Ium\mtiim\ t*ltin Htttiil.'iWitfrfrtfX Am? M'tWXl 122(21155-6: 55 VVI IIIN-NKST BEHAVIOUR IN A EUSOCIAL AUSTRALIAN I \LI OIUPINE BEE EXONEVRA (EXONEi'KEUA) TKIOENTATA HOUSTON {APIDAK: XVLOCOPINAE) by Zm\Sti:j\'4(S; Mk iimi P Sc iiw.\k/:,: Summary M Silln. /. at StiiWM'/, I* | llMN) \Viihiivne*t behaviour m a eusocial /Vtistialian allodjpuic bee iiufHritrn if\oncttrrltu} Irnlvnlala Houston JApicbe: \yloeopm;ie), 'Ihittx.K. So..S \ti\i 1220.95*03, -'' May, !'>l>8. I'ndeisiaudin^ the processes involved in the evolution of .soeial behaviour kus become oik- of' ihe uintf challenging areas ol modem l*ioloi»_y. Since bees and wasps exhibit j \ariei_v nl micijI 'M^jiusalions ihey are panicukirlv useful lor aiklrvssutu social cvolintunur. qUctriOiK. Mlodapuic hces are especially useful lot .Ajininmu mvial evolution,since speciesdisplay varying formsi>l social oryunis.ilion from solitary meusueial. IInsstudyexaminesvvilhin nest bcHlUViftOI of lAo/ttmu \I:M>/u-nnfio)iritlrutttui, ;i nativeAusitalum.illmlapnie |V\'. rhis -.penes has Ifjc latest known w1t)n> tftc* o| any allodapinc bec and exhibits sinking ni/c \analion antone female Herniates suggesting lhal sociality UttlV be resided .is highly eusouuf Ileiv we assemble a olHt-hteiravniulnlio.dialptijliieasl,otihmicssjfpole'ctiheissdsipHeeciressbuyni(theshmoawrkdeidalparlatchtoiucgehomfaonwynhcuhgugvricosiriircsinaidvisspilmaiyleadrIidnt(hheosfeorremcoolrdheitdilnocr Ovtillj a*_'uiushe tvluoioitts have not1'ceii recorded "01 oilierAu«ai;ilian allodapmcs and have been recorded only lately m oilu-t allodapinc hnina. Jhuwititt (riif'^tttttn appears to diller from olhoi highly eusocial species where there is usually littleor no agression bin instead 'penile despotism". Kl v Wouns. fwmlint irn/toiatu social behaviour, -illodupine bee*, agression Inlrodiiclion aitangevl in a NOtjiiciiec Of piumtive l»» advanced' has been qucslioncd (KuKuk W95J but lew studies I'bfl allodapinc bees provide opportunities for have explicitly invesliyaled whethei prmiitivc 01 Miiiiparaiivu approaches lo the evolution oi social 'advanced' lorms of sociality within ta\a correspond bchaviotu because ol ihe wide range: o( social to hasal tvrdistal positions wilhiti phylogenclic lo-e-, organisation within and hclween species and ^eucru t.y'OUHHi Irittcnlttht isan Australian allodapinc bee One -onull and endemic Australian subgenus lhal lives in .^cmi-arid environmenis. This species has f strnt'tmlht. contains1 fours specie* lhal titttgp IVom ihe tatjiesl known volciiy swes ol any allodapinc bee tilt predominantly solitary l\\o>t<'iiru tnw so/// and exhibits nun-pholopicul dilTereuiiatiou be!ween kaymeut { Vlichenei |V0?| ^ iho etist.rial /'. pUUlfVc i-astes (Houston 1*-J77. Hursl cV Schwar/ ttitkflliflttfHotfStOn l'>77; Hursi & .Schwat/ \$%), |*9*Jfil Much of the informalion about social In most comparative slticjies of insect social organisation has hcen inferreil from disseelion of evolution tlTora is an implicit assumption that lies! occupants and brief observations o! lemales small colony sr/c is a.-.socialed with flexible and ouiside ol their nests (Mntision 1077; Muini beltav iourally medialeil reproduciivi- skew. I he oupub) M is suspevted thai this species exhibits m.imleuauec ol dominance hieiarehtes via physical casie dilTerenlialion, where large females ac'oiiisiii is cortsHJered u pnmili\e trail (Wilson (termed 'Majors') are citieeii like ami smaller l'»7li C'orrespoiulinydv, ItirgC colony si/e> with lemales i'Minors') acl as workers within the sliou^i repiocluctivc skew and non-aLonislically colonies (Houston 1077- Hursl |*)%). Ilowevei, niainiuincd hiciatebie-s aiv Usually reijiirtled as mou- W|l mi oi->.i heha\iour;il studies Imvc not he-encarried derived trait-. Wilson (1^71) su^yested that less olil to assess whrlhci lln:sr lwi> nn>iphs really are sophisticated funns ij| siut;il niyailisatioii would lichavioiually dislmcl. Colony sueandiheassocialion Invoke physical mechanisms ol control such £& Ixlwcen morphology and repnHluclivc slaltis su^^esi a.L'^'iession wilhin acolony, hul thai IhiN is icplacctl by thai this species more closely approaches the highly "gentle ilcspi^lism'1 in more advanceil lorms cri eusocial loim oforganisation characTcrisiic olapiue- sotiabtv- It is* also generally assumed that a hojh rueliponme and highly eusocial halicline bees, than level orbehavioitral spe.ialisalion isa more derived any other allodapinc hee. hail ami iluti this can leadlohigher levels ol colony I In- study invesicates Within-Mesl heluoioiii in cllicieiicy I.Icmuuc (Wo. H<tvvevcr, the idea thai ijbservaliou c<m>nics oi I,, fritit'ntitht. A icitcilouv ol thllcrcnl foims of social organisation can be behaviours is presented here in the loriu id* a behavioural catalogue inul eompaicd with other Sdtoal mI BiOll^liVll Vii-mo l-liiuk'^ Inivfrsilv l>l S.