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Hybridization between the Great and Spotted Bowerbird Chlamydera nuchalis and C. maculata: an authenticated hybrid bowerbird (Ptilonorhynchidae) PDF

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Preview Hybridization between the Great and Spotted Bowerbird Chlamydera nuchalis and C. maculata: an authenticated hybrid bowerbird (Ptilonorhynchidae)

HYBRIDIZATIONBETWEENTHEGREAT AND SPOTTEDBOWERBIRD CHLAMYDERA NUCHAUSAND C MACVLATA AN AUTHENTICATED HYBRID : BOWERBIRD(PTILONORHYNCH1DAE). CLIFFORDB FRITH ANDDAWN W. FRITH Frith, C.B. & Frillu D.W. 1995 12 01: Hybud»/.atiofi between U»e (foal ftfld Spoiled Bowcrbird CMamydrra nuchalis and C, wwcWtfW: an authenticated hybrid bowcrbird (ftilonorhvnchidac>.Afe*mt/>rAoftheQueenslandMuseum38(21: 47M76. Brisbane. ISSN 0079-R835, EvidenceuihybridizationinthewildbetweenlheGreatandSpottedBowerbirdsChUmydern tmcholisandC.maitilani(Plilonorhynchidac)isdescribedandillustratedfromaQueensland Museum specimen. Another Queensland Museum specimen posvibly represents such a hybridand isalsodetailed Bowrbirds, hybrid. CliffordB.FHrh&DuwnW*Frith. Fnonodura'.P.O.BoxS&l,MaLtnda,Queensland4883, Australia; IAugust I99X TheAusiraJo-PapuanPtilonorhynchidae(bow- Two Australian monogamous bowerbirds, the erbirds)consistsof19specie* Sibley& Monroe, GreenAUuroedut crassirostritand Spoiled Cat- ( 1990; Frith, 1991; Frith et al., 1995a) and was bird A, ttxehmotis. were reported to have once long considered closely related to birds ofpara- hybridized in captivity (lnedale-,1950). but no disc,Parasidaeidae, (Gould, 1869; Sharpe,1891- specimen apparently exists to confirm theevent. 98: EII.OIJ873; Gilliard. 1969; Cooper & ,n lhe New Guinea genus Ambtyornis. the F^havv, 1077; ^Sejhhooddddee,, 11997766)).. RReecceenntt ssttuuddiieess Sfrcakcd .4, .aiw/rfrt^hJ-Macgi Bwvc dcmvtlst^Vthat thetwogroupsare, how-1" hm\A ™te*' 'T^l^l **^*?^?&^!*l*?^*cln3 N .scr.pbyiugcnUally disparate among passer- "***> u,• IntlK^fc Frith & Frith, 1990 1994 A"-*1;^^, -^; .^^, £Kt/wcrmwerWsskr,j ^#«ail.n.;19S93^;leCyhns&tidMiosn&roeScho1d99d0e;., [Jic (wo s . _ cfnt, As they were supposedly related, and po id nip 1- -' I a i' anticipated that hybrids might ,,,. „u hvhriti , ' rVj ^HiVc the occur betweI'en> -vvp'eorlbiyrgdysnpoecuiseslbaexcaausweitshuicnh euonipUurbtjhutduaut,a)..-C,he.n the editor '.Ftii-h &• Fri ith Jrssffldae4MayrJ941J962;FullerJ979). , ;,,,,,,;>,. idun LKcveraJ hybridbowerbirds have been reported tlfeai and Spoiled R but none aiftbetrtitfated. A supposed hybrid be- ufrc' n tween the Satin Ptilonnrhynrhus violaceux and (rntti PI al.'. r)LJi Regent BowcrbirdSericuluschrysocephaluswasKr^" represented by a single adult male in the irides- METHODS cent blue plumage ofthe formerspecies but with :v,L lu fL/ extensive yellow wing patches and lipping to tail feathers apparently expressing Regent Bovver- '^1!°^^^^^!^ qhis unique speciPmen^,rkntow^ndaskuM^^ fWrbird ,,vtS/.h^I,a-r/(pVDviy,gg1j*ic9?1,-9i188o;6r7nI;rcGrd-oaluel,1d4,19i15o80/6)l9o;w'caEislil-itoht*e. *, '.?r;h-u,nd ho\f\ on rlr0$leCr^^Uc\hivnins.i<l\edHlXl»krn SSL (>r ,,,.. ,.,, , , ,. Ptllh et ,, ' isCuWbsj^bcotlom^Joimlr>r6^i>y;crasnydciotsncseurbnsienqgueitnstaudtihseanptpiceiatry njaytgcsdh>> ti a,ii-d:e|uoc»aJvoopleSnwwiothodviianengthdicdkmetvs fordoubtaboui itsorigins(Mar- ulTjnl Hush , n-. I! I .:-J; 64>md, 1969). Hybridization must < f7,-MV,; hwn-nhiht, henealii vvhicji most bowi r • itwcdtd^ipossibility. however, between were Nvatcd • : conv'n's nicies were crc< thrte nv.vLh.selv related (Frith Ik Frith. 