Corella,2005, 29(2\t 254A A LONG.TERMBIRD BANDING STUDY IN UPLANDTROPICAL RAINFOREST,PALUMA RANGE, NORTH-EASTERN QUEENSLAND WITHNOTES ON BREEDING

CLIFFORDB. FRITH andDAWN W. FRITH 'Pdonodum', PO Box 581, Malanda,Queensland 4885

Received:30 September2003

This study is the result of a 2o-year bjrd banding project in upland rainforeston the Paluma Rangs, north- eastern Queensland. During the first 13 years ('1978-90) banding was carried out evsry year but thereafter onfy during 1995 and 1997, mist-netting being largely (74y" ol lolal netting hours) carried out during the first three seasons. Results provids the first long-term information on birds living in Australian upland rainforests, with emphasis on species endemic to the wet tropics or Atherton Region. Species banded were primarily those that forage w:thin the forest subcanopy,understorey, or on the ground. Peak breeding extends lrom late August or September to December or early January when tgmperaturesand rainfall incroaso and relatively moro food (fruits, flyjng insects and leal littsr invertebrates)is availabl€.Wing flight feather (primaries and secondaries including the tertials) and tail moult mostly occurs during iate Nov€mber-March,with poaks during January- March (i.€. after br€eding activity has declined and wel season rains hav6 start€d). Data are presented on biometrics plumages, solt part colours, characters for ageing and sexing, breeding, moult, survival, longevity,territoriality, sito lidelity of individual birds and seasonal movem€nts. Seasonal and annual variationjn captur€ rales are given and rocapturedata prssentedfor most trequentlycaptured individuals.

INTRODUCTION The Atherton Scrubwren Sericornis &eri. endemic to the Atherton Region, is reported to The avifauna of the upland rainforestsof the Paluma breed on the Paluma Range (Griffin 1995) but was not record during this Range, located at the southern end of the wet tropics study. Regiont {Alherton of north-easlernQueensland. has been Some other bird species endemic to Australia have documentedpreviously (1974, by Grifhn 1995) and Frith populations isolated within the wet tropics rainforest that (1984). Data presented herein result from general a represent subspeciesdistinctive from conspecifics further avifaunal banding project carried out during 1978-1990, south (e.g. the Yellow-throatedScrubwren Sericornis citreo- 1995, and 1997 and representrhe first of their kind to be gularis caimsi, Pale-yellow Robin Tregellasiacaptto nanq published for an Australian tropical rainforest.Several and Satin Bowerbird Ptilonorhynchus violaceus minor). long-term avifaunal banding studies (i.e. >10 years) have Populations of some other species, also recognized as been carried out within sclerophyll woodlands and forests: distinctive north-eastern Australian subsoecies. extend on the Brindabella Range, Australian Capital Territory beyond{north or sourh) the Atherton Region tfor further (Lamm and Wilson 1966; Horey and Wilson 1971; details see Schodde and Mason 1999). Tidemann et al. 1988), in south-westernAustralia (Brown et al. 1990), and at various New South Wales localities Recent genetic studies have presentedevidence of sub- (Lane 1969; Hardy and Farrell 1990; Marchant 1992; speciation between the northern and southe.n wet tropics Leishman 2000). Some of these long-term projectsalso populations of a few bird speciesthat occur either side of provided 'barrier' the basis for more intensive studies on most the Black Mountain betweenCaims and Mossman commonly banded species, including information on (Josephand Moritz 1994; Josephet al. 1995; Schoddeand survival rates (e.g. Morris 1975; Boehm 1974, 1977, 19'18, Mason 1999). For example, the Chowchilla is represented 1982; McFarland and Ford 1987; Robertson and Woodall in the north of its range by the subspeciesO, s. melasmenus 1987; Nicholls and Woinarski 1988; Brown et al. 199Q., and in the south (for which data are presentedhere) by O. Farell and Hardy 1993;Wilson 1994). s. spald.ingii.Similarly, the northem population of the Pied Several speciesbanded during this study are endemic to Monarch is of the subspeciesA. k. terraereginae atd, the the Atherton Region, including the Fe.nwren Oreoscopus southern population is of A. k. kaupi. A similar situation gufturalis, Mountain Thornbill Acanthiza kqtherina, Brrdled, may also be true of the Grey-headedRobin Poeciloclryas HoneyeaterLichenostomus frenat4J, Bower's Shrike-thrush (Heteromyias) albispecularis (and the Black-eared Catbird Collu icincla bowe ri, Tooth-billed Bowerbird Scenopoeetes - see Frith and Frith 2004: 238) but present data for the dentirostris and Golden Bowerbird Prionodura newtoniana northem population are inadequate to make comparisons that typically occur within upland rainforest (i.e. above c. (Schoddeand Mason 1999).Australian Cney-headedRobins 400 m asl), and Macleay's Honeyeater Xanthotis are presently treated as a subspeciesdistinct from those macleayana, Chowchilla Orthonyx spaldingii, Pied, occurring in New Guinea. Populations of Bower's Shrike- Monarch Arses kaupi, Yictor\a's Riflebird Ptiloris victoriae thrusheson either side of the Black Mountain 'barrier' may and Black-eared Catbird Ailuroedus nelqnotis that occur also differ slightly, but supposeddifferences between their within both lowland and upland rainforests of the region. measurementsare conflicting (Schoddeand Mason 1999),

25 Frith and Frilh:Bitd banding study in paluma Range Corella29(2) The aim oflhis . study was to obrain informarionon the Drometncs.plumage. STUDY AREA soft part colours. charactersfor "l.o si.rdy.was carried sexing._breeding,moult. survivat, tongeviry. ^.This our in upland raioforest. at approximaretv850- :-C:i19 vJU merres above paluma terntonahty, site fidelity and seasonal sea levrl. on the Range. troptcal DoflI;allem moueme-ntsof Ouee_qs]l^a]_aOgJ9fmately 80 kilomebes north of rownsvile. ttre stuav lnorvtduals.lhe speciesmost frequently ar€a fl9000S. caught were 146.10E) wa\ Iocared srveo Ulorn.u., fro^ rf,._rl*orili inevitably those primarily insectivorous consisred species-foraging :lljTi 9:o of.rwo50 hec*" pl"o .^"h;;;;;;;i ;'b.i within the oyir"d srudy subcanopy, understorey or ground teaf litter. l1,::1".:r,llo1 siredescriprion see Frirh atrd Frirh-2001c). A nafro$dtrl roadbiseckd lhe mainsiudv Some data for the Black-eared Catbird, Tooth_billel areatSA I t. wheremost nenitrg was carried out. The area was marked with a grid of metal Bowerbird, Golden Bowerbird, and Victoria,s stakes.Elevei fin"UiiJ nermatreDll)ma-rked shDdard nel sites{sile oumbfrsl_5 aodg-15) *ere appear elsewhere(Frith esaDrsDeo and Beehler l99g; Frith and Frith ln ratDtdresteidle. side of rheroad dlrough SAI (Fig.Il A 1998, ?001a, p.t of our mist-nenrng 2001b, 2001c, 2004) and are not .epeureJ flc-.. j9*9" wa5performed ai siLest+. iites t, h-erein r ano4 weretocated io disturbfdforesr wrlh a deDle except to be included in totals of net capture rates. Calarnrr.dominakd uodergrowthwherea! site 2 wa! wtlhiD a more opeDunderrlorev. Nomenclature follows Christidis and Boles second The ltdl+; untess {uogndded)srudy area rSA2) \ as of the samet;;;;i,j^, ;, amendedby Schodde :rnowas and Mason ( I999) in which casethe con0guou!wiri. SAI andexrended DonlwestwarG up a txll fatter approxrmately ro rs lof lowed. The BIack-eared Catbird Ailuroedus 960 meIre,above sea levrl. An old forestryf"e'gi""i;i"- track thar brsecteda narrow ridge provided me-lqnorisis retained as a species rather than access.F"; ;;;'";;;"" ai a rnarxeoslanoard oet sites(sile numbers5_g) were subspeciesof the GreenCatbird establishedto eirhe.r A. crassirostris(Frith and side of lhis track within an areaof 250 metresx Frith 2004). 100metr€s (Fig. l). ,ui"flll aDd lemperaluresshow markedleasonality oD rhe raruma--,.1j|1":l Notewonhyresuhs from this the firsr long_term KaDge.lhe dry seasoDexteoding from April_November,with bandine June-August study published beiDgthe driesrand coldesrmonrhs (Ag. 2a). RainfallaDd for upland rainforesr oi the erhertoi lernp€ralures iDcreaseduring Seplember-OcloberaDd decrease during Region arc briefly discussedwithin the April-May. species accounts, The hot wet seasoDis December_March,*irt mosrraii but only. if faifing data differ from those published fo, th",;;i;; duriogJaouar,_-February Meao monrhty rainfalt (F,g 2") i;;rh; summarized raturnal{ange (based years in pertinent volumes of the Handboof of os 1978_90)is presenredaiwe as the acrualmonlhty data ibr the firll ralian, New Zealand and Antarctic Birds (HANZAB). threeseasoos of the study(August 1978-Februaryl98l) when most standardmrsr,nerllng was performe(

\/ Figrrrc L Ifu loeation of stardard net sites 14 and 9_15 in study area I, and S_4 in stud, area 2, on,ha Eal:hstudy a,ea ii","ai . '.ttu4a'(r.!1::,!,":t::-:"::!:y::i!,.oi:,:b'd osltlmetes (= 50 hapc, trorc: Fncr stnqr? ttnes thow creek systens. double patu cl lirzs rcprctcn, the din road Irom PatumaTowtl,hip knren;s.at southtto patumaor^ r, .,"lni _i,i ii,*'i,*i,n-"sn ttnesshow contourc SAt:botda, tahitudcsin m); thedotkd line showsa s"ti-tr*t tn oigi ii2 Soo"r. June,2005 Frith and ftith: Bid banding study in paluma Rangg

Over the sameperiod mean monthly maximumtemperatures ranged A total of 2 523 hoursof mishetting \ras performedover 302 days ftom 16.7-26.6"C(Fig. 2a) and minima from 9.6_19.8.C.Availability at 15 staDdardner sitesin SAI and SA2 (Table 1). At eachof the 15 of insectsand litter invertebrates each month were monitoredfrom standardlret sitestwo 9.15x 2.74metre and two 12.2x 2.74metre nets Augustl978-February (Frilh l98t andFrirh 1985;D. Frith andC. Fritlr w-ereercckd eachday, to an effecliveheighl of 2.5 metres.From Augusr 1990;Fig. 2b). lg78-November1982 mist-Deniog was caniedour initially al two s]rcs simultaleouslyover two days (Augus! 1978-Marchl98O), then at two METHODS sitesfor one day oDly (mid March lg8o-November1982). Ners were opened between060G-0700 hours Durirg the firsl 13 years(t97E-90) bandingwas caried ou! every and closed berween 1700_1800 hours.During subsequentyears Detting was less yearbut thereafteronly during 1995and 1997.A bandinqyear was take; frequentat two sites simultaneouslyover two days.Nets wereopened as from t Augujl lo 30 July Misl.ne[ingwas nor pierformedevery oDIyduring momings until capturerates declined, month of each bandingyear, but mosrly (85%) auring the avifaunal usuallybetween 1030-1lm hours. breedingseason of August-Januar)and (74%) largely duritrg rhe first Net sites were rotatedto avoid the developmetrtof net-shvDessiD threeyears.ol the study,liom 8 Augusr l97g lo I Januarylqgl apart birdr: lhus mist-netti0goccured ar eachsite allorervalsoI at le;st $ree from June-July1979 (Table l). Misr-rettirg was carriedout at standard to four weeks.The numberof net sitesused varied from veat to vear net sitesand randomlyplaced ones (Table 2). siresl-4 withrnSAI wererhe otresmosr frequenrly rj+q of iorat 2201 1000

600 -{ zuf 1sq

t .^ d 200

ASOND J FMAMJLJJASONDJFMAMJJASONDJ August1g78-February 1981 400 2500 1l o 350 2000

1500 I 9 b rso 1000 3 !t 500 5 ii(! 50 0 0 ASONDJ FMAMJJ ASOND J FMAMJJ ASOND J August1g78-February 1981

o 1.0 E

o 0.4

= 0.0 ASONDAsUNUJ FMAMIMAM J J ASONDASONIJJ FMAMJ J ASONDJ F August1978-February t 9el Figure 2 (a). Monhly tainJall (")hite columns)and meannonthly tempercture(a) du nE Augus, 1978- February 1981 on the Palurna Range, north-eastern eueensland. For comparotive purpos"i the mean monthlt rainfa| for 1978-90 (black colunns) are also given. (b) Mean nonthty narnberof ditonal ilsec,s (stiped columns),of li$er iwenebrates (white columns)and of noctunal insects(a) duing August 1978_ February 1981, on th2 PaLumaRange, nofih-eastem eueenslanil. (c) Mean nunber oJ bitd ciptures pet how (dataJot all standatd net sitescombineal and basedon a fult bandingday Utom 0600470b to IZ'00- 1800 h) during August 1978-January 1981on the poluna Range,north-easkrn eueensland.Note: mist netting wos nol pedonvd during most of January-Februarydue to exceptionalmigatl (Fig. 2a). 28 Frith and Frith: Bird banding study in patuma Range Corella29(2)

TABLE 1 Periodsof mist-DettinSduring different montlD of €ach avian breeding seasoDat sta.ndardnet sites in study areas 1 and 2 dudng 1978-1990, lgg5 and 1997on the PalumaRange, north-eastern eueensland.

