Corella,2004, 28(1): 11-19

BREEDINGBEHAVIOUR, VOCALIZATIONS AND PLUMAGEOF THE RED-CAPPEDROBIN goodenovii lN CAPERTEEVALLEY NEWSOUTH WALES

VICKI POWYS

''Rocklards", Glen Davis Road, Capertee Valley, New South Wal€s 2846

Received: 26 lune 2002

This paperdescribes the nestingbehaviour, vocalizations and plumageol 12 pairsol Red'cappedRobins Petroicagoiodenovii studied during the breedingseason trom September 2000 to March2001 in 12 hectaresol dry woodiandin CaperteeVa1ey, t':ew South Wates. Individuals were reliablyidentified by ,song and Giritory.four brown-plumagedbreeding males had rust-colouredcaps. Fourteen distinct vocalizations were recognized(1 1 for adults,three for young).

INTRODUCTION Male RedcapPed Robins called often prior to nestrngand were located aurally. Vocalizations were oped opportunistically and The Red-cappedRobin Petroica goodenovii occurs in the asso,riatedbehaviour notcd, from Septemberto March. Sub_songfrom at the end of April 2001 Samplesof drier regionsof all mainlandAustralian states(Blakers e/ al. an adult male was recorded songwere recorded from ail the breedingmales in this study, and conspicuous territorial 1984) but despite being a widespread at varioustimes of the day, and were later analysedbolh aurally and species,information on breedingbehaviour and vocalizations sonagraphicallyto checkfor individual variation. is very limited and often anecdotal.The most complete Voice was tape recordedwilh a Sony TCD DlO Pro DAT and was made by Coventry behaviouralstudy of the species SennheiserME 67 gun microphone,and analysed on a MacintoshiMac (1988),but was limited to just thrce pairsand did not include computerusing sound processing software Peak LE v2.10(O BIAS sonagrams.This paper attemptsto describeand illustratethe 199?-1999)and soundspectrograph software Audiocraph v.3 2 (O completevocal repertoireof the speciesand to match voice RalphSutherland 2000, Canberra, Australian Capital Territory) with behaviour.Adult malesare brightly plumagedin red, black and white but brown malesmay also pair and breed RESULTS successfully.There is confusionin the literatureregarding Teff ito ry estab I ishnte nt atrd sottg the plumageof brown malesand femaleswhich I hope to partiallycorfect. George (1950, l95la, 195lb) and Hutton Eight red plumaged males and four brown-plumaged (1991) studiedcaptive nestingbehaviour. Disney (1974), males paired with females and establishedadjoining Hall (197,1)and Hobbs (1986) have studied plumage. territoriesthat averaged1.02 hectares,in vegetationthat Limited intbrmation on Red-cappedRobins is fbund in was dominated by Motherumbah acacia (Powys 2004). field guidcs (Stewart 1976; Frith 1984; Simpson 1986; From Septemberto Decembermale Red-cappedRobins Boles 1988;Schodde and Tidemann1988; Slater 1989; advertisedfrom prominentperches, giving frequentbouts Pizzey 1997 Morcombe 2000). In a concurrentpaper I of Territorial Song dikerdreeya-dikerdreeya(Fig. la) during describethe 2000-2001breeding biology fbr the same l2 daylight hours until paired and nesting, usually within pairs of Red-cappedRobins ref'erredto in this presentstudy l-2 weeks.One brown male sang intermittentlyfor three (Powys2004). weeks in November Decembgrbefore attractinga mate and nesting. lf a nest failed or if fledglings were nearly METHODS independent,males would again give the Tenitorial Song for one or two days prior to a subsequentnest att€mpt.This trilled TerritorialSong was not loud but carriedwell at a hectare in Crpcrtcc Valley is located 18 kilometes Tbe 12 study sitc (kHz) within a east of Cnpenee (33'07'5, 150"08E) and 380 metres above sea level, on relativelylow frequency and was audible the westernslopcs of lhe Great Dividing Range in New South wales. Mean distanceof 150 metres. annual rainfall is 500 millmetres and summcrs are oflen hot and dry with low humidily. Aspect is north-eas(with stony soils suppoflingdry woodland Femalesdid not give the TerritorialSong prior to nesting, including Motherumbah A.ada .heelii. Black Cypress Pine Callilris but fbur different females used the Territorial Song sn..//i(/re.i. Narrow-leavedIronbirk trdllpr,rr .rrfa, Tumbledown Red- occasionallywhen feeding nestlingsand fledglings(Fig. g'rm wifi an undersloreyof native herbs and gasses. E. duLbata, Ib). One female with fledglingsused the TerritorialSong ll,lat( rial:i u'u| t'l4ltods when disputingterritory with a neighbouringmale. H male and H female had similar Territorial Songs; the female were not banded but individuals were reliably identified by their plumage irregularities and terrilory; they were given a code name and consistentlygave a slighter weaker version (Figs Ia-b). tcrrilorics were mapped with data being compiled for 29 nesting Sonagraphicanalysis showed consistent individual varirtion attempts by 12 pairs, as outlined in (Powys 200,1).Twenty-five robins in maleTerritorial Song which was sometimesdiscernible were colour-sketchedfrom October to Decembcr Moulting, brown males in the tield. The Scolding call heh-heltheh-heft(Fig. lc) were re'skctchcd in January. February and March to monitor their progressivechangc ro full adult plumaBe (Table l). replacedthe males'TerritorialSong when the fem.rl?was

