Systematics and Biodiversity 3 (4): 433?453 Issued 24 November 2005
doi:10.1017/S1477200005001702 Printed in the United Kingdom C? The Natural History Museum
Kristofer M. Helgen
School of Earth and
Environmental Sciences,
University of Adelaide, Adelaide
SA 5005, and *South Australian
Museum, North Terrace,
Adelaide SA 5000, Australia
submitted March 2004
accepted December 2004
Systematics of the Paci?c monkey-faced
bats (Chiroptera: Pteropodidae), with a
new species of Pteralopex and a new
Fijian genus
Abstract The fruit-bat genus Pteralopex comprises the monkey-faced bats, a group
of six endangered species found only in old-growth forests on certain islands in
the south-west Paci?c (the Solomon Islands and Fiji). The taxonomy of the genus
is reviewed in detail and updated accordingly. Two ?cryptic? biological species are
shown to occur in sympatry on both Bougainville and Choiseul in the northern
Solomon Islands (corresponding to Pteralopex anceps Andersen, 1909 and a previ-
ously undescribed species) and each is accordingly described and reviewed. A new
genus (Mirimiri) is erected for the Fijian monkey-faced bat (formerly Pteralopex
acrodonta), which differs greatly both morphologically and genetically from spe-
cies of Pteralopex in the Solomon Islands. Ecomorphological differences between
sympatric Pteralopex species are brie?y reviewed, including potential differences in
functional morphology and feeding ecology. Geographic patterns of occurrence and
future survey priorities for monkey-faced bats are also discussed.
Key words biogeography, cryptic species, Fiji, morphometrics, ecomorphology,
Melanesia, Papua New Guinea, Pteralopex, Pteropodidae, Solomon Islands
Introduction
The monkey-faced bats are a highly distinctive group of
medium-sized fruit-bat species endemic to the Solomon
Islands and Fiji, previously classified in the single genus
Pteralopex. In recent years, aspects of the taxonomy, geograph-
ical distribution, phylogeny, basic biology and conservation
status of Pteralopex species have attracted study (Flannery,
1991, 1995; Colgan & Flannery, 1995; Fisher & Tasker, 1997;
Bonaccorso, 1998; Parnaby, 2002b; Colgan & da Costa, 2002;
Ingleby & Colgan, 2003). However, detailed information re-
mains unavailable for most of these subjects. In particular,
species boundaries and taxonomic definitions within the genus
(the subject of the current report) have varied over time, ob-
scuring the identity and basic biological attributes of various
monkey-faced bat species. On the basis of all known mu-
seum material and all available field observations regarding
the genus, taxonomic boundaries, geographic distribution and
basic biology of the monkey-faced bats are reviewed here.
*Address for correspondence.
Email: kristofer.helgen@adelaide.edu.au
Materials and methods
Specimens cited here are deposited in the collections of the
Australian Museum, Sydney (AM), Bernice P. Bishop Mu-
seum, Honolulu (BBM), Natural History Museum, London
(BMNH), Field Museum of Natural History, Chicago
(FMNH), Museum of Vertebrate Zoology, University of
California, Berkeley (MVZ), National Museum and Art
Gallery of Papua New Guinea, Port Moresby (PNGM),
SenckenbergMuseum, Frankfurt (SMF), and the United States
National Museum of Natural History, Smithsonian Institution,
Washington, DC (USNM).
Measurements are abbreviated (and, where necessary,
defined) as follows: FA (forearm length); HB (head and
body length); E (ear length); WT (mass); CBL (condylobasal
length); ONL (orbitonasal length, here defined as the distance
from anterior edge of the orbit to the midpoint of the premax-
illae); ZYG (greatest bizygomatic width); MTR (length of
maxillary toothrow, C ? M2); BBM (breadth of braincase at
intersection with zygomata); MH (coronoid height of mand-
ible); PL (length from posterior midpoint of palate to anterior
midpoint of premaxillae). For wing measurements, digit is ab-
breviated as ?D?, metacarpal as ?M? and phalanx as ?P?; thus,
D2P2 refers to the second phalanx of the second digit, etc. For
433
434 K. M. Helgen
dental measurements, terminology and abbreviations follow
Hill &Beckon (1978), with incisor abbreviated as ?I?, canine as
?C?, premolar as ?PM?, and molar as ?M?, but with upper teeth
designated by superscript (e.g. M2) and lower teeth by sub-
script (e.g. M2). For more details on the features of soft palate
anatomy referred to here, see Andersen (1912). External meas-
urements for museum specimens were recorded by the original
collectors in the field. Craniodental variables were measured
by the author with hand-held calipers to the nearest 0.01mm.
Single-tooth measurements are measured on the crown. All
measurements of length are in millimetres, and measurements
of mass are given in grams. In the principal component ana-
lyses featured in this paper, components are extracted from a
covariance matrix of log-transformed craniodental variables.