Referencenumbers Total number of Detting Total number Months of of standard days hours Number of birds of mlst net sitesr'. at sites at sites banded t978t79 Aug-Mar I to8 68 631.25 3'76 90 466 t979t80 Aug-July to 4,9 to 15 94 990.75 231 224 461 1980/81 Aug-Jan to4,9to15 249.75 76 12 148 t98r/82 Oct-Jan, May to4,9to14 l6 148.5 59 64 t23 1983 Sept-Nov to4,9to12 8 79.0 22 36 l9E3/84 58 Sept-Dec,Mar to4,9to14 22 102 75 t3'7 1984/85 Aug-Nov,JaD to4,9to14 20 94.25 52 1985 89 Sepr-Oct to4 8 3E.5 25 l8 43 19E6 Nov-Dec to4 8 35.75 33 1987 SeprOct to4 8 33.25 55 66 1988 Nov to4 8 32.5 30 t'7 1989 ocl, Dec to4 8 36.5 l9 t4 1990 33 Sept 3and4 2 l5.5 9 4 1995 l3 Oct 3and4 4 l6 l5 8 1997 Oct 3and4 4 t9.'75 t'l Total 3 20 302 2 523.25 r 100 684 t't84 ra bandiog yrar *as from Augusr lo Juty.'see rsee Tabte 3 for the oumber of hours per monlh. Figure l..four nets at each standard-net-site:two 9.1 x 2.?4 meirs aod two 12.2 x 2 ?4 met-resoersl see Methods

TABLE 2 A list of all bird speciesand numbers bandedand recapfuredat standardand raodom net sites in study arcas I and 2 duriog different months, of the year,during 1978_1990, paluma 1995,and 1997on the Range,north_ea;rcm eueensland. Numberof birds Total number Numberof capturesper SPecies month bandedrecaprured of caprures J F M A M J J A S O N D chrysococtj.l :y,:,lrr_,1:X1i"-,"*u.o tucidus I 0 I 0 0 0 0 0 0 0 0 I 0 0 0 Ncedopusitla I 0 I 0 0 o 0 0 o 0 0 0 I ::l*"Il_*fi*:.B!ff-brc_asted Paradrse, 0 o Tanysiptemsllyia I 0 t 0 0 0 I 0 0 Kingfish€r 0 0 0 0 0 0 Noisy Pitta Pita ve\icolor 3 0 3 I 0 0 O 0 0 0 {J 0 0 I 1 White-tkoatedTreecreeper Comobates leucophaeus g .7 19 2.1 4 O 2 2 O 0 0 2 8 2 0 Oreoscopusgu urutis L"T**l 58 50 t08 g 6 1 3 I 4 4 ll t3 t3 2.t t4 Yelfow,lhroatedScrubwren Se.i{orris (itrcogularis g ll3 ll4 22i Zt 6 4 2 | 8 Zi 45 3t 41 32 Large-bifledscrubwreD s magnircstr[r 130 6 246 8 13 4 g 8 2 36 49 48 B_rownCerygone Gerygone 2i 31 mouki 8 0 8 I 2 O 0 0 0 0 Mountair Thornbill 0 2 3 0 0 Acanthizakatherina 30 7 3,1 0 0 0 0 g g Lewin'slloneyearer 2 O 1 4 E 4 Metiphdgalewinii 6 I j I O I 0 0 Bridled Honeyeater 0 0 3 I 0 0 2 Lbhenosto,tlus.frenotus 60 23 83 2 1 i 3 2 White-ch€eted Horcyeater ph)'lidonyus I o 5 26 12 tB 6 nigrc I 0 1 0 O 1 0 0 0 0 0 FastemSpinebill Acanrhorynchus .t 0 0 0 0 tenuimstris 22 29 3 5 3 0 O 0 I z 9 Pale-yellowRobin Tregellasiacapbo | 3 z 32 O ft 2 3 0 0 | O I l0 l0 ? I Grev-headedRobin poedrodryasarbispecularb 3 265 231 5oi 28 16 21 6 4 2 g 66 t|,z o h,,nyx.\patdiniii r20 82 46 :l:".nil,:.... s z il 0 0 0 0 0 0 t 3 4 0 I 2 b35rem Psophdde\utivaceus _w]'ipbird 1 4 ll I 0 0 0 0 O Z 0 4 I 3 GoldenWhistler pa.hycephatapectoratis .t 0 28 t6 44 8 4 I 0 I | 2 2 lO 6 Bower'sShrike-thrush Colluricindn z bo*eri 5,1 33 90 S 2 9 I I 0 0 Z 28 Yellow breastedBoarbill Machaeti j t2 1.7 t3 rynchusJtayiventer 3 0 0 O O 0 O I 0 I | Black'facedMonarch Monarchanplanopsis 0 0 O 3 0 3 1 I 0 0 0 0 O 0 Sp€ctacledMonarch M. ttivir|atus 0 0 0 I 19 Z 21 3 4 0 0 I 0 O O Pied Monarch Arseskaubi 2 3 3 5 I O I O 0 O I 0 O 0 RufousFanrair Rhioiduri 0 0 0 O 0 rufifrons 63 46 loi 15 18 21 2 0 0 0 u R. 0 z 34 1.1 9g f":{l^.. . tutisinosL 64 26 e0 8 6 5 3 o 2 3 e 16 victoria's Riflebird hitotis vicb.iae s 20 s 5 l o o 0 r 0 0 0 l l 0 2 Black-earedCatbird Ailuroedusmetanotisl 0 r S.t 24 ti f 4 2 I I O O 1r 27 t? Tooth-biuedBowerbird scenopeetesdenrirostrisr 10 s 23 14 ll I 0 I 0 0 0 0 0 8 colden Bowerbird prionodun newtoniana. 8 7 10 22 S iO I O 3 0 I 0 0 I 5 g Sajin Bowerbird pitonorhynchusviolaceus 4 6 2 O 2 O O 0 0 O I O 0 I Mistfetoobird Dicaeumhitundina.eum 0 O 0 | 0 1 0 0 O O 0 0 O BassianThrush Zoo rcra lunutato' 0 0 0 O I 6 2 i Z O 0 0 0 O Totalnumbe. 33 0 0 3 0 0 3 1 120 741 186t t3l 89 98 3t 26 2t 'lack 35 l9s 3El 317 317 221 of capturesduring a monthis not .""pt"i excluded:Y"1,'"..T':,::":::"",l,HliJ,"^l-**o:,1:"t1y.,q1.to'^s*::1ycl35ihe courtsare-excruoea. aut ioi[*" *oliilt m*"," andnesrs are possibilitvrhar captures arso incruded the no.."t-turi"? rr,,o.oi;;,;;;; ;i";#;d;";"H:"J,t'"llfiilJ$":: June,2005 Frith and Frith:Bid banding study in paluma Rangs 29

