Biometrics of the birds of paradise (Aves: Paradisaeidae): with observations on variation and sexual dimorphism

53 Pages·1997·19.8 MB·

by  Clifford B & Frith Frith

BIOMETRICS OFTHE BIRDS OFPARADISE AVES: PARADISAEIDAE): WITH
(
OBSERVATIONS ON VARIATION AND SEXUALDIMORPHISM
CLIFFORD B. FRITH AND DAWN W. FRITH
Frith, C.B. & Frith, D.W. 1997 06 30: Biometrics of the birds of paradise (Aves:
Paradisaeidae): with observations on variation and sexual dimorphism. Memoirs ofthe
QueenslandMuseum42(1): 159-21 1. Brisbane. ISSN0079-8835.
TheAustralasianbirdofparadise familyParadisaeidae ispresentlyconsideredtoconsistof
42 species. The family includes species groups and species exhibiting morphological,
ecological, zoogeographical and behavioural characters of fundamental interest to many
disciplines ofscience. Over recent years there has been an exponentially rapid increase in
interestinbirdofparadisebiologyand systematicsatalltaxonomiclevels. Biometricaldata
presentedhereareby farthe mostcomprehensiveanddirectlycomparativegatheredforthe
Paradisaeidaeatthefamily,generic,specificandsubspecificlevels.Meanvaluesandranges
of measurements of each subspecies are given, and interspecific and intraspecific size
variation and sexual dimorphism discussed. These data and others are used to assess the
validityofbirdofparadisesubspeciesdiscussedby recentauthors. Paradisaeidae,Birds
ofParadise, systematics, sexualdimorphism, morphometries.
Clifford B. Frith and Dawn W. Frith, Honorary Research Fellows of the Queensland
Museum, 'Prionodura',P.O.Box581,Malanda, Queensland4885,Australia;1JFebruaryJ997.
The Australasian bird of paradise family paradise show remarkable interspecific diversity
Paradisaeidae is presently considered to consist of highly colourful and ornate plumages, which
of42 species constituting 16 genera and involv- they usein highlyritualizedand complexsolitary
ing a total of approximately 100 subspecies or communal (lek or exploded lek) courtship
(Gilliard, 1969), to which a few havebeen added displays at traditional sites, perches or courts
more recently. Birds of paradise include species (Diamond, 1986; Beehler, 1989). A remarkable
groups and species exhibiting morphological, feature of these birds, and stressing the close
ecological, zoogeographical and behavioural geneticrelationshipsbetweenthem,isthemarked
characters of fundamental interest to many dis- diversity of intra- and inter-generic hybrids
ciplines of science (Diamond, 1981, 1986; (Stresemann, 1923, 1930, 1934; Mayr, 1941;
Beehler, 1989; Johnsgard, 1994). Thus there is a Gilliard, 1969; Fuller, 1979, 1995). The vast ma-
large literature about birds ofparadise including jority of known hybrid specimens are in full or
at least nine major monographs, in addition to subadult male plumage (Fuller, 1979), but re-
more general works and hundreds of scientific cently twohybridshave been describedin female
publications (Frith, 1979). Moreover, overrecent plumage (Frith & Frith, 1996a, b). One of the
years there has been an exponentially rapid in- latter, a unique hybrid between Lophohna
crease in interest in theirbiology and systematics superba and Parotia carolae, was until recently
at all taxonomic levels (Bock, 1963; Gilliard, erroneously known as Lophorina superba
1969; Diamond, 1972, 1986; Pruell-Jones & pseudoparotia.
Pruett-Jones, 1988, 1990; Beehler, 1989; Sibley
& Birds ofparadise are so conspicuously signifi-
Ahlquist, 1990; Clench, 1985, 1992; Cracraft,
1992; Christidis & Schodde, 1991, 1992; Frith, cant to sexual selection theory that they were
& & described and discussed at length by Wallace
1992; Frith Cooper, 1996; Frith Frith, 1990, (1869) and Darwin (1871), and innumerable au-
1992, 1993a,b, 1994, 1995, 1996a,b, in press).
thors since (see references in Frith, 1979; Dia-
Birdsofparadiseattractconsiderable attention mond. 1981; Beehler, 1989;Johnsgard, 1994). In
because of the elaborate to bizarre plumage of attemptingtounderstandtheevolutionaryorigins
adult males that are related to the polygynous and functionsofpolygynous mating systems and
mating system ofmost sexually dimorphic (and the role and influence ofsexual selection within
some sexually rnonomorphic) species and prom- them, it is important to know the physical attri-
iscuity in males. For details of their general ap- butes ofthe species concerned. In particular one
pearanceandbiology seeGilliard(1969),Cooper must be aware of fundamental sexual dimor-
& Forshaw (1977), Beehler et al. (1986) and phism (in characters otherthan secondary sexual
Coates (1990). Promiscuous adult male birds of onessuch asadultmale plumage). Sexualdimor-
.
160 MEMOIRS OFTHEQUEENSLAND MUSEUM
phism in size is particularly significant as it is validbutthatmorecomparativematerial,orother
usuallyconspicuous inpolygynousvertebratesin types ofevidence, wererequired forfirmconclu-
whichmalesaretypicallylargerthanfemales,but sions. While larger series of skins of some taxa
is reversed in polyandrous species in which fe- wereavailabletousinasinglecollection,orwere
males may be larger than their multiple mates gathered together ata single institution, forcom-
(Darwin 1871). Notwithstanding the enormous parativereview ofplumage morphology this was
literature on the birds ofparadise, no study has not possible for the majority of the subspecies.
been devoted to comprehensively demonstrating Thus, we seekhere toprimarily usetheconsider-
and reviewing variation in size between the spe- able biometrical data we accumulated to assess
cies, sexes and their subspecies. No publication the validity of bird of paradise subspecies pre-
presents more than the vaguestofmeasurements sented by Gilliard (1969), Diamond (1972) and
foronly the smallest ofsamples. Most ofthem in subsequent authors (Cracraft, 1992; and refer-
fact fail to indicate any sample sizes. encestherein).Thevalueofobserveddifferences
The monograph by Gilliard (1969) is the stan- in average size between populations has limita-
dardmoderntextforcomparativebirdofparadise tions given that size is a continuously varying
measurements at the species level, but only size character. It is more useful to plot individual
ranges for some basic characters are provided specimensoverthegeographicalrangeofataxon
therein.Nosamplesizesaregivenandfew,ifany, in order to perceive any clinal variation in size,
are presented forthevast majority ofsubspecies. but this approach is most rarely applied and is
Cooper & Forshaw (1977) presented measure- well beyond the scope ofthe present study. It is
ments for 'five or more specimens' of nominate clear,however,thatbiometricdataavailableprior
subspecies of each species only. The only sub- to this study are grossly inadequate and have led
groups for which reasonabledata have been pre- tonumerouserroneousstatementsconcerningthe
sented in recent years are those of the sizes ofsome characters ofsome taxaandothers
Glossy-mantled Manucode Manucodia ater concerning supposed sexual dimorphism in size
(Gilliard, 1956) and of all Paradisaea species (see below).
