopom) andes saermtsmnenme Ge oniitiat aeeasent a etm wows Sn ameter ‘Smith, JL. & MKC Rust 1990, Tunneling response and mortalty of the western Siesta onl thi nace genes ini os eater Entomol Soe.Amer. 65: 756-757. tenes 7 ona er npennegtnarsmne cee a eee ong apn Te a Eee creer enn mes oa ECT a er ny ‘populations of he Formosan subterranean trmta woptera Rhnotermtaae) in 6a, BRN Borman. Socnbalegy 12: 200304 ) fcc ee cart mree een ten emer veg cet eo earrans Seigitint ternary ageing Symposiamonthe FomosanSustoranean Terme, enh istngofthe Pte an Conese eicamsae arerer or one ) Etholagy of the Robber Stingloss Bee, Lestrimelitta lmao (Hymenoptera: Apidae) by ‘Shi F, Sakagams, David W. Rouble, & Ronalfo Zucch" assTRAcT ‘Over 9 eda by colonies of robber singlass boos Loses nao were ‘bsoned n Bre! and Panama, n most eatas involving a known robber obey or obsrvatin hive Petal hots inludedatioat nave sings bee species an Aticanized Apis matfra ntoal, 17 spocies hosted ads \Wereporthere studies made ofthe robbing proces focusngon bthhost and paras Robbing cloiesomoved brood coll provcons, same honey, wax End resin and pen from meliponine nat, seldom exterminating colonies. Geena wre not atacked, while provisions in brood cals were totaly removed honey as pllaged tomanaybee andeausedeaiony ascondng Prelored hosts nad long, arumen entrance tubes, moderato toarge colony Size, high quay Brood cot provisons and were no larger than Lima. “Seaptotigenaand NannotigenaresstodLlmaoreavely stony and w itequent host bul avayeravested robbers successulyInvadod. Aube moray during ride usualy was sigh, lasses of immeblo host adults ‘emalned neat pupee and propupae, ft untovched by robbers. Host bees ‘Sudony resumed scvty and nest dfense near tho trminatin of ais, robbers gradually wehdraw. ‘Guarding L.Imao wore unagresshve a thir own net ut dung raids formed ating around the host nest entrance and defended fom al other insocs, Guarting robbers oeceslonalelvatedthe abdomen and exposed ‘hts ntereogmental gion between hid erga whe fanning the wings. Exeaptatthe stato ond ofa, when host and robber ough (dacerible release of tal odor rom robber mandbuar ands. No robber fdr was noted Ista nests thvugh most of tho aid. Won rads were ‘ill intomuptd by rolosse of . ao mandibular gland substances, Fobbers uly departed andhost bees rived king robbers that remained. Repeating this experiment not eays produce the same result. Comex timing inthe lease of hoat and robber chmicals ely contol the robbing “umes fw Tengen So Hea ry Sp Ow se ies ‘Solar apo enh tan Sz Sabo Parana rUntO@APOAAMECSeeder dette ed ze Seen Vt 21, Na 189 process Hos nests wore occupod by Lestimelta for upto 8 days bu usualy ony ‘one day for less than 4s mp clonias were ak aided exdtaneo Raids occured mero requety in wot season hon Hower abundance and ‘colony food stores wore lw thus rad Wequeney was invereay related to potential harvest per raid. Mass aids ncorpvated up 1 600 boos Willd ‘ais for nest materia were made by ony afew bes. Large groups arived at hostnestsolowinglane scouts and emallgroups, Throughout aad, workers few nan olipso at the L lmao nest entance. Raising boos were guar, colectrs a ling mate orfod caletors, pearing nto specials ‘coating tag, KEY WORDS: social bees, stinoss bees, robbing behavior, Lestrinelt, ‘nest usurpation, troplel bee communes, Atianized honeybee itRopucTion Evolution oftowervstinghabisin bees edt storage ofnctarandpoln Inthe nostsasprovisons orl. This mirobaly processed and eniched foodatvactsmanytypeso! predsiors and parasites, butfow aroso specialized land prevalent as paresite boos Bahar 1070; Michenes 1974: Wes 1967; Rubik 1960). Obligate robbers, somtimes trmed "lptabate™ bees ae ‘nope and eusocial, consisting