1l U Powys:Behaviour, \^ccalizations and plumageof Red,cappedRobin Corella28(1)

TABLE I Table showingplurnage and moult details for fou.rbrown males November ro March. BROWN-PLUMAGEDMALES BB MALE November Grey brown uppcr, strongly rust-coloured forehead cap, pale breast, whirish stripe on low€r wing. December Unchanged. 2l January 3 red patches on breast and red mark on srdc of forehead, black smudge on head, grey brown upper. 30 January cap almos( comptelc, bhck reptrcrns brown.i r,*,r, u".f""o w,nss, white wins mark Y.:::-11..:"exlen0s to coverts.!:"::,red 7 February Fairly even red on breast.black Dlunase almost complete bul a fcw brown flecks, whire srnpe In wings dcvelop€dfunher. 20 Fcbruary Moxlt complere, strong cadmium red-corour on cap and brcast, dcep marl brack on heacr,back and wings. fulr length wing slripe, ^white shiny black legs, dark beak. AB MALE November urey brown upper' stronglv rusl coloured forehcad cap. pale buff-washcd breasr wirh a doubtc band of fain( pink spots. Obvious whirish wing slripe extends !o coverB. December More faint pink spots on breasr. orherwise unchanscd. 23 Jafluary R-ed cap dcveloping bul still brown sE€aked, 3 fainr rJ smudgcs on brcrsr, somc black flecks around eyes and bclow beak. 29 January More black on face, red cap prominenr, crescent of red on"middtc Ur"".t, ii"V+."*" back and wings, white wing stripe cxtends to covcrts. 3 February a:]lr-"T.:^*::Pttlt of red and black, wirh parchy black and brown on back. head and wrngs, whilc wins slrjpes, rco crp, ano pilehy reu on brcdsr. 2 March Moull complele, strong orange-red breast smaller than BB male, red cap has I brown fleck on it, thin brown ring around eye, rvhite wing stripc comptc!e, dark beak. NH MALE Novcmber O-Y" upper with pale neck. srreaky rust coloured 9:l forehead cap, palc beak. p:rle breasr, thin whitish slripe on Decembcr Unchanged. 0 January Black mrrks on necI and irroundr]e5. olhLr$i.e unchrnged. 7 February r_O.,qh red fe, hers in cap. shght butf wash on ] ::, uppir breast, btack developing on face and wings. 13 February r rrrnr sp'rs on hre.sl red f'nK mirk on cap, mixed black aDdbrown on head,back and wings,no white in wing. By 16 Feb pink spors changed to red spots. 2l February ":mptere. hrtfred. 2 red,muJse, on bred.r. rult tengrh qhire 5kipe apfearinS winrs. 3 March I'rcK3ii:f prumJgcll::li: :,,1-":, (omprcte jr"cap on armos( dusryroohng. red cJp armosr (omprere. red on brea.te.rrnjs ro inrn uir ineomierc 13 March Moult almostcomplete, dark black headand back, orange-red breasr, stjll a few brown sFcaKson cap. MB MALE November Grey_brownuppcr. stronglyruslcoloured forehead cap. pile preybreast wilh 3-4 pink spots,whitish stripe on lowerwing. December Unchanged. late-Dec Abandoncdterritory and mouft detailsnot known.

incubating; nestingmales answered neighbours' Tenitorial December-January after repeated nest failures. Juveniles Songswith the Scoldingcall ratherrhan the Sons. sometimes stayed within their parents' territory after independence, Pai ng while the parentswere re-nesting.Assuming that brown males were less than two yearsold (Boles 1988), Red-cappedRobin pairs stayed together fbr the duration and that femaleswith indistinct or no rust-colouredcap were of the breedingseason and were apparentlymonogamous, less than two years old (George 1950), mixed-agepairings both birds defending their rerritory while they were nesting occurred when older coloured males paired with younger and feedingfledglings. One pair continuedto defendtheii females, and two younger brown males paired with older tenitory fbr three months without re-nesting,following a females (those females had extensive rust-coloured caos). nest failure. Five pairs deserted their territories in Other pairs appearedsimilarly aged.