(LeCroy, 1981). As the latter two studies were The present re-evaluation of bird of paradise
based predominantly upon specimens in the taxa in the light of significant biometrical data
American Museum of Natural History, they un- setsis timely bothin termsofcomplimentingand
derstandably present relatively small sample assessing Cracraft's (1992) revision ofplumage
sizes compared to those presented here. morphology. Moreover, it provides basic evi-
Several authors have discussed the validity of dence supporting the subspecific taxonomy tobe
various bird ofparadise subspecies with respect followed in a forthcoming monograph ofthe bi-
&
to plumage morphology and relative sizes ology of the group (Frith Beehler, in prep.).
(Gilliard, 1969;Schodde&McKean, 1972, 1973; This is desirable as it will not be possible to
Diamond, 1972; Coates, 1990; Cracraft, 1992). present and discuss the significant supporting
Cracraft dramatically revised the systematics of biometrical data in that forthcoming work. Ac-
the family by applying the 'phylogenetic species cepted genera and their sequence are those of
&
concept* (Cracraft, 1992 and references therein). Beehler Finch (1985), with the addition of
This inlargepartinvolvedCracraftreviewingthe those taxaextralimital toNew Guinea and there-
subspecies acknowledged by Gilliard (1969)and fore not dealt with in that publication. As taxa
elevating the vast majority of them to 'species' accepted by Gilliard (1969) form the basis for
level. Cracraft (1992) considered 25 of the bird contemporarystudies, we followherehissystem-
of paradise subspecies recognised by Gilliard atics and scientific names at the species and sub-
(1969)tobenot 'diagnosablydistinct',and there- species level (more recently described taxa
forereducedthemtosynonomyinaccordwithhis accepted). The latter are presented in chronolog-
phylogenetic speciesconcept. Herecognisedone ical orderofdescription.
'species' named since Gilliard's monograph Data sets herein provide a significant resource
(Epimachusfastuosus ultimus, Diamond, 1969) to students ofthe systematics and biology ofthe
and named two new 'species' himself{Manuco- groupand alsotothoseinterested insexual selec-
dia [keraudrenii] aruensis and M. [keraudrenii] tion and size dimorphism in an avian group (in-
diamond!) cluding both polygynous and monogamous
Alloftheaboveauthorsexpressedthe viewthat mating systems) or avian biometrics in general.
anumberofsubspecies(validorinvalid 'species' Beyond that, a data set ofthis magnitude for an
ofCracraft, 1992) were possibly or probably in- entire avian family will prove ofvalue tobiology
HIOMETRJCS OFTHE BIRDS OFPARADISE 161
students wiih an interest in moregeneral studies small steel decimal rule. When the central and
of .systematica speciation. sexual and enher di- outer tail reclriecs arc the same length, ornearly
morphism. so (<3mm difference), only the longer mcasuu
men! (usually tail length) is presented as 'tail
METHODS length', unlessthere is mtrageneric variation, and
thenboth 'tailcentrals*and 'tall length*arcgiven.
During arecent worldtouroflarger collections Othermeasurementsweretaken with steelelec-
ofbirdofparadise skins (be authors examinednil tronicdigitat verniercaliperstothenearerwhale
sexed specimens from a recorded locality. Mu- decimalpointandcheckedandzeroeddaily. "Bill
seum specimens were examined at or from (on length1 is from the union of ihe bill with the
loantous)the followinginstitutions; Queensland fore&kull to iheupoflheupperraandible,and "bill
Museum, Brisbane; Australian National Wildlife v. nkh iron (he -interior nostriledge. 'Total head
Collection, CSiRO, Canberra; Australian Mu- length' isthemaximumdistancefromthebackof
seum, Sydney; Museum ofVictoria, Melbourne* the Skull lo the lip ofthe uppermandible It was
South Australian Museum, Adelaide; Western measured only from specimens retaining sufii
Australian Museum. Perth.AmericanMusei> cienl ofthe rear skull to permit it. It is possible
Natural History, New York; The Natural History some specimens that appeared complete in thfa
Museum, London; Museum of Vertebrate Zool- respectmaynotinfacthavebeensoandtherefore
ogy, University of California, Berkeley; The the total head length figures presented must be
Field Museum, Chicago; Museum ofCompara- ieredminimumones. 'Tarsallength'isfront
tiveZoology,Cambridge; BernicePauahi Bishop the itttertam] joint 10 the lower edge of the last
Museum, Honolulu; Royal OntarioMuseum,To- undivided scute (scale) before the toes diverge.
ronto; Academy of Natural Sciences, Philadel- Where possible, all ofi1ve.se measurements wetc
phia, the Carnegie Museum of Natural History. taken from a total of3677 museum specimens
Pittsburgh; Peabody Museum. Yale University, Weights, recorded upon death of the- bird, were
New Haven; Delaware Museum of Natural His- noted from the labelsot measured specimens
tory. Wilmington, National Museums & Galler- alsofromthoseofanadditional408 skins thatwe
ies. Liverpool; Nationaal Natuurhistorisch didnotmeasure(totalspeamensbeing6085).We
Museum. Leiden. Swedish Museum of Natural also include additional weights in = 298} ob-
History, Stockholm, Zoologist Museum tained from living birds by ourselves or others
K.0hcnhavns Univcrsitct, K0benhaven; Museum trapping and releasing them.
national D'Hisloire Naturelle, Paris: Staatliches in certain bird ofparadisegroupsspecieshave
Museum ftir Ticrkunde. Dresden, Zoologische peculiar or unique feathers that provided addi-
sSteauamtssBaemrmlilnu;ng,MuMsuenuchmenA:leZxoaonldoegirseKhoeenMiug-. ptoiosnsailblmeeaisnulrraesmpeencitfsicusevfaurliaUti\otna.rt eIxnamMicnmaiuitiodiu
Staatliches Museum fur Naturkunde, Stuttgart; keraudreniiwe measured the lengthof'eartufK
Forschungsinstitut und Naturmuseum offeathers from theirposteriorbase to the lipol
Senckenberg> Frankfurt; Zoologischcs Institut the longest with a small unstopped steel rule.
