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

6 Pages·1995·0.44 MB·

by  Clifford B Frith

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