Fig^utelgpJosite.s,'la8raNsho||i|'s.atIsfrnaduhRet|tappedRobins'a'Te.ri|oriaISot1e:2h*edphnses'dikerdreeya-cl|ker(lreeya'}1d/e, 2 00. 0717 hours Each phrase .Der. tlpi..aLLJbesan with 2-3 ii'k\ Iouowe(l tryitun ,iii,n uuauy ining ;, a," iiiiii i^a "ai"s ||irt a tick. Tlt? doubk phrase w&t aLtrdls about 2s dutation, at 24 kH., and with nor tha 2 seon6 beth,eencaiLs. b. ienitorial Song. dika&eey;-dikadreeya, II f?nmle 25 Jan 0t 0629 hour.\' (H Dnle and H 'whulneJs fennle had sinitat longs: the.feDwte Bat,e a sti1htet iit", ,"^nil (. s.ordine catt: t\ro ftont a rcbuted ,nale (in rcsponse to a goanna), 7 Feb. 00, lBa hours. This cil ,onpriia " harsh whinnelt tike .a 6 o5-t lercnd drl.r/ion, HEH-hch-heh-heh.,ln a -l ninute sequen::eof calls pait Jron a rea(tinl to a goanna neat nest, ,nate and fe,nlile .onsistentty Savedifepnl ve.sions: the nnl.'s (all (Fis lc) t^- norc emplntic at irc stan, began wi* i ticl, ana tle pnrase wat laskr ant jtiorter than ttrc jenate;i. ntit call v'&t hanno irull! conpler at 14 kHz, with 5-12 notes per phrasi jc 'vhnmcf! uuered ar a rate oJ t0-16 notes pei setond. (tn Figs and ld gapsbe6,,)een lnve been tthonened 'rrhinneys'.frun fron nore tha l second.) d. Scoldine Call. t\'to a fenaie (in responseto i goanna), 7 feb 00, t330 hour'\' e. Aisression Rattle: squeak aM mtt.lefron H nale (in delence o.fJledgling), 2i bct. 00,'17ll hours. Tltis call cimprised a loud staccato r|ie'dfft|"0Jaboutlsennddu.a|ionandwas3i\'enbymalesandsonetinesfeiale.,'often|hecaIlwa:extendedintoaseiieso1ra||les..lipiCall the..rcttling call in(oryorated a high-pil(hed 9 kHa squeak anl several tilct. Tlrc tuttling notestaere utercd at 22 notesper secon(1.ln ttrcfield, nf c.all rounded rlnryer ond faster trdn the Scolding call. f. Ti&ina: thrce ti.k! fro,n a coloured 'note (edited to shonen gaps beth,een ti.k), j7 Juty gg, 1013 houts A single slnry tickof 24 kHz $)asgive at wrling 'chittetury'fton inter'.ak by nates andfernks in alitation, (ontact,'rnd as a wantng. g. chiuerinp: H nnle (.hoosing nest site wih Ienale), 1 Nov. 00, 1708 hours. This call tnnpised a pntlonged undulating, soft ciittering coil, tntltllttf Itur cdll reached 12 kHz arul was hannoni.allJ conplex. Therc werc 12-24 notesper sicond witir high"notes u ercd at a fa:ter rai. (Fig. lg was edited at 3.3 seunds to include high-pitched notes at 8 kHz). (.ourtin8-fe;dins lt. Sltott call: wo risiig squeakJ J-Eeek! calls frcm H nale (choosing ttest site withfenale),4 Nov 1816 lrcurs. i. Sho courting-feeding .00, Mil cornbined with Short ion'p: jai,aiieroreeya Umm nale H before 24 Nov 00, 0837 hours. j. Ieedins fenale)' sho song: dikerdree, t/ nat;1at before feeding Jl;dgtin lg'ort. 00, t62E houts. This caII rcmPised a slbrtened version of the Terito al Song. k. Short .ourting-feedine .ali: rtsing jeeit catl it,t-lni yottowea by sofi rutle morc ttan I se.oM oJ Nest'feedine soft ruule from H.male (.ofering food to incubating fenate before finale accepts.foodnear tn" n"st1, n"n Female Beeaing Saueal with three 4-10 kHz down-sluned squeals, skeeeeett, fnm H femali a: $e is fed iy H nate. i,t io" oo, 0844 houn. (gaps beween sqi"ols werc edited and shortened slightly). 13 March, 2004 y Powys.Behaviour, vocalizations and plumageol Red-cappedRobin