und Zoologishes Museum, Hamburg; Museum Likewise we measured the structurally similar
Zoologicum Bogoriense. Bogor: National Mu- tuft of feathers at the base of the "flag' tipped
seumandArtGalleryofPapuaNewGuinea, Port occipital plumes in adult male Parotic species
Moresby. and the analagous single elongate 'ear feather
Asample ofatleast25 individualsforeachscx found in female-piumaged Pteridophora alheriL
and age class for each subspecies accepted by The maximum length of the longest occipital
Gilliard (1969) was measured where possible. plumesofbothPamtiaspeciesandPt.albrrtiand
Measurements were all taken in the same stan- Ihe WBSt length ofCnemophihtsmacgrvgoriiwere
dard way with the same instruments by CBF. likewise measured. The maximum length of the
'Wing length* is the flattened and Straightened modified upper wing coverts or ^standards' of
chord and was measured with a stopped Steel Semioptera wattacei are also presented. The
decimal rule. 'Tail centrals' is the maximum length oi pectoral flank plumes (of Seleucidis*
lengthofthelongestofthecentral pairol rectnees Paradisaea, etc.) are rather more subjective and
from the pointoffeatherentry into the skin to its giveonly an indicationofrelative flankplumesize.
tip, and 'tail length' the same but tothe lipofthe Tins involved laying the male specimen on in
longest tail featherotherthanthecentralpair.Tail backwiththeendofasteel ruleatthethetipofthe
measurements were made with an unstopped tailandsubjectivelyassessingtheaveragelengthof
,
162 MEMOIRS OFTHEQUEENSLAND MUSEUM
!hc bulk of plumes iexcluding odd finer and no longercoverHie culmen to the bill tip. Thisis
longer feathertips)prowling beyond the tailtip. not a measurement ofa discrete structure and is
In Tables 1-42 we present mean values, sun far more subjective (and variable) than 'bill
darddeviations, rangesand sample .sizes loreach length' (from anterior margin of the skull to the
measured character ofeach sex and age class of bill tip),ThustheculmenmeasurementsofGilliard
all subspecies, and for each species as a whole. (1969)andCooper& Forshaw ( 1977)areconsis-
Data for markedly sexual dimorphic species in tently shorter than ours and we will not allude to
which males may have an adult, subaduJt (trace them again unless to note their inherently mis-
to almost completeadull maleplumageintruding leading nature tsee Lophorina \nperhu below),
minalloe-fteympael)eplpulmuamaggeea]reapnrdesiemnmteadtusreeparmaatleely(ffeo-r =Tahdueltf;olSlAow=ingsuabbabdruehvi:atIi=onsimamraetuusreed;bMelWowL; A=
each male age group- Data for females of those mean wing length; MTL - mean tail length.
species whose plumage is generally similarat all MTCL= mean tail central length; MLL = mean
ages, are presented collectively. The latter are tarsal length, THLM=BWtotal head length; MBMLW=
referred 10 as adult females despite the fact that mean bill length; = mean bill width.
sampleswill inevitably include some individuals - mean weight; METL = mean ear lufi length,
lessthanadultgiven thegreat similarity in plum- MOPL = mean occipital plume length; MFPL =
age, but data for individuals smaller than adult mean flank plume length.
size (i.e. juveniles to immature*) arc excluded.
Data forf basically monomorphic species, in RESULTS AND DISCUSSION
IVhidiadultmalesandadultfemalesaregenerally
similarhut have adisecrnihly different immature PARADISAEIDAE
plumage, aregiven separately foreach immature CNEMOPH1LINAE
sex. Asingle exception isLoboparadisea sericea
for which we combine data for immaturcs ;nni (hemophilus macgregoriiCrested Bird of
subadultsofbothsexesastheverylewspecimens
ofbirds inthelatterplumagedidnotwarranttheir Paradise (Table 1).
separation. A9 similarly sized to or slightly smaller than
In the species accounts we cite the original Ad. and of similar proportions, MWL and MW
description and type locality of each taxon after hy 4 & 7% respectively. MTLsimilarin A2,Ac?
the species name, if monotypic. or after each and \6. Thus, <$ do not acquire a progressively
subspecies name. Early synonyms are not pre- shorter tail with age (as in C. loriae), MTI
sented. We discuss \hc biometric data for the proportion of MWL similar in both sexes, 80 &
species as a whide, and forsubspecieswhere itis i respectively. MLLandMBLofA? 3 & y%
necessary to point out differences in size and Shorter than In A3 respectively, but similar in
proportions between them, We do not here fully proportion to MWL in both sexes, Crest lengths
describe plumages of nominates ordescribe and ofAd and SA<3 average 34 ±4mm (n =51) &
discuss distributions of ihe various Utta unless 32±3mm(n=9)respectively,butshorterin(I)J
pertinent as these arc widelv available (Gilliard, and A2, averaging 19 ± 6mm (n = 22} & 17 ±
196*); Cooper & Fnrshaw. 1977; CracraJf 1992) 4mm (n 25) respectively.
amnodnowgirlalpbhe(Fprrietshe&nteBedeJatlneerw* ininpraep,f)o.rthcoming Re1p)orCt.oMfl.BnntiiasehgtNegeowriiGuDieneVais1,88188-9809.:A6n2n.uMall
Because bird of paradise measurements pre- Knutsford (erroneously MtMusgrave in Gilli.uu
sented in Gilliard (1969) and Cooper& Forshaw 1969), Owen Stanley Mis. Wing lengths ol 116-
(1977) are those rcfered to in the contemporary 121mm for AJ and 1 15-120mm for A9 pre-
IuHogieal literature we make specific obser- sented by Cooper & Forshaw (1977) are
vations upon them as and when required. Our inexplicably long given that our measurements
relatively large sample sizes enable us to criti- are ofmaximised wing lengths.
C
cally evaluate previously published assessments 2) m, sanguineus Ircdale, 1948. Australian
od several size-related characters and have in Zoologist 11 162. Kumdi, Ml Hagen District-
some cases found Ihemmisleadingorerroneous. Like nominate but generally more red, being or-
Gilliard (1969) and Cooper & Forshaw (1977) ange-red about head, less red on back to orange
present 'culmen length'asopposedtobilllength. rump, underparts with less copper-red SUffitsion
Culmen length is thai oftheexposedculmen and MWL slighuj iorger. but MTL slightly shorter,
is taken from the point where forehead leathers than nominate.
1
BIOMETRICSOFTHE BIRDS OFPARADISE
163
3) C m. kutortwto Mayr & CJilliard, 1954. 3] Cnernophilus l. inexpectata Junge, 1939.