of af least 7 netopical Lestimolitaandone ‘Aan Cloptorgona (Sakagary & Laroca TOS3; Wile 1070, Camargo & ‘Moure 1989; Ayala 1002),Therobbingeyndome rosolndependentyinhese _2lneages,indeated most by divergent gentle charactor Mishene 199), [Al species but Lmao F. Smith apparently have resctod dtibutona, ‘though upto 3 are sympatie (Camargo & Mowe 1065). Wo have deter: mined, by exairing males, that we sudled asihgl species whch ranges at least beteen Panama and southem Braz, earesponding to Liao interpoted by Schwarz (1949), Nonethooss, he ehlogy of robbing bees remains te studied due to arya the parasxe and dfeutesin observing colonies. Oly 7 nests of Lastimelita were loeatod in 3 communty-nie ‘suveys producing 507 nests of 64 melponine species (Hubbell & Johnson 1977; Roubl 1978, 1989). Robeerboos may usu enrehostnestsbut more commonly, sreprted herein, selectively harvest rom host colonies without ‘exterminating thom. In bth Ronayboos and stgles boos, robbing soars faitycommon, witht origratc exception of Astanstinglessboos Noguela Noo 1949; Mlchener 1578; Dyer & Seeley 1087, andpersonalebeewalgns), Certain ans rob fod rom other at species ests cari back tothe most (son 1971). Such ants have boon termed coptobitlo but donot usurp the host nest or remave ts ood. Robber bees totefore appear unique among boos andthe social insect, ‘Wheress occasional pllaging behavior has been shown for unspecilized hag Pet i — Petar a ne retin singeosboos the genera Oxtigona,Somatigonsan elpona (Row 18h ootaete is ovr ane stn ont ants rao. ‘Shao’ ard Chal ecology sony carte ispecies oxsterc as ‘hotles parser tues cies Raceg beer indo gnu hat Sean epost ono tl nvr aaah etal Tella amemre r wit ua toewaton hres Qltenr 1" Nagano T6210, owe tat o88 Wa tts Sutopan Lace begat 07 roca Oh 380. Ware SPC ao oe le cehaie pnd reste ca nd ceria oma apart roel netrates frre (Crs 1 Canc ‘cor un f005 Sumer 197) Adogesoodceercmaynte Crop. the bee posers ris eon 9 dy lk cea on Afb og, ote pole fdr carts (enandm & Roca : Sc arpa Kr (1961 thas been sages song men tsa by a opt bes soso th ie me Secret wi ect. ny sgt carnage aupponedy rn 0 ornate pase Howey tngicsbesmshes ina (Teapot) rush tiga rnd cr hao wih eno cay detonve cho tan 008: tnt et. 1900, Sgaes boo sede tak Sate ea Letina bares nts at hay ‘Choe tehoorts oduct besasnte parse Nepors 1079 trovsedta olny etostboesmghpreacethareaut Tene Eputees re east n gt sow dla rca hte, Gur ppl Choate ooo teeta otasenats rao on apart tela stay asthe mia tnt ard bok rere MeTH00s 4) Studies in Braz Observations were made during wots ta subtropical lal, Fibro Preto, at 218, aude, When fomperatures topo 5" boo actly stats well ‘terbees openthol nest entrance, Bazin Limaovers sudiedinacoiony ‘dmicled na las lopped observation ive (Sakagar 1966). The nest snd ‘ony were caloctod at Aragaria in ho sat of Goias and were cbseved ina mslponay onthe Ribera Preto campus of ts Uivesty of So Paulo, during 2 ly 102 October, 1871. vinyl pipe corridor hough th wal of he bosang mwhientheeclony waskept provided a nest envance tube. Oly on ‘olny of Lastimelitsxited atthe. Intisargatre were approximaly {7S colonies of 17 spaces of ther stngloss boos, and about 50 hives of ‘Aicanizod Apia malfea Larall, the fra Neovopiel honeybees sil Interred ocasionaly with imparted European apary bees (Ruiner 1988;2 seston Vt 21, Naa. 180 kno Rich ac Pe Bos Foutk 198; Rout ota in pes). inspections of the robber colony and ts actvty were mado daly for 2 ‘months at Ribas Preto, Bursts of acy signaled Yad. Meliponary . colonies were chocked to confi whic, I ary, hosted a rad, On some ‘ceasions Lesimaitapllaged caionies asi he malpenary, Seated by the return of boos earyng corimen nestng material onthe Ned