-ulh belntviuiiral studies ol allodapmes. I hesc data will MKiralui 'i''*) ('flu 'i'"1 v-t'-i.ni. s \n,( viu i m also be used i\u- specific analysis til behavioural /(I,ISlvHtlf"IlilHlct^AllH.OI specialisation which will appear in a futureseries of V. / Sll-J NX M V S( flWAK/ publications, hi addition, the idea that morphological had approximately 12 hours in the artificial nest m caste differentiation and large colony si/e mq allow theirodours to permeate the nesl before n was associated Willi low levels H agonisiu in Colcftty possihk- dn them to leave tlhe tic\t morningr Sheks mi* vr.ilion is discussed in relation lo the social were haphazardly plaeed near nesls lo ael as visual oieanisalion ol /.. trirfi'iUitht cues foe returning beus, Materials and Melhmls tit-htividumf nhscrvLttitMS Oitcc obsetvalion ncMs were set up Ives wetv Stmh •i/t'S allowed lo adjusl lo their new environment lor one f\\<>natnt Iritlcrtfftta nests were collected from week het'ore observations began. Data colleehou Lake Gilles Conservation Paris (13ft 48" \\. 32 34' Si involved 'scan* and 'foeal' sampling techniques AluosctatreadliainItnhethinsnrafrleia.eaFs.twoildmKlvurentPenensitnssuwlea.reSporninI-h tIlAieJlpmoasnililol>n74o).f Seaccahn isnadmipvliidnuagliinnvotlhveedobsreercovradtiinogn cipally »n disused beetle burrowsexcavated \uAtttittt nesi. jiMnji a 5 mm scale along Ilie glass and was l><tf>\n<ittriw IJcnth (Weslern Myall) and Huhrlrytw conducted immediately belbte and after lotal oUtiifulimn iDcM'.K (Hulloek Bushr Deail branches sampling. Tins was done to determine whether ol both tree .•.peeics were examined lot nest certain bees were spending more Ihue than olhei in .-Mi.mt.es i.e. the exit holes made by the original certain areas of the nest, lor example, near Ilie hcelle occupants Intact colonies wetv collected entranee or near the brood. Pocal sampling involved during lchruar> 19^5, t tvkl collection ol nests 2 mill observations ofeach bee in a nest. Nests .nui took phtee when temperatures were eool f 12 C-20 inihviduals were raiulomly seleeted eaeh dav (t>i O. to ensure thai all ueeupants weie present. »>rderol t>bseivations. A headband magmbci [^5 Once an entrance hole was loeated, iKli branch was m.tgnifieation) was used lo observe the behaviourolI removed, entrances were blocked with tissue paper, individuals. All hchaviours pelformed in a 2 mm the brunch was placed in a walerproot bag anil period lor each individual were recorded into a stoied in an nisulaied containet with iuc [qr voice opeialed tecoidei Observations were linn transport lo Hinders University. scribed on to data sheets at a later dale These Al Hinders University the nests were stored in a behaviiitir.il data were used to eonslruel Ihe hchav eonslanl temperature room al apptovhnalclv 10 ( toiiral catalogue and later lo examine behavioural lor processing. Nests were opened using a kmle ^VLialisalion. and all nest occupants, including blood and nest Hchavioural observations took place m ihc aln-i dCornht.entAsduslutcsh uwseproelleni,ndwievirdeuatlrlaynsfemrarrekdeldo auPseitnrgi Cnooann.d( Ibse0e0s- 1w7e0r0eh)a,ctwihveen Ilnemtpoeiraaklur1e0s nweesrles :_we2r1*e IJtMntifoM*1 and TcslorsIM enamel paints applied to observed with up to four nesls ftifljlg ob-.eived ill any ilie thorax and metasoma liee colonies were Ihen oiv session, fable provides information alioui I Iraiislerrcd lo artilleial ohservation nests, which nests were observed, when Ihes were obseiveil Artificial nesls were similar in design to those and how many minutes ol observation each bee per deserihed |>> Sehwavv A Overboil t |y^3} hut were IfliM teccived. In addiiion, the numbers ofbees ih n ina<le ot ptne wood instead ol halsa, Bach nest weic present lor the initial a\k\ final obsciwitmn consisted ni .« rectangular piece ot iinireaLed pine peliods ale giveu. wood 210 \ 9j\ ^ |.s mill A gTttftyp was gouged into one longitudinal lace o mm dfWfl \ 200 mm lengthi 'I he groove was smooihed out wilh a metal rod to Ktsoits remove any sphnlcrs ol wood A piece ol glass. 210 \ 2<i mm. was placed flush against ilk groove and secured al both ends wilh insulation 'ape. A blai4. cardboard cover was plaeed ovci the gla-^ loexclude The contenls of nest> collected in bebrtiary l1*"^ IhJil hetweeu observation periods. are summarised m 'lable 1 Uunug lhc--c samphnv t >l>sei\ulionnests wen: set upon suhhonvontal trivs peniKU. cotomes used for behaviouial observations ina shade house/it HindersUniversity.Oneeiul olThe were rearing brood. In early bebruarv colonies shade house was open so that Ives could loraee contained brood Hi all developmental stages, i.e. heeiy outside, Nest enhances lacvd the open c^h\ ol r\';_'-., larvae, prepupae and pupae. By late bebiuaiy the shade house, A maximum ol lour nesls wu- female bees in Ihe colonics had almosl c^ascit ci"j plaeeil on each tray with approximate!) 