Beehler et al.
1000 [Auk, Vol. 124
Frtee. Melipotes carolae, sp. nov. (Wattled Smoky Honeyeater). Original watercolor
painting by John Anderton.
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A NEW SPECIES OF SMOKY HONEYEATER (MELIPHAGIDAE:
MELIPOTES) FROM WESTERN NEW GUINEA
Bruce M. Beehler,1,2,6 Dewi M. Prawiradilaga,3 Yance de Fretes,4
and Neville Kemp4,5
1Melanesia Center for Biodiversity Conservation, Conservation International, 1919 M Street NW, Suite 600,
Washington, D.C. 20036, USA;
2Division of Birds, Smithsonian Institution, Washington, D.C. 20013, USA;
3Ornithology Department, Division of Zoology, Research Centre for Biology-LIPI, Jalan Raya Bogor Km 46,
Cibinong 16911, Indonesia;
4Conservation International Indonesia, Jalan Pejaten Barat 16a, Kemang, Jakarta 12550, Indonesia; and
5IndoPaciﬁ c Conservation Alliance, c/o the Bernice P. Bishop Museum, 1525 Bernice Street, Honolulu, Hawaii 96817, USA
Abstract.—We describe a new species of smoky honeyeater from the Foja
Mountains, an isolated range in western New Guinea (Papua Province, Indonesia).
A common inhabitant of montane forest and forest-edge in this li le-known north
coastal range, this species possesses a ﬂ eshy pendant suborbital wa le, unique in
the genus Melipotes, among other characteristics that distinguish it from all conge-
ners. This ﬂ eshy wa le provides a morphological link between Melipotes and the
monotypic genus Macgregoria, an alpine inhabitant of the Central Ranges of New
Guinea, traditionally treated as a bird of paradise (e.g., Frith and Beehler 1998) but
now regarded as the sister genus to Melipotes (Cracra and Feinstein 2000). The pres-
ence of an endemic meliphagid species in the Foja Mountains highlights the biogeo-
graphic signiﬁ cance and conservation importance of this geographically isolated
upland forest tract, which is also home to the endemic Golden-fronted Bowerbird
(Amblyornis ﬂ avifrons) and Berlepsch’s Parotia (Parotia berlepschi), a distinctive,
recently rediscovered species of six-wired bird of paradise (B. M. Beehler unpubl.
data). Received 7 June 2006, accepted 4 November 2006.
Key words: Foja Mountains, Indonesia, Meliphagidae, Melipotes carolae, New
Guinea, new species, Papua, Wa led Smoky Honeyeater.
Una Especie Nueva de Melipotes (Meliphagidae) del Oeste de Nueva Guinea
Resumen.—Describimos una especie nueva de Meliphagidae de las montañas Foja,
una cadena aislada del oeste de Nueva Guinea (Provincia de Papua, Indonesia). Esta
especie es un habitante común de los bosques montanos y los bordes de bosque de
esta cadena montañosa poco conocida de la costa norte. Una de las características
que distinguen a esta especie de las demás especies del género Melipotes, es que ésta
presenta una carúncula carnosa colgante suborbital que es única en el género. La
carúncula carnosa representa una conexión morfológica entre Melipotes y el género
monotípico Macgregoria, que habita las zonas alpinas de las cordilleras centrales de
Nueva Guinea y que tradicionalmente había sido tratado como un ave del paraíso
(e.g., Frith y Beehler 1998), pero que ahora se considera como el género hermano de
Melipotes (Cracra y Feinstein 2000). La presencia de una especie de Meliphagidae
endémica en las montañas de Foja resalta la importancia biogeográﬁ ca y para la
conservación de este bosque de tierras altas aislado, en el que también habitan la
The Auk 124(3):1000–1009, 2007
© The American Ornithologists’ Union, 2007.
Printed in USA.
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Beehler et al.