tae = (Fig) .Inmary instances, however, rlsed bee colonies wer observed (Fg 4), andthe attacks had begun So mln po datcton, “Theos ealny was obsorved intermittent unt whdraval of he rang bes In adation fo observing robber behavior and host uid ofthe net, {he ids of hived host colories were occasional removed or replaced wih a sheet of glase to record nronest behavior, Robbers somstinesInvaded ‘oones in lass ppd cbservaton hives, lowing continuous observaton ofthe entre rating proceas 2) Studies in Panama inPanama, near soaleveland et Nt, cunalboe foraging actly I begins at eat by 05:20 thvoughot the year A pronounced dry season fats | ‘fomJanutry ul Apo ay- The destmenthis Marchandralfalishoayy In October end May (Rand & and 198) Fou cloras of Lestmalita wera ‘Maltainod in sections otros trunk n vis they were found and 4 natal ‘esting colonies were observed in lowand forest Seven of those colons \were foundin Back mangrove forest near tho Pactie Panama Canal ointhe lentand wel forest 60k east of Panama Cy (Rouble 1985, Four natural ‘oon in Woes, and 3 rais af Lmao onthe nase of ther apscieg, ere ‘bservedin Soberania National Park narhf Gamboa hint Canal are, at the above fd ses, and alo on Barro Colorado sland (BC), a natire ‘reserve mainiained bythe Smithsorian Tropial Research Intute. Other ‘aids, constidng most of toa in the study, wore noted dung bit daly Inspections of stingloss bee ness ding 1982101902 st melpanrios kept ih ' {resent area approximatoly 150m tom mixed socendary and remnant ‘maryland forest al Curndu, onthe Pace side ofthe Panama Canal | ‘area Mefpontiescoriainod om 400 60 colonies of 151030 species, most in natural og nests found In lonfand forests (Rout 1999) The ype and umber of clones changed fom ment to ment as new coloiee were boughtinerremaved. Some weremaintsnediawooden ives ve difarent colonies of L limao were kept for observation at diferent times, Fld | ‘bservatins of colonies were made ding a bu 3 oars between 1520 and | ‘902, Experiments performed athe melpanay dering rade are deacrbed ‘elon the Results seton, Curundu rads, and those a fold sts, restited ftom th acuty of natural clones % o00 , 7% Xrw ceservation interrntet oo 00 ate — 5h A 1 aotanse neta with carne Toss Templeton ctrg chy Leena oe nt eer rnd onan eg 367 Soa trans oa Ss oO (© very cal, vo FTghts o00 0 Aus XXX OO OO OOO DOO Oe e , oo x00 a— — 8 i # ooeeo00 oyooyy Odetive MiQhts@ cote tow Mighte Yito, tact tnfometon present 1 Moning wit carmen Tote SeSe ne Sony We, No 808 RESULTS 1) Activity at Nest Entrances — Guarding Despite thor aggresive robbing behav, workers of Lmao wate un- _agqressvelonardobserversorinsctsepproachngthesnesia Thorax lan ‘est was conta vsted by honeybee workers and alo by stingese bees trom the melponay, such aa Scgptorgena posica Laelia, S Dipuncata Lepelte,Matpona pico chanel Grbodo, M quadifaracataLepeltio, M.nuvenis Lepeleter and Tigana spnpes Fabricius (Tabla 1 & Fg) Esch colectedcarumen femtho ot protuberances ofthe nt nance (F, 26). Guarding Lestimelta caked no response besides extending the artennsotowardthese bees, TubosvoresortmesbulloywerkarLectimalta ‘pon entrances of colonia bang aided, but the natural nest eirance fs ‘sual larger, projecting up to 20 2 30cm thom the nest of Lestimelta (Schwarz 19:79; Wile &ichoner 1978Fg.2) Colany debs eectod am hess of Lesbomita accumulates under the onranea (lg. 26) and was afte visted by workars of the stnglss bee Partamona in Panama, Wich 7 {Unoe morehost bows than robbers (range = 31040, N= 10 colonies). none Instancotherebbercolonylstateas 148 workersinraidsonS,postea dug ‘early Smo, while various colonies ofthe host species lst a combined 1310 ‘workers duet these rats, When the hovering be began to land duing ber wahsrawal fanning theiwings th emaring rebar wiazow within ‘Stow minutes. 