Bulletin of the American Museum of Natural NovaGuinea, new series 3 77.Bijenkorf,Oranje
History 103: 361. Mt Grata, Kubur Mis. Up- Mts. Iridescence of temals of Ad more groen
perpaxtsofA d very like sanguineus but slightly than The blue of loriae or the violet-purple uf
paler, less saturatedwilhredand undcrpartsmore amethvstim butA? asothersubspecies. Simil.u
black, less suffused with red-brown. Characters in site to nominate, with MTL only slightly
supposedly distinguising this subspecies from shorterthannominateand(moreso)amethyuina.
sanguineus have been doubted (Diamond, 1972,
Bechler in Coates, 1990; Cracraft, 1992); pre I ulw»paradLsea sericea Yt IK a-breamed Bird
foundaMt GiluwespecimenofC in. Sanguineus ol Paradise (Table 3
MWL
showing these character and MTL. i if A9
kuboretms slightly shorter than sanguineus, but (uniquely in the Paradisaeidae) slightly
sMaomrpplhesoltooogysmadloleslorsumgeagneisntg,fuhlowcoemvpearr,isotnhsa.t MlarWgeLr tahnadn,MbuWt si(msialmaprllye psrmoaplolr)tiboyne2d t&o, .
kuborensis is invalidand should be synonomised respectively. MTLofA? and Id slightly lonI ger
with sanguineus (4 6l 7% respectively)thanAd .Thus, d acquire
cbforr4roo)dumegdhtC,ht.eawKnhmriA.acedhtskusvebspaseMspctesipc,amileeewnsrheitnrh(etaohnltelhiWfeaiaueshllpgdeics(ai1iHe9dsi6gt9ihos)lcuaonwnrmadeess- aMMtiTvWpeLrlLoyg5riM9neLs&bsoLit$vhseWilsmyei*loesa-flrsiMigMnhWtlBlLeyLngistonhhfoAAratdne9drai5nnt%MdapirlAWloo?pwLnoigrtretherisotpanhegecat.-no
birds. He felt that ifthe specimen was not faded inAd, but similar in proportion to in both
it might represent an unknown subspecies. An sexes.
Ad specimen in Zoologishe Museum, Hamburg 1)L. s. sericw Rothschild, 1896.Bulletinnfthe
from Malingdam, nearMtGoliath, Irian Jaya, we British OrnithologiM-' Club fv l ft. 'Dutch New
examined also fits this description but could pos- Guinea'. Restricted to the Wcylund Mts (M .
sibly be the result of immersion in alcohol, In 194 1 ) v said to be larger than d (age UnSpcd
wing, tail, tarsalandbill lengthsmeasure 115, 86, lied) in the Weyland Mis bul not so on Mt
43.6 & 29.4mm respectively; bill width 6.3mm Karimui (Diamond, 1972). Certainly 9 fromthe
and weight 90g. These measurements are within Weyland Mts have longer wings and tails, than
the ranges ofothersubspecies. those on Mt Karimui but Diamond's samples are
toosmall (Ad x 1; Id x 1 8 x 2] formeamnglul
comparisons.
Cnemophilus loriaeLoria's Birdof Paradis* 2) L s. aurora Mayr, 1930. Omithologische
(Table _ Monatsberichte 38 147 Dawong, Herzog Mts,
Plumage of upperpans significantly brighter
A?
AMdW,a5ns%dimoiflliasgirhmtteinrl.oarrMpoTrnolLpyorfotrfiaocAntsi?,onMaalWnldyLsImsdailmlisellriagrthhbtaulnty sa(ipnamdlielmrao)r,rtemoogsrreeereinbciresoah.wnaL-naydreglleelsroswtb,hraotnhwenn,ocmrbiuontwanutnefd,acrrMppaWarltLesr
longer (3 & 5% respectively) than Ad. Thus, d and MTLofAd by 3 & 1% respectively. While
MacTquLir7e1aapnrdog1re4s%sivoeflyMsWhoLrteirn tAaidl waitnhd AaAg?e9. rneojtecdtiDsicuasmsoinndg'sit,(1C9r7a2c)raaftttri(b1u9t9i2o)n oafpbpieradrsefdrotmo
MreLspLectainAvdedlMy,BLsoopfroApo?rtinoengaltiegliyblylo(n2ge%r)MisWnhoLrter nthoemiMntatKear(iwmhueirearweae, hPaavpeuaalNseowinGculiundeeda (tohetmh)e
than in , but similarin proportion to in andconsidered them to be L s. aurora If. how-
both sexes. ever, they are aurora, theirmeasurements w
1) Cnernophilus I. loriae Salvadori, 1894. An- be at die lower end of the si/e range fen
nals Museo Civico Genova. ser 2. 14: 15 1. subspecies.
Moroka, Owen Stanley Mts. Iridescence on ter-
Ad
lialsof blue PARADISAEINAE
2) Cnemophihts i amethysttna Stresemann,
1934. Ornithologisehc Monatsberichte 42: 144. Mftcgregoria pulchra Macgresor's Bird of
Schraderberg, Septic Mis. Iridescent upper sur Paradise (Table 4T
facesoftertialsofAd deepviolet-purple,butA?
abssuutbostMpheeWcriLessu.abnsdpeMciTesLsSliimgihltalry ilnonsgie/re tthoannomininoatthee,r psomarAltlit)onbmeyadr1t1ko..edA11ldy&,s2Mma6Wl%lLer.restMpheaTcntL,ivbaeunltyd.sMiImdiWlaarirslnaylmSpprAlode-
164 MEMOIRS OFTHEQUEENSLAND MUSEUM
have notbeen collected or are difficult to differ- inate but distinct in being much darker overall,
entiate and so it is unknown ifMTLincreases or much more glossy blue-green, darker and more
decreases with age. MTL 76% ofMWL in both blue-black oncrown and uppertail. Farmore dif-
sexes.MLLandMBLofA9 8 &6% shorterthan ferent from nominate than morotensis and possi-
Ad
in respectively, but similar in proportion to bly approaching species status (Cracraft, 1992).
MWL
in both sexes. Concealed bases ofprimaries with trace ofwhite
Thissexualsizedimorphismisdistinctlydiffer- only. Oddly Lambert (1994) incorrectly wrote of
ent from all three members of the subfamily obiensis that birds have a whitish streak above
Cnemophilinaeandisgreaterthanthemajorityof and behind the eye. Larger than nominate in all
polygynousParadisaeinae.Itsextentisunusualin measured characters. MTL71% ofMWLin both
a monogamous bird of paradise (Rand, 1940; sexes and thusproportionately shorterthan nom-
Beehler in Coates 1990). inate.
1) M. p. pulchra De Vis, 1897. Ibis 1897: 251,
Wpii.n7g. MletngStchrsatocfhl1e8y8,-s1o9ut3hm-emasftoerrAn?Ne(wCoGoupienrea&. Manucodia atra Glossy-mantled Manucode
(Table 6)
Forshaw, 1977) are inexplicably large given our
measurements are maximised wing lengths.