1002 [Auk, Vol. 124
For more than seven decades, New Guinea’s
greatest ornithological mystery surrounded the
unknown location of the native breeding haunts
of the Golden-fronted Bowerbird (Amblyornis
ﬂ avifrons). This species was described by Lord
Walter Rothschild from a single trade skin (an
adult male) obtained from an unknown location
in western (then Dutch) New Guinea by indig-
enous collectors apparently in the employ of the
plume merchant Duivenbode (Rothschild 1895,
Gilliard 1969). Two additional specimens made
their way to Rothschild, presumably from the
same feather merchant, once the species’ nov-
elty was determined.
A general belief held that the forests that were
home to this “missing” bowerbird would also
support additional avian novelties, so there was
considerable subsequent eﬀ ort to locate the pre-
sumably montane homeland of this bowerbird
in the hope of discovering an additional trove of
ornithological novelties. In spite of diligent inves-
tigative ﬁ eld work throughout the ﬁ rst half of the
20th century, the mystery bowerbird eluded an
array of western ﬁ eld ornithologists, including F.
de Bruĳ n, Ernst Mayr, Fred Shaw-Mayer, Crown
Prince Leopold, S. Dillon Ripley, Sten Bergman,
Max Thompson, Phil Temple, and E. Thomas
Gilliard (Mayr 1941, Gilliard 1969). Ripley (1964),
in particular, was convinced that the unexplored
Foja (= Gau ier) Range of northwestern New
Guinea was the home of the bowerbird, but his
a empt in 1960 to penetrate this range from the
north via the Tor River failed because of the logis-
tical hurdles of hiking into this remote, trackless
area of diﬃ cult terrain. Later in the decade, E.
Thomas Gilliard proposed to get into the Foja
uplands by helicopter, but these plans were cut
short by his untimely death (M. L. LeCroy pers.
comm.). It was not until 1979, and the ready avail-
ability of helicopter transport, that Jared Diamond
was able to carry out a pioneering observational
reconnaissance of the Foja Mountains (followed
by a repeat visit in 1981). He succeeded in locat-
ing the “lost” bowerbird and provided an initial
characterization of the range’s montane avifauna
(Diamond 1982, 1985). However, because
Diamond collected no specimens for museum
study, the Foja Range has remained the most sig-
niﬁ cant mountain massif in New Guinea without
a bird survey documented by museum specimens
(Diamond 1985, Helgen 2005).
The Foja Mountains lie just east of the
Mamberamo River, and north of the Taritatu
(Idenburg) River. They are entirely isolated
from the Central Range as well as from all
other adjacent upland areas. The range is linear
(west-northwest–east-southeast) with several
summits above 2,000 m, the highest reaching
2,200 m. The range is very young, composed
of recently upli ed deep-sea sediments that
have been sutured to New Guinea’s north coast
by the tectonic collision of the Australian and
Paciﬁ c plates (D. A. Polhemus pers. comm.).
The entire massif, encompassing ~300,000 ha,
is apparently unroaded and perhaps untrailed,
with no villages except in the lowland fringes.
Informants from Papasena and Kwerba state
that local people do not enter the uplands, in
part because of inaccessibility, but also because
the summits are considered sacred.
From 15 November through 9 December
2005, an international team organized by
Conservation International (CI) and the
Indonesian Institute of Sciences (LIPI) carried
out a 25-day-long Rapid Assessment (RAP) of
birds, mammals, amphibians, reptiles, bu er-
ﬂ ies, and plants in the western part of the Foja
Mountains (Fig. 1). On 21 November 2005, the
CI RAP team helicoptered into a highland peat
bog, one of very few points of helicopter access
to the interior of the Fojas. Within minutes of
arrival at this site, members of the ﬁ eld team
noticed a wa led honeyeater foraging unwarily
in forest at the edge of the bog. It was notable
for its red-orange facial skin and the pendant
wa le that hangs down from each side of the
face (Fig. 2). This bird represents a new spe-
cies of smoky honeyeater (genus Melipotes), for
which we propose the following name:
Melipotes carolae, sp. nov.