416) nterspocte Conflets Related to Robber Attacks ‘Colonie near thoes beng raid occasional responded by attacking host, ‘or robber bees. When Lastimalta sacked colony of Szapiorigona lttapannis Face, afow dozen workers From a nest ot Mopona eburnea_ 2 seceatgy l., o. 1083 separated by 1m rom the Scaptorigna tacked hovering host workers with hele mans. Intrspocsc response to alam odor, presumably al et ‘mandibular gland orgin, were soon several times among T.angustua, & baro-coloradonsi and Mt. ebumee panamica al of whch began to tack ‘each othr and Lostimelta stra id of Scapotigona wasted within ‘few meters ofthe nost of another specie. Sir alarm bahar th as seen for Paratigona emaicops Swaz, Tina mizoonslsSlvere, ‘and Nannotiganapenlampoties. When Lastielta attacked nests lostspeces. the Paratigonaand Tigonaasoatackedand torpedo erat ‘enestofhannotigona, Toy alsoattacked eachother andthe Lestimolta, “These shirishes lasted far ess than an hour thas tf Wkely many Sear ‘nents occured that woreunabsared,Malpora bicolor resctodtaaraion Scaptctiganaby gathering atthe nest enrancs of Lestrinta se raping with quardboes, Pesably some workers! Lesrimotaretring omtehive ‘ot Scaptnigonainadvertenty entered he nest Maipona separstedy los than few meters, Workers of Ht colar hovered in front of eaonis oth (quactfascata ae Apis, grappling in the al nd fasing tothe ground with defenders ofthese colores. Som of tho 2 Malponarelumed tate nests wth dead workers of Lingo attachod to lgs. Afr such rete, exons ‘esponded sty andmany workers ected em te neat, Masses of hovering ‘M-ebumea wore formed arly by presenting one oc 2 rushed worker Lestinlita at nest errancos. Such hovering goups of bess ware need several times on other occasions, and they persisted approximately 20min [Athaugh no attack by Liao was observed, Ray have Been a cause, 16) Ratd Duration Raids lastod a fw ours to 6 days, but the most common sustained raid lose des during a slnge ay. aids nga ingle day ranged tom {to 8 ts (mode average ahs, 8 of 23 cass). "Tho tng of ai aon and termination in the Brazil study fs given in Fig. 8. Rad intiation showed ‘bimodality; te peak occurence was t 10.000 1:00hr and 1220001800 hrs. Reds ended whan al Lestimetita let the host nest, which oocured marly between 1600 and 17:00hrs. Temperal acy of Leste in Panama confimedtrendsrocordedinSra2l Foraging Lestimoliaworasoen the nests of other bees ang every hot of tha day Debreen 0800 and 18:00, wihamaximamnoteabetween 1:00 and 1200, total ot raid was \tnessed, 6 of wich inolved conspecie coleries of Lestinelita. Seven ‘cases of simultanoous rai, probaly eigiating fem the same colony ot Lestimelita volved 2 host colonies hs same species or subgenus Pour wore of 2enlonis of Scaporigonabarocolradensi Onsofthe dave rais ‘fected a colony of Scaptotigonaluspennis and 1 of § pectoral. Party callow Lestimalita, having a uly black Need and propodeum bit whi Sng Pat 8 2a Paar Be a ‘melasomal sgmants found only in young bees, paripted in the rads Suggesting throng rm the same colony. Char double ralds were of 2 olnisol 7. Teragenseaangustaaand2 colonise Paraigonacmatcas. "None of the colony pars were grouped close togeher. Several raids wer ‘bterved at ha Ieepon o rich, but only int Instance wore bath the Bopinnng and ending sbsorvod for @ given host coon. In 11 eases, raids ‘Began fem 0815 to 13:00krs and fished Bebween 11/30 and 15:00. A {otal ald duration of 2.5, for 13.00 fo 15:90, was recorded at 1 nest of ‘Scaptotigona petra: Rass ht lastod mule days occured for 2 days tone nest of Namatiganatostacatamis and for 2 days al a nost of T. [atragoisca) angusia, Nest appeared