2) M. p. carolinae Junge, 1939. Nova Guinea A9 similar to or slightly smaller than AS in
d(iNfefewreSnetrfierso)m3n:om8i2n.atOer,anMjeWMLtsa.ndPrMopTorLtisohnoarltleyr MsiTzeL, parnodpoMrtWionbsy a4,nd6 n&ot1ab3l%y rweespiegchtti,veMlyW.LI,S
by 5 & 6% and 17 & 16% and MLL longerby 5 and 19 smaller than respective adults in MWL
& 4% in AS and A9 respectively. Weights of (6%) & MTL (7 & 5%), and lighter (sample
carolinae heavier than pulchra, but samples too small).Thus,taillengthincreaseswithageinboth
small for meaningful comparisons. sexes. MTL 84 & 83% ofMWL in AS and A9
respectively. The tail/wing index for both sexes
Lycocorax pyrrhopterus Paradise Crow combined ranges from 84-89%, these figures
similar to those given (82-88%) by Coates
(Table 5) (1990). MTL longer than other Manucodia spe-
sMiTzAeL,9parnsoidpmoiMrlatWriotnobsyora3,nsdl4ign&hotlty1ab2slm%yalrwleeespriegtchthtia,vnelMAyW.dLI<i,Jn c3siiem&si,lae5rx%cilnuspdhrioonrpgtoerMrt.itochnoamntroiiMinWMALLdLinraebnsodpteMhctBsievLxeelosy.,fAbu9t
have not been collected or are difficult to differ- Since a thorough study by Gilliard (1956), re-
entiate and so it is unknown ifMTLincreases or sulting in the acceptance ofthe three subspecies,
decreases with age. MTL 73% ofMWL in both onlyCracraft(1992)hasreviewedthe species. In
sexes.MLLandMBLofA9 2 &5%shorterthan assessing plumage variation, from the phyloge-
AS
in respectively, but similar in proportion to netic speciesconceptpointofview,Cracraftcon-
MWL
in both sexes. sidered there to be no diagnostic characters
1) L p. pyrrhopterus (Bonaparte, 1851). Con- within M. atra populations. We agree with
spectus Generum Avium 1 (1850): 384. Gilolo I. Gilliard, however, that it would be misleading to
Concealedbasesofprimarieswithnowhite.MTL include all populations in a single taxon in the
75% ofMWLin both sexes. face of what in fact are quite gross plumage
2) L. p. morotensis Schlegel, 1863. Ibis 1863: differences apparentat the extremes ofthe range
119. 4Mortag'. Much like nominate but paler, of this species and the clear differences in size
slightly more brownish abovebut slightlydarker demonstrated here. We therefore follow
below. Concealed bases ofprimariesextensively Gilliard's treatment until genetic studies provide
white. Significantly larger than the other two definitive answers to this complex problem. Our
6su9b%speocfiMesWinLalilnmeAta?suarneddcAha9racrteesrpse.ctMivTelLy,7a1n&d ameawshuorleemeangtrseeofclwoisneglyanwditthailtfhoorsetheofspeGcilileisaarsd
thussimilartoobiensisbutproportionallyshorter (1956);buthisexposedculmenisnotcomparable
than nominate. MWLand MTLofAS 15 & 9% with our bill length and his tarsal length range is
and7 & 8% longerthanin Ac? pyrrhopterusand larger than ours, indicating differing measuring
obiensis respectively; these figures slightly less techniques.
(12 & 4%, 6 & 3%) forA?. 1) A/, a. atra (Lesson, 1830). Voyage of the
3) L p. obiensis Bernstein, 1864. Journal fiir Coquille, Zoology I: 638. Dorey, north-western
Ornithologie 12:410. Obi Is.Generallylikenom- New Guinea.
) 1
BIOMETRICSOFTHE BIRDS OFPARADISE 165
2) M. a alter Rothschild & Harlerl, 1903. but samples too small for meaningful com-
NovitatesZoologicae 10: 84. Sudest I. Flanksand panions.MTL71c7c ofMWLinboth A3 andA9
belly Markedly more violet than atra, Notably Tne adult Uul/wing index for both sexes com-
largerthan nominate. Ad by 12 & 15% and A1?' binedrangesfrom70-76% MLLandMBLofA9
9 & 12% in MWL and MIL respectively. MLL. 3 ft &t shorter than in .Ad respectively, but
THL and MBL of A 6 exclusively larger than similarin proportion toMWLin both se
nominate and nearly so in A9. GilLianfs 1969) Ad tarsal lenath of57 mm
3) M. a. subalter Rothschild & Hartcrt, 1929. andCooper&1 Forshaws*H977) of54-59 mm fa
Bulletin of the British Ornithologists' Club 49; both sexes are very long.
1 1U Dobbo, Am Is. Said to average much more 1 M. c. comrh ScfaMer, PL, 1876. Proceedings
purple and violet with oil-green rare on adults of the Zoological Society ofLondon 1876: 459.
(GUIiurd, 1969). On average larger than nomi- Huon Gulfin errorfor Fcrgusson 1
nate, but smaller than alter An analysis more 2) Al c twbriandiMayr. 1936. American Mu-
ttohotresotugthhethpaonssiIsbiploistsyibtlheathebirredissarreequcilriendalininorsdiezre sPeluummaNgoeviatsaiensom8i6n9ate3 wKiatHheuMnaW;LTraonbrdiaMndTLL
hoin larger ones in the extreme south-east of Shone* by 7 & 9% and 6 & 7 % hi A d ami
Papua New Guinea to smallct ones toward the respectively. Thus, this subspecies ra on average
cureinc souiheaMe.m-most part of the range Of smallerthanthe nominate with limitedoverlapin
the generally smaller nominate on the Gulf of size ranges of wing and tail in each aduJl
Papua. Takingthisandthetwo island populations' all', p-
atry into account it seemson balance moie useful
Manucudia chalybataCrinkle-collared to recognise their differences than to conceal
MsUUCOde (Table 7) them, notwithstanding their possibly qnjesil
ahte status(Cracraft, 1992).