numberof hours)used over the period sludy as a whole, sites5_g in b€ entirelyruled out. For sexuallydimorphic species the term .female- SA2 wereused during rhe first season (ll%) only andsites 9_15 in plumaged' is used in r€ferring to possible immature males whose SAI from th€ sccondto lhe seventhseason (35%; Table l). plumageis identical to tha! of adulr females.Tbe terms .aalult'and 'irrunature'ar€ not necessarily As the periodof nettingat eachstandard net site varied.caDture data indicativeof sexualmaturity, as inrmature malesof somespecies (e.9. padycephala wereslandirdrzed b) exprrjsingresuks ds lhe meannumbers of nfl the ColdenWhistler pe.toralis) might breedin femaleplurnaSe (Marchatrl area-noursper captureand the meannumbers of capturesDer 10.net and Higgins 1990j. area.hours.Ncr areahour. fot eachnet werecalculated usinp effective A bird is definedas femaleby ils plurnage,size, and/or the presence nerheighl muhrplied by rolalner tenglh and the numbersJf hoursa o[ a broodpalch if it is esrablishedt]at only the femaleof rh; species net area was open (cl Tidemanner al. l98g). Diurnal variationsin incubalesbecause a review of brood patchesstates that .thoughboth caprurerates between0630_1830 hours are expressedas lhe number gendersrnay have lhe potentialto developa broodpdlch, il is a geDeral of capturesper two,hourperiod. Caprure raresfor eachmotrth and for oDservaUonthat lt only occursin the genderactually itrvolved i0 the entirestudy appear in Table 3. Variationin capturerates during the incubation.'(Lea and Klandorf 2002, p. 100). Brood patcheswere fiIst thr€eavifaunal breeding seasons are discussed in relationto clinate recordedon a subjectivescale of l-3, with regardto rieir size and atrdthe relativeavailability of inverrebrates(Fig. 2a, 2b). degreeof vascularization-'f indicatinga small brood patchand.3' a largeone (Rogerset al. 1986i Nettingat randomsires was canied out duringvarious months of each Rogers1989). For sexuaitydimorphic specieswhose firsr yefi immature year (for untimedperiods) at courtsof Tooth,billedBowerbirds. bowers maleplumage resembhs ihar of adulr lemales(e.9. Yellow-rhroaled Scrubwren) of Golder Bowerbirds,and at nestsof both Black-earedCatbirds aod age was rakenitrrc accoutrr. A bird was assumedto be femaleif Golder Bowerbirds.Data for theserandom captur€s of bowerbirdsat recapluredmore thaoa yearafrer rls lasl cap(ureand sdll \ eariDg plunuge. suchprediclable focal points of acri!il) areexcluded from our analysis. female For rhesedetdils see Frith and (20OIc) Frith Dam for Blac*.eared Birds werenot methodicallyexamined for body moult, but if visibly Catbirdscaugh! by Ilets set at courlsand bowers(as opposed!o their active was recorded as such_ Moult in the wing flight nests),and for other (non-bowerbird) feathers bird speciescaught at all random (primaries,secosdaries) and tail were recordeditr more detail.but not Detsltes that were locatedless than 300 mehesfrom sBndardnet sites, the.sequeoceof moull rherein.Io rhespecies accousrs, the moothdunng are iDcluded.Resuhs for theseindividuals are itrcorDoraredbecause eachseasotr lhat body.wing. asd lail mouh werefirsl recordedaod !h; some,which werefirst bandedat sbndard net siles,were subsequentlv month it was completed,or nearly so, were noted. Moult cycles for retrapprdat randomnet sttes.or vice verca. Saodardand randomnei somespecies are definedand presentedbut our dataare oot iompared sites werc plotted accuratelyso tbat distancesindividual birds movedfrom with those in HANZAB unlessthey diffe. from Atherton Reqion records their origilal point of banding io recapture sites could be measured.For llereio. Subcuratreousfal depostrswere subjeclivelyscored- on a scale the 15 standardnet sites the ceDtralpoint betweenthe four nets was of 1-4 (as detailedin Rogers1989). usedin order to measuredistalces between them but for the random Recapture net srtesthe coury'bowerhestskuclure was lhe focal poinl becauselhe rates were calculatedfot those speciesmost frequetrtly (>20 locatiooof lhesewere precisely plotred (Frilh and Frrdr 20OIc). laptures) recapruredin SA1 ov€r rhe whole study periodiTabb; 2 and 4). Recapturedata do not include birds caughi twice during a During July and August l98l StephenGarDett mist_netled for two singleday. The recapturerate is definedas th€ numberof individuils momings(ll h 15 min) at randomwithin SAL His resultanldata are recapturedas a percentageof the lotal numberbanded. combioedherein. Other banders also misfneltedrandomly wirhin SAI_ Survtvalrates were duriog30 Nowmber-2 December1990 {S. c Lane,t. fu. Uardl ana . calculatedfor speciesmost frequentlyrecaplured in SAI F vanGessel) and 3-5 May ts92{D. Rogers).bur ooly rheirreciprure from Augusl lg78-Decembrr 1989wheo bandinswas iarried out everyyear (Tables data for iDdivrdualbirds origioally batrdedby us are ured herein 8-12). IDcalculariog survivat esumaies only birds 139 recaptured of the lolal of 74? recaptufes,Table 2J Conversely.we rerrappednioe at leastonce in yearsfollowing bandingare us€d,b€cause other long-term individualsoriginally banded by others.These other banders piaierl nets bandiDg studies have showo that factors such as movements young aloog.or close to. the edgeof rhe road rhal bisecledSAl, ;ear ro our of birds. mortality, and even net shynessmay te respoosible slatrdardnet sites,but their exact locationswere nol plolted. Because for the post-baodidgdisappearance of some individuals. Mean annual of this the dislance from where we banded itrdividual birds was survivalrate was calculatedin two waysi first, for mean annualsurvival estimated !o the nearestpoint oo the road or tack. By doing rates,the oumber(a) of bandedbirds presentand the .directly number preseot this th€ distancethe recapturedindividuals had moved sinc; bein; tbt srill ooe year later were calculated(i.e. Method I of Nicholls, baodedwere possibly underestimaled. but as so few birdsare involvei and Woinarski1988, based upon Lack 1954);secondty, aD such a potential discrepancyis iosignificant.previously published estrmateot mean monthly survival rates was obtaitredby calculating the percentage Ioo8evnyrecords for iodividualbirds originally baDded during ihis srudy of individualsknown to be alive at successivevearli .Recovery iotervalsfrom aoo retrappeclby orhers(i.e. as Aoon uoder Round_uDin the dale of their original capture(based on Merho; 3 oi Cotpllot $e only referredto if rheyexceed our longeviiyrecords Nicholls and Woinarski 1988). From mean annual survival an expectancyof furlher life was then calculated(cl Fry 1980).Standard Each bird capturedwas bandedwith a metal baDdprovided bv the and random net sites are referred to collectively "Birds 'capture' as capture sites herein. AusEalianBrrd and Bat BaodtDg Scheme. Envirotrment Austsalia. The t€nn refersto all bids caught,including recaptures. wereplaced in blackcottoD bags. taken to an irhmediatelylocal banding sution for processingaDd retumed to the point of capturefor releasei RESULTSAND DISCUSSION Biometricdata are presetrredfor eachspeci€s in addition to sex and age group, whetr plurnage and/or soft part differences permitted lheir Thirty-onespecies were caughtat the 15 standardnet identification.Body weight was recordedwith a pesolabalance to 0.1 sitgsover 302 degreeof accuracy.Wing, tail, netting-days,involving 2 523 hoursduring 8 tarsus,bill andtoral head lenEths were 1978-1997 measuredin a staodardizedway {Rogers1989: Friih aod Frith 2001c). . A total of I 784 captureswas recordedwhich Otrly body weight, wing length, and tail length were rneasuedand includedI 100individuals and 684 recapturcs(Table l). An recordedfrom recapturedindividuals. Studen!'s two-lailed ,_tests wete additional83 captures(20 individualsand 63 recaptures) used for statistical comparisons.Means are given + one standard weremade at randomnet sites,plus a furthertwo species deviation.As few juvenile iDdividualswor€ nettedtheir measuoments (Buff-breasted are ercludedfrom considemtion. Pmadise-Kingfisher Thnysipterasylvia ar'd, Pied Monarch)that were not caughtin standardnet sites. Nomeoclaturefor ages aDd plumagesdefined in Marchant and Thus a total of 33 specieswas netted at both standard Higgins-(1990)and Higgios ?, ai. (2001)is followed. and Ajuvenite plurnage randomnet sitesduring this study,involving is the first lost-nestlitrg otre, which is €ither replaced(pariially or I 867 captures completelyJ.by oDeor morerecog zableimnature plunages or by adull (l 120 individuals and 747 recaptures).Capture rates prunxrge.t trst rmDvrtures:ue typicallyrecognized by lheir relentio0of appearin Table 2. Thirty of the 33 speciesnetted were some Juvenile plumage characters,pa icularly those of the flight passerines(1 864 captures). feathe$, but some immaturesmay be indislinguishablefrom aduits. Thus,the possibilitylhat someindividual birds coosideredto havebeen At least 25 of the 33 speciesbanded are breeding adulls might ioclude some first ir nahrres,lacking juvenile tsaits,cannot residentson the PalumaRange and are presentthroughout 30 Fith and Fith: Bird banding study in Paluma Range Corella29(2)

TABLE 3 Mont y periods of mist-netting at 15 standardnet sites io study areas 1 and 2 during 1978-1990, 1995 and 1997 oo the PalumaRadge, north-eastem Queensland. MONTHS Apr May June July Aug Sepr Oct Nov Dec Tota) Neftidg period Numberof days 16 t5 664 44062 52 40 32 302 42.5 358.0 501.5 354.0 288.3 246.5 2523.1 Total numbcr o{ hours 224.4 169.5 166.8 6',t.3 61.3 43.0 '7.2 't.'7 Mean number of hours 94 10.6 10.4 I i.2 10.2 10.8 10.6 9.0 E.l 6.8 8.4 Totai numberof oets' 96 64 64 24 24 lb 16 160 24tl 208 160 128 1208 Capture rates Numberof captures t29 88 91 25 24 t3 10 179 382 329 283 231 1',784 Total number of 23981 l8 088 l7 797 7 t80 6 53-? 4 59(t 4 539 3 82ti 53 544 3'7802 30787 26 303 26 93s3

Mean net-area-hours 185.9 205.5 195.6 28i.2 212.2 353.1 453.9 213.5 140.2 114.9 108.8 I13.9 151.0 per capture Meao capturesper 53.3 48.'| 5i.l 34.3 3tr.1 28.3 22.O 46.8 71.3 8?.0 91.9 8?.8 66.2 nct'area-hours(10')

,four dets (two 9.15 x2.14 m atrd rwo 12.2 x 2.74 m) were erected each day at each standard net-sile; see Methods. het-area-hours ate effective del height (2.5 m) x net lengths (9.1 m or 12.2 m) x mean dumt€r of hours nets were open x number of nets (after Trdemann ?t al. 1988; see Methods)

TABLE 4 Recapturemtes for more frequently caught bird spcies (>20 captues) at standatdand random Det siles in study arca 1' duriDg 1978-1990, 1995 and 1997oo thc PalumaRange, north-eartem Queensland.

Numb€r of birds Total Dumber Number of times indrviduals recaptured Recaptu'c Specie:r baoded recaptured of capnuee I 2 3 ii 5 6 I 8 9 l0 rate(qo)l Comobaei leucophoeus 14 8 22 t i -15.7 Oreoscopusguttutul 43 102 7 4 2 -'\ 2 Sericomiscitreogularis 92 103 t95 22 2A E 31 58.7 S. maSnirostrk tli 114 225 7 9 32 44.t '7 Acanthizakalhetita 23 30 4 21.i Uchenostomusfrenatus 52 23 ,1i l1 1 2 30.8 Acanthorynchustenuiroslrk 18 4 I 21.8 Trcgelktsiacopito 2l 6 21 4 I 23.8 PoeciLodrya[albispecula s 244 229 4',t3 42 l8 lt J6. ) Pochycephalapectoralis 24 IJ 39 2 I I 20.8 '79 ColLuricinclnboweri 50 29 t2 6 38.0 Monarchatrivirgatus l9 2 2l 2 I1.0 Rhipdura rulftuns 58 45 103 9 0 2rl32.8 R. falisinosa 59 26 85 8 I 25.4 Totaypercenlage 828 662 I 500 t54 3122 106 23 22

'data for study ar€a 2 arc excluded from this arnlysis as we only rnist-ne(ed therc during our first banding yeari see Methods. lbowerbirds are excluded; see l troduction. rthe number of individuals recaptured as a percentageof the number banded. the year. The rarely sighted Little KtngftsherAlcedo pusilla every month of the year some individuals may move into may also be so but requires confirmation (Grifhn 1995). adjacent wet sclerophyll forest on the western edge of the Buff-breasted Paradise-Kingfishers arrive (at lower Paluma Range during colder months (Dettman 1995: elevationsthan our study area) in mid November to breed, Higgins et aI 2001). (pers. and depart during late February-early March obs). Excluding the honeyeatersand Victoria's Riflebird, which While most Noisy Pittas Pitta versicolor move to lower are mixed feeders, and the bowerbirds and Mistletoebird altitudes during winter months, some individuals remain Dicaeum hirundinaceum which are mainly frugivorous, all (see species account). Black-faced Monarchs Monarcha other passerines(85% of captures- Table 2) caught were melanopsis are absent from the Paluma Range in winter 'insectivorous',feeding mostly upon arthropods.Most (Griffin 1995), although some individuals (probably capture/recapturerecords were of ilsectivorous birds that immatures) have been seen in the Atherton Region during forageprimarily in theunderstorey/subcanopy (e.g.Large- this season (Blakers er al. 1984). Spectacled Monarchs billed Scrubwren Sericornis magnirostris, Pale-yellow Monarcha trivirgatus and Rufous Fantails Rhipid.ura Robin,Rufous Fantail) or upon or nearto the forestfloor rufifrons Ne also absentduring winter and possibly move (e.g. Femwren,Yellow-throated Scrubwren, Grey-headed to lower altitudes and./ordrier open-forest (Bravery 1970; Robin- Table2). Whilst otherfrequently caught species, Griffin 1995; Higgins and Peter 2005). On the Atherton suchas White-thrcatedTrerrrer-ry Cornbates leucophaeus, Tableland, Yellow-breasted Boatbills and Pied Monarchs Bower'sShrike-Thrush, Golden Whistler, and Grey Fantail may move to open-forest during March-June (Bravery Rhipidura fuliginosa forage mostly in the subcanopy/ 1970), but this is unrecordedfor the Paluma Rangc (Griffin canopy they also do so in lower strata,particularly in 1995). Whilst Bridled Honeyeaters were re.orded during wettermonths Grith 1984). June,2005 Frithand Fith: Bid bandingstudy in PalumaRangs

The number of net area-hoursper capture at 15 standard Many Bridled Honey€aters were captured because, net sites averaged 151 (range 116-205). Conversely, the although they are mixed feeders and most frequently seen number of captures per lOa net area-hours averaged66.2 in the canopy foraging upon flowers, they descendto lower (range 47-80). Less than two birds per hour were caught strata in searchof arthropods(Frith 1984). In constrast,the at each standard net site. A comparison between capture Macleay's Honeyeater that was common in the study area rates at sites l-4 in SAl, where the larger proportion was not netted once as it forages predominantly in the (547o) of netting was performed, showed that nets at sites forest canopy (Crome 1978; Frith 1984; Griffin 1995; pers. l, 3 and 4 caught more birds than nets at site 2. The obs.). The recapturerates of breeding summer visiting number of capturesper lOa net area-hoursaveraged 70, 75 Spectacled Monarchs (ll%o) and Rufous Fantails (3370), and 87 at sites l, 3 and 4 respectively but only 50 at site are surprisingly different. This possibly reflects the fact that 2. This was probably becausesites l, 3 and 4 had dense Rufous Fantails more frequently forage just above the Calamus-dominatedundergrowth about them while site 2 ground (animal A comparison of capture rates between each of the first food was relatively sparse,an adult was found on the road three seasonsof the study, when mist-netting was carried weighing 56.3 grams. This is well below the averase out regularly throughout each month and day, showed they weight of 9l gramsrrange 86-97 g) (Tabte5). varied considerably between seasons,particularly during breedingmonths. From August-Januaryduring the breeding A nest with three eggs was found on 27 November; on seasonsof 1978179,19'19180, and 1980/81the numberof 9 December it still contained three eggs but they had capturesper hour averaged0.8,0.5, and 0.6 respectively hatched by 14 December.An adult with a fledgling (c. 2 (Fig. 2c). Lower captue ratesduling the lattert*o seasons weeks out of nest) was seen on 20 January, and another relatedto the monthly August-Novemberrainfall being well adult with a fledgling on 26 January. No brood patch or below the seasonalaverage l32vo tor l9i9 and 55% for fat was recorded.One individual had some flisha feather 1980 - Fig. 2a). Drier conditionsresulted in relativelv moult in January. sparseavailability of arthropodsduring thoset*o ,.uron, (Fig 2b; Frith and Frith 1985; D. Frith and C. Frith 1990). White-throated Theecreeper Cormobatesleucaphaeus Jv.Ioleoyer,it was exceptionally dry during 1979 and many birds failed to breed as a result (Frith and Frith 199d, A total of 27 captures: 19 individuals (7 2001b;D. Frith and C. Frith 2000). adult males and 12 adult females) and eight recaptures.Distances between Distances between capture points of species more capture sites of individual adults averaged139 t 75 (range frequently re{aptued in mist nets in SAI indicite that many 56-294) metres. Recapture rate was 36 per cent, with no of these are relatively sedentary.Recaptures rates ranged fro; individual being caught more than threc times (Table4). llEo to 59% between speciesin SAI (Table 4). Recaphfe The longest period between banding and recovery was for rates were nohbly high (>407o) for ground-foragingspecies a male (a.t'lt when banded) of 2 years, 11 months, 9 days with the exception of Chowchillas which are Aimcutt to caugbt at four different net sites, at an ayeragedistance of capture in mist nets (Jansen1993; Fith et al. 1997). 98 (56_l4l) metresapart. suggesting sedentiriness. 32 Ftilh and Fnth: Bitd banding study in patuma Rano€ Corella29(2) -F:e pE'i:F-$is:-3 - -5 eF- 3s-* ggs.: ; i 3E i fr:: F: ::::;;: g,H 3 E i E:E;; € q 33 E E i i ii: 3:r; -;;: i Fs;;; I i $HE I::: !i E; ss 2 g g: Fi; { E F3 3! !:: gS C g-q gg; t kE 3:: !;;;;;i, ;E!;FFiSiaE;3;3;i3;d;!!EiEjHgg;iqlHqEii=u