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Defettceof territory Nest butldhg Both prior to and during nesting,male-male encounters Nest building was observed for 16 of the 29 nests were observed on at least 23 occasions,involving eight studied, and females alone built the nest. They collected different neighbouring males. The Aggression Rattle cill cobweb and constructeda nest baseby wrapping web back (Fig. le) would be given by a male while in fluttering direct- and tbrth across the tree fork with wide-reachinshead line flighr rowardsan opponent.In perched.n.6unr..r. movements.Males sometimesstrlddled the site in fhe opposing males would face one another while oerched a female's absence,giving the Chittering call with head few cenlimetresapart. gire the AggresstonRaLll; with the movements.The femalethen collectedshort strands of dry beakwide open,spread their wings or fluff out their body tussockygrass and cobweband built the nestwalls around feathers,and advance or retreat along the perch for shoit her while sitting on the nest base.The femalecompacted distances.Occasionally one male might make pecking a the nest material with a vibrating prodding beak action, action toward the other, with no actual contact.Durins pushedher body down into the nest for shaping, and made male.maledisputes. femlles where present stayed in lh; trips every few minutes (often accompaniedby the male) brckgruundgiving Scolding and Ticking calls rFig. ld. ll). bringing fresh beakfuls of nesting material. A brown male On one occasiontwo pairs were seendisputing and one after feeding his female at the nest then inspectedand sat male physicallyattacked the neighbouringfemale, forcing in the nest for 30 seconds.pushed down Into ir and her to the ground while she was in a retreat flisht and then prodded the rim, but had not brought nest material. On€ peckingat her.On anotheroccasion male H d]ve-bombed colouredmale brought a small piece of nestingmaterial and pecked at female BB during an inter-pair dispute and addedit to a nest.Males often inspectednests and sat lasting5-10 minutes. for 5 15 secondsin them. Femalessomelrmes recycled a recently failed nest (rhis was observedfor 12 nests), Boundary disputes occasionally occuned between tearing at the old nest and carrying large beakfuls neighboudngfemales, these involved Scolding and Ticking of material to the new nest site. Only Calls and flying up in the air at eachother two or threetimesl one nest was re-used irl.rl/r. Femalescollected Usually one female would have a begging fledgling nearby. nest lining material and addedlichen and moss to Femaleswith beggingfledglings sometimesdisputed tenitory the nest exterior. Males often fed females durins with neighbouringmales using face-to-faceencounters plus nestbuilding at or nearthe nest. Scolding,Ticking, Aggressron Rarrling and TerritorialSbng Egg laying, incub.ttiorr and broodng from both, and brief aerial disputes.One brown male chased a neighbouringfemale with fledglings for two minutesfrom Nestingchronology is describedin (Powys2004). After perchto perchwith Ticking, Scoldingand beaksnapping from incubating the eggs, the female parent then brooded, both, before the brown male retreated. preened and shaded her nestlings for decreasingamounts of time until they fledged. During prolonged or heavy rain Courting and copulation sittingfemales protected their eggsor nestlingsby angling Nest site selectionbehaviour was observedon at least the body and flicking their wings to shed rain from the l8 occasions, involving ten different pairs during nest.When eggsor nestlingswere exposedto hot sun,the November and December.When a pair of Red-capped temalewould sit or standon the nest rim with her wines Robins were selectinga nestsite the male would straddle partly openedro provide shade.In cool weatherfemal;s a suitabletree fork and move its head from side to side brooded in a more horizontal position while flufling their while giving a prolongedsoft Chiueringcall (Fig. lg) for feathersto cover the nestlings.A male parent continued to l5-60 secondsduration. This Chitterinccall could onlv be feed his nestlingsduring quite heavyrain while the female heard for a disranceof up to 50 mirres. in optimum was shelteringthem. conditions.The female would then sit in and straddlethe Foraging and of young tree fork as if to test it. Several potential sites would be feeding examinedin this way. The following day nest building Food items and feeding chronology are describedin usually commencedat one of thescsites. Courtins males (Powys 2004). Adult males and females hawked for air- typicallygave an empharicsqueaky Jak call combined borne ,gleaned in foliage, or watchedfrom low with Short Song, or a squeakyJ-geek, call (Fig. lh-k), perches and hovered low before pouncing on ground- then ted the female on or near the newly-commencednesti dwelling insects and spiders.Adults sometimesused wina- (male Short courting-feedingcalls were heardat least70 manlling{lowering of both wingsr to flush out -s. times on l8 occasions from ten different males). Larger items were tenderizedby thrashing or passing Copulationwas observed on sevenoccasions for four oairs. sideways through the beak both fbr self-feedins and the both whenthe pair wereselecting a nestsite and also durins feedingof young.One male parentseveral times-retrieved the nestbuilding period. The femrle immediarelyprior t6 a large insect dropped by a fledgling (two weeks out of being fed by the male typrcallygrve rhe Femali Begging the nest),and when the item jammed in the beak of the Squeal (Fig. lk) while quivering her wings. After feeain; fledgling, the male re-positioned the item and pushedit in. the perchedfemale, the male then flew in severaltieht circlei Winged insects were always pushed head-firstinto the aroundher. and during rhe I 3 secondmatinq c;ntact the throat of nestlingsand fledglings. Older nestlingscould female perchedin a submissiveposrure while the male manageto swallow quite large items wholei a l3 day-old flutteredbehind her. Nest selectionbehaviour and coDulation nestling took several minutes to swallow by degreesa 5 was llso observedfor pair H jusr one day afterdepiedution cenlimetre long grasshopperthat had been fed to it by the of their nestlings;the pair immediately re-nested. male parent. 15 March,2004 V PorvysrBehaviour, vocatizations and plumage of Red-cappedRobin