M. cintlyhaiu Pennant. 1781), Speeium
t
Faunula Indica. in Forster's Indian Zoology
1781: 40 (based on Daubentou, Planches Manucodiajobiensts Jobi Manucode
Lnluminccs. pi. 634). New Guinea, restricted to ' -Table9
the Arfak Mts, Monotypic, A? similar to or A9 Ad
swrleeisigpghehtctlt,yivseMmlaWyl.Lle1.r6tMhaaTnndLA19aondsinmaMslilzWee,rpbtryhoapn3o.rrte4isopn&esctaiMnved sMiT/eL. parnsoidpmoiMrlatWriotnobsyora4,nsdl5ign&hotlt2yab1sl%myalrlweeesrpieg[chtbtia,vnMelMWyW.LLII,dD
adults in MWL(2&4%)and MTL(6&4*),and a&nd2%19) samnadllMerTtLha(n4r&esp6e%ct)i.vebuatduslatsmpilnes smal(l3
lighter (samplesmall).Thus, tail lengthincreases
iwnitAhdagaenidn Abo9thrseesxpeesc.tiMveTlyL. 8T2he&a8du1l%t otafiMl/WwiLng TMhTusL,7lu6il&le7n5gt%hoifncMreWasLesiwniAth6agaendinAbo9thressepxeecs-.
index for both sexes combined ranges from 81- - Tic adull tail/wing index For both sexes
combined ranges from 72-76%, these figures
64^ these figures within the rangeofthose (7X-
ob8uf6tA%sS.)impir2leas&reni5tne%pdrbosyhpooCrrottaeirtoentShta(on1M9i9nW0)AL.dMinLrLbesoptaehncdtsieMvxeeBlsyL., CSwhoioatrthteemSrt(hIf1et9air9la0)nfM;gaeWMAoLLdfLthroaessnepde1c6Mt9iBv-eL7l8y,%of)bAuptr9essie1mnit&leadr5bi%yn
The tarsal lengths of41-45mm for A6 and of proportion to in both sexes
3&-46rtun for A9 ofCooper & Forshaw 1977) We pan geWl8ll] IfcCOtJCilft our measurements
do notagree withourmeasurements.Thiss(pecies with those forthe species by Gilliard ( L969 .. bur
abeisiulatmdloiltfsvhfteiscialulmallittlteeatxrrotseadprniendfcaflioeevrsmeeonrdtrloiapaephtseoihlnfaomrvgooeysmtaaMsm(.e5ta*hJs6eoufy%rr)aieermsnehesOnhgftels&nl,- onlkofoi*rj1Ata6at8rd-salu1las7nwld3iotmr1hm6A5Cf-9ooo1r7opwfe2irnmnog&mmiafnFnoaodrtre4ws,hi2an-wwg4hsi'4acnm(hdm19a4-fr7ole7r)Metx5ascrimlzsueum-ss
toil proportionate to wing length. sively too small (wing) or too large. (DitfSUS).
1) A/ / johwnsts Salvador!, 1876. Annals
Manucodia comrii Curl-crested Manucode Museo Civico Genova 7 (1875): 969 WWi.ipi.
(Table 8 Jobi {- Yapen) I Only known from Yapen I ;n
j Gcelvink Bay, west Irian Java.
A9 slightly smaller than Ac and of similar 2) A/. ; rnbiensis Meyer. 1885. Zeitschrifi Ittr
proportions. MWL and MTL by 5 cV 4'v respec- die gesammte Ornithologic 2: 374. Rxfu. Geet-
tively Id and19 smallerthan respective aduHs. rinkBfr) Now known from the mainland
IM MEMOIRS OFTHEQUEENSLAND MUSEUM
of northern New Guinea adjacent to Yapen 1 sjbspecific diagnosis (Gilliard, 1969; Cmci
eastwardtotheAstrolabe Bay areaofPapuaNew 1992) Datafor 1 subspeciesarepresented,eight
Guineaandsouth inthewesttotheupperSetekwa as acknowledged by Gilliard (1969) and the two
R and in the east the upper Rarnu R. MWL and described by Cracraft (1992).
MTLshorterthan nominate, by 5 &6% and 3 & M L keraudrenii (Lesson & Garnoi), 826
ZL; in A <$ andA? respectively. Thus, the main- BulIl)etin Scientifiques Naturelles (Ferussa1u 8:
kind birds are on average smaller than those on 110. Dorey, Vogelkop, north-western New
Yapen I(Gilliard. 1969). As samplesofthe latter Guinea METL 19 ±4mm (n * 24) & 18 ± 2mm
mall (n=6),andwingar/Jtail lengthsranges (n =15i in Ac? and A? respectively. MTL 78%
oi mainlandbirdsin factentirely overlapthose of ofMWL in adults.
the nominate island form, the validity of this 2)M.LgouidI(Qrey^ 1859, Proceedings ofthe
subspecies remains in doubt- Gilliard ( 1969) Jo Zoological Society ofLondon note, p. 158. Cape
scribed rubiensis as similariojobiensis (<3 wing York. Like nominate but plumage iridescence
length of latter 177-]79mra, tail !32-I42mnU more green, less purple (particularly so on the
hut smaller (6 rubiensls wing length 168- upperwing and tail). Ear tuft feathering more
e1q7u4amtem,totjauilst1if2y7-t1he33rmetme)n.tioPnreQsternutbideantsaiasrcanidnaodn- 2na9r±ro4wmlympo(innt=ed1,6)MiEnTAL<53a4n±d Abr9nmre(snpe=cti3v3e>lyf.t
the basis of this we agree with Cracruft (1992) This is much longer than in keraudrenii, but not
thatit should be synononused vA\hjobiensisl but longerthan inpurpureoviolaceus{paceCm
more dataare clearly required. 1992). MTL proportionately long. %l% ofMWL
in adults
Manmrodia keraudreniiTrumpetManucodc 3 M. Ljamesii (Sharpei. IS77. Catalogue of
i
(Table 10) Birds in the British Museum 3 181 Aleva, Wall
Suund, British New Guinea. Throat and breast
AS similar to or slightly smaller than Ao in dark metallic blue, washed with green, lacking
Msi/T.eL. parnodpoMrtWionbsy a4.nd5 n&ota1b2l%v rweesipgechtti,veMlyWLI,o ipnuartpel.eMoEfTn-LjMP2n7.i±ir5,mEami(cnuf=is25l)oomgemj l&haj2i4I±n B7OmItmh
and J5 smaller in MWL (5 & 3%) and lighter (n =MW25L) in AS and A9 respectively. MTL 78%
(sample small) than respective adults, but only of in adults.
IdMTLnegligiblyshorter(2%).Thus,tail length 4)A/. k hutt&etoi($h»rvsy i882. Journalofthe
increasesonly slightly withagein 6,thoseof19 Linnean Socie mdon 16: 442- East Cape,
andA9 beingsimilar. MTL78*. ofMWLinboth NewGuineainenor forNorma,nbj I. Back, wrop
sexes The adult tail/wing Index tor both sexes and uppenail are dark bluish-purple (less green)
combined ranges from 73-78*%% these figures and the ear tuft feathers are less blue and n
being within the range of those (70-81%) pre green than nominate. Generally like nominate,
s3cn&icd5%bysChwurtteesr [t1h9a9n0]inMALdLarensdpeMctBivLeloyl, hut b1u2t%muancdh2l0ar&ger1o5ve%railnl,AMSWaLndaAnd?MrTesLpecbtyiv1e8ly&,
Simjbir in proportion to MWL in both sexes. Lai tufts also longer, METL 26 ±_3mm (n = II)
Proportionate bilJ length is remarkably consis- & 22 ± 3mm (n =S) in Ad andA9 respectively.
tent, MBL21% ofMWLin all subspecies except MTL 74% ofMWLinaduhs SO propOrtKmaJteiy
ktmsiemi andgouidi for which it is 20%. theshortest ofall subspecies.