1993) al bowers Ci\ and ohsernations l%m\ geftera. pHdtographs made <Fnlh et al., 1995b). 472 MEMOIRSOFTHEQUEENSLANDMUSEUM FIhGe.re1d.eAs,carnibaeddul(tremaarl)easGraenaatdBuoltwemrablierSdp(octetnetdreB)odwiesrpblaiyrsdh(ifsocrreogwronulnod)thleomoaklseonh.ybBr,iddoGrrseaalt&xCS,povtetnetdraBlopwleurmbaigrde detailofthelivehybridGreatx SpottedBowerbirddescribedhere,bandedwithABBBSband#081 71976when csapuegchitmeonns1Q1MS0ep6t4e2mb8er&19Q9M10.2D7,2v6en3t,ralth&e Eh,yFb,riddorGsraelaatppxeaSrpaontcteedofB(o1w.etrobri.)rtdwoQMSOpo3t0t0ed59B,owearnbditrhdremeusGreeuamt Bowerbirds QMO30058,QM027527 &QM027576. HYBRIDBOWERBIRD 473 TABLE 1. Measurementsofa live hybrid Greatx Spotted Bowerbird andthose oflive both the bower- bowerbirdscaughtatbowersinthesameareaofnorthQueenslandwiththeappearance owning Great and Oftypical GreatandSpottedBowerbirds. Measurementsinmm. several Spotted Weight Wing (Tail Tarsus Bill THL Bboowweerrbtiordwastcaht atnhde GreatBowerbird follow them as they Mean 205 172 129 47.2 38.5 67.5 decorated or dis- Range (187-219) (165-175) (121-134) (45.M8.7) (37.3-39.8) (66.1-68.4) played (Fig. 1A). n _ 6 6 6 6 6 3 On 1 1 September Hybridbowerbird" 170 167 124 45.6 37.5 65.4 1991 a bird identi- SpottedBowerbird3 fied as the same in- dividual was mist !Mean 138 149 108 41.0 [32.6 ib0.4 netted ai the same iR-ange (124-166) (140-157) (101-120) (38.1-44.4) 01 0-36.5) (57.6-63.4) Great Bowerbird 31 31 24 18 |l7 29 bower, photo- 1 Data from threebirdsat the Bruslee studyareaandthreefromTownsville graphed(Fig. 1B,C) - Dataaretheaveragefrc>mdietwoexaminationsofthelivebird (1 1 Sept 1991 & 1 Sept and measured (Ta- 3iD1a9t94a)artheatfriosmno3w1 bsipredcsiamtetnheQBMrOu3sl0e0e5s9t.udyarea. btliemse1w&e 2t)h.oAugthtthaitt to be hybrid but us Bird skin specimens examined are in the the possibility it was an abnormally plumagcd CSIRO Australian National Wildlife Collection immatureGreatBowerbirdremained,webanded (ANWC)andtheAustralian (AM),National Mu- and released it (ABBBS band 081 71976). Its seum of Victoria (MV), Queensland (QM) and pinkcrestwasthenonethirddeveloped,itsmouth South Australian (SAM) museums. Capitilized yellowanditsgapepink-yellow.On 1 September descriptivewords forcoloursarethoseofSmithe 1994thisindividualwasagaincaughtatthesame (1975)unlessweconsiderthemtobeinappropri- bower. At this time the bird, now known to be at ate, in which case we use uneapitalized words. least four years old, was fully crested with a Colour reference numbers (in parenthesis) are flesh-yellow mouth and plumage character* of Ihitse ofSmithe (1975) closest to that being de- both Great and Spotted Bowerbirds. As we now scribed but not necessarily ofthe precise colour had no doubt the bird was hybrid, we collected observed. Colours considered standard forGreat and prepared it as a skin for the QuecnsJjrid Bowerbirds in the part of tropical Queensland Museum (QMO30059). It had a fully ossified concerned (18-23'S and 138-14S6E) are taken skull and enlarged (10 x 7mm) gonads. from a subadult male (QMO3O058) collected at the same location as the hybrid, and from adult specimens QM027575 and QM027576. Speci- Descriptionofthe GreatxSpottedHy- mensofSpottedBowerbirdsusedasstandardsfor