Spellings for island names follow Mayr & Diamond
(2001). Within the Solomon Archipelago (referred to through-
out this paper as the ?Solomon Islands?), the islands of Buka
and Bougainville lie within the political boundaries of Papua
New Guinea (Northern Solomons Province), while Choiseul,
Ysabel, New Georgia and Guadalcanal are politically part of
the Solomon Islands.
Previous taxonomic treatments
of Pteralopex
In 1888, Oldfield Thomas described Pteralopex atrata, a new
genus and species of medium-sized fruit-bat, based on a series
of specimens sent to the British Museum (now the Natural
History Museum, London) from Guadalcanal in the central
Solomon Islands (Thomas, 1888). Though superficially sim-
ilar to species of flying-foxes (genus Pteropus) in external
form, notable traits of this highly distinctive new genus in-
cluded an unusually robust skull, massive bicuspidate canines,
cheekteeth with complex, cuspidate surfaces, and wing mem-
branes attached along the mid-back near the spine, rather than
on the sides of the body. Soon after, Matschie (1899) uncrit-
ically included Pteralopex as a synonym of Pteropus (as did
Simpson [1945]), but all other authors have correctly acknow-
ledged the distinctiveness of this genus since Thomas? (1888)
original description.
Based on the skin and skull of a subadult specimen col-
lected by A.S.Meek in 1904, Knud Andersen (1909b) later de-
scribed a second species of the genus, Pteralopex anceps, from
Bougainville in the northern Solomons. Soon after, Andersen
(1912) reviewed the morphological attributes of P. anceps and
P. atrata in greater detail. He observed that although the two
species are similar in size and coloration (both are black or
blackish-brown), they differ both in trenchant aspects of dental
morphology and in the texture, length, and distribution of the
pelage. Sanborn (1931) later identified a monkey-faced bat
specimen from Ysabel as P. atrata, presumably on the basis of
Andersen?s (1912) diagnostic characters.
Nothing was subsequently reported about Pteralopex
until Laurie & Hill (1954) united P. atrata and P. anceps
as conspecific subspecies in their authoritative checklist of
Wallacean and Melanesian mammals. This was presumably
done because the two taxa, superficially similar in appearance,
were at the time known by very few specimens and were
thought to be allopatric (with P. atrata on Guadalcanal and
Ysabel, andP. anceps onBougainville). However, this arrange-
ment ignored Andersen?s (1912) review of the outstanding
dental and pelage differences that distinguish the two taxa
(Andersen, 1912).
In 1968, Phillips (1968) reported on newly collected
Pteralopex specimens from Bougainville and Choiseul, ob-
tained during expeditions by the Smithsonian Institution in
Washington and the Bishop Museum in Honolulu. Influenced
by Laurie & Hill (1954), he identified these as ?P. atrata an-
ceps? ? apparently on geographic grounds ? but he noted that
they were more similar in colour and in some dental traits to
P. atrata (as opposed to P. anceps) according to Andersen?s
(1912) descriptions.
Hill & Beckon (1978) described a newly discovered,
small-bodied species of Pteralopex (P. acrodonta) from the
Fijian island of Taveuni, the first record of the genus out-
side of the Solomon Islands. These authors reviewed in detail
how this species differs considerably in cranial, dental and ex-
ternal aspects from the other Pteralopex species known at that
time. On the basis of their examinations of the type series of
P. atrata and the holotype of P. anceps, Hill & Beckon (1978)
again recognized P. anceps and P. atrata as separate species
from Bougainville and Guadalcanal, respectively, though they
did not examine themuseum specimens fromYsabel, Choiseul
or Bougainville that had been reported by Sanborn (1931) or
Phillips (1968).
Flannery (1991) described a distinctive new small-bodied
species, Pteralopex pulchra, from cloud forest at 1230m on
Mt.Makarakomburu,Guadalcanal. He also reported additional
monkey-faced bat specimens from Bougainville and Buka
(which he identified as Pteralopex anceps) and from lowland
Guadalcanal (which he identified as P. atrata). Subsequently
Flannery (1995) reviewed the biology of all species of Pter-
alopex then described (P. atrata, P. anceps, P. acrodonta and
P. pulchra) as well as that of a newly discovered, unnamed
species from the islands of New Georgia and Vangunu in the
western Solomons.
Soon after, Fisher & Tasker (1997) reported the results
of a detailed field study of the ecology and life history of
this then-unnamed species. Parnaby (2002b) named this new
species from New Georgia and Vangunu (as Pteralopex taki)
in a review of the entire genus. He contrasted the attributes
of P. taki most closely with P. pulchra, which he apparently
considered to be its closest relative (cf. Ingleby & Colgan,
2003: 21). He also allied P. atrata (which he considered to
be a Guadalcanal endemic) and P. anceps (which he con-
sidered to be the only Pteralopex species recorded from Buka,
Bougainville, Choiseul and Ysabel; contra Sanborn, 1931) as
immediately related species that differed primarily in body
size. Notably, Parnaby (2002b) criticized features used by
Andersen (1912) and Hill & Beckon (1978) to diagnose
P. anceps and P. atrata, arguing that these characters are in-
consistent. Finally, on the basis of then-unpublished genetic
data (Ingleby & Colgan, 2003), Parnaby (2002b) considered
the phylogenetic relationship between P. acrodonta (from Fiji)
and all other Pteralopex (from the Solomon Islands) to be
rather remote.