15 um laving and hrood rnosllv comprised larvae. pfeptyVK hetweeu each nest. Ohservation nests were fiisl and |>upae, There was a great deal ol variation in the plaeed in the shade house ill dusk 5-h days atiei number ol adult lernales present in a nesl. rammer eolleelion and opening- Thin eiisuied that the bees ftom I-1X ibi- I BEHAVIOUR IN AN ALLODAPINE BEE -' TABLE I- Detailsfarnests aj Hxoneura tridentata observedin thissludv. Nest First Last tots1 nun ber Total minutes Initial no. of Final no. of observations observations of of Individuals Individuals observation observation penoIs pel nest per bee pernest 1 7 Mar. 14Apr. 15 3d K 5a 6 7 Mar. 14 Apr. 15 M) 9 5U 9 7 Mar. 14Apr. 15 30 4 I2h 12 7 Mar. 14Apr, 1? M30 13 !5fl 3 5 Apr. 4 May IL> 3 5 4 5 Apr 4 May 1? AS ; 5h :o > Apr 4 May 14 3? 4 5* 30 26Apr. 10 May 20 40 (i 5* 43 26Apr. 16 May 20 40 3 4'1 56 29Apt In May 20 40 4 4 Decreases in thenumber of indivduals were probably due Lo death whilst foraging ordispersal toother nests'1, Increases weredue to the addition ofnewly eclosed bees'1, orintruders which swapped nestsc. Tabu 2. Summaryofnest contents farcolonies oj"Exoneuru tridentata colletledin iehnutry 1995-fttym Lake (lilies. SouthAustnilia. Nesl eontents Mean value (±S.E.) Mean valtie <+S.E.) forearly February forlate February (N=24) fN=B] Hggs 1.21 (0.57) 04)8 (0.08) Larvae 1.75 (0.63) 0.02(0.27) hvpupae ()_(>7<0.2N) 0.31 (0 13) I'upae 2.42(0,72) 2.23(0.7% Majors 1.17(0.1(11 1.23(0.34) Minors 4.17 (0,K3. 4.3XH.30) Males 0.33<(U3> 0.3K(0.2I) heluicitntntl repertoire In Ihe following section behaviours observed during the study are presented us a behavioural catalogue. Observed behaviours ateclassified into four hinciional groups (often iuler-eonneetcd or overlapping): (i| self maintenance, hehaviours. (ii) nest maintenance behaviours, (in) inter-adult behaviours, and fiv) adult-brood interactions. Mmnm Si i.i NAWCI til ii wiDiiks inv nvi: Sees were reeordeLl as being "inactive" when no Oilier behaviour was bciue. performed, hiuelivily often occurred within a behavioural sequence. I;or evample. a bee eould slap grooming, be inaelive Tot some lime, and then navel forward in the nest Bees (v!S I eould either he standing 'upright1 or the> could be lying 'upsidedown' on the floor o\ the nesl. Maeia at Females per nesl cdtefscr(iIp9t9i2'o|n ioufeluadesdimislliagrhtbemhoavvieomuern,ts"Rienstitnhge"ir Fign.esIl)Hoifsthosgoureaumilott"teiniliho-nnytiistiazecso(llneucmtbeedrforlomleLmaakleesCipiellrcs However, in this study bees were only recorded a SouthAustralia. Ivbruyrv l-995> ^S Z STTjEft* M. I*. SCIIWAKZ inaelive when ihev were motionless. Exoncilrti Ct ratithi spp. which have U turning burrow irtthnittift spent ii largo amount of lifftC inactive enlargement near (he nesi emrance (Macia a ul Since inactivity can occur wiihm ami between 1992), Turningotlcnoccurrc<laspart ofa sequenceof bch.oioill'Ul sequences il is difficult ttl show behaviours during interaction* between liulividllllki numerically Lite mmaim ul tirtic upeui jiiuwive i.e. it UTUld occur during sequences which involved because oi ihc wav die data were collected "nudging" "passing" or "avoidance" (see belowT l| -> (ieiicialty. lh0Ugh( lIlC bCCfs wcie inoic active when bee appioaclicd but axoided anolhcr bcc. il inti'lii temperatures were >,!0 (* aiul/or when a loragci cither "travel" up to Hie bee, and then back away iiv il returned might "turuH and "Travel11 ill the opposite direction. si 1 1 tU*i)OMINU NM'IAK l>l nVOKATlMN "(iroommg" was observed lrci|iicutlv. ami included Individuals were observed flexing ami bending the .inv iciivily where Ihc body suilace was cleaned proboscis anil although dropteis ot ueeiai could not Sct|ucnccs (or cleaning dillcrcni areas li die body he seen with ai die magnifications used,, it wax wivrc similar to those reported Tor fihttt/i<(<j>t.\ hrw/n assumed tlliU bbCJ WCl'c dehulraiing ik\tai as has Cameron au«l (intfithi spp., (Mactar/ ni. I('92t.The been observed in olhcr allodapmes alter Iceding mosi common sequences were: U) head cleaned In iMkheuer V>72; Maeiaw,// IW) Some bees slow I iniliaflj Wiping a forcing with the proboscis then ly fully extended and ictiacled the whole pro- loivlcg used to wipe the length oi die antennae. boscis without hendint! it. The proboscis was hi i.mini- HI the htwc; I'orcley a-'.tin Wiped Willi tllC extended and held out lor aboui 20 sec Hi * proboscis, lollowcd h> Ilic wiping of the head rehacicd beIore beiag extendedmmagam Some with the (orders, (h) the rnctasonia wascleaned hy iiulividnals spent die whole u\n observation Using die ithial spins on the luudlcgs |o sciapc oil period pcrlorming this behaviour. JijNl/pcllcn. (t1) die thora\ was cleaned with the mid |Cy£ (die imTisoma and the thorax were often Nt SI AUSI N ISM 1I I Luoimicd at the same tune with the diflcrcnt logs), (d) When individuals were regularly abseni from Ihc Ihc winy surfaces were groomed by drat'i'tiiri ihe nest it was assumed that they