Males (including brown males) sometimesfed newly Behaviourof young hatched nestlings and also fed the female who passedthe Nestlings more than one week old self-preenedand food to the nestling. Males often used the Short Song flapped their wings and in hot conditions gaped and sat a female or young and diker-dree (Fig. tj) prior to feeding trigh in the nest. Nestlings and fledglings called -and females sometimes used the Short Song prior to feeding quivered their wings while begging and receivingfood her young. Males som€timesinstead gave the Nest-feeding fiom a parent.Nestlings when more than one week old and Soft Rattle call (Fig. lk), just prior to feeding a female on for the tirst few days after 11edging,gave the trilled or near the nest, or just prior to feeding the nestlings. NestlingBegging Call (Fig. 2a) when about to be fed by Sometimesno call was given by a parentprior to feeding parent.Older fledglings gave the FledglingBegging Call the nestlings but the nestlings were alerted to the parent's peep (Fig.2c), which was also used as a contact call. approachby the sound of fluttering wings. Nest-sitting Fledglings also gave the Fledgling Flight CaU tiko-tika-tik(l females often quivered their wings and gave the Female (Fig. 2b) when in flutte ng flight and this call was used Begging Squeal as the male approachedthe nest with food extensivelysoon after fledging, then less ofien as the 'jak'and 'j-geek'calls tbr the nestlings.Emphatic squeaky fledgling approached independence,and functioned as a were not used by females, nor by males feeding young, contactcall. This call was also heardfrom nestlingswhen and were only usedby courtingmales. exercising their wings severaldays before fledging Both parents removed faecal sacs immediately after First tlight distancefor three fledglingsat nestH6 was feeding a nestling;the feeding parent would inspectthe 10-20 metres,and after two days the fledglings were active rear of the nestling after giving a food item and if a faecal tliers. Fledglings used wing mantling when foraging, seven sac was produced,the parentwould collect the sac in its days after fledging. They hawked, pounced,caught and ate beak and fly (sometimeswith a fluttering fljght) about small insectsand caterpillars10 14 days after fledging, 20-30 metres before dropping it. while also begging and receiving food items from the

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Fig\re 2. Sonagrams itowing ca\s fron )roung Red capped Robn$. a. N?stling Beggh! Call: rrillt.t'ron lll nesling (l day b{ore Jledgittg), 28 Sept.00, 1105 houts. A pulsed tiLL at 7-3 kH., trrrrrrrj trrrrrrj tntntr. There v,,ercabout lU12 notes per till, and 2 ttills per se.o|!|. b. Fledpline Flipht Call: A high pik hed 6-9 kHz r?rier ofrn l.r tika-tika-tik^ frcn H6 Jledgling (10 days alet iedging), 25 Jan. 01, 0621 hours. Delbery rate was about 7 ti(kas per second, duration of ca| d?pendedon length of|lisht. c. FledpLi e Bepsinp Call: peep rclls fron 116fledtlinq (10 dars afto fedging), 25 Jan. 01,0621 hours. A rtmttg peep, fairly level in pitch at 7 kll. wa.t giien frequently by ledglings &\ a be\ging and co tact call. Each peep wos about 0.5 sercn^ in letrytll, Ba betueen peep! werc wtiabk. V Powys: Behaviour,\ocalizations and plumage of Red-cappedRobin Corella28(1) parents,and they followed afier the foraging parents.When Bathing and preening parents had a food item they called to their young with If water was the Short Song or the Tenitorial Song; if the youig did available adult Red-capped Robins would bathe not come to the parent then the parent went to the young. beforepreening. and young bathid threeweeks afrer fledging. Some lerrilories did not have water unless it Independence rained. Where a water source was shared between tenitodes, disputesoccuned. Ground sun-bathingsometimes Parentalfeeding of the young decreasedas the fledglings occurredwrth aduils:rhe would squat,fluf,irs feathers approachedindependence. At five to six weeks after and spreadits taii and remajn motionlesslor uD to one fledging,a parentwould sometimesfly aggressivelyat its minuteor morewhile sunnrng itsel[. beggrngyoung and give the AggressionRattle. The vouns then ceasedro give lhe Fledgling negging Cait Uui Nestsuccess and reection to predators continued to forage with their parents in earlv N€st successrate and potential predators are outlined in independence.Young from nesr NH2 used Ticking und (Powys 2004). When female H flew to the nest to incubate Scoldingcalls when newly independent.and usedthe but saw a hawk, she perchedinstead on the rim of the nest. AggressionRattle prior to independence. ready for flight. Incubaring and brooding females If two young had fledged from a nest, both parents fed constantly watchedfor predators.When a goannawas near bothyoung non-erclusrvely bul somettmessepaiately. Borh a nest both adults gave Scolding calls and occasional parentsequally fed the fledglingsro independenci.wirh Ticking and fluttered around the goanna.When a wallaby and two exceptions.In Octoberfemale H spurnedtwo voune Joey were close to an incubating female, the male gave the l5 drys rlrer lledgingand builra neu nesr.Male i rhei Scolding call with occasional Tickinq. A female fluttered solo fed the two young for another l8 days to aruund her nest fur sevelal m-lnutes,once following independence.In Januaryfemale AB solo fed one fledgling the too close approach of an observer to a nest with eggs and again while maleAB was moultinginto adultplumage. The time when she had nestlingsthe femaie llutleredaround from fledging to independencewas 33 40 days when the the nest for eight mrnuteisoonafter a nearby Brown young were aged7-8 weeks(see Powys 2004). Falcon had flown away. In the latter case, when the f-emaleeventually returned to brood her young, Sub-songand Whisper-song she kept looking skywards. The fluttering behaviour seemedto indicatedistress rather than being a calculated A scarcely audible Whisper-song,prrrrt-pit, pitti pit-pit, disracliondisplayl rhe behariuurdreu attentionto the nest repeated,was heard from two different brown males with and it continued long after the danger had passed.Constant fledglings,from a six-week-old fledgling in answerto the Ticking wasused by parentsas a warningcall whena bird- male and from an unrelatedadult female.A iuveniiefrom of-prey was perched nearby and the nestlings would nest AB2. neuly rndependenl.gave J rery simrlarbut respond by keeping still and quiet. The robins always somewhatlouder versionwhile fbraging in companywith reactedaggressively towards . the female parentin March. The call was noted as beins shortbursrs of a high-pirchedreeling chrrrer wirh somi Plumage of adubs Ticking but was not recorded. A very similar song tiom Plumage for adult Red-cappedRobins is well described an adult male (H) was recorded in April 2001 (Sub-song in field guides;adult males in this study were rypical in Fig.3). Male H's Sub-songdid nor appearto be dir€cted havingred breastand cap, black head,neck and chin, and at anotherrobin and was given while the male preenedand white flanks, belly and wing stripe.However, there were foraged. Whisper-songwas audible within a distanceof individual differencesin plumage, describedelsewhere three metres;Sub-song within a distanceof l0 metres;sons (Powys 2004). Adult females had grey-brown head, back structurewas similar for both. and wings, with paler chin, breastand belly, but the overall