Wa led Smoky Honeyeater
Holotype.—Museum Zoologicum Bogoriense
(MZB), Cibinong, Indonesia, MZB number
30628, male (testis 4 × 6 mm), Bog Camp, Sarmi
District, Papua Province, Indonesia, elevation
1,650 m, 2°34.5’S, 138°34.9’E, mist ne ed on
especie endémica Amblyornis ﬂ avifrons y la recientemente redescubierta ave del
paraíso Parotia berlepschi (B. M. Beehler datos no publicados).
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New Honeyeater from Western New GuineaJuly 2007] 1003
28 November 2005, prepared by B. M. Beehler,
ﬁ eld catalogue number BBFM-54.
Diagnosis.—The new species is readily
assignable to the genus Melipotes by the large
circumorbital patch of bright facial skin, the
overall sooty-gray plumage, and the short black
bill. Each of the following four characters dis-
tinguishes M. carolae (Fig. 2A, C) from all con-
geners: (1) elaboration of the circumorbital skin
patch into a so , loose, and ﬂ eshy pendant wat-
tle at the base of the jaw on each side of the face;
(2) the deep red-orange (rather than orange-
yellow) coloration of the facial skin patch; (3)
the nonﬂ ushing condition of the circumorbital
patch (see below); and (4) the dull gray throat
patch, only slightly paler than adjacent upper
The color of the circumorbital patch merits
comment. Individuals of all previously known
species of Melipotes can “ﬂ ush” this patch a
reddish or deep-orange color when agitated.
Melipotes carolae diﬀ ers by exhibiting a circum-
orbital patch that, at all times, is substantially
redder than the nonﬂ ushed state of those of the
other species. This was visible and obvious at a
distance in all free-ranging birds observed in the
ﬁ eld (n = 15 sightings by B.M.B. and additional
reported sightings by J. M. Diamond). Of appar-
ent signiﬁ cance is that M. carolae appears to
exhibit a circumorbital patch whose permanent
state is much like the “ﬂ ushed” state of the other
species of Melipotes. Our many photographs of
M. carolae indicate some mo ling of facial color
at times, but no change in state from unﬂ ushed
to ﬂ ushed such as that found in ater, gymnops,
and fumigatus (e.g., Fig. 2B, D).
The three other species of Melipotes diﬀ er
from carolae in the following ways: M. ater (Fig.
2F), which inhabits the upland forests of the
Huon Peninsula (Fig. 1), is substantially larger
(more than twice the mass of all other Melipotes),
with a yellow circumorbital patch, satin-black
plumage, white spo ing on the breast, and a
nonpendant caruncle-like “sub-wa le” on the
lower edge of the yellow circumorbital patch.
Melipotes ater is considerably more sociable and
noisy than any other member of the genus (J. M.
Diamond pers. comm.). Melipotes gymnops (see
Coates 2001), which inhabits the upland forests
of the Vogelkop and Bird’s Neck Peninsula of
westernmost New Guinea (Fig. 1), has a dark
throat, distinctive pale streaks on the lower
breast and belly, ochre-washed undertail coverts,
and a yellow circumorbital patch. Melipotes
fumigatus (Fig. 2B, D), a widespread upland spe-
cies of the New Guinean central cordillera, the
Cyclops, Bewani, and Kumawa ranges (Fig. 1),
is in most ways similar to M. carolae but exhibits
an orange-yellow (not red-orange) circumorbital
face patch, lacks the pendant wa le, and exhibits
a distinctly paler chin patch that contrasts with
the dark breast plumage.
Description of holotype.—Bill and legs black.
Iris dark brown. Plumage coloration ( following
Fig. 1. Map of New Guinea, showing ranges of all species of Melipotes and Macgregoria pulchra.
Beehler et al.
1004 [Auk, Vol. 124
color terminology of Smithe 1975): crown,
darker than jet black (89); mantle between jet
black (89) and blackish neutral gray (82); chin
and upper throat between blackish neutral
gray (82) and dark neutral gray (83), under-
parts darker than plumbeus (78), with light
neutral gray (85) edging to contour feathers
on breast; scapulars and primaries paler than
dusky brown (19); undertail coverts darker
than smoke gray (45).