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Adult males averagedslightly larger than tbmales (sexed The two young birds capturedwere readily distinguishable by diagnostic plumage and by the presence of a brood from adults by their duller, browner plumage and lacking patch as only femaies - incubate Higgins et al. 2O0l) an eyebrow. One caught in December was beginning to notably in wing length, but differences between the sexes moult fiom juvenile into immature plumage; is iris was dark tor any measurements including weights were not grey brown. The plumage of the second young individual significant (P > 0.10; Table 5). However, differences in caught in February, was worn and more like that of adults wing lengths for 16 males of this subspeciesmeasured by but its iris was dark red brown unlike the dark brown of Bgle-s-11dLongmore (t983) and analysedby Higgins ", adults.Thus, juveniles moult a few months after fledging into al. (2O01) were significantly (p < 0.01) longer than those a first immatureplumage that is similar to adults,the timins of four females. of this beingpreviously unknown (Higgjns and perer2002;.- Little is known aboutthe breedinghabits of this subspecies. Breeding was reco.ded during August-February. Nests One female was recordedwith a scale 2 patch brood on 14 were found under construction or complete but empty October.Active wing flight feather,tail, and body moult was during AugusrSeptember (n = 7), nests contained eggi recorded_during January-March,moult being near-complete during August-November(n = 12) and nestlingsduring or complete by April; no moult was noted durine Aususf September-February(n = 6). Eight birds had a brood patch November,Highest fat scales were noted durins itou.ilba, (sexesundetermined) when caught;two in October andJanuary. but monthlysamples were small. lscale 3), two in November (scales2,3), three in December (scales l, 2, 3) Fernwren Oreo sc op us g uttu ralis* and one in January (scale 2). Fledged juveniles/immatures were seen during October (n = 2), A total of 108 captures:58 individuals(56 adults.and December(n = l) and February (n = 2). Wing flight feather 2 immatures) and 50 recaptures(all adults). Distances moult started during October and was complete, or nearly between capture sites of individual adults averased 137 t so, by April-May. Tail moult started dudng September_ 6l trange44-350) metres.The recapturerare was high October and was complete, or nearly so, by April-May. (44Vo). Of 43 individuals banded in SAI only 19 weie Flight feather and tail moult peaked during December- recaptured:I I individuals were retrappedonce or twice February. Body moult occurred during September-June. and the remainderthree ro six rimes (Table 4t. The Most individuals carried some fat throughout the year, but individualrerrapped six times was eaughtduring rwo some monthly samples were small. seasonsat three net sites averaging 123 t 45 metres apad. Y€llow-throated Scrubwren Sericomis citreopularis The longest period between banding of an adult and its cairnsi* recovery was 8 years, 2 months, 14 days. Over this period the individual concerned was caught five times, the first A total of 227 captures: I 13 individuals (59 adult male, being at a net site 113 metres from the one it was 53 female-plumagedof which 28 were confirmed females, subsequentlyretrapped at four times. Another individual, and I immature) and I 14 recaptures(all adults). Distances also adult when banded,was last caught 6 years, 9 months, between capture sites of individual adults averaged l 14 t 26 days later.This individual was retrappedonce at 150 49 (range 3l-244) met es, A female-plumaged bird (not metres from its original point of capture. These data included in the above calculations) banded in September indicate a sedentaryspecies that holds relatively small 1978 at SA2 was recapruredan exceprional770 metres territories,at least when breeding.Moreover, one adult distant in January 1980. As this distancewas so much captured five times during April 1980-October 1983 and greaterthan all others pcrhapsthis individual was immatue anothercaught six times during August 1980-October 19g5 when banded (although juvenile traits were not noted) ard were caught on four occasions at the same four net sites was thus not territorial. When recaptured aeain in 81 metresapart, On two of the four occasions,in November December1980. still in female-plumage,ir *as 214-metres 1982and October1983, both were caushtin rhe samenet from its previous capture. The recapture rate was high together.Thus. a mared pair appearedto hold a terrilory (59Eo). Of 92 individuals banded in SAI 54 were over severalseasons, During October 1983 one bird of the recaptured: 42 individuals were retrapped once or twice, pair was recordedwith a large white eyebrow and the other and 12 individuals three to five times (Table 4). a smaller, duller, one with a scale 3 brood patch. Seven males were caught at the same net site over a In earlier literature females were describedas beine period of two (n = 3), three (n = 3) and seven (n = l) duller,having less black on rhe breasr,being slighrly palei seasons;and five females were similarly recapturedat the on the crown, and/or having a less distinctive white same net site over a period of two (n = 3), three (n = l), eyebrow than males, but these traits are now considered five (n = 1) and nine (n = 1) seasons.The latter female peter to express individual variation (Higgins and 2002). had a brood patch when first caught, suggesting she was That said, of pairs photographed visiting nests, observed resident in the area. The banding to recovery period of this nest building or feeding young, one individual (assumed female was 7 years, l0 rnonths, 8 days. That of one adult to be male) appeared to have a more distinctive white male was 6 years, 10 months, 3 days. This individual was eyebrow than its mate (pers. obs.). Whether one or both caught at three net sites during this period, at a mean sexesincubated was not observed.Thus, caDtureswere not distance of 143 metres apart. The number of times that sexedbecause. alrhough there is a slighr diiferencein size some individuals were recaptured at the same net site or between sexes with adult males being significantly larger at a nearby site(s) indicates a sedentary life style and than females in wing length, there is considerableoverlap suggests that these scrubwrens occupy relatively small in size ranges (Higgins and Peter 2002). home ranges or territories, at least when breeding. Fith ahd Frilh: Bid banding study in Paluma Range Corella29(2)

Calculated mean annual survival rate of 93 individuals representedthe roplacementof accidentally lost feathersit using Method I is 58 per cent, and using Method 2 is 6l could have been the start of tail moult as this has been per cent, indicating a mean expectancyof further life after recorded as eady as July in this subspecies(Hall 1974). banding of 1.9 and 2.1 years respectively (Table 8). These That said, six of 36 captures started tail moult in are possibly underestimates,however, resulting from few September, suggesting that tail moult starts before wing marked birds being recapturedduring the latter part of the flight feather moult. Flight feather and tail moult were sruoy. most active during January-March, and were complete, or nearly so, by April-May. Some body moult was All captures were of birds in adult plumage save one noted during August-May. Birds female-plumagedindividual that had remnants of juvenile carried some fat during most months of the year, but had more (scale plumage and a brown iris (as opposed to the darker red- 3) during October-December. brown of adults). Although adults are sexually dimorphic, first year immature male plumage resembles that of adult Large-billed Scrubwen Sericornis magnirostris vlridior* females so that it is possible some captures identified as female may have bcrn first immatue males. Measurernents A total of 246 captures: 130 individuals (104 unsexed in Table 5 are only for adult females whose sex was adults, 18 adult females, 8 juveniles to filst immatures)and confirmed by the presencea brood patch (as only females 116 recaptures (111 adults and 5 first immatures). incubate- Huggett 2000), or by age. Analysis of the large Distances between capture sites of individual adults data set in Table 5 clearly shows a signihcant difference averaged 110 t 48 (range 25-230) metres. The recapture between the adult sexes, males being larger than females rate was 44 per cent. Of lll individualsbanded in SAl, and significantly so in wing length, tarsus length and body 49 were recaptured:24 individuals were retapped once, 24 weight (P < 0.001) and tail length (P < 0.01). Moreover, individuals two to six times and one nine times (Table 4). analysisof adult and iust immature museum specimen data Eighteen individuals were caught at the same net site showeda signilicanrdifference (P < O.O5)in wing lengths over a period of three (n = 7), four (n = 4), five (n = 4), and body weights between males and females, based on nine (n = l) and 11 (n = 2) seasons.The four longestJived 10-13 birds of each sex (Higgins and Peter 2002). Because individuals were banded as adults. The longest period of considerableoverlap in measulementsand weights, between banding and last recapture of an individual was however, these characters were not used to differentiate ll years,2 months,23 days, it being caught l0 times at between first immature males and females. six different net sites ayoraging I 19 a 51 (56-175) metres Two nests (each being built by a pair of birds) were apart. Another individual, last recaptured 10 years 10 found, on 24 October and 3 November. Seven females had months 9 days after banding (Anon 1991), was nettedthree a brood patch when caught: one in November (scale l), times at two net sites approximately 25 metres apart. three in December (scales2,2,3), and three in January Similarly a third individual, a female, was last recaptured (scales 1,3,3). An immature was caught in a net together l0 years 27 days after banding, being netted four times at with an adult female on 28 February,another immatue was two net sites 138 metles apart. A fourth individual, last in the company of both parentj on 22 February, and recaptured 10 years 11 days after banding (Anon 1992), another was similarly escorted on 2l March. Active flisht was caught five times at four net sites at an average of feathermoult was not observedin any of the 62 individu-als l0Jt60(31-220) metres apart. One first immirure examined during September-October.The first signs of this individual, first caught in September 1983, was recaptured moult occured in November One individual showed sisns at the same net site the following year in adult plumage of tail moult as early as 8 August. Whilst this may hive and was last recaptured in October 1989 at a net site l0O

TABLE 8 Survivalof 93 individuallymarked Yellow-throaled Scrubwrcns captured from August l978-Decemberl9E9 in study area 1 on the palumaRange, oonh-eastemQue€nsland.