were foraging wiirjs under die mctasoma with the hind legs, However, il lhey were absent tor more than 5 obs.r Wiping ihem t»CtWCn the mctasoma aiul hind Icy-. vation sessions In a iow. il was assumed lliat \\\>'\ and then flicking tlicm back inlo position, were either dead or had dispersed. Absenteeism (or (booming did not occur as one |0ng uninlenupled foraging activity! was only observed when temper sequence as has been Clbfcervnd lot $. hcwiiti alines were > 25 C. Poragcrs were idcitldied when (Macia tt oi I1W1) Grooming could Iv briel' or they were seen reluming to the nest, I 'poii rvuiW- la,i lor |lw whole 2 min observation peflod irtjj, Ionisers usually worked their >vo> down the nest passing and interacting with other unh si nail Koln M«>\ i.Mi Mis \nluals. ul'ten bavin;.' 'buccal ermtweP with othci This was intermitlenl hehaviour. which was nftttfl individuals, pivsuntably provitliuy ihcm with iicl- obseiwd during long bouts ol inactivity, and behav- lai (sec inter-adit11 bebavioursi. Often slilIi a iour comprised slight movements of'head, body or bcc Would then leave ihc Ucst auaiu and rcluin leg*, which did nol involve any olhcr type ol behav- lalei. foragers were nol observed IcLdniv tai iour vae. rM Wl>i i im; Nt:si VImnii nvnci Hmwin, i.:s li:i\clling" involved nioMiiy lotwant-. or backwards up or down the ncsi lor 1-20 cm Bees ihal were ia klthW! liavvllme were Usually very active bill Ihc Uavclliug A ticc was recorded as guarding" when il occupied speed vaued. Iiavclliug forward otlcn rcsullctl m a the position closest lo the nestentrance with its bodv bee coming inlo contact with others and was usually oriented so that its bead svas facing away from die followed by "passing" (see below) cnirance. Such a position allows the niciasouiu lo block the nest entrance from intruders, as recorded I! K-'SI'Si. for olhcr allodapine bees H hcuitri iMaola <t ttl 'liiruim'" was used lo describe a change of IM2), />'- MLWii (liatra ft ul. IVOii ami /•. i»n>h» directum in the nest. Turning involved curling l|lC "Mi Im. ,v Sdtwaiv l'^3). Ilurmg cnardugj Ibc bcc body and sonu-isaidiing. icsulting m Ihc bcc facing was inaclivc citheron ils back or standing upright, If the opposite ilireelton. Both Majors and Minors a bee was closest to anil facing the nest enhance, it appeared lo lurn with equal case. This behaviour was mn recorded as guarding, since bees in ilus occurred anywhere in the nesr. unlike that in position would often be III ihc process ol leavim* sy BKllAVfOUK tN AN AI.LOHAI'IM; BIT *J the tu'si. Minors wore tyfl&n scon guarding and ir< other's metasoma 01 lace. II 'individuals were lace some nests, Majors, pur"ieulurly egg layers, were not to face the two individuals tapped each othci seen t« guard ill all. antennae, Guarding Jul HOI always occur near the nesi enhance, In some nesls the "guard" was ^tutiopifd rvssisa; '/4-1/* »>l ide way down the nesi but was Hit- bet "Passing" is the exchange ol positions by nesi closes! 10 Hit" nest enhance. These guards were males. Passing occurred when individuals Were sometimes seen 10 'pair;*!' (he nest from that section either facing each other or the "passer" was luciug up to the entrance. This involved the bee rapidly Ihe mclusoma ol'lhe individual she intended lo pass. "travelling" forward, whilst rapidly antennaim-1 In each case, iiuhwduaK oriented (hemselvcs \cn ("inspecting") the ties! luiuen bcfoie leluntiny to Ihe ter lo-venler. csscnhally walking over each other. A guard position. In some nesls it also appeared thai pa>s was either simple or complex. "Simple pass Iwo individuals would guard alternately or one in ing" involved ihe smooih exchange ol positions, front of the other Although ihere were times when with individuals usually llaltening their bodies moiv than one individual was seen in Hie gUtitil a;jaiusl Ihe nest wall. "Complex passing" involved posilion. (here were individuals who never imc individual foiling a) anothei individual's Inm I "imarded". Owing ihe study, wo othet invctlcbiate-. pans., and/or simygliny and yrasping each other wore observed entering the nesls. Since there was no with Ihe logs. Hi her one or both individuals would l inleHcieucc liom ttthitr inverichrale predators in thr int.