Measurements of holotype.—Wing chord
(unﬂ a ened): 124 mm; tarsus: 35 mm; tail:
102 mm; bill from base of skull: 25 mm; bill from
feathers: 19 mm; weight: 52.5 g.
Specimens examined.—Five apparently adult
individuals (the holotype and four paratypes)
were mist ne ed at the Bog Camp between 28
November and 4 December 2005. Two were pre-
pared as study skins (MZB 30628 and 30629), and
three as whole anatomical specimens in alcohol
(MZB 30630, 30631, and 30632). All specimens
are held at the MZB in Cibinong, Indonesia.
Tissue samples of all were preserved in 70%
ETOH. Comparative study skins of the spe-
cies ater (n = 3), gymnops (n = 11), and fumigatus
(n = 29) were examined from the collections of
the National Collection, Cibinong (Museum
Zoologicum Bogoriense), and the National
Fig. 2. (A, C) Melipotes carolae, (B, D) Melipotes fumigatus, (E) Macgregoria pulchra, and (F)
New Honeyeater from Western New GuineaJuly 2007] 1005
Museum of Natural History, Smithsonian
Institution, Washington, D.C. The measurements
of wing, tail, tarsus, and bill for M. carolae did not
diﬀ er substantially from those for its putative
sister form, M. fumigatus (Table 1).
Distribution, ecology, and behavior.—Recorded
only above 1,150 m elevation in the Foja
Mountains of western (Indonesian) New
Guinea, M. carolae was found by the ﬁ eld team
to be common in forest interior and forest-edge.
It was seen repeatedly foraging unwarily in
the middle and upper levels of the vegetation,
especially at plants producing small fruits.
The Foja Mountains support a large expanse of
continuous, closed, humid tropical submontane
forest. From the forest physiognomy, we esti-
mate that this mountain range receives what
appears to be a moderate amount of rainfall
annually (about 3–4 m per annum, with a dry
season that apparently peaks in August, accord-
ing to local informants). The type locality is a
sphagnum bog ~500 m in diameter surrounded
by mossy forest of moderate stature (20–25 m)
dominated by gymnosperms (e.g., Dacrydium,
Dacrycarpus) and montane angiosperm taxa
(e.g., Nothofagus, Lithocarpus). The montane veg-
etation of this isolated mountain massif exhibits
the Massenerhebung Eﬀ ect—thus, the forest at
1,650 m in the Foja Mountains is similar to a for-
est of higher elevation in New Guinea’s central
Melipotes carolae exhibited behavior typical of
the genus Melipotes (Beehler et al. 1986, Coates
1990), with one exception: the individuals
encountered on 15 occasions during the ﬁ eld
trip were silent. We never heard or recorded this
species making a sound. We encountered no evi-
dence of nesting or other reproductive behavior
during the ﬁ eld period (late November–early
December). The two dissected individuals had
Etymology.—The speciﬁ c epithet honors
Carol Beehler, wife of the senior author,
acknowledging her long and unstinting sup-
port and her personal commitment to biodi-
versity studies in New Guinea. The English
name highlights the most distinctive feature of
this novel species.
The genus Melipotes is an important member
of the upland forest bird fauna of New Guinea
Table 1. Measurements for specimens of three species of Melipotes (sample sizes > 1 in parentheses).
Wing chord (mm) Tarsus (mm) Tail (mm) Culmen (mm) Mass (g)
Male 124 35 102 25 52.5
Female 112.5 30.5 95 23.5 54
Unsexed 123, 125(2), 125.5 33, 33.5, 34, – 99(2), 100, – 21, 24.5, 26, – 59, 60, 63, 65
M. fumigatus goliathi
Male 109, 118, 119, 122, 123(2) 29(3), 30, 34.5 103, 108, 109, 111, 113 21, 22, 23, 25 62
Female 105, 107, 108, 110(2), 111, 114 29(6), 30, 31 95, 101, 103(2), 104, 108(2), 20(2), 22(3) –
M. f. fumigatus
Male 104, 107, 113,114, 115(2),117, 121 27, 28(2) , 29, 33.5,34(2),35 91, 93, 94 96,100,101, 21,24,24.3,25.5 46, 47, 49, 60
Female 108, 109, 112, 109, 114 28(2), 29(3), 31,34 93(2),96, 103, 106,109 20, 22(4), 24, 24.5 42, 48, 54
Male 100, 108, 109, 121(2) 27(3) 94, 97, 111 18, 21(2), 22(3), 24
Female 103, 108, 106, 107, 113, 115 26, 27(2), 28(3), 91(2), 96(2), 97, 99 19(2), 20(2), 21, 22 49, 58
Unsexed 109, 119 29, 32 93.5, 100 22.5, 23.5 –
Beehler et al.