nota. Nest entrances wore omplete and occupied by host bees, on the nat day fr Nl tostacecortsand ‘day ater for. angustula ‘17, Most Colony Selection ‘Summarizing the Iypes of bees stacked by Lostimlita in Braz and Panama, the group having the highest prbabiy of hosting a aid was ‘Scaplotigona (Table 1) Intenso atlack was ls relatvel igh on some ‘Prebola, Nannotigona, Molpana, and Pavagona, Numbers of rads cb- Served on ach species are reiatodto the toll onl, Thorato of these 2 hurmbors fs used as an dex ot host preference. Relative preference orhosts ‘MiberS rol, calculated inthismanne,sS,postca > P.sp.>P.cronana Sith quadiascata > N estacecomi> T.angustus >» A melifers, Nive Spaclsof singles bee, of 14 clones, never hosted ais (Table 1). The Droference index ocenat apply whan a clany was axerinatederis iors ‘ere completly depleted by aug atack Because twasthon unavalabio {orth ropeatedatactsobseredatmany colonies. AtacksonN.fstacelcoris ‘were conf only trie bt the actual requoncy must have bosn high. Many forages of Lmao etumed to the nest rom ost eaonies that were ‘ot acated (Fig. 1)- These ikl returned rom natural olonies of. tosace- Itami and P.droryana and T, angus nes the cerumen ls pale brown, ‘98 wae cormen cared by those Lesiinlta In Panama, the raiding intently, in descending oder, was Seapotigona barocolradonsis > 3, pectoais > Paratigona omatceps > Nannotigona perlanpoides = S. {tejponns = Lesimeliaa lmao» > P(Scaura lattarss > T.(Pieseomat lra> 7 (Tetagonisca) angustula> M,ebumea Amostalolnies tacked by Lestineltta in Carundu were in natural nesting caves. Albough the ‘ier of specas var ths ste, the nests coneidred hore were Kop feast ano. "Raids inthe fort of Parque Nacional Saberania and elsswhore wero ‘seanon2 nesta Melon oburea (Roubk & Whooor 1982, anc most on BC}, Moipona favasa (trai), Aas meliforascutlata Lato (8 Yad), i_ a Seong Vl, Na. 199 ‘Scapttigona baracolradensis (on 8Ci and Soberanla Park, 2 rise), Partamonasp. (ald, Pebaa fant tad and Tigona muzeensis(on®Ci, 1 raid. Evidence of 2 raids of Mefpons ebunea was oblaned by examining ost ntances whore those of L lmao had baen bu, which were unmistakt by ofthe hoe speci, 18) Spectos Not Attacked ‘Colonies never ralded i Braz incuded 4 epoces of Malpona, 3 species ‘of Plbeia, and the fofoning species groups or genera: Dolchttigona, Lowotigena, Tigenisea, Celetigona, Tigona, Tevagens, Paramons, CCephatatigona sna Gaotigona Fe obsorvaonsinPanama dmorsirated ‘hat succosstl ae affcted some ofthe above spcte thal have highyy ‘aggressive deenaing wotkrs, TigenamuzcensianaPartamana Datatom (Curundyiseate thet 18 species o ingles boos wore never atacked by Losuimetta but 4 of thea were raided elesuhere in Panama: Mona {avosa,Poboatanki,Tigonamuzoensisand Paramanaat cupraF.Srith “The spovadic natura of observations of th sngless be clones at Curund may acount ferthis discrepancy. Colonies ctsoveralmore species. nluing ‘Neguetaps mandala Cckere,Tigona amaien siveztiana Vaca, T. (Teragenses) buchwald Fes, snd Oxyrigona, were kept ore Tew months 1 Cun bit were nt aaced. Bae cleiee Both in eudoor hives and hatralnosts wore raided by Leste, 3s were hives Kopin bukaings at Ribera Preto, where mos colores were n hives. Fr al specie, there Ineutfciort nrmaton an te atactvenas to Lestimalita Non@tss3 ‘ll eaionies present at Curd wore observed wih appronmatly equal {requeney, thus the colenios in placefor soveral months establish tne foraging hate af Lina, twas uncertain whether a specs nevor succeseuly attacked was voided, merely missed by scouing Lestimelta or percelved as fr less ‘Sulatl a ahost han ater accessible specie. Further nates weaken ont tho posable ozourence of avoidance, Considering the amas species nat tacked, to