- Sometimes one iimIivulual MajM bite the othci captive situation, guarding hi 'his study may not on Ihe venlral side between ihe metasoma and lite rcl'lcel nalural behaviourof this specks. thorax, near the articulation between the tiochaniei and the thorax. Passing sometimes involved bite! INSI'U'JINt. "buccal contact" between die iwo individual-,, This behaviour involved a hoc abcrnalrly aim\t\ although il Was olieil dill'icull lo determine clearly ualni;: ohieets. lor example Ihe nesi wall or whether buccal contact had actually occurred. Ii brood. |yjg>i were frequently anleimated in this was noi always easy io distinguish between lhc way, Sometimes bees travelled up and dow u the pas-.er and the "passed", except when one was nest inspecting the lumen wall Owing ihis hehav initially stationary *m*l anothei was travelling iour Ives moved their heads slightly and rapidlv omm moved antennae. MiK-VAL l r Individuals were olteii observed to much each MOVING lUIJRIS other's open mandibles with their own open Debris in Ihe nesi was moved by passing it mulei mandibles; this was lermed "buccal contact". When ihe hody with ihe forelegs to (he hind legs then individuals were involved in such interactions, one pushing backwards with the hind legs or tnelasoma individual was sianding upright and the other was This behaviour was nuv iappiov, (),>'; of the positioned upside down. Individuals also cn^a;-ed ill obseivaliou tiiuei. since the nests were in haul, brief huceal couiact during passiue. Ilaritt^ liiic-giained wood w lich required little mam- approximately 5% o1 buccal contact inici'aclains. U-nancr. Debits ohsctved in the nest included neelar flow between the ni'.»u(h parls ol individuals exuviae and. occasionally, dead uulividuaIs was observed and individuals were observed placing "Moving debris" \\t(s not usuullv observed utiless then proboscis between the mandibles of another temperatures were 3 2& ( individual. lVoHennv ol ydobulesol neelar(Media & Sehwar/, \fy)M, was not otv.cived m E, liii/ctihiia. Inti k'- \i)t i r Ht iiAVioi r;s k\ MfM^l W \N1 i "Nudginj:" involved one individual tisinjj ils face 'Avoidance", it combnalion ol oilier behaviours, to nudge oi hull Ihe melasoma or lace ol anolher involved one individual travelling towards another individual. The bee that "nudged" was usually individual and "auicuuaiing" cnhei ihe metasoma upright Nudging usually resulted in tme ol' the 01 lace ol'ihat individual and then suddenly hackim- followjne: away Of milling and liavelliuv it. the opposite at The nudyetl individual lurncd and the nudgci direction. rctrealed. which someliiues invoKed the nmkvd bee opening ils manilihles. \\\) NfSAI CUNl Ml b» If nudged from behind, the bee being nudged "Antennal contact" accompanied most inter-adult would sometimes posilion ils antennae laterally behaviours. When an individual came in to contact (out lo ihe side)..Ihen il nntlged again il might open will) another individual n either " mtcunaied" Ihe ils mandibles Thi> eventually ivsulted in ihe bee N> A. S'lLLN & M P NCIIWAR/. turning, investigating the "nttdgcr", and then simple MOVINC BROOri til complex passing ami/or buccal contact Older brood (laic mslar larvae, prepnpae and O The nudged or the nudger passing and "biting" pupae) were usually moved in a w;iy simikir to Mil cadi other. way debris was moved w the nesl. In /:'. tridentala. similar to fi hi' okV (P, S. llurst pers. eomm, l()95)- MANOtm I.ATIM. the bee initially held the brood with Ihe fore tarsi Mandibulatiug, i.e. the opening and closing of the then passed them under the body and pushed them mandibles not associated with eating, appeared lo backwards using the hind legs. Repositioning of occur before biting encounters. In some cases a brood occurred ollen within the nests ol fi irnhnfalti. appeared that mandibulaling was a signal that one Sometimes a bee would move each pupa until il individmil was rejecting an approach from another reached the end ol the nesl. then it would move thetii individual- lor example VV approached 'B\ 'A' all back again; seconds later another individual nudged 'B , K then opened mandibles, 'A' thru sometimes did the same thing. Some (Minors which retreated. "Biling" encounters sometimes fallowed. consistently stayed near the brood were often Similarly, il mi individual was nudged From behind observed perlorming this behaviour, In addition, it sometimes opened its mandibles and/orturned and bees sometimes simply handled the pupae with the laced tbe nudgeroften opening the mandibles again fine legs but did not actually reposition Iheim In addition, flattening ofthe antennae lateralis often i»uurred during mandibulaliug. This sometimes i.KOUMINri RWKHl occurred when individual* eame face to race or tf Bees tKcu^inialU extended the proboscis lo the one was nudged from behind. brood or bit gently al the brood wilh theii mandibles, such beha\lour was categorised as BIIINti "grooming brood". This behaviour was rarely In this study aggressive encounteis were observed observed. Grooming may have occurred during tot /. ttidt'iihtni. These involved biting laf mourn: or wilh handling but it was difficult h» mandibles, anteniuu . neck, legs, ihe ventral side of ohserve Ihe liner movements of such behaviour the thorax., iiround Ihe eo.xae ami melasoma. Often because ol die speed ol movement U the probosus when one individual tried to eseape from such an and the limited magnification. eucouniet the oilier bee would pull it back using its forelegs- "Biting" cucoiiuicin were often complex. oVH'oMIl"-.' Fur example. '\ fc&cd it* ftlCC IP midge B s lace. When "Ovipositing'1, the female ortenled hetsell so Then one or both bees opened the mandibles ami .1 drat the head pointed mwauK the nest euuame complicated pass followed. Whilst ihe bees were IHiring egg laying bees wereobserved in one ol thtee venter 10 venter and struggling 1holding each oilier position'-.: ventral -airlate lacing upwards, dorsal with legs) "iK would bite the othet on the ventral siulace lacing upwards and laleral surface lacing side ol Ihe thorax. Alter ii struggle- lh» bftlCJI bee upwards, Prior 10 diui during "0\'iposiiion*' Ihe slim- was \)W'\) observed on its back while Ihe bilei held Wis extended. Once an egg had been deposited the othet bee's atiicnutie in its mandihles. in a "lug dte floor of the nesl. the hec reiiactcd the Hil [\\ wat" eueounier. This tug-of war could last hn ,\ppro\intalel\ inm passed before the female 10 JO -ec. I'oIIovmh;- tl me ol Wfll ^TViOHIlW me mined around andI inspected theegg with the antennae. individual w hich hud initialed ihe imv (tin- buiem Oviposilion occurred close lo Ihe nesl \^\ KJ .1 sometimes attempted |vi pUKl n^nn and ollen tj mini mid was observed lot 4 Majors and I Minor (*> simple pass would follow. separate colonies) Individuals took approximately \ ft mm to discharge an cug Howevct one Mn|>:r Aln I I im« INIJ.KV tK'NS look \X nun 10 I,in .||1 1 \ \-.\.liln.\nM.iItUN <M1»tIIbKUr'o|o|dI was LlCC I'li.hed wilh ihe Discussion antennae, and to a lessci CXKM |hfi mouth p.io-. (openmy and elosme mandible-, on 'In- bl'OvXi) Ueliavioni lias previouslv been stuitttftl in dentil lor Individual'- Itfpptal pupae, larvae oregt-s. with each ft. iwwiiii (Maela at ill W91). /<. mishi, it. ktlttm anieunn Mlaiia tt ttl. iWh, A{k*firtpp <\<.tr>itm iSnand) ( VI..son 14>XH) and I hu tthn (Vlelni.V Si h;» .-\ NI ixa'-ai iii-amo lyy?}, B.Kunttm tthfcnttihi gC(WVi\) ^pv^pul jj t»fu* Hioo.l wvtv \otnelimes nudged heloir Hiey Wiu amonul ol tune inactive, siiiukit tooih.i In es (M,i.-»,. moved tjUl Mlttt ioUl did lift FC*Ull llOWCVfl nn! \W2\T\\\\ru*fiitti l('°M Ae|i\-tv tended to be Ihi brood appt-M iHv cbam:iiig. position gteater on davs when the lempei.itnrcs weu aboxr Ue'HAVIOL K IN AN At I ODAPINi: Ul.i; fl| 2?> Ci SinulaiU. when (he1 temperature was wanner mcJuded stuiymi! iliaira <t ol. \&$$\ Mason I^NKI, bees lcn<Jcd to forage more and. especially aficr Auonislie behaviout between ;t host and its paralle- iviuin of a forauci to Ihc nesl. general activity ls not uncommon ,vn\ often results in either host 01 appeared lo increase parasite heme removed bom Ihe nest tBatra cl at. Exec ttni uui,in,ihi was. nut observed lo exhibit ISWl). \yyressi\e interactions are also Icumd in tin: Lvpcs »>r ucM mainteitamJO behaviours found in social species ol the bee tnbes Halidmi ami nfhorallodapities. probably due lo Ihebard nature of \stoc.ipim flkced it til. I07S; Michener \WU), die nesl substrate Most allodapines excavale their However. /:. {rithniutu was «d'ien observed to own lu'sts in pithy simstrate material., wheicas /•„ engage in aggressive encounters which involved a in<fr,rtui.t do not. Alihough observation nests greaj deal ot rtidng and Mniggflrtgi wilh some provided no opporlunity \\x nest walls to dc^av encounters becomiiiy quite savage. Such encounters dming the course ol the study, natural nestr are were otieri picceded b\ imdLMU" and lollowed by also unlikely to rei|iurc tepuiis W MlC nest wall or passing The mandibujatmy dial occurred sonn miranee, since Lbey alsooccur in line grained Wood limes, either prior lo or in response lo nudeiue and "Jilts contra^ with /• hu olttr which pcrTovms biting. Ini^lil also be aeeacssivc in nature Tanc \ \anous nesl inainlenance activities such Etf Michener tll)K.^) toutni that some r.\"i<cnfti spp. cleariur ami larnpini' (removing loose- material produce iriitanh which elivti vigorous gniomine from llic nesl wall and shaping nesl lumen). responses in prcdaioiy ants. Halra ti w, |Iw3) csictnliHg ihe nest lunieii (excavating rear ol the descttbed mainlibulalinj.1 dtinue ayy.ie^sion bmiow). eollat cunsltucti*>n (Uiu)ptng wood into a between ft. uuMn and its social parasite ti. ttufniRn uillar neai nest entrance) arul removing debris and su^^csted th-il manLlibulai vectviioris wea (wood strand^i l-.wnaim irt/lvtiftitti may exhibit involvctl. Il may Iheivfore be sii,L'.L:csted ituu v\licn / nest maintenance netivlics ttj a ereatci debtee when rtfili'tuuiu mandibulaic at each »i!hef, Ihcv also nt'iv msis are tounded and Lheie is a iKVd to remove release clicnucal secretions which may be ucoriNic bass kit behind by beetle luiwic. or relay information about dominance sUKis. Ihis sludy is the firs! lo descriheegg laving to an Thcayonistic behaviours observed m /:. trkltlifilHi Iwotumtt species. £gj« laying ^as only ohsctved sii-ecsl that domiiiansc hiemivbies may be pieseui durum, (he day alihomdi it