bridBowerbirdSpecimenQMO30059 comparison withthe Great andhybrid specimens are QM06428, QMO10669,QMO10670 and The hybrid differs ventrally from adult-plum- QM027263, all in richly-coloured and largely aged Great Bowerbirds in not having the tbrotf, unwornplumage(Fit-.. IA). breast and flanks uniform buff-grey (Light Drab 1I9C)butadarkercolour,closertoGrayishHorn RESULTS (91), because ofthe influence ofheavy mottling and spotting that develops into barring on the sides ofthe chest and continuesdown the flanks At one Great Bowerbird bower (bower 'A" of and onto the thighs. A few of the most heavily IrUhetal., 1995b),wenotedSpottedBowtrbnJs barred feathers 10 either side of the chest are also visiting, displaying and decorating. On 23 washed with pale Cinnamon (39) which is September 1989andduring late September 1990 stronger on the concealed, basal, part of them. CBF watched and photographed a bird, consid- The centra! abdomen and belly is cream-yellow ered to be the same individual, that looked the I[541 as in Greatsbut itextendsfurtherup toward size of a Great Bowerbird but which had a few the chest as in Spotted Bowerbirds. Undertail dorsal feathers therich cinnamoncolourofSpot- coverts and undertail are as in typical Greats but ted Bowerbirdplumage and heavy ventral mark- these differ little from those ofSpotted Bower- ing and barring. This individual associated with bints. 474 MEMOIRSOFTHEQUEENSLANDMlISFUM TABLE 2, Measurement* ofGreat and SpottedBowerbird skin specimens from north crest is a discrete Queensland localities most nearthe Bruslee study area in Australian collections and area of con- thoseofahybridindividualfromtheseparentspecies.MeasurmenJsin mm. trastinelv uniform brownish Glaucous Weight Win- Tail Tarsus Bill |TKL | (79). GreatBowerbird Mantle, back and Mean 212 170 132 45.3 57.6 653 rump feathers show Range (175-317) (162-180) (125-141) (42.2-49.4) <35.2-4I.7> (63.0-70.6) characters of both B= 13 12 12 12 12 5 putativeparentsbut QMO27590 276 168 125 45.3 138.5 are more like those ofSpottedsinbeing Hybridbowerbird" !70 [67 174 45.6 137.5 65.4 more blackish, al- SpoiledBou-erbird most black on the Mean 140 149 112 40.3 32.7 59.6 centralback, unlike Ranee (117-158) (144-154;. i;ioi-ii7) (36.9-43.1) (29.9-35.4) (57.7-61.6> the dark grey- n= * 12 12 12 '2 9 brown (221) of 1 Specimens are from: 25 km S ofTownsville (CSIRO 41509c?. 11510?, 41513<5), Great Bowerbird Gilbert River, 74km W of Georgetown (CSIRO 374929, 41626$, 41627?), near back feathers. EsmereldaHomestead(QM0275756\ 27576o\ 27590(5 [possible hybrid -seetext]). These feathers on lSkro SofClermont (CSIRO43179c?}. 31fcm N nf Lyrni Junction (CSIRO 43141 9) the hybrid do have and 19kmNEofInvcrlieghHomestead(QM0275156*). the broad pale-grey 3Dataarcaveragedfromthe 1 i Sept 1991 and 1 Sept 1994examinationsofthe live hmi i,ppjn<> (hat gives SptHhueagcthiicsmnnedoncwsn(saCrpSeeIcfRir0mo4em3n:1cQ5M.7OlcT3O,0O40k35m195N.8W?)of,WCalveerrmloentyC(rQeMeOkcIT0o6s6s9idn\g,B1o0u6l7i0a?-\Mt6.7IksmaHN^ho-f P„tne~aar„^a„ini"cne"nfFO„-I?H fimnf.