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-l III I o seconds I 34

Egr\re 3. sonagram showing sub'song frdn an alull male Red-capped Robin. sub-sonp: from H male, 28 ApiI 0]. part of a one-minute sequence oI very soft, hiSh-pbched reeling chattea including elements of the Aggression Raule, Ticking, and Tenito al Song, rcndered in pat dr pritti-PREE-pritti-prit-PRRT-pit Mrcl of this song was 6-a kHz with ol,eton;s hot shou'n) at 12-14 kHz Robin 17 March,2004 V Porvs.Behaviour. vocalizations and plumageof Red-capped

brown males in this hue varied with individuals. Darker plumaged females had but strongly rust-colouredcaps of three (the rust caps ot a larger rust-coloured cap; paler females had a very slight, study may be diagnostic of gender or no, rusty cap. females, where obvious, tended to be larger and duller)' and the full-length white wing striPe in an otherwisebrown Plumage of fledglings bird also proved to be diagnostic of gender. Pink spots not a definite male indicator as one female had a pink All fledglings had pale unmarkedthroats and pale breasts were Strongly teritorial behaviour and long flecked with darker grey-brown markings, but the density spot on her breast. prior nesting were also diagnostic of of flecking varied in same-agefledglings and some birds bouts of singing to females sometimes briefly gave the also had buff-washedflanks. The flecking on the upper gender, although after nesting. body was more uniform betweenindividuals. The breast Territorial Song flecking was less obvious as fledglings approached Confusion in plumage descriptions for brown males has independence.Wing stripeswere tan-colouredat fledging, led to some inaccuratevoice and behaviourai descriptions, 'dance' but paler buff at independence.Both fledglingsfrom nest for exampleStewart (1976) describeda between AD3 had a pale rust wash on the foreheadat independence male and female Red-cappedRobins but this may have plus an un-tlecked grey-beige breast.Both tledglings fiom been an interaction between rival coloured and brown 'dance' nest NH2 were still strongly flecked on the breast at males. In CaperteeValley such action always independenceand did not have a rust wash on the indicatedaggressive bluff encounterson perchesor on the forehead.Of three fledglings from nest H6, two were ground between rival males (coloured-to-colouredand lightly tleckedon the breastat independencewhile a third brown-to-coloured),rival females,and rival male-to-female. had no llecks. and none had a rust-washedforehead. One Dance action never occurred between mated pairs. l7-weeksold fledglingstill had pale eye ring. Hall (i974) reported post-moult transitional plumage for PLunage of browt nales two young males as being a mixture of grey and adult plumage. (1991) noted that second-yearbrown males Table I details plumage changesin fbur brown males Hutton plumage Novemberto March. Threebrown malesmoulted into full in captivrtymoulted ro full ldult in one slageover in CaperteeValley adult plumage in one stageJanuary-March, but their mature 2-3 months. Three brown males studied plumagewas acquiredin diflerentways. AB rnaleretained moulted to full adult plumage in one stageover 2-3 months. pink breast spots and full-length wing stripes while Colour terminology for plumage describedby Hobbs breeding then developed partial red plumage before black (1986) and Disney (1969, 1974) is unclear;both authors 'red' (moult completed 2 March). BB plain-breastedmale referred to brown plumaged Red-cappedRobins with developed partial red plumage before black (moult caps,which probablyindicated reddish-brown caps. Hobbs completed20 February).NH plain-breastedmale developed further noted that some females at Dareton,New South partial black plumage before red (moult completedlate Wales, had a red wash on the breast which appearedpink, March). MB male with pink-spottedbreast abandoned his and Disneymentioned traces of pink on the breastof both territorymid-December and moult detailsare not known. brown males and females. Capertee Valley pale females developeda buff or brownish wash on the breast alier DISCUSSION moulting, but perhaps this was not the pink colour as describedby Hobbs. Pluntoge (1999) George(l95lb) claimedthere was a plumagedifference Schoddeand Mason describeda reddishwash on (1991) betweenmale and female fledglings in captivitybut did not the breastsof immature males. Hutton described say what it was. Hutton (1991) describedfledglings in young males (but not females) at 4-5 months having a pink captivity;in a clutch of three,females showed more dark blushon the breast.It is worth notingthat Hutton's captiv€ grey stipplingon the breastthan males.At nesrH6 in this birds were fed a colour enhancing food supplement each study, two fledglings aged four weeks were densely January. Considering the number of hue variations stippled on the breastwhile a third fledgling had much less describedin the literaturefor wild birds, perhapsthere is stippling, perhaps indicating two females and a male. a dietary link to the developmentof red-pink coloration for Comparingfledglings from other nestsin this study,there brown plumagedRed-capped Robins in wild populations. were more than two distinct variationsof plumage,so it All of Hutton's young birds had rusty caps, but several was diftlcult to sex the young by their plumage in the field. breeding females in this present study lacked rusty caps, and brown males did not develop pink-washedbreasts. George (1950) found that captive females less than two Conversely,Hobbs'(1986) finding of femaleswith pink- yearsof age lacked rusty caps and severalbreeding females washedbreasts does not match with Hutton's captive pale- in this presentstudy also lacked rusty caps but Hutton breasted females, perhaps also indicating individual and/ (1991) found in captivebirds aged 5 6 months that borh or genetic variations for plumage characteristics in wild sexeshad rusty caps, the males'cap being strongerin birds. Morcombe (2000) described brown birds in coastal colour than the females'. Field guides (Stewart 1976; populations as having a slight brown tint along the flanks Simpson1986; Schodde and Tidemann1988: Slater 1989: compared to paler inland birds, and Schodde and Mason Pizzey 1991) wrongly state that brown male Red-capped (1999) also found a trend towards paler inland birds. In Robins lack rust-coloured caps, although Coventry ( 1988) Capertee Valley, plumage of brown birds was variable described a banded irnmature male with a russet caD that according to age and stage of moult, and did not indicate latermoulted in one stagero full adult plumage.The small a tlend one way or the other. 18 V Powys: Behaviout,vocalizations and plumageol Red-cappedRobin Corella28(1)