Number of Number of iDdividuals survivins Yearr individuals banded ?9/80 80/El Al/82 82 83/84 84 85 86 87 88 89 Torals 78n9 26 '7glao rE 13 8 4 4 2 I I 0 0051 2'l t'tt26622100046 E0/81 l4 430000000 El/82 7 5 310000004 E2 I 1000000I 83/84 6 0000000 84/85 0 000000 85 I 86 00000 I ll3 8? 6 tl2 88 3 0 0 89 0 0 Tbtal lumber of birds prese 93 18 30 24 16 t2 4 Ttttal 3 21 2 112 numbcr of thesebirG 5 t3 20 t3 14 3 2 ptesenl ode year tatier o I 2 65 AnNal survivat (%) 72 67 54 43 33 15 61 o 100 100 6l '. baodinSyear was from Augusr !o Julyi seeThble L June,2005 Fith and Fith: Bid bandingstudy in PalumaRange

metres from its first capture point. Two individuals banded Nineteen presumed females had a brood patch when by us were reportedas being retrappedby others some four caught: one in September(scale l), five in Novenber (all kilometresfrom our bandingsite (Anon 1991, 1992),but scale 2), eight in December (scales1,1,1,2,2,2,3,3) and five this is erroneousand results from a discrepancyin latitude in January (scales2,3,3,3,3). Juveniles/immatures were and longitude recorded by the two parties (19"00'5, mist-nettedin December (n = 1), January (n = 1), February 146"10'E by us and l8'50'5, 146"08'E by others).The (n = 3) and March (n = 3). Parents of two young gave a number of times that some individuals were recapturedat disfaction display close to the net. A juvenile/immature the same net sites or at a nearby net site(s) indicates their was also seen in the company of one or two adults on 2l sedentarylife style and suggeststhat thesescrubwrens, Ianuary,22 February, and one on 7 April was being fed which forage mainly in the understoreyand subcanopy(cJ by its parent(s).A nest in Decemb€rcontained an estimated Frith 1984), hold relatively small territories at least when week-old Fan-tailed Cuckoo Cacomantisflabellifurmis - breeding. a common brood parasite of this scrubwren (Brooker and Brooker 1989; Jansen 1990). Calculated mean annual survival rate of 108 individuals by Method 1 is 74 per cent and by Method 2 is 77 per Wing flight feather moult was not observed in the 117 cent, indicating a mean expectancy of further life after captures examined during September-December,although banding of 3.4 and 3.9 years respectively (Table 9). The in January l0 birds were all actively moulting them (Table latter figures are possibly underestimates, however, 6). Tail moult started earlier than wing moult with l0 (of resulting from few marked birds being recaptured during I 17) individuals doing so during August-December.Wing the latter part of the study. flight feather and tail moult were most active during January-March,and complete,or nearly Eighteen of 122 adults banded were sexed as female by so, by April. Some body moult occurred during September-June. the presence of a brood patch because, although the Individuals carried no fat during most of the year, but did (at incubation regime remains unknown for this species,only scale 3) during September-January. females of some other Sericornis do so (Higgins and Peter 2002). Although males have a longer wing, tail, bill and Brown Gerygone Gerygone mouki mouki+ tarsusthan do females (differencesbetween the sexesbeing significanlfor somedata sets for the subspeciesviridior). othel- A total of eight captues: eight individuals, no recaptues. captures were not sexed becausethere is considerable overlao All individuals had adult plumage bur were not sexed. between (Higgins peter the sexesin size ranges and 20021. Although there is no difference in plumage, adult males of the subspecies richmond.i have a significantly longer First imrnatures are difficult to impossible to separate wing and tail than adult females but it is not known if this from adults, although some may be recognized by their is the case in other subspecies,including the nominate relention ofjuvenile plumage characters(Higgins and peter form, for which data are too few for meaningful analysis 2002). The plumage of the eight young birds banded (Higgins and Peter 2Q02). ranged frorn clearly having a juvenile plumage with softer feathering than that of adults, a conspicuoui gape and/or Two adults were observed at a nest enfance on 30 a grey iris, through to more of an adult-like first immature January but nest contents were unconfirmed, and on 20 plumage with juvenile some remnantsof plumage and with March an immatue was seenbeing fed by a parent as well a grey-brown to brown through to the more red-brown iris as feeding itself. None of our captures had an apparent of adults. brood patch. Wing flight feather,tail, and body mouli were

TABLE 9 survival of 108 individuallyrnarked LarS€-billedscrubwrens captured during August 1978-Decembcrl9E9 in study area I on the palumaRange, north-eastemQue€Nland.

Numb€r of Numb€rof i ividMls surviving iDdividuals banded ?9/80 80/81 8l/E2 82 83/84 84 85 86 8? 78n9 l8 16127 s 4 79lao 2 I 169 1l 5 4 2 I I I r28 80/81 8 ll11I I 08 8tn2 E 1000 0 0l 82 4 22 t l9 83/84 l0 I 0 04 84/8s 5 85 I l< 5 0 86 I 02 0 00 8 2 88 I 24 89 2 00 Total nunber of birds presedt 0 l0E rE 27 18 t3 l0 9 Total dumbcr of ftese 7 81 1 124 birds t6t7 12 8 8 617 presentone year later 5 6 92 ADDualsurvival (%) 89 63 61 62 80 67 100 EE 71 86 'a bandingyear was from Augustto July; seeTable l. Fith and Frith: gitd banding study in Paluma Range Corella29(2) recorded in two individuals in Februarv. One adult carried Bridled Honeyeater Lichenostomusfrenatus* fat (scale 2) in January. A total of 83 captues: 60 individuals (47 unsexedadults, Mountain Thornbill Acanthizo katherina+ 13 adult females) and 23 recaptures. Distances between capture sites of individual adults averaged110 t 73 (range A total of 37 captures:30 individuals (27 unsexedadults, 75-300) metres, Recapture rate was 3l per cent. Of 52 3 adult females) and seven recaptures.Distances between individuals banded in SAI only 16 were recaptured: ll capture sites of individual adults averaged93 t 50 (range were retrapped once, and the rest two or three times (Table 31-128) metres. The recapture rate was low (229o), with 4). The longest period between banding and.ecovery of four of the five recapturesbeing recaught only once and an individual banded as an adult was 7 years, 6 months one individual three times (Table 4). One adult female first and 22 days,this being longer than previously recordedfor captured with a brood patch in December 1986, was next the species(Higgins et al. 2001). The individual concemed, recaptured at the same net site 8 years 9 months 21 days possibly a male (based on weight), was caught four times later. Three individuals banded during season 1979 were (March 1980, April 1980, November 1982, and October refapped during season 1983, each at the same net site 1987) at two net sites 81 metres apart. A period of 5 years where banded.Our data indicate this is a sedentaryspecies, l0 months was also rccorded for a female banded as an as was suggestedin the absenceof evidence by Higgins adult. It was caught at the same net site four times during and Peter (2002). November 1978 (with a scale I brood patch), November 1979 (with a scale2 brood patch),May 1980,and September There is no discernible differences in plumage between 1984. Thus data indicate sedentarinessin this species. the sexes.Only three of 30 adult individuals banded were sexed as females based upon their brood patch because, Calculated mean annual suryival rate of 52 individuals although the incubation regime remains unknown for this using Method I was 73 per cent and using Method 2 was species, only females of some other Acanthizq species 64 per cent, indicating a mean expectancy of further life incubate(Higgins and Peter 2002). Other captureswere not after banding of 3.2 ar,d2.3 years respectively (Table l0). sexed, becausethere is considerable overlap between the The life expectancy resulting from Method 2 is possibly sexesin size ranges(Higgins and Peter 2002), even though an underestimation, however, resulting frorn few marked there appearsto be a slight difference in size between the birds being recapturedduring the latter part of the study. sexes,with adult males being significantly larger than This speciesis considereda sedentaryresident, especially females in wing length. above 600 metles ab()ve sea level, although a proportion of Two presumed females had a brood patch in October the populationis said to descendto lower altitudesin winter, (both at scale 2) and one in December (scale l). Two nests or westward into drier areas(Higgins et al.2OOl). On the were found in November each with large young being fed Paluma Range, Bridled Honeyeaters may move west into by two adults. An adult was seenfeeding a bronze cuckoo, ecotone habitat (rainforest and wet sclerophyll forest Chalcites sp., on l3 October. Little moult was recorded in dominatedby the Flooded Gum Eucalyptusgraadts) during the 34 individuals examined, partly becauseno birds were colder months (Dettman 1995). That birds do rnove captured during January-Februarywhen moult is most between rainforest and ecotone habitats was substantiated active (Higgins and Peter 2002). No wing flight feather by the fact that an individual hrst bandediri ecotonehabitat moult was recorded, but one had new primaries in May. on L4 June ).982,by a Townsville banding group, was One individual had started tail moult in Decemberand retrappedin SAI on 2 December 1986 some 2.5 kilomet es another was completing it in May, and body moult was from is point of banding. Bridled Honeyeaten changetheir recorded in November and in May. Most birds carried diet from a predominantly insectivorousone during wetter some fat (scales2, 3) during November-December,but months to a predominantly nectarivorousone during colder 5umemunthly samplcs werc small. drier months, when fewer insects are available (Frith 1984; Fig. 2b). Thus the movement of at least some birds from Lewin's Honeyeater Meliphaga le'vtinii mab rainforestinto the westernecotone du ng winter may occur becauseof a greater abundance of nectar in the latter at -A total of sevencapturesr six individuals (adults)and one that time. recapture two seasons later. Few birds were captured, possibly becausethis honeyeatermostly forages in the There is no discernible difference in plumage between canopy (Frith 1984). There is no discernible difference in the sexes.Thirteen of 60 adult individuals were sexed plumage between the sexes.Thus, captures were unsexed female on brood patch development. Although the because,although adult males of this subspeciesare incubation regime remains unknown for this species,only signit-rcantlylarger in wing tail, bill, and tarsus, there is females of some other Lichenostomus species develop a considerableoverlap between the sexes in size ranges brood patch (Higgins et al. 2001). Other captureswere not (Higgins et at. 2OOl). sexedbecause, although males are significantly larger than females in wing, tail, bill and tarsus lengths and in body Little is known about the incubation regime in Meliphaga weight, there is considerableoverlap between the sexesin species,and none of our birds showeda brood patch. Single size (Higgins et al. 200I). birds were seennest building as early as 3l August and as late as 24 March, and a bird was sitting on a nest in On 26 September a bird was seen carrying lichen to a November. Four birds were recorded in active wing flight half completed, suspendednest approximately 11 metres feather, tail, and body moult in January. Two individuals above ground. On 14 October lichens and tendrils were during August, and one in March, had fat at scale2. beins collected as nest material from below one metre June,2005 Fdth and Fith: Bid banding study In Paluma Range

TABLE IO Survivalof 52 individuallymarked Bridled Honeyeatersduring Augus! I978-Decemberlg89 in study area I on tle palumaRange, north-eastem Queensland.

Number of Number of individuals survivins- Season' individualsbanded 79lE0 80/81 El/E2 82 83/84 84 85 86 87 88 89 Torals

78n9 1t 3221110000 0 10 19/80 19 l0 8 7 5 4 2 2 | 0039 80/81 0000000000 8l/82 lll110005 82 2 00000000 E3/84 llll105 84/85 2 000000 85 00000 86 I 0 000 8? 3 000 88 2 00 89 0 0 Total oumbcrof birds present 52 3t2 109',t144 2 159 Total numberof thes€birds 2r08?54421043 prcseorone yeiir later 't8 Annualsurvival (%) 61 83 80 7l 5? 100 50 50 064 ra banding year was ftom August to July; see Table l.

above ground near a road edge, and on 16 October another Nesting of this subspecies is mostly recorded for individual was collecting nest matedal. Fourteenpresumed November-Decemberbut two females each had a scale 1 females had a brood patch when caughr: one in Siptember brood patch in September and another a scale 2 in January. (scale l), two in October (scales 2,2), nine in November Wing flight feather and tail moult occurred during (scalesI,1,1,2,2,2,2,2,3) and two in December(scales 2,2). December-March,peaking in January-Februaryand being On 19 October two fledglings were perched 3.6 metres mostly to fully complete by March. No moult was recorded high in a sapling with at least two adults feedinq them at during July-November. Most birds carried fat (at scale 3) a fast rate, and on 29 Ocrobera young bird estimatedto during Noyember and January,but some monthly samples have been out of the nest one week was observed.Moult were small. data were few. Wing or tail moult was not observed in 54 captues examined from September-November.Wing moult Pale-yellow Robin Tregellasia capito nana* was recorded for only J. individual in December and tail A total of 38 captures:32 individuals (28 unsexedadults, moult for another in February.Flight feather and tail moult 2 adult females and 2 juvenileVimmatures) and six re.aphfes was complete, or nearly so, in April-May. Body moult (all adults). Distances between capture sites of individual appearsto start before that of wing or tail. One bird had adults averaged119 r 13 (range 113-137) mehes.Recapnre some body moult as early as August, but sevenothers had rate was 24 p€r cent, with four individuals being retrapped body moult during December-May. Most captures canied once and one individual twice within two yearsof banding. some fat (scales2-3) during September-December. There is no difference in plumage between the sex€sbut Eastern Spinebill A canthorhynchustenuirostis cairnsensis* as only females incubate two of the 30 adults were contirmed temale the presence A total of 29 captures:22 individuals (20 unsexedadults, by of a brood patch. Other captues were not sexedbecause, although males 2 females) and seven recaptures. Distances between the are larger than females, significantly so in wing, tail, capture sites of individual adults averagedl4l t 39 (range and tarsus lengths, there is considerableoverlap in size ranges 100-188) metres.Recapture rate was 28 per cent, with four (Higgins anri Peter 2002). individuals being recaught once and one individual twice within two years of banding. Few birds were captured, One young bird, captured in February possesseda rnainly becausethey forage in rhe canopy and particularly blackish bill grading to orange-yellow at its cutting edges, in the upperhalf of it (Ffith 1984). a dark grey iris and a distinct yellow gape as is typical of juveniles while another Although females do differ slightly from males in head, bird captured in January was in immature plumage, with rufor.ls feathering grey- neck, and underpart plumage, captures were unsexedsave and a (unlike for two presumedfemales wearing a brood patch. Although brown iris the darker brown of adults), and some rufous fernales typically incubate one male is reported to have head feathering. Neither bird was moulting. done so (Higgins et al.2OQl).In the three other subspecies Nests containing eggs were found during Septemberand (tenuirostris, dubius and halnaturinis) adult males are October (n = 3). One capture had a scale I brood patch significantly larger (P < 0.01) in all body measurements in October and another a scale 2 one in November. An and weight than adult females and presumably this is the adult was observed feeding a fledgling on 26 December same fot ccirnsensis.Although unsexed,our sample of 29 and another doing so on l7 February. Two young were captures adds considerably to the previously available caught during January and February (see above), and a biometrics of six individuals of this northern subspecies. young bird was seen feeding itself on 27 February. Only Fith dnd ftith: Bird banding study in patuma Range Corctta29(2)

