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Field observations of the behaviour of free-ranging eastern barred bandicoots, Perameles gunni, at Hamilton, Victoria PDF

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Research Reports Field Observations ofthe Behaviour ofFree-ranging Eastern Barred Bandicoots, Perameles gunnii, at Hamilton, Victoria. Anthony C. Dufty* Abstract that the recording ofunique behavioural A total of 23 individual and 10 social acts often increase considerably with in- behavioural acts were observed in afree- creased time devoted to observation of ranging population of Eastern Barred behaviour. Thus, less frequently-ex- Bandicoots at Hamilton. Individual be- hibited behaviours may not be observed haviour involved acts relating to body withoutconsiderableeffort,anditislikely posture duringfeeding,groomingand in- that the complete behavioural repertoire vestigation. Mating behaviour was of P. gunnii is yet to be revealed. Hein- promiscuous. Copulations between in- sohn (1966) in Tasmania, and Brown diintveirdmuiatltsentlwyerfoerupratpoid45 mainndutesr.epIenattreod- c(1o9m8m9e)natneddDufbtryie(fl1y991oan)inoVbiscetrovreiadhaivne- missions occurred when the female dividual and social behaviours of P. carried advanced pouch young. Many gunnii but the only systematic research males gathered and mated with females so farreported is that ofMoloney (1982) odlufraicntgiotnheiirnrtehceepltoicvaetipoenrioadn.dMpaulressuiutseodf 6a5ndhColuurnsieof(1o9b8s7e)rvwahtoiounndreerstpeocotkiv1e6l5yaonnd receptive females. Mutual avoidance be- captive bandicoots in Tasmania. haviour was often maintained between This paperreports on fieldobservations bandicoots although several antagonistic atHamiltonandassistsinthecompilation interactions were observed. These inter- ofa behavioural reportoire for free-rang- actionsusuallyresultedintheflightofthe ing Eastern Barred Bandicoots, Pera- subordinate after strike, chase and/or meles gunnii. Also, the paper discussed threat vocalisation occurred. the implicationsofP. gunniibehaviourto Introduction the management ofboth the free-ranging Although relatively widespread in Tas- andcaptive populations in Victoria. mania, Eastern Barred Bandicoots Methods hcuaravrd(1PainegnoeS9aeLvoailidSei8mnrniinctle4ngaontatsbp;sSeaumlsnreoeoerrnttiorccnBaeftcsfdikiledriaue,nxpslcoffs(ftlterew1ioasdbsetn9Sirnteeacn9ogmaut-thts0aunlSdori1aHt)ni(oa9va1inesnnS^8im9oise,ger9o9inivieme)0uletnbavrh;rrthgiee3aiaioaenDlcgSennrnomaukefw,sldalrfbpeaTiodV1ettednaacnii9hcceysdsaclne9rinttmgiase0a1itoeas)ns9oPirivpndsnc9oineheAiasa1alrdcaulafpaaiiilss(oeo)lvysetgnMpishrbuudoaaeTebaultlenoodheerahaind,n-nnlede-a- dgHhadbsCmb1iaroianpioe4cOtcoumodthn2ottboryiuatoelo3ssln0vtrihte0td2ygiolrwnoh'ofbySvweatneueatsashErshtpHptMs.apaiiaotuvotlnomewnuitAogniwmioeslgslouiecebargtvgrdiren1eeoeho.perd0ddntfraa0ti,Alfl1bbnioon9WgyenorituT9Vatdokga0nisiaittwhge.depcavtteidsetli1Anesorwd2tgoqinrtuaoooiufraaVshpnaoaal5kao4tsbirebul2isstJina(tswemzetu3neidsa-ral7gtddplthvye°aohaedl4rtitaslu5elton1bobotri*a9catayggiiti8nhonnpehnSha-e9-tfe-ngel uFVr.agcgetenontnaannadeneGddolcodfnomrcalnud(ed1tea9di7lt7ehda)tdtienhmfeororenwTSaatrsodann distouiccaittaeeldboiebmhsmaeveridvoiarat.itoeOnlbsysearofvftaertiidnoudnissvkiwdaeunradlewceaornne-d 3SST2 Lrment ?* De<~ * couotnfoifnesdigthoto(nuseuailnldyivbiedutawleeunnti2l-3ithmoourvse)d, bandicootsappearedtobeunperturbedby thespotlightandtheobservers' presence 54 TheVictorian Naturalist Research Reports That is, no escape behaviour that could Table1.Individual behaviouralactsobservedin have been attributed to the observers' free-ranging Eastern Barred Bandicoots at presence and no erratic or unusual be- Hamilton. haviour (excessive grooming or sniffing Act Description ofthe air) was observed. Information on Quadrupedal Manusandpesbothrestingonsubsiraie.body behavioural acts observed and general