*p way (QM027263<5), WernadJng Homestead jQM0273739). near Mt. Isa (CSIRO °/ea< • dorsal 6621'?,417569,417579.SAMB 205059>,CoIlulamHomesteadi'QMO27303(J)and plumage but this is SW Barcaldine(CSIRO373218>. renderedinconspic- uous by large cen- Unlike adult, subadult and immature Great tral featherspotting Bowerbirdplumagesthehybrid's malar, loreand oftheSpotted Bowerbirdtypeofmarking. These ear coverts are distinctly mottled with the same spots, notas largeorsymetricalasintheSpotted GrayishHorn(91)asthethroaiandbreast.Unlike hut nothing like Great plumage, vary in colour GreatsbutastypicalofSpottedBowerbirdsthere from a paJe almost creamy off-white (92) to, in isanindistinctbutclearlyvisiblepaleoff-whitish few places, the Cinnamon {123A) and rich Mi- malarstripe (onlyjustapparent in Fig. IC). kado Brown (121C) typical of Spotted Bower- The hybrid is dorsally different from typical birds (Fig. ID-F). Table 3 comparatively Greats ofany age. The wingsare generally sim- summarizescharacters. ilar to those of an adult Great except that the Primary feathercolouringand markingismore extensivepalerips toprimary covertfeathersare like thatofthe Greatthanthemorebrownishand notoff-whiteorwhitishbutare washedwiih pale extensively paler-marked Spotted Bowerbird. Cinnamon (123A) as in, but paler than, Spotted Rump feathers have a conspicuous dark sub- Bowerbirds. The crown feathering is intermedi- terminalbandtypicalofSpottedsbutnotofGreat atebetween thatofthe twospecies. The feathers Bowerbirds. Upper tail colour and markings are are not simply tipped with pale Buff (124), or whitish as in Greats, or streaked with rich Raw noidiscernahle fiom those ofGreat Bowerbirds. Adult Greats have a clear yellow mouth and Sienna (136) conspicuously edged with dark grey-brown (1 19A) as in Spotteds, but are cen- Spotteds a fleshy one, the mouth of the hybrid trally sported Pale Hom (92) washed to varying beingintermediatebetweenthem. Measurements intensity, but stronger nearer their bases, with ofthe hybrid specimen are compared with those pale(39) todarker(123A)Cinnamon(Fig ID-F). of live (Table 1) and skin Great and Spotted Crest colour in the hybrid is Rose pink (9) but Bowerbirds(Table2). DNA maybeMagenta (2)orbe shot withPurple (1)in Mitochondria] studyofmaterial fromthe some lightasis true ofboth putative parent spe- trunk of the hybrid specimen confirms that the cies. Some forwardmost and hindmost crest female parent was a Great Bowerbird and the feathersarctippedsilvery-white. Posteriortothe father, therefore, a Spotted Bowerbird HYBRIDBOWERBIRD 475 TABLE3. Somecharacters ofadult Great and Spotted Bowerbirds in centra] east Queenslandcompared with thoseofahybridbetween thetwoofthem Character Great Hybrid Spotted Throat,breast&flanks Uniformlightdrab(W9C) Greyishhorn(91) Cstrreeaamk-eydeblllaockwi(s5h4) Abdomen,belly&chest Cchreesatml-iygehtlldorwab(5(411)9bCut) Cream-yellow(54) Cream-yellow(54) Palemalarstripe Absent Present Present Lore&earcoverts Uniform lightdrab(119C) MottledGreyishhom(91) (Mo1t2t3lAe)dpalecinnamon Paleprimarycoverttips Off-white Verypalecinnamon(123A) Cinnamon(123A) Crownfeathering Tippedpalebuff(124)to (C9e2n)trwaallsyhespdoptatleedp(a3l9e)thoorn &Streedagkeedddraarwksgireenyn-ab(r1o3w6n) whitish darker(123A)cinnamon (USA) Subterminaldarkbandonrump Absent Present Present feathers Central hack feathering Darkgrey-brown(221) Blackish Black withbroadpalescalloping Present Prencin Absent and largecentralfeatherspotting Absent Present Present Mouthcolour YelJov Yellowish-pink Flesh-pink (Christidis,pers.comm;Christidis,Frith& Firth, Mention should be made of a bird unpubl. data). (QMO27590) collected by Julian Ford 10km N of Esmeralda Homestead, north Queensland on 18 September 1986 with unenlarged gonads but