A difficult spe iiqileX Within which 5) M. k. purpureoviolaceus (Meyer, 1885).
Gilliard (1969) acknowledged eight subspecies, ZeitschriftfurdiegesammteOrniibohtgie2: 375.
which Craerafi (1992) reduced 10 seven pi. 15. Astrolabe Mts. Generally like nominate
('Specks')bysynonomising\t.Ltuuyen with M, but larger with back, breast and belly intensely
trpureovwlaceus, To these, Cfacraft 1981) -cent violet-purplish. Eartufts notahls long
a(Id9d9e2dj LefWfOorttstenwoltywidtehssctrainbdeidng,fomrmosr.e Ccroalcl1reacftti'nsg MinEATc'Lu3n8d±A6?mrmes(pie)c=ti3ve5l)y&. M3T4L±746n^ino(tnM=W29L)
andstudy ofthivspee.esarerequiredto meaning- in adults.
fully understand distributions and lntraspeedTc 6) M. k. neutnanni (Reiehenow, 19J8). Jew
variation given the paichiness of collecting on furOrnithologie66: 438. Lordberg. Litesimilar-
n:ainiand New Guinea and the subjeciivc nature sued nominate but lower back, rump, uppeitail
i -imagecharacters used(colourandquality of and wings dark black-bluish washed iridescent
retracted light in the form of iridesenee). The deep violet-purple, not moregreenish. Bfea$l
a ofthe laceolale 'earlulls*arc also used in belly dark metallic blue. METL 13 ± Imm I
2
BIOMETRICS OF THH BIRDS OFPARADISE 167
9)& IJ ±lmm<n=8)inAd andA9 respectively Paradigallacarunculata Lone-tailed
andsimilarto thoseofanotherwiseoverall larger Paradigalla (Table 11)
adelberti. MTL 80% ofMWLin adults.
oMNfeo7tar)hroeM-b.Nciekd.weDnmiEtsantiygrcrliaiaal,n(dGrnZoeoireontnlhwo-ageyias.csiat/l1e9erC4/nl2pu)lN.buePr1wr9no:ac5gGe1cu.eidWniaenuagt.,s0 tss(ihu1voAreBre!St%n>eXireTmtmaTahsihra,luknsseMM,odfWlTdyILCdsLamacsainqinludmliiAeMrlrae9Trta(Lih5nap%nbrl)yoe,Agni1g1d*1tdehs#is(tni7ov5%meA%Jloys9art.nledMosmpnAeTegaceLd-r
purpurecviolaceus and ear tufts also notahiv
Ml=uWn4g;.LiMnEATdLa3n8d±A19lrrnersnp(enct=ive7l)y&. M37TL± O7r8n'rn (o tatahbiellywriientmhalieanngdgeet,rh.t(hMe20cTe%Ln)tra7a1nldp&aa.itr7a6ifn%acrrogefraesMaitnWegrcLroanlisenitdAhe*arn
in adults. The supposed evidence of a and AV respectively (contra P. brevicauda\, so
phti4grhpeurreowviinogla-ctcanisl (Giinldleiuxrd,in1m9a69y)riistmheaangrien porfoApo?rti7on&at2el%y slhoonrgteerritnhatnhein9A. MdLrLespaencdtiMveBlyL.
MWL
given limited numbers of specimens o\' the for- but as a proportion of negligibly (2 *N:
mer. As no other characters including ear lufl longer.
length appear to differentiate them, we concur Wine lengthsof 152-157mm forAd and
with Cracraft (1992) that mayri should be 159rnmlorA9 ofCooper& Forshaw (1977)arc
onomised withpurpureoviolacvus, dramatic«Hy/exdu$ive > shorten thwitmre (bribe
8) M. k. adelberti (Gilliard & LcCroy, I9i formerand near exclusively shorter lor the latter
Bulletin of the American Museum Natural His
lory 138: 72. Nawawu, Adelberl Mts. Generally 1) P, v, canmculata Lesson, 1835, Htstotae
similartonominate but likeA/, k neumutmi. with naturelledesoiscauxdcparadisctdescpirnaqucs
upper wings and tail more green (not, orfar 1835: 242. Arfak Mts.
blue to purple) Ear tufis shorter than other foIrms 2\Pc\ intermediaOgilvie-Grant, 1913. Bulle-
eIx4cempmt (nnea-ma1)nnlio,ngMiEnTALd1a3n±d A2m9mre(snpe=cti1v0e)ly&. tUiunkowfatheR.BrNiatsisshauOrRnai,thaotlo5g.i5s0t0s'ft.CTluhbe sJtla!tu1s0o5f
MTLM80% ofMWL in adults. btheeenlornegcepnrtolbyledmiastciucsaslefdoirnmdPe.taci.l,inatnedrmewdcianhoatse
9) k antensts (Cracraft, 1992). Cludistics8:
10 Wiinnem Bay. Kobror I, Am 1, West Irian. thherreeeoknlnyowtnhat it is annsinhvaalviedbteaexnonshasowtnhetoonbley
aMdujcachenltesmsagirneleanndthNaenwnoGmuiinnaetaejaamnedsidiifbfeyrbferionmg relativ&ely young individuals of P. brevicauda
(Frith Frith, mi press] Its biometrical dnta ate
generally much darker and bluer, lessgreen. Up- therefore included within P. brevicauda.
pcrparis, particularly back, suffused purple, and This species is distinctively different from P.
lanceolate head feathering deeper, more purple, da in si/e, relative proportions and in
cobalt blue than in jamesii. Ac? slightly I; sexual dimorphism of these characters
thannominateinMWL(5%)and more so in MTL individual? of< wuncalatalMLVcashortertail than
! 10%), Ear tuft length most similar to nominate, adults, whereas in P. bm-icuuda younger birds
METL 22 ± 4mm (n = 4) in Ad and thus much I considerably longerone than adults.
MTL
shorter than jamesii. proportionately the
MWL
longest, 82% of in adults. Paradigalla brevicauda Short-tailed
10)M. k. duummJi(Cr^n\i\- 1992) Cladisfics Paradigalla (Table 12)
K. 12. Awandc, neat Okapa, Easiero Highlands &
U.sirici PftpUfi New Guinea. Similar to P. brevicauda Rothschild Hartert. 1911.