rbaoidsye.dandheadslightlylowerthanparallelto notes on the frequency ofthese acts was Tripedal Asabove,hutonetorepawretractedtobody Crouch Completemanusandpesrestingonsubstrate, recorded. Other information recorded (if backarched,headraisedandforequarters possible) during encounters included: in- lowered. Asforcrouch,buthindquartersrelaxedand dividual identification, sex, ageclass and headloweredtosubstrateandmanus reproductive state (presence/absence of sometimestilledtooneside. Rear Hindlimbsincontactwithsubstrate,head pouch youngoryoung at foot). verticallyextended. Sit Asforrearbutheadnotextended Walk Slowquadrupedallocomotion,cursorial Results motionofpectoralgirdlewhilesaltatorial Individualbehaviour motionofpelvicgirdle, Run Fasterquadrupedallocomotion A total of 23 distinct individual be- Gallop Rapidlocomotion,whereinforelegsare havioural acts was observed. They are rpeotwrearcftuedltsoimbuoldtyanwehoiulsethhirnudstlse.gsexhibit listed in Table 1, and described using the Leap Verticalspring(upto1m),usingsudden terminology of Moloney (1982) and Hook eLxotuednsgirounnto,frheipnedalteegds,upto5times. Clunie (1987) as reviewed by Coulson Nose Lateralmovementofsnoutacrossthe (1990). Three behaviours ('bipedal sta- Dig Ksuxbesatvraatteinwgitshubasutdriabtleewisntihfff.orelimbsand nce', 'prancing' and 'climbing') which insertingsnoutinhole. were recorded by Moloney (1982) and Peed hIenlgedsotninsgubfsotorda;telawrhgielietesmmsal(lee.rg.inteecmtsa.rines) Clunie (1987) were not observed during (e.g.insects)aremanipulatedwhilesilting Manipulate Claspingandrotatingitems(generally this study. food)withforepaws. Foraging areas at the Hamilton Push Hindlegsandtailincontactwithsubstrate. forelimpsbroughtdownandawayfrom Municipal Tip were associated with hard bodyduringextension. shelter and located in a patch arrange- Scrape Pullingnestingmaterialsbackwardswith ment. Travelling within open areas Scratch fRoarkeipnagwsm.ovementsofhindfeettogroomfur between foragingpatchesoccurredrapid- (especiallyheadandneck). ly, and foraging was observed only when LWiicpke RLiucbkbiinnggatnhdecshnoeuwtinwgitfhurliocnkebdofdoyre(peaxcwesp.t individuals were within 30m of shelter. headandneck). Shake Vigorousshakingofbody,sometimeswhile Foraging ('dig', 'feed') was maintained inmotion. forbetween 5 and 25 minutes in a single Streich Forepawsextendedunderhead,body elongated,accompaniedbyyawning. foraging patch before the bandicoot's Rest Lyingonsideinloosecoil. *run' to the next patch. Bandicoots ap- peared to search for food in a random movement, although two individuals underthebodyduringdigging.The snout wereobservedfollowingfencelinesfor25 probedtheholeperiodicallyandasniffing and 17 minutes respectively. Nosing the orsnuffling noise was audible. When the ground with a lateral movement of the food item wassecured, the individual fed snoutandloudsnufflingnoisespreceeded immediately, while adopting a quad- allobserved 'dig'behaviour.Diggingwas rupedalorsitting stance. Ahigh foraging similar between all individuals and in- success was observed and often ban- volvedthrustingtheforeclawsintoavery dicoots would probe and sniff foraging localised area ofthe ground quickly and holes that had been dug on other nights, alternately and withdrawing them up- sometimes enlarging these excavations. wards and backwards. During digging, The excavations made during bandicmoomt the hindquarters were raised, the backar- foraging are conical pits, up to 80 ched and the snout pointed downwards deep. It is possible that they trap small towards the excavation. No attempt was invertebrates as Mr N. Gunn (pers, made to remove soil which collected comm., 1989) has observed bandicoots 55 Vol. Ill (2) 1994 Research Reports searching and removing invertebrates Aggression ('chasing' and 'striking') from foraging holesdugpreviously. was observed on three occasions. During Occasionally during foraging a ban- one interaction, the