1006 [Auk, Vol. 124
(Salomonsen 1967; Beehler et al. 1986; Coates
1990, 2001). Virtually all upland forests in New
Guinea support a single species of Melipotes. In
each montane habitat, the Melipotes tends to be
one of the most common and visible members
of the local montane forest community.
Systematics.—A comprehensive systematic
review of the genus must await analysis of tissue
samples of M. carolae, which currently remain
in Indonesia. Therefore, we here oﬀ er some
preliminary thoughts on systematics and bio-
geography of the lineage. First, using molecular
analysis, Cracra and Feinstein (2000) recently
demonstrated that the distinctive subalpine fru-
givore Macgregoria pulchra (“Macgregor’s Bird
of Paradise” in Frith and Beehler 1998) is the sis-
ter form to Melipotes. We provide a provisional
cladogram for the Macgregoria + Melipotes clade,
based on external morphology and natural-
history traits (Fig. 3 and Appendices 1 and 2).
We did not believe it necessary to run the small
data set through a cladistic algorithm (e.g.,
PAUP) because only a single instance of homo-
plasy appears (transformation of character state
8.1 to 8.0 for M. gymnops). In addition, note
that we included basal-derived characters that
unite both the Macgregoria and Melipotes clades.
These are not informative in terms of tree
generation but provide additional evidence of
the monophyly of the two lineages (cf. Cracra
and Feinstein 2000). We note that the spe-
cies concept employed here approximates the
polytypic–biological species of Mayr (1963) but
that we consider geographically well-delimited
and morphologically well-deﬁ ned allotaxa as
species rather than subspecies.
Figure 3 supports the divergence of the
Melipotes lineage from a Macgregoria-like sister
lineage, with a endant character evolution from
a large-bodied black montane songbird with a
ﬂ eshy orbital wa le to a smaller sooty songbird
with a circumorbital patch of bare skin. In this
regard, Melipotes ater appears to be basal within
Melipotes and the gymnops–fumigatus–carolae
species-group appears to be derived. Within
the la er clade, fumigatus and carolae appear
to be sister forms with many shared derived
We thus see an interesting pa ern of geographic
diﬀ erentiation in three distinct steps. First, there
is a Central Cordillera–Huon Peninsula split
(proto-Macgregoria–proto-Melipotes basal lin-
eages). Second, we postulate a colonization of
the Central Cordillera range and Bird’s Head
Uplands by basal Melipotes from the Huon
Peninsula, followed by east–west diﬀ erentiation
of this lineage into a western–Bird’s Head form
(gymnops) and a central range vicariant (fumiga-
tus). Finally, we propose a colonization of the
north coastal ranges by M. fumigatus, followed by
isolation and diﬀ erentiation of the Foja Mountain
isolate into M. carolae. We suggest that speciation
of carolae in the Foja Mountains is a product of the
geographic extent and height of the range and its
Fig. 3. Cladogram of the Macgregoria + Melipotes lineage.
New Honeyeater from Western New GuineaJuly 2007] 1007
physical isolation. Thus, the presence of typical
fumigatus in the Cyclops, Bewani, and Kumawa
ranges may be evidence that these ranges are
too small to foster the diﬀ erentiation of a species
vicariant of Melipotes (because of periodic extirpa-
tion and recolonization).