ina> also have occutred within colonies II appears that sonic lutlividuals u.ght li'bsetvatnnts were only made during ihe enea^e m certain types nfbehaviour which could be •jl\ day) Leg laying wassimilar lo thai described ha />'. interpreted as assertion ofdominance. Bees that are tnuttb (Kara vfttl- IWmirI t> /irnnVnMnci.i | , if/. often midyed or hitii-n -iinlthose thatexhibit avoulanee \W>2\. However two ol the three /:, ttitlrtiiuhi behaviour may have more suboidiuale toles in l|n- majors thai were observed ovipositing were rarely or nesl. Differences in the way individuals icspond to ticvei seen guarding. Che thud fllthUI was seen U\ othei individuals 111 terms ol these behaviours mas y_\u\v\\ luil she was usually 5 em from the base ot (lie be iclated lodominance (i.e. when >ome individuals not and nut near the entrance. This differs Ironi l< arc nudged lhe> euya^c in a simple pass, wherca.- lu'ivttii (Maeia <H tii- IWi and I ItnnUn whenother individualsarenudgedand/orbiileti UfteJ (llogcudoorn & Sehwar/ WK: hull .-/ ttt. in ptess) eu/aee m acomplicated pass). Brothers lx: Michener wIn re reproductive dominants are guards Rjig I Iv74t found lhat qneens' id Ui*it/%ltt\\tM\ laying In this species appears to be a vers \\iw\ :sf>fivf(ii>t ^cK' die maxima! nudyers in the colony processcompared wul othei bees (58 sec. H. In ivif T'he> suggested thai ninigfn^ behaviom indicnlc ti) (Maeta r< «// IW/. in term-, til" both the time dominance siniihir lo thai observed in other primi- taken LO depos(| *fl ry,:j ;nul the frequency ftI tfgg tively eusocia! wasps ;urd bees Hioiheis X laying, Oik- female, in pjiOLulac spent 3S mill Micheuei (l')74i cxperimenlalK showed, lot /. depositing an fcgg which ma> ha\e hecn related td _t{>l>vnw!. that nutlymy by ihe queen plays a role ill Hie 1'act thai Ihe temperature was tow ihul duv \< thedivisionol'labourauiouu the workersby inhibiting .'0 ( i. and bees were eetieially less active at ovarian development, lower icmpetatuies However, these obscivaitous During this ,>mdy ^uardine, behaviour wa> not llic <lid not eo\er the period ol maximal ggg prodttelMHi same us thatobserved in Held studiesol h >ritkiitahi. and slundtl be treated vvith eautiim. i.e. with (he abdomen curled anil used lo block Ihe Ai!.yiessi\e 'u-havioi i has ma been reported ftw entrance bom predators such as ane> 11hirst unpub.i othet allodapine bees except iarcl> bnwceti /; This may be related lo the I'ael that there was no nu\hi and its sivial pufitsilcH. htliasi**-(Halta i I <'' piodalion prepare- in die shade house environmeiiu l'*o :a and inlreLiuently lot A, &-\viiWHi and />'. fmvithi unlike studies 011 /•.. bfytyfai coudueicd in shade (Mastnt IVSX). The agonistic behaviour described houses where anK were a problem *Mli1M;1liustrj loi these species mainly consisted ofnudpiiiL!. buine However, females lhat were •juardiii'j ueie alw-ivs t»l legs and hndies an.l blocking passa.u^ bin alsi: facine the boitom ol Ihe nest which su^'cMs rh.ii f>i / STBKNa M.r sciiwak/ they wan* in a position 10 block the ncsi if the need all similui lothose lound lorotherspecies, su;jeesiimj arose. lhal such behaviours are likely lo be ancestral and I lophallavis to altruistic behaviour. foraiUTN that development ol' novel behavioural elements is ungate in energetically cosily ttitd risky behaviour Id not necessary Tor social ot^amsalion lo evolve bom obtain tooJ which they relinquish to others. small family groups,lolaoiegroups with uioipholoyical Trophulluxis is important in the social organisation ililleieniiahnii amoiiL! colony metnbers. ol many social insects (Wilson ls>7! ). In allodapines However, unlike oilier allodapines. I triJathtta llictv may he differences in the way in which exhibits frequent and overt agonistic behaviours ttophallavis is performed. £xfiWtiUM ht<i>l<>r have auKtin; nest mates Such agonistic behaviour has been observed to engage in solicitation behaviour often been associated with more prinitlivcly social lietore Irophallasis occurs (Melna& Sehwaty l*W.H species, According lo Wilson's i 1971 1 criteria, E Sulii,ilUlion mw>kcd individuals rapidly slrokiny triifi'iuttrn can be classed as highly eusocial because each olhci's antennae prior 10 huccal contact. dierc is female morphological dimorphism as.sociaud Trophallasis in E, hia>io? can also involve one with reproduclive division of laboni Ihctcloic. /:. (inMdeilvniadua&l pSrocfhfvcrviatiijv.' Wa$gBl|o,bull'erototletilui,qiu>idoltogalnoobtuhleesr nthualtlntaHghrtedsosesino'nt cwointhlionnualcooWliolnsyoncsan( 1b**e71rtespul;a±cueesdliho\n was not observed in £ IrittwUtiUt and ifsolicitation "penile despotism" as sociality involves larger jjtoup occurred, il was loo last to be identified. However, tl size and requires a greater decree of intCettlUOll is likely that individuals which enc-aged in "blk'etiJ Most i>ihcr highly eosoeutl species display diMinct couiael" where nectar flow was observed, weiv moi'phs which are directly associated with diseivle Irequcntly emj.aimj in tiophallaxts. Ttophallaxis bchavioural castes, involving minimal ornoagression allows ietuales lo Iced without leaving llie nesL. The Considering lite presence of aijeressive interactions presence ol trophalla*is in li tmlrntata thereioie within /.. iri</cnnifti colonies, it would seem (bat allows behavioural specialisation where only some taCt&JtftfCl colony si/c and the dcvclopmciil o| moi ol ihe lemalcs have lo lotape and other females can |tholi'tgiiii1 dilferenliation amone colons members perform oilierduties ill lite nest. need not be accompanied by decreased levels .