pair of bandicoots dicootwasobservedto 'rear' inanupright faced each other in the 'arched' stance stance while smelling the air. When un- with their mouths agape revealing their usual stimuli were encountered, a incisor, canine and premolar teeth. Eye 'bipedal' or 'crouch' stance wasadopted, contact was maintained between ban- often followed by a rapid 'gallop' or dicootsandafterabouttwominutesinthis 'sprint', On three occasions after these posture. The subordinate then slowly responses,thebandicootwas observed to turned to move away and the aggressor 'leap1 upto 1 mintotheairandthen 'run' was observed 'striking' the subordinate away in the direction it faced upon land- on its hindquarters. The subordinate fled ing. Thedirection appeared tobe random 'honking' with the aggressor in 'chase'. asallthreebandicootsturnedinawidearc Onanotheroccasiona 'chase' hadentered of up to 180 degrees, passed cover and the area where a third bandicoot (male) returned to shelter, close to where they was being observed. The 'chase' passed were originally. the third bandicoot within 4 m and he Drinking was observed only once. The responded by pursuing the pair for a dis- bandicoot spent four minutes lapping tance of about 20 m. The male then waterthat had collected in a foldofplas- returned and continued foraging. The tic, with a forward and upward motion of third occasion where aggression was ob- thetongue. served occurred when an aggressor Nest construction behaviour was ob- entered a foraging patch occupied by a served once. Dead grass within 1.5 m of subordinate.Theaggressorforaged about thenest sitewasscraped from theground 15 mawayfromthesubordinateforabout aroundperennial tussocksanddraggedor 30 minutes, apparently unaware of the pulled backwards into an upturned gal- subordinate's presence. However, on vanised iron roof gutter by the forelegs movingdownwindofthesubordinateand (for descriptions of nests see Duftv sensingitspresence(indicatedbysniffing 1991a), the air in a reared stance), the aggressor Fourbandicoots were observed emerg- 'chased' thesubordinate,causingthesub- ing fromdiurnal nests. Movement within ordinate to 'spit' and run for shelter. The the nest increased during the 15 minutes aggressor stopped the 'chase' in the area prior to emergence. Immediately after where the subordinate was foraging and emergence, bandicoots were observed to continued to forage in that area. 'shake' beforetheyranorwalkedtonear- by foraging areas. Grooming ('scratch', okwbiseprev\ed(Hcafkt'e)rbneehstaveimoeurrgweanscemowshtenofttehne Tfraebei-cra2n.giSnogciaElasbteehranvioBuarrarledacBtsanodbisceorovtesd aitn individual wasaway fromthenest within Hamilton. a foraging area. Upon retiring to the nest (observedby MrK. DrinkellorMrN.Gunn). alternocturnal foraging,bandicoots were aobnsdesrmvaeldltsolicmkos)veacrnoesssttmhaetenresitalesnt(rgarnacses A•*FiPeolrlilnoewal PNDeoersssciirnsitgepntpteirofinonlelaolwairnegaoofffceomnaslpeecbifyimc.ale. with theirsnouts. *Mourn Malerearsonhindlimbsandinclinesbody forwardoverfemale. •Thrust Pelvicthrustingduring;mourning. Socialbehaviour *Biie Malebilesfemale. Mutual avoidance behaviour predo- Arched aAnsdfboracqkuaadrrcuhpeedd.albutheadslightlylowered minated throughout the observation Sirikmg Sftorriekpianwgs.conspecificonthehackwith apeeroiondsswbeurteonofbosuerrvoecdc.asSioocnisalsobceiahlavinitoeurr CHohnakw CHhoanskiinnggrveotcraelaitsiangtisounbogridviennatbey.the actsare summarised inTable2. Spit rSeptirtetaitnignvgoscuabloirsdaitniaotneg.ivenhytheretreating subordinate, 56 The Victorian Naturalist Research Reports Notall interactionswere aggressive: on exactly and males were observed bump- one occasion three bandicoots were ob- ing intoobjectsplaced by MrDrinkellon servedforagingwithinabout 10mofeach the female scent path. Up to ten males other for about 20 minutes. During this wereobservedtocopulate with a