TABLE2 ThblecomparinB descriptions of vocalizationsfor the Red-cappedRobin from rhis studywilh lhoseof covenry (1988)and Hutron(199t).

COVENTRY HUTTON Territorial Song (m, sometimes 0 Song (m) Terriloriayidenliry catl (m) Scolding - agitation (m and f) Scolding (m and 0 Tcking - agitation/waming (m and - 0 Ticking contacr call (m and t) Ticking - warning call (m and t) Aggression Rartle (m, f and juv) Chittering - nesl site selection call (m) - Short squeaks courting-feeding (m) - Short squeaks communicarion (m and 0 Female Begging Squeal (l) 'Rubbing glasscall'(m and 0 Short squeaks - communication (m and Short song (m 0 and f) Short song (m) Short song (f) Nest,feeding sofi rattle (m) Rapid clicks - ncsrfeeding (m) Sub-song^vhisper Song (m, f and juv) Nestl'ng Begging - Call trill Nestling bcgging chirrup N€stling harsh squeak Flcdgling Be8ging Call - peep Fledgling call - ibin piping whisrtc FledglingFlight Call - tika rika-rika

Vocal izqtions and behav io ur was very similar to an aggressioncall givenby the paper progress]. Behavioural observationsfor CaperteeValley mostlv lPowys, in corroboratewith rhoseof CoventryitSA8; *to iescribei Squealsfrom the female Scarlet Robin are said to be the sevenvocalizations for Red-cappedRobins at Cooma.The same or similar to those of Red-cappedRobin females fourteenvocalizations found in this presentstudy expand (Coventry 1989). Robinson (1990) noted 'distinctive on Coventry'sfindings, and differ only slightlyin matching wheezingcalls' during nestsite selectionfrom maleFlame voice with behaviour.Coventry (1988) found that male Petroicaphoenicea and Scarlet Robins, and thesecalls may Red-cappedRobins at Cooma gave a shortenedversion of parallelthe Chitteringof the Red-cappedRobin male,but the Territorial Song before feeding the nest-buildingfemale, sonagraphicanalyses are needed to confirm these two but only gave a short series of five or six rapid clicks comparisons. beforefeeding the nestlings;the females'approach to the This paper gives inforrnation for only one region in New nest was silent.At CaperteeValley, males gave the Short South Wales and it would be interesting to compare these Song betbre feeding the nest-building female but also often findings with vocalizations,behaviour and plumage for gavethe Short Song belbre feedingthe nestlingsand only Red-cappedRobins in other areasof . occasionally gave the rapid clicking call. Females sometimesgave the Shon Song before feeding the nestlings ACKNOWLEDGMENTS and both males and females sometimes save the full I wish Io thank Syd Curtis for reading the first draft of this papcr, TerrirorialSong beforeand afrerrhe nesrling;had fledged. and Ederic Slater and Petcr Fullagar for their helpful advice on Both male and femaleoften approachedthe nest without computcr sound-processing.The manuscript was improved by comments calling but the wing flutter sound alertedthe young. All provided by Ian Rowley, Richard Noske, and an anonymous rcicree. vocalizationsfor Red-cappedRobins in this study were diagnosticof behaviour. REFERENCES Boles, Waltcr E. (1988). 'The Table 2 comparesvocalizations described in this study, Robins and Flycatch€rsof Ausrralia'. (Angus and Robenson: Sydney.) with those by Coventry (1988) and Hutron (1991). Blakers,M, Davies,S. J. J. F rnd Reilly, P N. (1984). .The Arlas of Previously undescribedcalls are the AggressionRattle, AustraliaDBirds'. (RAOU and MelbourneUniversiry Press.) Chittering, two short courting-feedingcalls, Sub-song/ Chisholm, A. H. (1949). Stray Fcalhers, Srrange Vocal Mimicry. fmr Whisper-song,and FledglingFlight Call. 49: 6(Hl. Chisholm. A. H. (1960). Remarks on Robins. 