Mypage’ ex Trgonscagraup, ast envances weretoo small to pormt the ortry of Lastimalia One atack of L.Hmao on Plbela 9p. romplod removal ofthis ealony, whereupon twas replaced ith anasto Tigona(Fesoomata) vara Lepletr. Eight minutos ar tho rid bogan, others of Fresoomelta defended tht nest fom Lestimaita, which Eon sbandoned the raid, avg one doad worker. On anche occasion, shorty sterthe nest entrance of Seaptotigana posta was oxoupiedby Lostimelta ‘honest wasremovedandrepiacedwithoneolFiasocrolta varia Ro 109 ‘Workers of Lestimelta sive near he est entrance, ch waa prom blocked bythe heads of Frescomelta. Lestimaa landed now tho nest ‘entrance but lanotappreacht Whonthohive wasslghty opened, numerous Saar et et — ter Ba 2 workers of Frieseometitalet the nest and tow tthe Lestat, taking thom {othe ground, locked in eombat. Fory-fve minutes afer Fisoomolta was Inodued, al Lstimelitadpattd, Two workiledy Fieseometa, which sustaned no morally. Two ar attacks on Scaptotigona wer nienionaly Interupted by suetion ofa clory of Frasoamelta varia, wih the came rest piscussion (ur observations of 83 aid by L.Imaoin widely separated areas include the ony deal tsa of theae events Conltency genet pir and onformtywitmanyebservaionsbyehorresearchers show thal Lestrimeta [S'a hgh specialzed paras ingot thal eondts aids nat ony i ge ‘t0upe but also a a fo indviut In gona, some workers are saci the begining of end of a raid (Kor 1983), and host species show ‘unistakablo signs ofmrimizing losses, The possi of fecve mechan. leat behavioraor chemical defense against Lestmeltaby specesthat were ‘ever stacked ean nly be resoved tough father stu. 1) Host Selection Preference of Lestat imaotor Seaplctigona was vet lar in Beall, land Panama, Since bah ata share ranges twoughous the lowland Toeats ‘fom northern Argentina to pethaps Meso, they should interact aquent ‘Sima’ verap in dttbusion exist between Lmao ands ether 2 pincipal hosts, Plaeis and Nennotigana Thesa genera cotan about 80 species, ‘hich sugges maols ptetaly quite ganerlze, oven consiorng that ‘only 17 of 40 closely-bseved specs were raided in Panama and Braxl (Table). Mlcane (1046) also winessadraeby .imaoin Panama, noting that 10 sie wor on Hamotgana arming 49 cis ete the presence of 82 clnies and 8 spocies of Pavtamons, Melina, Tig TTeragensca, Peseta and Scmpcrgona. Nogusts eto (1970) tb served 29 aac by lmao vitin ameliponay. These were upon Pebela land Nennotigona, wha lesa abundant colonies ot Melpona, Tetragonisea fd Scoporigona wore untouched. Fuormere, Laroca & Orth (1064) ‘eportad arid on Pebea and Isted 10 host species o singles boos, ‘uggeatng hat Lmao changes sang abt, 0 somo degree, probably ‘ccotngto regional avalabity of spocos,Otwornfrmtion sunmarized by {Ceroca& Orth (1984) confems te ropeatod association of Lestimelita wah host Scaptetigana, Nannotigon, Pela and Melpona “The preloronce that we notod for Seaptogens had nat been suggestad ‘noe Figo (1951), nar Ne interacton wth Tevagonisca boon noted sino Noguoira.Neto (1970), andWile (1961. Inadditonto our suits J. Gonzdlez (Personal communieston) has observed an attack of Presoomalta by aey seceelg/Vl, . 1088 Moscan Lostnatacta cron west spaces yl 102) where th ober ln feguoy rads ness fNerovigra nba Pots, tacingeatny a imotvidedeisingenatelPeseometavara whch Iey neat on ange lesa ty eer nes tats a 1 nis Eig obare apeiron ce ry 3 aad by ima Somevrionnnet anon an gras st ei “eee specs an aupraapesis ups a ae aac eqn, ‘noo tackdesaequrthy spent station oo Stach Anmoeataycame sch aange of ee ar Fata hig clr dey rar bea pad {Fra eal conte evan eed peters, parapet i te ropoioal atc Sequanen guise ay ale Sut wetmepaci shoo espacio mate asve) pot oretea, sccalnay fect eee, coo al