-! f,\cnrurn iiHiciiialu exhibits asimilarrepertoire ot overt intra colony agression. behaviours lo other allodapines (Macta ci nl. WW; Hana «•/ at. IVW; Melna & Sehwar/ 1993). Arknowlt'dijiiu'ni's Hcbaviours reeouled in this study, including adult adult interactions and adull-biood interactions, are We would like to thank friends and laboraloiv members who assisted with field work and N. Bull. 1 lIiJti aMn.ANn.Ot-I.ahIaMnW4A)lloI.'tuaspoiunaelihloyc,mitfakWltiCelattilVilaMnidwfpoolp'ulStaltliilolnl Sr.efeRreeyeess.forI*.adIvluircsel.onJ.thettimradnuasncrdipttw.oThainsorneysmeaorucsh (ApislUtt. \vIneopnuH11. BSl (lions) thesi*. I linuViy was partially funded by grants from Lhe Australian 1 IaI'Inl*likvAemrussitlr)ValoifaSnSouA[tlhlWoAtdliustpiiaRnlekepirloluduuuop.cuht)i/..vetythhif"til'nUrchhiufriSth'iun DRUeIseatrvfcthh Tto>ncrnieudisslioonM,fFr'.oSm. lIihceldSwoourtkh wAausslcraarhraiietd Strutli (Amliophoinlae, XyTOvOpMflC}, US' illnnM Dcparlrncnl ol linviioiimenl ami Natund Resources, iin'M', I a fwihu I Into' rally innimh), permit no. Q23256-fljf. issued lo M. I'. S. KWi'UMCCS A!ilauiFHinu;,anihtnairs,pSil.itMu.1iu.\iVmC^1rnli|W'i7n+tth1lVeKi.kiaI/)hhnL>dMh«ii<tatmVn\M.il<i>a<iSlHnltofVJltKl.u.pi2uiSd3iKJ7tl\j\-)k<:\>w-7't.»'i-lUlimVduKitiM[vii|'fcMiiJusi1 Iti(.I1.Jtn1aianlsiHVii.imaciJSr.t<,e.eMMWittA»nhiMie/Ui.s.epJiAi>V.i'In.ettli-.I^J.hN'iloa(vKjpiiinvsaio.Alp.ih'rsiicainlvi...jlWY\.\t.iorvJpfiiuf.-.n."ti^'JvlsuyM.A'oi"n*iH,> (ktltlkvHngiat,nopateisao:ciAarlitlp1oaprla1Mi>treidnein)..U/k-K}iink-wv.I\sliit/lhnHuntmmS",m'- CvnS»ir., ISfI(I,-tS,\vltiMpks fufliisiiv. I' t) i l<WM Itcmistiy and GivAo-iloiHMstic11 BSciflULLL-v.-miaounh. TrsoVcIi.al$ 1iUnl(Ieia\eVuunJs..i p)a7nKd) (lhlSnlBlMlii.cfitVio.lt'i..Ii\5Uo_i\hisiim-ittasiin;Hliiv,linnielaMiop;ijeliaan:d Apnri.lKiloupchlsoioildaeheii;/..oClhntiun,. /K/.KlfKtAWM.nVjrSatlxH'iPi'n-.em/lVi/s/avl>i.o2n5,ini5a|pMriUitHiivelv etisocial l^-e ll<»i,i \oookn. K $c S' uw\u/. M. J' tin pivs.vi Ctiiariliiii- Hroiheis, H. I, £i MitliL-nn. <\ I) lll>74) Inieraeliuns in spatatisalioninptv-ivpmducuvi M^lonu^ot iliealloLlapine loIoiiii:s iif potitiliselv sot-ial hc*?« III F.llinloniy tl| bet h..\<>ntitr<i hit >*!", Kitti/l t.t'l l:\.>l 10 idilvlivsiinocnnoptnc|ra (Haallliowc*liJaion. 7L.ilfSti»MNpg,lPMlJWhi\,i\'<'t:tlpHhlvmPiUn llohohtMVOSnt liFt.ehlU{Wh|ceMl7i?n)sNrcivltimlll)l;ttm»loMliocvh>ei"t'ei,.l,,/r/L'rm0Mv.iHlhf\t(JSJVI'f.t! S l/n/_ 101 I I ; BEHAVIOUR IN AN ALLODAPINE BEE 63 Hurst, P. S. & Schwarz, M. P. (1996) Morphological MiCHENER,C.D.(1965)Thelifecycleandsocialorganisation differences among females of the eusocial allodapine ofbeesofthegenusExoneura andtheirparasite.Inquilina. bee, Exoneura tridentata (Hymenoptera: Apidae). Univ. KansasSci. Bull 46, 317-358. Proceedings of the XX International Congress of (1972) Activities within artificial nests of an Entomology, Firenze Italy 25-31 August, p. 413. allodapine bee. ./. KansasEntomol. Soc. 45, 263-268. Jeanne, R. L. (1986)The evolution ofthe organisation of (1974) "The social behaviour of bees: a Maewtoar,kYi.n,sSoacikaalgainmsie,ctsS..MEon&itMokrehktnooelr,,JtaCl..D2.0,(1191992-)133. cUnoimvpearrsaittiyvePresstsu,dCy"amb(rTihdegeB,eMlaksnsaapchuPsreetstss)o.f Harvard Laboratory studies on the behaviour and colony (1975) A taxonomic study of African allodapine Tstariuwcatunre(oHfymBernaoupntsearpai:s hAenwtihttoip,hoariXdyaleo)c,opUinniev.beKeanfsraosm- bees. Bull Amer. Mus. Nat. Hist. 155, 67-240. Sri. Bull. 54, 289-333. Schwarz, M. P. (1994) Female biased sex ratios in a Mason.C.A (1988) Divisionoflabourandadultinteractions facultatively social bee and their implications forsocial i(nHyemuesnoocpitaelrac:olAonntihoepshoorfidtaweo).aJl.lKoadanpsianseEnbteoemoslp.ecSioec.s evoluti&on.OvEevrolhuotlito,n L4.8,A.16(8149-9136)97M.ethods for rearing 61,477-491. allodapine bees in artificial nests (Hymenoptera: Melna, P. A. & Schwarz, M. P. (1993) Behavioural Anthophoridae). Aust. ent. Soc. 32, 357-363. specialisation inpre-reproductivecoloniesoftheallodapine Wilson, E. O. (1971) "The insect societies" (The Belknap bee Exoneura bicolor(Hymenoptera: Anthophoridae). Press of Harvard University Press, Cambridge, Insect. Soc. 38, 1-18. Massachusetts).