DISCUSSION C a fully ossified 'adult* skull and identified as nuchalts becauseit maybehybrid. Thistentative suggestion is made because the characters now CharactersinthehybridpeculiartotheSpotted obvious in the Bruslee hybnd (Fig. 1A)werefar Bowerbirdare thepalemalarstripe, heavilymot- less sothreeyearspriortoitscollection (Fig.1A- liedandbarredbreastand flanks, sienna-streaked C). At that lime its back feathers appeared less crown and spotted back feathers. Notwithstand- black,its whitishprimary covertslackedthecin^ ing these unequivocal traits, the hybrid is far namon wash, and its erest was only one third larger in all measurements than any Spotted, developed. being thesizeofasmall Great Bowerbird(Table WhileQMO27590hasthedorsalplumageofa I). Peculiar to the Great is dorsal spotting paler Great, it has several lower back and upperrump hthyabnritdheisrmiacrhkceidnlnyasmmoanllteyrpitchaalnomfosSptomttaeldes.GrTehaet afeadtahrekrsswuabstheermdiwniatlhpbaalnedCiansnfaomuonnd(1i2n3SAp)otwtietdh 2B)o.weItrsbwieridgsh,tpoarnti1c1ulSaerplyteimnbweerigwhats, (1T6a8bgleasnd1o8nc sBtorwiepreboirrhdesa.vyItdsahrokwersmnototsliignngoonfetahrecpoavleertmsaalnadr 1 September 1994 172g.Thusitsaverageweight side of neck typical ofSpotted Bowerbirds and of 170g is lighter than that of all six Greats torfap1p2edtaakteints(sipmemciefdiicatleoclaytiaofnte(rTacbollleec1t)eadn)dotvheart pmroetstelnetdainndthbearBrreudsolneethheytbhrrioda,t,bcuhtesitt aisndhefalvainlkys thaiareaofnorthQueensland (Table 2). Itswing much like the hybrid. Notwithstanding an OftSi issmallerthanmostGreatBowerbirdsanditstail fied skull, the lack of gonad activity at peak more so (Tables 1 & 2). From its plumage at the courting season and only one pink nuchal crest time wearereasonablyconfidentthehybrid bird feather indicates it to be a subadult. Weight and wasat least two, probably three, years old when measurementsofFord'sspecimen(Table2)pro- banded on 11 September 1991. Thus it was at vide inconclusive evidence of hybridization, least five years old when collected. These facts however, and it is therefore treated here as a C and observations leading to the collection of subadult nuchalis.Shouldbothbowerbirdspe- specimen QMO30059 clearly indicate it is the cies prove to occur where it was collected a productofhybridizationbetweenGreatandSpoi- hybridoriginbecomes,however, a strongerpos- led Bowerbirds. sibility. Hybridsmaynotalwaysbeasobviousas 476 MFMOtRSOFTHEQUEENSLAND MUSEUM the specimen recorded here, and may more re- 1993.CourtshipdisplayoftheTooth-billed Bowerbird sembleoneorotheroftheputativeparents.Civen Seenopoeetes dentirostris ands its behavioural the potential extent ofsympatry ofbower-main- andsystematicsignificance. Emu93: 129-136. taining Spotted and Great Bowerbirds in central 1994, The nesting biology of Archbold's Bowerbird Archboldiapapuensisandareviewofthatofother northeasternQueensland(Frithe*al., 1995b)itis bowerbirds (Ptilonorhynchidae). Ibis 136: 153- possible that occasional hybridization between 160. the two is more widespread. FRITH C.B., FRITH. D.W. & MeCULLOUGH. M. 1995b. Great and Spotted Bowerbirds ACKNOWLEDGEMENTS Chlamydera nuchalix and C. maculate (Ptilonorhynchidae) sympatric and interacting at eachothersbowers. Australian Bird Watehef 16: This.studywouldhavebeenimpossiblewithout 4957. the kind support, interest, friendship and hospi- FRrTH C.B„GIBBS, D.St