eon sv sal Seen and poses ula cpus we ie at oar, ose {er woud poate dco sivas 2) Nest Entrance of Hoste Chances ha fst et etn sam prodary mptart to Lostineita i stent hat os bona Sapotgene cote Pb ‘edacme Tigonscr axe in so be shaped nears ‘helsaeothecane rns Tetapose Somasandsone Partgon Atel ien woap mere plage by Laci curse, In cast theseat moa ter singles bose sor aroma ards erat Thettree el entances ta pforednoaol astnatar at {ny eras ty provi corsen na Crson on perhaps mote imporay fo ang a ear thea Dror ass ‘aston onanos ota hoe ey we uray oad pis freshest te prince (a) any arom antes test ontancas ar madeby Hap, optaltgera Sctigone Uae Sonen) and Taragoe wen do toast tan Sec ses ay tow deters and Gopi tas crane need fe alr nok 3) Lara Poisons of Hosts Larval poo wert dels conser tony and pation vesltintot rats We cheat uch povions re tpt ot Sstetony Cessna taidbe sup seine et ey ‘cla te tien aac. Host eae war nt mecetary es esesaig te mst sted fod, evn ergot oy options ot Eras, hnca tessa one sults plage Pr eal ‘obrieot Anarene scsi. pts goss taney and pot, satya ne Tet tasted ag Poth it — ear Bae ae provisons needed forthe arvaectstingless bees sca tholarva of Apis are {ec diet by workers, The complex of glandular and oe: prepared {ood upon which be larvae fed iatheeforeunavadable honeybee ness Lage colonies of rive singless boos olen are maintained by highly ‘gressive spats ofthe gona Tiga ss Such clories appear tobe ‘tveded by Linas ahough ene fad on a colony ofthe aggressive T. Imuzoensis was observed in Panama, The robber bee attacked esdent bee ‘Colonies wihin most cltheabeanation renin Brazil Panama, Searching “nd foraging acy were spparentyconinoous and proceeded almost on a daly basi during faorabio weather conditions "The quay of larval provions(Hatflder& Engols 1080; Roubik 1080), and alao resin of corumen (Sakagams & Camargo 1968 sled in nosts of ingle bees vary mich among species. Cabohytates cru protein, and ‘symbiotic microbes vary widely between the su boos. The preference of maotor Seaplotigona, Nannovgonsand Peboia landsome Malponamay be song iniuencedy such charactors. These fre complemented bythe quay of source avalable snd he defensive behavior of pote host. ts noteworthy that Sestorigona maintains Colne which ae amen tha latgost of thewe found in eanral Panama and ‘nay have several thourand brood col at glen time (Roubic 198), ‘Gotonies of Seaptotigana pasion ae als aang the argos ound in Braz ‘hlichoner 1874). Tose large colonies ave detended by aggressive Quads, utter defense is weskonod and detested by th spec communication ‘heromaneandazomone of Lesvunelita. Nannerigona,whilonct amongthe [ergot colonies of stinglesa bees, has larval provistons wit high concer tions of protein and both they and Seaplotgona have high energy cotont (Partelder& Engels 1989; Buchmann & Rubi, qutodin Rou 1969). The Concentrations of hase ners ae frm 2 to 10 fines higher than those ‘recorded for ater singles bees. The spol fo which Nannotrgona snd ‘Scapotigonahave been compare, bethinPanama and raz are igona, otpona Partamon, Toragensca, Tettagona,Tigonecaand Leste ‘Ths rango of specioesnilarta that ofthe experimental mlpenerios where raiding by Lostimelta was caer, indete tat robber prterencels aly intuoncod bythe quay and quant of hos larval provisions. {4} Compettion Between Robber Colonies Intense confit ensued between conspecc Losin Panama and confit wth Tetragonica also were ekng Fatal combat betwoon tess ‘olnies has many mpleaons that require furs stay. Arai Lmao TFetragoniac,edminatng in usrpation ofthe robbers ast by tho ralded olny was reportedty Weyraush (quoted in Schwarz 1948). Thoobservation InPanama confms this phenomenon, and als hor ropors thal rettvey