female, observationforagingwastheprincipalac- threeorfourtimeseach.Copulationlasted tivity and no overt social interaction about 20-30 seconds and was repealed (mating, vocalisation, or chase) was ob- every few minutes formorethananhour. servedeven when twobandicoots moved During one period, R41 was observed to within 2 m ofeachother. Unfortunate- mating with male L8 R53 fourtimesand ly, the sex ofonly one ofthe bandicoots male L61 R8 six times as well as with (a female with advanced pouch young) othermaleswhoseidentitywasunknown. was determined and the group gradually After mating with one male, the female dispersed indifferent directions. was observed to move away while the Two local residents at Hamilton (Mr male was foraging and sometimes mated Ken Drinkell and Mr Noel Gunn) have shortly afterwards with anothermale. recordedmatingbehaviouronseveraloc- Mr Gunn has maintained a captive casions. Their descriptions are reported breedingpairofbandicootsonhisproper- below. ty outside Hamilton since 1984. The Mr Drinkell has lived adjacent to the bandicoots are maintained within 20 m" Hamilton Municipal Tip for many years enclosuresandgivensupplementaryfood andhasrecordedinformationonbreeding every second night. This has provided activity since 1988. Two individual MrGunn with the opportunity forcasual females were resident on his property observations of captive bandicoot be- during the study, individuals numbered haviour. During orbefore supplementary R41 and L8 R7. Female R41 was first feeding, MrGunn has recordedP. gunnii marked on 7 May 1988, south of the mating activity on fouroccasions (9 July Hamilton Municipal Tip when she was 1988 for 15 minutes, 4 October 1989 for about four months old, while L8 R7 was 45 minutes, 29July 1990 for 17 minutes, first marked on 6 February 1990 on Mr 25 September 1990for20minutes). DrinkelKs property when she was also All observations of mating were made fMoruDrrimnoknetlhls'solpdr.oRpe4rt1yw(aasbofiurtst4t0r0apmpefdroomn ediutrhienrgdluartiengafoterranfotoenr,lbieghftorreains.unCsoeptualnad- where she was first marked) on 13 Sep- tionwasinitiatedbythemalecheckingthe tember 1988 and was regularly observed femaleby a 'bite' tothe looseskinonher there subsequently. R 41 was observed hindquarters. As the male 'mounted', the mating with males about every 9 weeks female lowered her forequarters and and was the only breeding female to be raised her hindquarters. On three of the observed on his property until March mating occasions, the female was carry- 1990, when female L8 R7 was observed ingadvancedpouchyoungandduringthe mating. fourth the dependent juveniles jumped Although many males gathered during around the copulating pair. Copulation the female's receptive period, little ag- involved rapid 'thrusts' and lasted be- gression appearedtooccurbetweenthem tween 5 and 30 seconds. As copulation andavoidancebehaviourwasmaintained. progressed, the mail curled his tail under No spatial organisation (e.g. lek) or ob- his body from an initial lateral position. viousdominancehierarchy wasapparent. Betweencopulationevents, the malewas Males spent muchoftheirtime searching observed to 'follow' the female's scent fmoernttsh.eTfheemsaelemsalienmroavpiedm,enertrsatiicncmreoavsee-d cilnogsewliythainnd1hemwaosfohfetren(tohbesepraviredusfuoarlalgy- when more males were present. When a avoided each other during non-mating male picked up the scent ofafemale, the periods).The male wasoftenobserved to movementsofthefemale were replicated smell the perineal region of the female 57 Vol. Ill (2) 1994 Research Reports prior to copulation. Intermissions be- 1980) and it is likely that this odour is tweencopulationsusually lastedbetween central to social interactions. Stoddart 5 and 40 seconds and on 29 July 1990. (1980) speculated that the odour has a copulation wasobservedto take place 17 calming effect which facilitates mating, timesin 17 minutes. while it may also be integral to the