TURNER, K. 1995a. The tality ofthe McCultoughfamilywhomwethank Uixonomic status of populations of Archbold's sincerely. We thank Philippa Horton, SAM; BowerbirdArchbolulapopuensisinNewGuinea Richard Schodde, Ian MasonandJohn Wombey Bulletinofthe British Ornithologists' Club 109- CS1RO. ANWC; WalterBoles.AM and Stephen 115. Van Dyck, Glen Ingram and Wayne Longmore, FULLER, E. 1979. Hybridization amongst the QM Paradisaeidae. Bulletin of the British for access to specimens in their care and Ornithologists' Club99: 145-152. otherhelp.Mary LcCroyprovidedSmithc(1975) GILLIARD, ET. 1969. Birds or Paradise and Bower forthisstudy.WalterBoles.LesChristidis.Rich- Buds. (Weidenfeld & Nicolson; London). ardSchoddeandananonymousrefereeprovided GOULD, J. 1869. The Birds ofAustralia, supplement constructivecommentonadraftmanuscript. Les (Theauthor: London). Christidisprovidedevidenceofthematernalpar- 1REDALE.T. 1950.BirdsofParadiseandBowerBirds. entageofthehybrid. Dedicatedtothememoryof (Georgian House: Melbourne). Julian R. Ford who contributed greatly to the KL'SMIERSKI, R., BORGIA, G., CROZIER. R.H. & understanding of distributions and radiations of CHAN, B.H.Y. 1993. Molecular information on bowerbird phylogeny and the evolution ofexag- Australian birds including both putative parents geratedmalecharacteristics.Journalofevolution- ofthe hybrid described herein. ary Biology6:737-752. MARSHALL, AJ. 1954. Bower-Birds: theirdisplays LITERATURECITED anil breeding cycles. Oxford University Press, Oxford. CH1S"HRaOwrLiMsl,ey'As.BoKw.erb1i9r6d6"., EHm.Cu65R*a2w3n4s-l23c5y. and MAYRMursE.eu19m41o.fLNiasttuorfalNHeiwstGoruyi:neNaeBwirYdosr.k()A.merican CHRISaTmIoDnIgS.thLe.B&udSs-CoHfO-PDaDraEd.isRc.(1P9a9r2a.dRieslaaetiidoanes)hainpds 196E2. &FaGmrielyenPwaaraydiJrs.a,eJi.dCa.e.19Pp6.21(8e1ds-)2,04C.hIenckMa-Lyirs,i ofBirds ofthe World Volume XV. (Museum of Bowerbirds (Palonorhynchidae): protein evi- Comparative Zoology; Cambridge Massachu- dence. Australian Journal of Zoology 40: 343- setts). 353. COOPER, W.T. & FORSHAW, J.M. 1977,The Birds SCIiODDH,R, 1976.Evolutioninthebirds-of-paradisc ofParadiseand BowerBirds. (Collins: Sydney). and bowerbirds, afesynthesis. Proceedingsofthe 16thInternationalOrnithologicalCongress. 1974: DIAMOND,J.M. 1972.AvifanuaoftheEastern High- 137449. landsofNewGuinea. Publicationsofthe Nuiial) SCHODDE, R. &. MCKEAN. J.L. 1973. Distribution, OrnithologicalClub, No. 12: 1-438. taxonomy and evolution ofthe gardener bower- DIGGLES.S. 1867.TheOrnithologyofAustralia.(The birdsAnibyornissop. ineasternNewGutr>eawith author Brisbane), descriptionsoftwonew subspecies, Emu 73:51- ELLIOT , D.G. 1873. A Monograph of the 60. Paradisaeidae, (The author; London). SHARPE, R.B. 1891-98. Monograph of the FRITH. C.B. 1991. BowerbirdsandBirdsofParadise. Paradisaeidae, or Birds ol Paradise, and In Forshaw,L. (ed.), Enclopedia of Birds. Ptilonorhynchidae, or BowerBirds. (Theauthor (Smiiftmark: New York). London). 1994. Egg laying fit long intervals in bowerbirds SIBLEY, C.G & MONROE, B.L. 1990. Distribution (Pii'JonOfhvochidae). Emu94: 60-61. andTaxonomyofBirdsoftheWorld.(Yale Uni- FRITH, CB. & FRITH. D.W, 1990. Ncsling biology versity Press: New Haven). and relationships of the Lesser Mclampitta SM1THE, F.B. 1975. Naturalist'sColorGuide.(Amer- Melampitia(u^ubrif, Hmu90 65-73 icanMuseumofNatural History: NewYork I

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