Gibbs (1994) recently observed a Melipotes in
the Fak Fak Mountains (Fig. 1) that exhibited
a pendant facial wa le. On the basis of pale
lower ventral plumage, he believed that this
population was a derivative of gymnops, not
fumigatus as Diamond (1985) had postulated.
It is certainly worth returning to the Fak
Fak Mountains to collect the honeyeater and
other novelties inhabiting that undersurveyed
New Guinean biogeography.—The genus
Melipotes assumes greater biogeographic signiﬁ -
cance with the discovery of M. carolae. Melipotes
can be added to the list of readily speciating
New Guinean montane-forest bird genera that
have produced allospecies-level vicariants on
several of New Guinea’s north coastal ranges
(other examples include Rallicula, Melidectes,
Ptiloprora, Amblyornis, Sericulus, Parotia, and
Astrapia; cf. Diamond 1969, 1985). These lin-
eages form the systematic “backbone” of the
New Guinean montane forest avifauna, and
ﬁ ve of these seven “core” genera are endemic to
New Guinea’s humid uplands.
This discovery conﬁ rms that the Foja
Mountains are New Guinea’s third-most-
signiﬁ cant north coastal range, when measured
by species-level avian endemism (Diamond
1985). The Tamrau–Arfak Mountains of the
Bird’s Head Peninsula of the far west support
nine montane endemic bird species, the mas-
sive and high Finisterre–Saruwaged Mountains
of the Huon Peninsula support four species,
and the Foja three species. Like the uplands of
the Huon and Bird’s Head, the Fojas harbor an
endemic species of Melipotes and an endemic
bird of paradise (Parotia). As with Bird’s Head,
the Foja possess an endemic gardener bower-
bird of the genus Amblyornis (remarkably
absent from the Huon, which instead possesses
a subspecies of the central cordilleran species A.
macgregoriae). The Fojas also possess endemic
populations–subspecies of Sericulus, Melidectes,
Ptiloprora, and Rallicula (Diamond 1985; B. M.
Beehler unpubl. data).
Conservation signiﬁ cance.—On the basis
of ornithological surveys in outlying New
Guinean mountain ranges, Diamond (1982,
1985) provided the initial indication of the
biological importance of the Foja Mountains
in a regional context. Our discovery of an
endemic meliphagid in the Foja Mountains
and rediscovery of Parotia berlepschi (the “lost”
six-wired bird of paradise; Kleinschmidt 1897,
B. M. Beehler unpubl. data) further conﬁ rm
the faunal distinctiveness of these mountains
and provide strong evidence for their long-
term biotic isolation (Fig. 1). Although the
Foja Range is o en united with other outly-
ing northern New Guinean mountain ranges
(especially the Cyclops, Bewani, Torricelli, and
Prince Alexander ranges) as part of a single
biogeographic unit (e.g., Sta ersﬁ eld et al.
1998, Helgen 2005), our results oﬀ er new evi-
dence of the Fojas’ faunal uniqueness compared
with even these closely associated ranges. This
is also indicated by the discovery of new
endemic species of bu erﬂ ies, mammals, frogs,
and plants during our recent Foja survey (van
Mastrigt 2006, S. J. Richards unpubl. data).
Given that we conducted surveys over rela-
tively limited elevational spans (100–900 m and
1,150–1,850 m, with a ention primarily focused
at 100, 350, and 1,650 m) and in only one part
of the range, we believe that further ﬁ eld work
may reveal additional exciting ornithological
discoveries in the Fojas. It is apparent from
Diamond’s work and ours that the end of the
calendar year is the nonbreeding “quiet” season
for birds. Field work undertaken in August–
September may thus encounter more avian song
and breeding activity and potentially reveal cer-
tain retiring species that have been overlooked
by Diamond and us.
Our ﬁ eld work in the Foja Mountains fur-
ther validates Diamond’s (1985) suggestion
that these mountains represent a signiﬁ cant
biodiversity resource that merits large-scale
conservation. The Fojas are currently desig-
nated as a wildlife sanctuary (suaka margasatwa),
but this remains in essence a “paper park.” We
suggest that the entire mountain range merits a
management plan and that additional research
should be invested in delineating the distribu-
tion of all globally restricted-range species
inhabiting the Fojas. In addition, and perhaps
more importantly, Conservation International
is working with indigenous forest-dwelling
communities inhabiting the fringes of the Fojas
to delineate a long-term comanagement plan
Beehler et al.