6aru 60t 221-236. (1988). Chisholm(1949, 1960)noted mimicry of a ScarletRobin Coventry, P Noles on the Breeding Rchaviour of the Red, capped Robin Petroica goodenoyii. Aust. Bird Watcher l2t 209-216. Petroica multicolor trill by a male Red-cappedRobin Covenlry, P (1989). Notes on the Scarlet Robin. Aust. Bird Waklrct l3l during October in Victoria, and Coventry (1988) also 68-69. 'The observed a breeding male Red-cappedRobin giving a Disney,H. J. de S. (1969). AusrralianBird Bander',March 1969. Pp.19.(Australian Bird BardersAssociation: Sydney.) ScarletRobin trill. Red,cappedRobins in CapterteeValley 'Bird Disney,H. J. de S. (Ed.) (1974). in the Hand'. (Ausralian Bird gave no mimicry during the study period. When a Speckle.d Banders Association: Sydney ) 'Birds Warbler Chthonicola sagittata mimicked the Tenitorial Frith, H. J. (Ed.) (1969). jn the AusualianHigh Counuy'. (Reed: Song of the Red-cappedRobin in December,the male Red- Sydney.) cappedRobin respondedaggressively. George.F. (1950).The Red CappedRobir, Parr l. AusI. Avi(:ulture, D€cember. (l95la). Coventry(1988, 1989)found that somecalls of the Red- George,F The Red CappedRobin, Parl 2. Aust. At,icuhurc, January. cappedRobins and Scarlet Robins are very similar. In Ceorge,F (l95lb). The RedCapped Robin, Pan 3. Aust.A|iulture, May. March 2001 for this present study, calls given by a Hall. B. P (Ed.) (1974). 'Birds of rhe Harold Halt Ausrralian wanderingmale ScarletRobin when in conflict with a pair Expeditions1962 70'. (British Museumof NaturalHisrory: London.) of Red-crppedRobins. were very similarro rheRed-capped Hobbs,J. N. (1986)-Pink-breasted females in a poputarionof Red, cappedRobins P?rrolca goodenovii. Robin calls. Ticking Ausi Bird Wak:herllt 150 152. calls of the two specieswere very Hutton,R. (1991).'Australian Sofrbill Managemenr'. (Singil press pry similarand the AggressionRattle of the Red-cappedRobin Ltd.: Sydney.) 19 March,2004 t4 Po[ys: Behaviour,vocalizations and plumage of Red-cappedRobin 'The Morcomb€.M. (2000).'Field Guideto AustralianBirds' (SteveParish Schodde,R. aodMason, L J. (1999) Directoryof Australianbirds: Publishing:BrisbaDe.) '.(CSIRO Publishing: Melboume.) 'Field 'Field Pizzey,C. andKnight, F (1997). Guideto the Bids of Australia'. Simpson,K. (Ed.) (1986). Guideto lhe Birds of Australia' (Harper (VikingO'Neill: Melboume.) CollinsrSydney.) 'The Powys,V. (2004).Breeding biology of the Red-cappedRobin Petroica Slater,P (1989). Slater Field Guide to AustralianBirds'. zEt 4-10 (Lansdowne:Sydney.) Bood.enoviiin CaperteeValley, New SouthWales. CorclLa 'Readcr's Robinson,D. (1990).The NestingEcology of SympatricScarlet Robin Stewart,F. (1976). Red-cappedRobin ln DigestComplete Pettui.a n lticolor and P phoeni(ea Populationsio Book of AustralianBirds', lst edition.(Ed H J Frith.)Pp 360. OpenEucalypt Forest. tnu 90: 40-52. (Reader'sDigesi: Sydn€Y.) 'Reader's Schodde.R. and Tidemann,S. C. (Eds.)(1988) DiSest Sutherland,R- (2000).Audiocraph v 3 2 3 (copynghtsoftwate), Sutherland CompleteBook of AusualianBirds', 2nd edition (Reader'sDigest: Studios,PO Box 3849, WestonCreek, Australian Capital Terrilory Svdnev-) 261t.