three of 33 captureshad active moult; one in flisht feather 19 days. ll years4 months4 days and l2 years3 months moult in December.one in tail moult the samerionth, and l5 days later Four of the abovesix individualswere last one in moult in January.Most individuals ^body caught retrapped by other banders (Anon 1991, 1992, 1993). We during September-Decembercarried fat of scales2 or 3l note that reports by other banders indicate birds were retrapped at least four kilometres Grry-headed Robin Poecilodryas ( H eteromyias from their bandins site ) but a discrepancy albispecularis c inereifrons* in lalitudesand longitudes(19rc0'S, 146"10'E by us and 18"50'S, 146"08,E bv orhers) is A total of 502 captures: 265 individuals (204 unsexed responsiblefor theseerroneously supposed blj movements. adults, 22 adult females, and 39 juveniles/irnmatures)and Calculated mean annual survival rate of 100 individuals 237 recaptures(170 unsexed adults,65 adult females. and using Methods I and 2 is 77.5 per cent and 7g per 2 immatures).Adult recapturesincluded five (of l5)birds cent, rndrcatrnga mean expectancyof further life after bandins bandedas nesrlings{D. Frith and C. Frirh 2000t. eighr in of 4.0 and 3.0 years.respectively (Table ll). These immature plumage, and the remainder as adults. Dis;nces ari possibly underestimations, however, resultins from few betweencaptwe sitesof individual adults averas€dl0g r 60 markedbirds being recapturedduring qilC:.2!?8]l metr€s.Recapture tut. the lanei part of the uu.rug"d-39 p., ..nt. sruoy. Of 244 individuals banded in SA1, 94 were recaDtured:42 individualswere retrapped There once. another42 two to four is no discernible difference in plumage-were between times, and the remainder up to l0 times (Table 4). the sexes.Twenty-two of 226 adult individuals sexed as female by the presence of a brood patch when first The distance that 22 adults rnoved over 6_13 seasons caught or recaptured. Others were not sexed because. averaged97 r 40 (range 4,1-175) metres and another six althoughadult malesaverage significantly Iarger than adult were recaptured at the same net site where they had been lemalesin wing. tail. bill. and tarsuslengrh and in body for seven, nine (n = 3), and li (n = 2) seasons.It would weight, therc-is much overlap between appearthat older individuals,having size ranges(Higgins once esrablisheda and,Peter 2002) . terntory, move little distance (at least when breeding).-idults,The distance-eight immatures.subsequently caught as The plumageof rhe 4l young birds caughtranged from movedtrom rheir poinr of first captureaveraged 139 t 43 a Juvenlte one with softer feathering. rufous_brown (range l0G-233) metres, but the mean distance the five feathering on the head, a gape and./or giey to dark grey nestlings had moved from their nests to be recaught as iris, through to a more adult-like first immature pluriagl adults was greater retaining juvenile (mean = 2OOt 9g; range-were 100_360 m). remiges, rectrices, and a few othir The number of times that some individuals reoeatediv characlers including distinctive brown tipping to retained juvenile recapturedat the same net sites, or a nearby onelsj. secondarycoverts (Higgins and Fitei 2002;. two emphasizesthe sedentarinessof thrs specres. young birds caught in January with obvious traces of ptumagewere longest period Juvenrte in activewing and body mouh. and _-The between banding and recovery was two others 13 years caught in February-March had active wing, tail, 5 months 13 days, for an individual banded as a and body moult, suggesting they were moulting inti firsi nestling (Anon 1993; D. Frith and C. F th 2000). Another inr.{n.atu.rep^lumage. One young bird caughton 15 Janu-y individual.banded as a nestling,was retrapped12 years still had rufous featheringon rhe head bui lacked when l5 days later. Four birds banded-inadult ptumage this wer! last retrappedon l9 March. Four first immatures retrapped 11 years 2 months 14 wore aduh days, li year-s2 months plumage the following season.

survivalof 100 individuallvmarked crev-headed *"0lo, *or"*;f.:l,lrt".j yrr-o*"-*, 1989in studyarea l otr the palumaRange, lorth-eastem Queensland. Number of -- Number of individuals surviving Seasonr individuals banded 79/80 E0r8r 8rt82 82 E3/84 84 85 86 18n9 il 3E 26 20 I8 t7 14 r2 t2 9 79t80 20 17t398644 7 4 t7',7 80/81 3 3 5 533111 269 Et/82 0 0 0 4 42000 14 82 I 0 0 06 E3/84 00000 0 00 5 421 84/E5 4 I I 110 44 85 2 3 2 2t5 86 2 2 I I E7 3 2 2 l5 88 I 3 89 2 lt Ttrtal number of bilds present 100 43 38 0 lbtal number of 1q 34 30 2s 23 24 t9 l9 tbes€Lirds 26 33 30 30 293 preseDtoDe year lat€r 2t 19 22 t't t6 l3 22'1 Annual survival (%) .t9 .t6 6E 77 88 70 96 !a 7t 84 68 batrdint year lpas ftom August to Julyi;e Table L June,2005 Frith and Frith:Bid banding study in paluma Rang6 41

Nesting begins on the Paluma Range in late August-early females incubate, three individuals (two later retrapped September, peak breeding months being Septembe;_ once) were confirmed as female by their brood Datch. November (D. Frith and C. Frith 2000). Only females Distances between capture sites of indiuidual adults incubate.Thirty one female caplureshad a brood patch averaged105 + 53 (range 3l-156) metres.The longest whencaught: (scales two in August l.l). rhreein September period between banding and recovery is 7 years, 2 month {scalesI,l.l). eighr (scales in October 1.1.1,1,r,2,3,3), and 17 days for a female first bandedin adult plumage and elevenin November(scales 1,1.1.1.1.2.2.3.3,3.31,five in rebapped by another bander (Anon 1991). The distance December (scales 1,2,2,2,2) (scales and two in January that this individual was recaptured from its point of 1,2). wing flighr fearhermoult staned in Novemberand banding was less than 150 metresand nor at l;ast four continued through to April, by which time it was complete kilomekes as reported, this error resulting from a or nearlyso. Tail moult started earlier,in September.and discrepancyin latitudesand longitudes(19"00'S, 146"10'E continuedthrough to April when completeor nearly so, by us and 18"50'5, 146'08'E by others). FIighr feather and wing moult peaked during January- March. Body moult was recordedduring September-Vay. There is no discemible difference in plumaqe between Birds carried some fat throughout most months but more adult sexes.Captures were not sexed becausJ,although so (scale 3) during AuguslNovember. adult males are significantly larger rhan femalesin tiil length and body weight, there is considerable overlap in Chowchilla Orthonyx spaldingii spaldingii+ sizeranges (Higgins and Peter2002J. A total of ll captures:nine individuals(2 adult males. A nest containing one egg was found on 23 September, 7 adult females) and two recaptures.Few of this common and anothercontaining two young with their primariesjust bird were caught because they are difficult to capture in bursting from pin on 12 September.Nest building was (Jansen mist nets I993: Frirh et at. l99j). Both reCaptured recordedonce in November Three times durins November individuals were caughrar the same ner site aj rheir newly-fledgedto half fully grown young tl-or 2) were original capture, the male 7 months 26 days later and the observed accompanying an adult(s), and during March a female2 years,8 monthsand 23 days later,emphasising large immature accompaniedan adult. Two captures had a the sedentarynature of older, territorial birds. brood patchduring September(scates 1,2) and anotherin October (scale 3). No moult = Adults are sexually dimorphic, with males beins was recorded in July (n 2) or during September-November(n = significantly(P < 0.01) larger than femalesin wing. taiii 7), but a single March bird had active wing bill_and tarsus lengths and exclusively so in body weight flight feather and rail moult. Fat was carried (scales 2 ar'd by being up to 50 gram heavier (Jansen 1993 Frith et al. 3) in March, and September- 1997;Table 5). November This speciesbreeds during most of the year, with paluma Golden Whistler Pachycephalapecto rql is pectoralis nesting peaking during July-December (F th et al. 1997). A total of 44 captures: 28 individuals (10 adult males. Thirty active nests were examined during May-January. 4 adult females, and 14 unsexedfemale-plumaged) and 16 The nestling of a January nest fledged on 3 March. recaptures(l adult male, 14 adult females and 1 unsexed Femalesonly incubate and individuals were recorded with female-plumaged).Distances between capture sites of a brood patch during August (scale 3), September (scale individualadultr averaged130 t 4l (range8i-220.1 metres. l), Novernber(scale 3), and December (scale 2). Juveniles. Recapturerate was 21 per cent. Of 24 individuals banded identified by their lack of adult throat and breast colour in SAI only five were recaptured:four were retrappedonly and rich variegatedrufous, mottled plumage with con- one to three times but the fifth bird, a female of unknown spicuousblacVrufous wing bars, were observedassociating age, was retrappedeight times, the last time 11 years after wilh an adulr pair or a flock during August (n = 2), banding (Table 4). During this pedod she was retrapped September(n = l), November (n = 2) and December (n = at four adjacent net sites at a mean distance of ll} ! Z7Z 3). Single immatures,being browner rhan aduhs and (range 8l-160) metres apart, and had a brood patch during retaining some juvenile wing and tail plumage and a few three breeding seasons. rufous-brown feathers on the head and./or distinct wins bars.were observedin flocks during February,Seprembei Adults are sexually dimorphic, but males do not attain and November Two of these immatures had the full white adult plumage until early in their third year and sornemay throat and breast of adult males and the third the rufous not do so until early in their fou h (Higgins and peter one of adult females. Little moult information is available 2002). As males sometimes breed in immature plumage and our data are few, with one male and one female and, because both sexes incubate and develoo brood showing some wing moult in December. Another female patches(Rogers et al. 1986), only 4 of rhe 18 female- had one new central tail feather 14 millimefes lons and plumaged captures were confirmed as female by their still in pin in August, but whetherrhis was due ro acciJenral possessingfemale plumage when recapturedduring a feather loss or the start of annual moult is unknown. Only subsequentseason(s). Other captures in female-plumage two females were noted carrying fat (scale 2), one in were unsexedbecaus€, although secondyear immature and August and the other in December adult males are significantly larger in wing and tail length than females of this subspecies, there is considerable Esstern Whipbird Psophotesoliyaceus lateralis* overlap in size ranges (Higgins and Peter 2OO2), A total of ll captures:seven individuals (4 unsexed Four females had a brood patch when caught: two in adults, 3 adult females) and four recaptures. As onlv November (scales 2,2) and two in January (scales 2,2). Tlvo Fith and Fith: Bid banding study in Paluma Range corella29(2)