1008 [Auk, Vol. 124
for this important forest block. We believe that
these communities are the local stewards of
one of New Guinea’s most remarkable natural
We thank the Indonesian Institute of Sciences
(LIPI), the Local Forestry Oﬃ ces (BKSDA) in
Papua, the Department of Forests, Directorate
General of Forest Protection and Nature
Conservation, and the Ministry of Interior, for
permission to conduct ﬁ eldwork in Indonesian
New Guinea. We gratefully thank D. Darnaedi,
Director of the Research Center for Biology
of LIPI, for his strong support of the scien-
tiﬁ c collaboration between the Conservation
International team and his Indonesian scientiﬁ c
team. We thank Conservation International
Indonesia for abundant assistance and support
of our ﬁ eld expedition, especially J. Supriatna,
I. Budiningrum, and the Mamberamo ﬁ eld team.
Our ﬁ eld work in the Foja Mountains could
not have proceeded without the hospitality
and partnership of the people of Kwerba and
Papasena villages, especially P. I. Tawani and
P. T. Kawena. HeliMission made its helicopter
available to us, without which our montane
survey could not have taken place. We salute
S. Richards, co-leader of this expedition, for all
that he did to ensure its success. We thank R. C.
Banks, J. M. Diamond, K. Helgen, M. LeCroy,
R. Pasquier, P. Rasmussen, D. Wilcove, and two
anonymous reviewers for technical and editorial
suggestions that improved this paper. The proj-
ect was generously supported by grants from the
Swi Foundation, the Gordon and Be y Moore
Foundation, the National Geographic Society,
and the Global Environment Project Institute.
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Associate Editor: N. S. Sodhi
Character Matrix for Melipotes and
1. Facial wa le: 0 = present.
2. Wa le type: 0 = semicircular and ocular, 1 =
circular and facial, 2 = circular and with pen-
3. Wa le color, at rest: 0 = yellow-orange, 1 =
4. Body size: 0 = large, 1 = medium.
5. Contour plumage background color: 0 =
black, 1 = gray.
6. Ventral plumage pa ern: 0 = plain, 1 = pale
spo ed–streaked, 2 = mo led–scalloped.
7. Undertail coverts plumage pa ern: 0 =
dark, 1 = ochre.
8. Chin plumage: 0 = dark, 1 = distinctly
paler, contrasting breast; 2 = slightly paler than
9. Wing patch: 0 = present, 1 = absent.
10. Sociability: 0 = sociable, 1 = solitary.
11. Vocalization: 0 = vocal, 1 = mildly vocal,
2 = silent.
12. Wing sound production: 0 = present, 1 =
13. Bill morphology: 0 = short and straight.
14. Diet: 0 = highly frugivorous.
Note: Characters 1, 13, and 14 are included to
demonstrate characters linking Macgregoria with
Melipotes (see Cracra and Feinstein 2000).
Appendix 2. Character matrix for the Macgregoria–Melipotes lineage.
Character Macgregoria Melipotes Melipotes Melipotes Melipotes
number Character pulchra ater gymnops fumigatus carolae
1 Facial wa le 0 0 0 0 0
2 Wa le type 0 1 1 1 2
3 Wa le color 0 0 0 0 1
4 Body size 0 0 1 1 1
5 Contour plumage color 0 0 1 1 1
6 Ventral plumage pa ern 0 1 1 2 2
7 Undertail coverts 0 0 1 0 0
8 Chin plumage 0 1 0 1 2
9 Wing patch 0 1 1 1 1
10 Sociability 0 0 1 1 1
11 Vocalization 0 0 1 1 2
12 Wing sound production 0 1 1 1 1
13 Bill morphology 0 0 0 0 0
14 Diet 0 0 0 0 0