main- tenance ofdominance hierarchy. Discussion Mating acts in Victoria described here During thisstudy and others (Heinsohn byMrDrinkellandMrGunnweresimilar 1966; Duffy 1991a) bandicoots spent tothose described by Heinsohn (1966) in much oftheirtime foraging and feeding, Tasmania, Heinsohn (1966) observed suggesting that dietary items sought are mating behaviour once in captivity and either low in energy or are hard to find. twice between free-ranging individuals. Thehighsuccessatsecuringsubterranean Onestrikingdifferencewasthehighcom- morsels observed during this study and petition for receptive females at thedepthofforagingholes(upto80mm), Hamilton.Drinkellreportedthatsmallag- suggest that bandicoots possess well gregations ofbandicoots occurred during developed olfaction. The dependence on the female's receptive period and that olfactiontodetectfoodresourceshaspre- several males mated with the female. viously been reported by Heinsohn Principally, the conservation of Pera- (1966),Moloney(1982)andQuin (1985) melesgunnii in Victoria has involved the in Tasmania, and Dufty (1991a) in Vic- establishmentofthree reintroduction and toria. The observation that bandicoots two captive breeding colonies (Seebeck utilisefencelinesduringforagingmaybe 1990; Dufty 1991b). The artificial nature duetothelackofstructuralcomplexityat of captive breeding may alter the in- Hamilton, although the higher floristic dividual and social behaviours observed diversity and lower compaction ofthese in free-ranging population and com- areas may also be important. The lackof promise the long-term viability of P. extensive fat deposits (Lenghaus et ai gunnii.Toavoidthis,threestrategieswere 1990), aggressive defence of foraging integral to the P, gunnii captive breeding patches and rare aggregations of ban- programme: avoid domestication (either dicoots suggest that food resources at through human association or selective HaAmgiglrteosnsiavreedleifmietnecde.offoraging resour- bmraeneadgieng fionr icnadpitviivdituya)l;s thmaitniarmeizeeasyant-o ces appeared to be the most common tagonistic interactions between colony cSd(sdfsssHtstumotoeeo9erercuarrmiim8aidilacvpnir4atynlgeosln.)egidrwoaernahhangblaDdtbacnvcselhoathboheyepnmaisaaydmsihodtvsi(ieilncoaitSc1inechacnotuto9grehnccd^auo6asryaeutrt6r.ry,spns)c.uytiheiAbwbibysxAnoteaenhdlm(wsrnrhgiitd1uadebavmhi9gisiivon7ogccftdisa7peuhoeuoottr)sgaodarueetshtlarsseasesogdedntdiiunthdfnwetrafegrxweinrraRhotnradonuhimgesaatmdsbrtisietkaetmntoaahheleonerbsailtysedd.-ats bfrmseeptpmefeearhenoeDfieeettplvmenoddeuwidcstbmalsertopretewaseeoomosrticdnnworittshiumifsoihea,ctencdoscntohnsa'ouiadro'tapmotltuatnneaeimbt(i,tsddosieatvt,nlu.retiiigwpaemic.tomcepnyiantpfclrcftmgooteePomoisilom.arsouiocettanstragnrneg,uialuyasteortgtnli)anvheenno.f.tiregddiowvieyTiireeos.tAvfodionfminridofdneatelouoateneayadaron-tsg,lardssnrsogaeasaenenentwamvdgilufaeouaetirtsnricchaenymdetdeerlg (S19U85K) Jardg0uemd!natthnactebuahino^dpaiercrcoatohetys,otloRfuamscastieinlol-n aclnaotuwasegeronirinesjpturrioicedsui,cnttieinrvcaercetoaiusotnepsu,stt.orenTslsoylomevnieenlismmaailnsede resided in each breeding pen, low den- wmch exudes apungent odour(Stoddan jsuitvieensiloefs wienrdievirdeumaolsvedwearsesomaoinntaasintehde,y 58 TheVictorian Naturalist . Research Reports became independent, food was supple- management of Eastern Barred Bandicoot, mented when needed, and shelter was Peramelesgunnii,inVictoria.Unpubl.M.Sc.Thesis. provided for fleeing subordinates. UniversityofMelbourne,Melbourne. Despite a promiscuous mating system Fagen, R.M. and Goldman, R.N, (1977). Behavioural catalogue analysis methods.AnimalBehaviour25: prevailing in thefree-ranging population, 261-274. P. gunnii were initially bred in pairs to Heinsohn,G.E.