pale nests were being built in November, each by a female- Adults are sexually dimorphic. Females have a bill plumaged individual, and on 7 December a nest containlng and retain the apparently juvenile characters of a rufous feathers,and rufous lores two eggs was found. These hatched the following day. A superciliary stripe, pale eye-ring no superciliary stripe and have pale female was feeding a young bird on 15 February. In July Adult males have greyish lores, grey eye-dng feathers, and a black bill (C' one indiyidual had two newly Partly-grown tail feathers, ftrith O. 1990). As only females incubate, 13 of possibly due to accidental feather loss. One individual had Friih and 39 female-plurnagedindividuals were confirmed as adult started tail moult in November and another in December' by the presence of a brood patch (when first caught or Wing flight feather and tail moult peaked during January- retrapped). HoweYer, 26 female-plumaged individuals could February and by May was comPlete. Most individuals not b; sexed as there is little vadation in sizes and body carried some fat throughout the year, with larger amounts weishts between sexes,or between birds of different ages (scale September-November,although samples 3) during (Hilgins and Peter 2002: Table 5). Moreover, immature lacking for some rnonths were small or males retain female plumage characters for more than a year,with the female-likegrey bill darkeningwiti age(C' Bower's Shrike-thrrrsh Colluricincla boweri* Frith and D. Frirh 1990tpace Higgins and Peler 2002) A total of 90 captures: 57 individuals (14 adult males, Four immature males had the bill black (as of adult males) 13 adult females, 26 unsexed female-plumaged, and 4 but also had traces of rufous in their brows and lores. One immature males) and 33 recaptures (adults). Distances of these was banded in October 1981 but when retrapped between captue sites of individual adults averaged130 t in October 1983,and againin September1984, still had a 53 (range 35-250) metres.Recapture rate was 38 per cent. slight rufous wash to the feathersfrom nostrils back to over Of 50 individuals banded 19 were recaptured: 18 thJeye. This latter situation is similar to that in Grey Shrike- not attain full individuals were retrapped only once or twice, but one thrushes C. harmonica, in which birds do year (Disney 1974). individual female was recaptured five times during four adult plumage until at least their third seasonsat four net sites averaging 131 t 53 (range 100- Breeding occurs from September-January(C. Frith and 175) metresapart. The longest Period betweenbanding and D. Frith 1990). Only females incubate. Founeen fernales recovery was 8 years 10 months 25 days for an adult had a brood patch when caught: one in September(scale female, which was caught at two net sites 138 metles apart' 2), one in October (scale 2), six in November (scales Another female was recaptured at the net where it was 1,2,2,2,2,3),five in December(scales 1,2,2,2,3) and one banded, seven years later. These results indicate the in January (scale 1). Nests (n = 15) were found containing sedentarynature of these individuals. eggs or young dudng October-January,fledglings being seen during January-February.One individual had started Calculated mean annual survival rate of 24 individuals tail moult in September,and in November another showed from 1978-1988, based upon four net sites in SAI only, tail moult and yet another some wing flight feather moult. was 50 per cent (C. Frith and D. Frith 1990) Updating Most birds showed wing flight feather and tail moult these figures to include captures for all net sites in SAl during December-March,this p€aking in January-February. (n = 50) increasesthis figure to 76 per cent using Method Slight body moult was recorded in April. Most individuals 1 and 59 per cent using Method 2, resulting in a mean carried some fat throughout the year, with larger arnounts expectancyof fudher life after banding of 2.4 and 1.9 years (scale 3) during September-November,but samples were respectively (Table 12). small or lacking for somemonths.

TABLE 12 Survivalof 50 individually markedBower's Shrike-tkushescaptured ftom August 1978-December1989 iD study area I otr the PalumaRange, north-east€mQueensland.

Numb€r of Number of individuals surviving Season' individualsbanded 79180 80/81 81/82 82 83/84 84 85 86

18n9 I 311000000005 79l80 25 54443ll00022 80/81 3 1100000002 8v82 2 1l1000003 82 o 0 0000000 83/84 I 0000000 84/85 000000 85 I l0001 86 I I 102 87 2 000 88 I 00 89 I 0 Tool oumber of birds prcsent 50 36665412r135 Total number of lhese birds 155541101o23 presetrtone yeiu iarcr Annual survival (%) 33 83 83 83 E0 25 100 0 100 059 ra bandinSyear wasfrom Augustto July; seeTable 1. June,2005 Fith and Filh: Bid banding study in Paluma Range

Yellow-br€ast€d Boatbill M achaerirhynchus flayiventer in November 19?8 was subsequentlycaught an exceptional secundus* 500 metres distant in January 1979, and again in March 1980. As this distance (not included in the above A total of three adult captures: a female in June, a male calculations)is considerablygreater than thoserecorded for in August, and a female in September. Adult sexes are other adults it may have been returning to the Paluma similar in plumage, but females are paler and duller than Range for the season (see Recapture males overall, and similar in size and weight, exceptfor below). rate was 33 per cent, and whilst l5 l9 individuals ta6us length which is significantly longer in males than of were recaptured in females in this subspecies(Higgins and Peter 2005), but once or twice, two were recaptured four times, one (Table our data are too few to confirm this Oable 5). individual seven and another l0 times 4). The individual captured 10 times involved five net sites at an Boatbills were rarely sightedand it could not thereforebe averagedistance of 156 t 59 metres apart (a banding and ascertainedwhether they moved to open-forest during March- recovery period of 5 years 2 months 6 days). Even within June, as occurs on the Atherton Tableland(Bravery 1970). one season the same individual was caught at three This subspeciesis known to breed during August-January,but adjacent net sites during October-Novemberat an average no nests were seen. No brood patch or moult was recorded, distanceof 189 t 75 metles apad. Another two individuals, but the Juneand August individualshad sornefat (scale2). also adult when banded, were recaptured4 years I month 27 days and 4 years I I months 4 days later. One of these Black-faced Monarch M ona rc ha melanops is individuals was capturedseven times at two net sites (being A total of three adult captures: in December, January and caught at both net sites during 2 seasons)only 113 metres February. Whilst there is no discernible difference in plumage apart, and the other five times also at two net sites 125 betweenadult sexes,male wing, bill and tail are significantly metres apart. The recaptures of some individuals clearly longer than those of females (Higgins and peter 2005) but emphasizesthe sedentarinessof the species,at least during oul data are too few to use this in sexing captured birds. the breeding season. This is a summer breeding visitor about paluma from There is no discemible difference in plumage between mid September-lateMarch/April (Grifhn 1995). Breeding adult sexes. Captures were not sexed because, although is known during November-February(Lavery et ql. 196g) wing and tail measurements of this subsDecies are but no nestswere seen,and captureslacked brood patches. significantlylonger in adult males than in adult females, None was moulting.bur December-Januaryindividuals there is considerable overlap in size ranges (Higgins and carried fat (scale 2). Peter 2005). As both sexes incubate (Boles 1988) the presenceof a brood patchis not a criterionfor sexing. Spectacled Monzrcln M onarcha trivirgat us melanorrhoa The three young birds captured on 18 February, 28 A total of 2l adult captures: 19 individuals (adults) and March and 3 April had a dark brown iris similar to that of two recaptures.Recapture rate was ll per cent, both adults.The March bird was juvenile but with its breastand individuals concernedbeing recapturedat their banding net abdomenstill downy and a tail two thirds grown. Its upper site within a single season. There is no discemible breast was rufous-grey and not mottled, legs pale purpiiih- difference in plumage between adult sexes. Captures were flesh, and bill blackish. The April bird had all the juvenile not sexedbecause. while rhe wing of malesis significantly characteristics,with distinctly rufous upperparts,blackish longerthan in lemalesand the tarsusof femalesis loneer mottling on the upper breast, and pale purplish-flesh legs. than in lhe malesin this subspecies.rhere is considera6le Its upper mandible was brownish and the lower one overlap in size ranges (Higgins and Peter 2005). The yellowish brown. The February bird was adultlike but with presenceof a brood patch is not a criterion for sexine remnants of rufous edging to feathers, greyish becauseincubarion dark bill, rolesof the sexesremain unconfirmed. and greyish-flesh legs. Both probably incubate, as in other monarchs for which data are available(Boles 1988). Paluma Range birds move to lower altitutudes in winter, whilst others may be passagemigrants. Birds are most Birds were mist-netted only during September-February, . frequently seen during November-December(pace Griffin becausethis species apparently moues io lower altitudes 1995) lhrough lo March. A brood parch tsiale I1 was in winter (Griffin 1995; herein). Nests were under recorded for four individuals in January and three in constuction during December (n = 3) and contained eggs February. A nest was being built in November, and four during November-December (n = 2). One caDture haa a times in March an adult together with a juvenile/immature brood patch in January and two in February.A bird in were seen.A young bird was caught in each of February, immature plumage (similar to adults but with a greyer face) March, and April (see above).Our moult data are few. One was seen on 22 February. Tail moult was noted in one indiyidual was commencing wing moult in November and individual in November and another in December. two more in December.One individual was in wins moult Individuals mostly carried fat (scale 2) from November- in January.ln April at least nine of l5 captur-eshad February, but monthly samples were small. completedmoult. Individualsmostly carriedfit lscale 2.1 from November-March. Rufous Fantail Rhipidura ruffrons intermedia A total of 109 captures:63 individuals (60 adults. 3 Grey Fant&il Rhipidura albiscapa (fuliginosa) keasti juvenile/first immatures) and 46 recaptures (adults). A total of 90 captures:64 individuals (52 adults. 12 Distances between capture sites of individuai adulis juvenilesto first immarures)and 26 recaptues(all adults). averaged l4l a 56 (31-269) metr€s. An adult first causht Distances between captue sites of individual adults averaeed Fith and Fith: Bid banding study in Paluma Rang€ Cotella 29(2)

133 a 38 (31-200) metles. Recapturerate was 26 per cent at least 15 years before being killed by a domestic cat in with 14 of 15 birds being recaptured only once or twice, Paluma. Becausethis species primarily forages in the although one, possibly a female, was recapturedsix times. subcanopy/canopy(Frith 1984; Grant and Litchfield 2003) The period between banding and recovery for the latter few were mist-netted. individual, banded in adult plumage, was 6 years 1 month 20 days. It was caught at four net sites during different Black-€ared Catbird AtLuroedusmelanotis maculosus+ seasonsat an average 147 ! 47 (100_200) metres apart. A total of 102 captures:75 individuals(16 adult males, Another individual, also banded as an adult, was caught 22 adult females and 37 unsexedadults) and 27 recaptues. three times during three seasonsat the same net site and Seventeennestlings were also banded but none recaptured was last caught 5 years I month 8 days after banding. (Frith and Frith 2001b). Excluding 21 capturesat neststhe Recapturesemphasize the sedentarinessof this species. remaining 81 (57 individuals a\d 24 recaptures)were There is no discemible difference in plumage between caught at standard or random net sites. At these, 14 the sexes.Captures were not sexed because,although the individuals were retrapped once and five twice. Six wing of adult males is significantly longer than that of recaptureswere originally banded as adults at nests. adult females in this subspecies,there is considerable Many catbirds were netted because they forage in the overlap in size ranges(Higgins and Peter 2005). Moreover, understoreyand on the ground and becausesometimes they according to Rogers et aL. (1986), only females, identified attack and eat small birds entangled in mist nets (Frith and on measurementsand cloacal condition, develop a brood Frith 2001b. 2004). Catbirds were also netted around patch but as both sexesdo incubate (Boles 1988; Higgins Paluma Township. The combined data is published and Peter 2005) this criterion was not used to sex birds. elsewhere(Frith and Fdth 2001a,2001b, 200lc, 2004). The plumage of the 12 young birds capturedranged from Tooth-billed Bowerbird Scenopeetesdentirostris* clearly juvenile, with conspicuous rufous feathering with grey legs and grey-brown iris, through to a more adulrlike A total of 74 captures:50 individuals(28 adult males, first immature plumage with rernnantsof rufous feathedng 9 immature males, 13 unsexed adults) and 24 recaptures and with darker g.ey to brownish-black legs and the dark (Frith and Frith 2001c). Of a total 74 captures only 37 (23 brown iris of adults (Higgins and Peter 2005). individuals and 14 recaptures) were at standard net sites, all others being caught at random nets erectedat the courts Grey Fantails resident throughout the year are very dark, of males. At the standard net sites six individuals were almost black, but paler birds (probably migrants from the captured once, one caught twice and two caught three south) winter in this area between April and August or times. Four recaptureswere of individuals bandedat courts. early September of most years (Griffin 1995). No Most recapturcswere male court owners at courts close to individuals were recorded as being paler during this study net of visitors to them. For further but the possibility that some captures may have been our standard sites, or information biometrics, moult, survival, migrants cannot be overlooked. Birds were caught every on banding, longevity, and seasonality see Frith and Frith month of the year except May, possibly due to fewer breeding (2001a, netting hours dudng that month. One capture had a brood 20OIc, 2OO4). patch (scale (scale during October l), four'in November Golden Bowerbird Prionodura newtoniana* 1,2,2,3) and one in December(scale 1). One newly-fledged juvenile was caught on 6 December and 11 other juveniles A total of 175 captues: 109 individuals (36 adult males, to first immatures from l0 January-l9 March. A bird was 4 subadult males, 27 immature males, L2 confirmed carrying nest matedal on 28 October, one bird was seen females, 30 unsexed female-plumaged birds) and 66 incubating/brooding on 16 November, a pair completing a recaptures.Of these only 30 captures (22 individuals and nest on 2 December, and an adult with an adult-sized 8 recaptures)were caught at standard net sites, all others fledgling was sighted on 3 February. One individual was being caught at bowers or nests.At standardnet sites five caught as early as August with only six tail feathers, all individuals were retrapped once and one retrapped three of which were worn, but whether it had staned to moult times. Three recaptureswere originally banded at bowers. or its feather loss was accidental was unknown. Five For further information on banding, biomerics, moult, captues had some tail moult in September(n = 1), October survival, longevity, and breeding see Frith and Frith (1998, (n = 3) and November (n = 1). Flight feather moult was 2OOIa, 2001c, 2004). not recordeddudng September-December.Body moult was recorded from August-March. All March captures were Satin Bowerbird Ptilonorhynchus violaceus minor* actively in tail, wing and body moult. Most individuals Two individuals caught: one adult male at a Golden's carried some fat during September-November. Bowerbird bower that it was visiting to steal lichen decoration for its own bower and a female-plumaged Victoria's Riflebird Ptiloris victoriae* individual caught at a standardnet site on 9 October 1985. A total of six captures:five individuals (l adult male, I The latter individual had begun moulting body plumage adult female and 3 unsexed female-plumaged birds) and one and had one secondary missing. It was retrapped in adult recapture with a brood patch and thus female. It was male plumage on I October 2@\ 16 years I I months 22 retrapped 100 metles from its banding location 3 years, 6 days after being banded, at a bower approximately two montis and 23 days later. The longest period of life was for kilometres from where it was originally banded (Anon a male that took a minimurn of, and doubtlessly much 2002). Although this is the longest period of life recorded longer than, 3.3 years to acquire adult plumage and surviyed for the northern subspeciesminor such longevity is not June,2005 Frith and Frith: Bid banding study in Paluma Range