Kates Zoutogicae 18: 159. Ml Goliath, cen-
purpwwvfolaceus but defined as distinct from it tralDutchNewGuinea. Monotypic. Synonym: P.
ivniohlaevt-ipnugrpblaecks,heuepnpse,rwthiengbsreaansdt taanildwhiotlhlystdraorngk Wb.>oirntsleirgmhetdlivasm(asleleePr.tchaninAtder,meMdiWa)L, AanSdMsiWmilhayr
metallic blue with little orno violet-purple, and J & 6% respectively. MTCL only 2 & Iturn
lanceolate head feathering bluish-green as op- shorter than MTL in Ad and A? respectively.
posed to greenish-blue washed violet-purple Ear A? MTL.however,isconsiderablylonger<
tufts slightly longer than in purpurtbwioiaCeuS, thanAd MTLofIdalsomuchlonger(47%)than
METL40 ±9mm in = 1 1 ) & 34 ± Tmm fd = 41 ill and. 19 also have a longer (31%) tail than
Ad andA9 teSpdCfivel) and longer-than moitioi AV sampie small). Thus, d acquire a progres-
|
>peeies MTI | it adults y shorter (32%) tail, the central parfi
) )
168 MLMOIRSOFTHEQUEENSLANDMUSEUM
dMeTcrLea3si4n%g oifnMreWlaLtivien Aleon\gthbui[3455K«I iwnitAh9.agseo mfoournpdhiFsomrian bdiilslcluesnsgitohn, soefe'Frreivtehrs(eidn'prseesxsu)a.l di
proportionately much longerin 9 MIL of a9
5M%WLlunger than MinBALc", bui as a proportion of Ptiloris magnificus Magnificent Riilehirtl
The same. MWLsimilar in both sexes, hut (Table 15)
ainsAfl?protphoarntAioon of negligibly (2%) longer A9 markedlysmallerthanA6 >MWLandMW
A stated average tail length of 90mm for this by 19 & 34% respectively. MWL of A9 exclu-
ipecieS is too lung (pace Cooper & Forshuw. sively shorterthanAd A9 considerably smaller
1977). in MWLandMT—L(12.& %%) than Id, and much
lighter (27%) more so than the other two
Ptiloris paradUeus Paradise Riflcbird Ptilorisspecies. MTLofA9 sMhoWrtLer(3%> ihan In
(Table 13) Ad. but as a proportion of much (10%)
longer. MTLofId longer |7'X) than Ad. MTCL
P. paradiseus Swainson, 1825. Zoological and MTL similar in A9 and Id. but 59t shorter
Journal 1; 479. No type locality (= northern New in Ad. Thus, d acquire a progressively shorter
South Wales). Monoiypte A9 average smaller (6%) tail, tail centralsdecreasing (1 1%) in length
thanA(J, MWLand MW(sample small) by 10& at a greaterrate with age. MLLand MBLofA9
iihnnutbAosdtmr,haelsMlpbeUWscIatLmiapvselaleany)d,,prAMMoTpTSoLLratltoishfooanAnslo?IifgdhsMthaluWnyidtLsemlfiangl1eh7lgtve;elrri)g(iit3bh^layn aa1l0pTmra&oorspstoIarlitl'l>iueonsnnghgeootrrhftsMaeornWfdLtChnaaoenrogepilneeirAXgC&il6ub&Frlloeyvsre(psle2jhc&taiwlv3o'e(*nl1gy9)e,7lr7bount)tghaeaarrnsc
longer. MTL of Id (sample small) also ours forAd and 9 respectively.
shortIer (4%) than Ad. MTCL4, 3 & 9% shorter 1) ft nu magnificus (VieilloL 1819). Ntmwau
than MTLinA9, Id andAd respectively.Thus, Dictionnaire d'Histoire Naturellc, nouvcau edi-
d acquire a slightly overall longer tail with age, tion. 28; 167. 'Nouvelle Guinee\ restricted to
unlike other Ptiloris species, but subsequent tail Dorcy. Vogelkop.
apcgerneot.proMarlLtsiLodonefctroAeaM9sWe3Ls%lisgsihhmtoilrlytaerrinitnhaacbntoutiahnlAsledexneg.stb.hutMwaBistLha ZAuos2otlroPag.liimca.a.lalMSboeWcritLettEylolofilotLA,odnIKd7ocn1.,.lPplr.om5cm8e3e.sdChioanrgptseeroYfoartnkhd.e
MofWAL9 78%%lloonnggeerr.than Ad and as a proportion Of ManTdL.conMsLpiLcuforuascltyionmaolrley ldeescs,uravnedd btihllannainrrootwheerr
Fordiscussionof'reversed*sexualdimorphism twosubspecies. Extentofculmen base feathering
in bill length, see Frith (in press). is intermediatebetween thatofnominatenutRnift-
cus and xntercedens. MTCL ol Ad relatively
Ptiloris victoriae Victoria's Rillebird shorter (8mm) than rest oftail than in other two
(Table 14) subspecies (both 5mm).
3) p m. intervedens Sharpc. t882. Journal of
ft victoriae Gould. 1850. Procceding6 ol die IheLmnean SocietyofLondon, Zoology 16:444
ZoologicalSociety i)fLondon I849: III.Barnard Milne Bay and East Cape, south-eastern New
sIsm,alNloerrtthhaQnueAedns.laMndW,LMoannodfyMpiWo,byAi1J0a&ver1a8g%e Gseuxienseaa.ndInaaillmaogsetcallalsmseesatnhimsesausbusrpeemcieenstsisoafllbobtuht
respectively. A9 also slightly smaller in MWL identical to nominate except for MBL being e
and MTL (3 & 4%) than Id and lights (7%) 4mm less Bill stiaighter than that oialbeni-
MTL of A9 shorter (4%) than in Ad. but as a Thcrc arc striking differences in 6 advertise-
poanilpyo3rt%tosnhoorfteMrWinLAsVliaghntdlyId(47.,bu|tl5on%gers,hoMrtTerCiLn mHuennsttecianll. bientwSeheanrptche1s8u0b1s)pebcuiLesal(sfiorswtintohtiend'b.hye
Ad- Thus, although very little variation in tail populations of/' m. aihvrti (pers. obs.). la /' m,
length between Ad and 13, tail centrals do de- magnificus and R m, alberti the culmen is uu
creaseslightlyin lengthwithage.MLLofA9 6% feathered along the ridge while in P. m. inter-
•iiiiilk-i in Ad, hUl as a proportion Of MWI. monlyasmallproportionoftheculmen base
Asidmilaanrdinasboathprsoepxoerst.iMonBLofoMlWA9LV5&% lloonnggeerr.than eissunafenadthP.crmc.d.aTlbheertHiaanrkcpllounmgeersitnhaPn. mt.hematgainlifhiu-l
( Soopet & Forshaws* (1977) A9 tarsal length in /! Oa. intereedens they arc equal DO or shorter
rangeof34-39 mm is exclusively longerthan we than the tail. In view of these differences it has