(1966).Ecologyandreproductionofthe maximise outbreeding and conserve low Tasmanian bandicoots (Perameles gunnii and density alleles. However, individuals Isoodon obesulus). University of California. were promiscuous in the reintroducled PublicationsinZoology80: 1-96. populations that were large, less in- Lenghaus,C,Obendorf,D.L.andWright,EH.(1990). VeterinaryaspectsofPeramelesgunniibiologywith fluenced by genetic stoichasticity, and special reference to species conservation. In regulated by naturally selection. 'Management and Conservation of Small The successful management of Pera- Populations'. Eds. T.W, Clark and J.H. Seebeck. meles gunnii captive breeding and (ChicagoZoologicalSociety:BrookfieldIllinois.) reintroductioninVictoriahasbeendue,in Moloney,D.J.(1982).Acomparisonof"thebehaviourand part, to the resolution ofmany problems ecology of the Tasmanian bandicoots, Perameles thatwereassociatedwithcaptivityandthe gNuontntieir(G1r7a9y7)!.838U)napnudblI.sooBd.oSnc.obes(uHloenss)(ShTahwesainsd. species' individual and social behaviour. UniversityofTasmania,Hobart. The application of behavioural informa- Moon, B.R. (1984). Currentdistributionof(heEastern tion hasaidedtheconservationofEastern Barred Bandicoot, Perameles gunnii, in Victoria. BarredBandicootsin Victoriaandshould National Parks and Wildlife Division, Victoria. be seen as an important component ofall ATretchhnuircaRlylaRhepIonrsttitSuetreiefso.rENnov.ir5o:nm(eDnetpaalrtRmeesnetarcohf wildlife management programmes. Conservation.ForestsandLands:Melbourne.) Acknowledgements Quilt, D.G. (1985). Aspects offeeding ecology ofthe bandicoots, Perameles gunnii (Grey 1838) and I wish to thank Mr Noel Gunn and Mr Isoodon obesulus (Shaw and Nodder 1797) Ken Drinkell for their many informative (Marsupialia: Peramelidae) in southern Tasmania. discussionsregardingbandicootbreeding Unpubl. B.Sc. (Hons) Thesis, University of behaviour and allowing me to use their Tasmania,Hobart. data in the manuscript. The manuscript Russell, E.M. (1984). Social behaviour and social wascriticallyreviewdbyMrJ.H.Seebeck organisation ofmarsupials. MammaliaReview 14: 101-154. and MrG. Coulson. Russell, E.M. (1985). The metatherians: order Marsupialia.InSocialOdoursinMammals.Eds.D. References Macdonald and R.E. Brown. (Oxford University Brown, P.R. (1989). Management plan for the Press:Oxford.) conservation of the Eastern Barred Bandicoot, Seebeck, J.H. (1990). Recovery management of the Perametesgunnii, in Victoria. National Parks and Eastern Barred Bandicoot in Victoria: statewide Wildlife Division, Victoria. ArthurRylah Institute strategy.In'ManagementandConservationofSmall for Environmental Research Technical Refwrt Populations'. Eds. T.W. Clark and J.H. Seebeck. Series,No.63.(DepartmentofConservation,Forests (ChicagoZoologicalSociety:BrookfieldIllinois.) andLands:Melbourne.) Seebeck, J.H., Bennett, A.E and Dufty, A.C. (1990). Clunie,P.(1987).Aspectsoftheecologyandbehaviour Status,distributionandbiogeographyoftheEastern ofIsoodon obesulus(Shaw and Nodder 1797)and BarredBandicoot,PeramelesgunniiinVictoria.In Perametes gunnii (Grey 1838). Unpubl. B.Sc. 'Management and Conservation of Small (Hons)Thesis. Department ofZoology, University Populations'. Eds. T.W. Clark and J.H. Seebeck. ofTasmania,Hobart. (ChicagoZoologicalSociety:BrookfieldIllinois.) Coulson, G. (1990). Applied behaviour: its role in Stodart,E.(1977).BreedingandbehaviourofAustralian conservation biology of the Eastern Barred bandicoots.In TheBiologyofMarsupials'.Eds.B. Bandicoot. In 'Management and Conservation of StonehouseandD.Gilmore.(UniversityParkPress: Small Populations'. Eds. T.W. Clark and J.H. London.) Seebeck. (Brookfield Illinois: Chicago Zoological Stoddart,D.M.(1980).Observationsofthestructureand Society.) functionofthe cephalic skinglands in bandicoots. Dufty.A.C.(1991a).Somepopulationcharacteristicsof AustralianJournalofZoology.28:33-41 PeramelesgunniiinVictoria. WildlifeResearch 18: 355-366. Dufty, A.C. (1991b). Conservation biology and 59 Vol. Ill (2) 1994

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