unusual for the nominate southern form. of which some sclerophyll habitats and, secondly, data are analysed by individuals survive greater than 20-30 years (Anon 2003; different methods to suit the specific aims of the studies Frith and Frith 2004). in question (e.9. those of Hardy and Farrell 1990 and The subspeciesminor breeds from August-February Leishman 2000). Becauseof variability in monthly and (Frith and Frith 2004). On the Paluma Range three nests annualnetting patterns,and resulting samples,standardising -22 (as werefound containing eggs, on 3 November. November, data did Tidemann et al. 1988) provides, however, an and 23 January. These nests had one to three day old accuratemeans of comparing capture rates. nestlings in them on l0 November, 4 December, and 30 Results of the present study demonstratethat (a) capture January respectively.Although the latter nest was not rates were higher at nets located at sites within relatively subsequentlyexamined its single nestling,if successful, dense undergrowth (sites 1, 3 and 4); (b) monthly capture would have fledged in mid-February.A female was tending rates were highest during September-December,when pairs a well-plumagedfledged immature on 20 April. establishor re-establishterritodes and breed (Table 3)l (c) daily capturerates were highest during the moming (0630- Bassian/Russet-tailed Thrush Zoothera species 1030) and afternoon (1630-1830), and (d) there was con- A total of eight captures;six individuals (adults) and two siderable va.riation in capture rates between years, particularly recaptures.Recaptures were caught l19 metres and 56 during breeding seasons,in relation to relative availabilitv metresdistant from their original point of capturewithin of anhropodfood resourcesthat correspondto *at ,auron two years of banding. At the start of this study the then rainsor the relativeseverity of dry seaions1Fig. 2t. White's (Scaly) Thrush Zoothera daumtz was considered the only Zoothera speciesin Australia (Schodde 1975), but Seasonality this was subsequentlysplit into the Russet-tailed Z lleinei Breeding seasonsof north-easternQueensland birds are and BassianThrush Z. Iunulata (Ford 1983; Holmes l9g4: well documented (Lavery et al. 1968; Lavery 1986; Schoddeand Mason 1999t.rhe peninent subspecies of each HAI.IZ AB volumes) with relatively small differencesin the in the Atherton Region being Z. h. heinei and Z. L cuneata. altitude of rainforest affecting the start of annual nestina, resp€ctiyely.Both specieshave been identified,by their calls, For example.on the Athenon Tableland,where seasonil in upland rainforests of the PalumaRange (Grifhn l9g5). temperaturesincrease earlier and heayier winter rains fall BassianThrushes have a notably longer wing and tail ensuring greater leaf litter invertebrateabundance (Jansen than Russett-railedThrushes (Ford 1983).Based on these 1993; Frith et al. 1.997),the nesting of birds such as Grey- criteria, in particular wing length, our eight captues would headed Robins and Black-eared Catbirds besins three to weeks paluma be Bassian Thrushes and it is, therefore, for this sDecies four earlierthan on rhe RangefD. Frith and that dataare given in Table5. However,rhe possibiliivthar C. Frith 2000; F th and Frirh 2001b). data include one or more Russet-railedThiushes cinnot, The Paluma Range avifaunal breeding season typically however,be ruled out becauseZaothera thrusheshaye b€en starts in late August-early September, as annual seen during all months paluma of the year on the Range, temperaturesinitially rise (Fig. 2a). Peak nesting is during and both species may breed there (cf Griffin 1995). late September-Decemberwhen temperaturesand rainfaf One individual mist-nettedduring Septemberhad a small increase and flowers, fruit, flying insect, and leaf litter brood patch. A nest was found on 30 October containins invertebrate foods are annually most abundant (Frith and two eggsthat harchedon approximatelyl3 November,ani Frith 1985;D. Frith and C. Frith 1990;Fig. 2a, 2b). As a another on 7 December containing two nestlings ready to generalisation,eggs are mainly laid during late September and October and nestlings are present fledge. Birds were also observed sitting-contents on nests on during November- 6 November and 8 January but nest were December. However, Chowchillas start annual egg laying unconfirmed. On 16 February a juvenile, one to two weeks as early as May (Fdth et al. 1997) and Femwrens in July out of nest, was accompaniedby its parents. (Higgins and Peter 2N2; herein). That these two species forage on the ground almost exclusively upon invertebrates. CONCLUSION which remain relatively abundant during the drier, colder months of May-July, may be significant (Frith 1984; D. Capturc rotes Frith and C. Frith 1990; Jansen 1993; Frith et at. 1997; It is widely acknowledged that it is difficult to compare Fig.2b). Ye ow-throatedScrubwrens, which also forase predominantly capture rates betueen long-term misrnetting slu-dies on the ground, do not. however,start io peter becauseof differing avifaunas,climates, habitats,numbers. breed until September (Higgins and 2002) but it is possible sizes and placement of nets, and periods that nets were their later nesting reflects foraging techniquesand erect each hour/day/month/year. For example, such diet. Heavy rains during January-Februaryreduce or bring difficulties in attempting to compare captureratei resulting to an end nesting activities, with most nestlings departing from a study in forestsof the Southem Highlands of papui nestsimmediately prior to or during early wet seasonrains. New Guinea with those performed in lowland rainforists Annual food resources typically peak as newly fledged of Brown River near Port Moresby in that country are offspring are provisioned by their parent(s), and birds are discussedby Frith and Frith (1993). Similarly it is equally moutlrng. diffrcult to compare capture rates from the present itudy Annual flight feather and wing moult predominantly with other long-term ones carried out elsewhere in occurred during Derember to March on the Paluma Range, Australia. Firstly, other published long.term banding it being most active during the wetter months of January projects in Australia have been carried out in drier to March (Frith and Frith 2001c; Fig. 3). For some species 29(2) 46 Fith and Ftith: Bitd banding study in Paluma Rang€ Corolla

Eggsand nestlings Fledglings 70 60 50 o) g40 a P30 {, &^^

10 0 ASONDJFMAM Months (Table 4), in 6gt$e-noult 3. The percentageof aduh bitds, involvinS the 14 mostfrequentll capturcd bitd species (whiti colunil, with a fat dcPosit scate 2 (blark coluttu) ond scale 3 (striped coLumn)during a banding year (August-July)on tlk Palwg Ran8e,no h'eosle Quee\slaid Tfu months thal e88satd nestlingswere nastly prcsent in nestsarc denoted by o solid black line, and the montl8 thot fled8linq| were mostryseen by o dotted line.

tail moult commencedbefore wing flight feather moult, as disperse from their natal territory or perish (Bell 1982a' early as September-October.Body moult began before that 1982b). Thus, newly-banded juvenile and immature birds of wing and tail, as early as August for some species, showed a lower recapture .ate than adult birds at our net peakedduring December-March,and was complete by late sites. Individual birds recaptured more frequently than May or June. others were invariably longer-lived ones. This emphasizes the sedentarynature of older and territorial birds. Subcutaneousfat increased in birds at the start of their breeding seasons,peaked during September-Octoberwhen Survival and longevity most species were incubating, and decreased during November-January when nestlings and fledglings were Based on long-term banding Projects, mostly in being provisioned (Fig. 3). Toward the end of the moult sclerophyll woodlandsand forests,several studies detail the fat loads increased, immediately prior to the annual dry survival rates of various Australian passerines Survival (leaner) winter. A similar fat load cycle is described for rates resulting from these studies are, however,pa icularly bowerbirds of the area (Frith and Frith 2001c). Some difficult to comDare when estimates are derived from species (e.g. Fernwrens, Yellow-throated Scrubwrens) different methodi of capture and analysis (Brown el al. (scale and, canied greater fat loads 3) during June-July 1990; Rowley and Russell 1991). Australian Passerinesare although monthly sampleswere small, this might be related typically often long lived (Fry 1980; Brown er al. 1990; to a relatively greater abundanceof leaf litter invertebrates Rowley and Russell 1991; Yom-Tov et al. 19921Baket et at that time of year. al. 1999).Results presentedherein further substantiatethis, as does work on bowerbirds (Frith and Frith 2001c, 2004). Sedentariness Such high survival rates are typical of Australian passerine Capture locations of species more frequently recaptured birds living in relatively stable environments with year- (Table 4) indicate that many are relatively sedentary,as has round availability of food (Woinarski 1985;Yom-Tov 1987; been found to be typical of many species by other long- Kalr et aL 1990; Yom-Tov et al. 1992; Martin 1996), term Australian banding studies in drier sclerophyll forests. It is clear from such studies,and other sourcessuch as the ACKNOWLEDGMENTS 'Recovery Round-up' pages in CoreLla ard records summarized in Baker et al. (1999), that many small Wethank all peopleliving in Palumaduring our work there, for help passerines, especially those of woodlands and forests, given itr various ways. We particularly thanl Dorothy atrd Johr Boyce, Kelly Davis, Atr&€e Criflid, and Linda and Bill Venn.we thaot Stephen exhibit high site-fidelity. For example scrubwrens, thombills, Gamett and Gay crowley for help and encoumSement.we thank staff robins, whistlen, and fantails may remain within a small area of Birds Australia (RAOU) coDcededwith HANZAB for supplying for many years (l\4archant 1982; Wilson 1994; Huggett 2000; unpublishedor in press rnaterial. We thatrk PeterWoodall, John Farrell, Higginsand Petet zWZ). Results of this studyalso clearly and an atronymousr€fe.ee for coostructive ctiticism of an earlier draft indicatethis. Rufous Fantails,present on the PalumaRange of this pape., which we dedicate to the memory of StephenMarcha gave only dudng the sumrnermonths, return to the samelocation who so much to Australian ornithology. thereto remainsedentary during eachbreeding season. REFERENCES High recapturerates were notably typical of speciesthat foragepredominantly upon, or nearto, the forestfloor and Anon. (1991).Recovery Routrd-up. Corella 15: 152-154. (Table AooD.(1992). Recovery Roudd-up. Corcua 16, ?9-3O,94-95. within the understorey 4). Given the sedentary Anon. (1993).Recovery Routrd-up. Corc a l7t 62-44. natureof most rainforestunderstorey species, and their Anoo (2002). Recovery Rouad-up. Corclla 26t 116. rclativelyhigh densities,most of eachyear's offspring must Anon (2003).Recovery Round-ip. CorcIIa 21t 3l-42. June, 2005 Frith and Frith: Bird banding study in Paluma Rangs 47

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