Paci?c monkey-faced bats 435
Figure 1 Recorded distribution of Pteralopex species in the Solomon Islands. Above, recorded distribution of P. ?anneryi (
) and P. atrata
(H17009). Below, recorded distribution of P. anceps ( [de?nite], ? [possible] ), P. taki (), and P. pulchra ().
The identity of Pteralopex atrata
and P. anceps
I have recently examined all museum material of Pteralopex,
including many specimens not seen by Parnaby (2002b). The
latter include specimens from Bougainville (at BBM, BMNH
and PNGM; see below), Choiseul (at BBM, see below),
Ysabel (at FMNH, see below), Guadalcanal (at BMNH, MVZ,
and SMF, see below), and New Georgia (at SMF, see below).
I have also drawn from field observations of Pteralopex re-
cently reported from Choiseul (Bowen-Jones et al., 1997) and
Ysabel (Leary & Pita, in litt.). Importantly, these examinations
clearly demonstrate that two distinct species of Pteralopex
occur sympatrically on Bougainville, contrary to Parnaby?s
(2002b) conclusions. These could be called ?cryptic? species
in that they are superficially similar in size and dorsal col-
oration. However, these taxa differ consistently in body size;
cranial size and shape; size, shape, relative proportions, and
important occlusal aspects of the teeth; and length, texture,
and distribution of the pelage. These two species correspond
essentially to diagnoses of P. anceps and P. atrata as presented
by Andersen (1912) and Hill & Beckon (1978), although the
atrata-like population of Bougainville is much larger-bodied
than that of Guadalcanal (the type locality of atrata), and dif-
fers also in certain other ways (see below). Parnaby (2002b)
incorrectly considered diagnostic features previously used to
distinguish between P. anceps and P. atrata (Andersen, 1912;
Hill & Beckon, 1978) to be inconsistent because his sample
iiar 
OS QB* 
-&:r 
10* 
5$ 
Bougainville 
o ChoiseU 
VMkUw#t<^ KAntmxykjX   ^^^ 
Malaita 
Makka 
10 # 
Buka 
Bougainville 
CholmeJ 
&   O/^New 
Mabita 
Makira 
436 K. M. Helgen
fromBougainville was composite, consisting of both P. anceps
and an undescribed species similar to P. atrata,1 described be-
low as Pteralopex flanneryi sp. nov.
Examinations reported here further demonstrate that all
museum specimens from Buka, Choiseul, and Ysabel repor-
ted in the literature as P. anceps (e.g. Flannery, 1991, 1995;
Bonaccorso, 1998; Parnaby, 2002b) also actually represent
Pteralopex flanneryi. These taxonomic distinctions and iden-
tifications are documented below. Notably, field observations
reported by Bowen-Jones et al. (1997) confirm that P. anceps
occurs in sympatry with P. flanneryi on Choiseul in addition
to Bougainville. Further, Pteralopex atrata is recorded for the
first time fromNewGeorgia (see below), where it is apparently
sympatric with P. taki.
The status of ?Pteralopex? acrodonta
?Pteralopex? acrodonta is a small-bodied species of monkey-
faced bat known only fromhigher altitudes onMt.Koroturanga
on the Fijian island of Taveuni; all other Pteralopex species
are endemic to the Solomon Islands. Together, Hill & Beckon
(1978) and Parnaby (2002b) documented a number of strik-
ing morphological differences that separate ?Pteralopex? acro-
donta from all other Pteralopex species. These distinctions led
Flannery (1995, p. 237) to presume that ?P.? acrodonta ?has
been isolated on Fiji for a very long time?.
An allozyme study (Ingleby & Colgan, 2003) recently
revealed that the electrophoretic distance between ?P.? acro-
donta and two Solomon Islands Pteralopex species (P. taki and
P. pulchra) is equivalent to that between some megachirop-
teran sister genera. Another recent study drawing from mi-
tochondrial and nuclear sequence data (Colgan & da Costa,
2002) documented considerable sequence divergence between
?P.? acrodonta and P. atrata (the only monkey-faced bats in-
cluded in the study) and was unable to confirm that these
species are more closely related to one another than they are
to species of other pteropodid genera. On the basis of these
studies, Parnaby (2002b) explicitly questioned the monophyly
of Pteralopex as currently defined.
Monkey-faced bats from the Solomon Islands and from
Fiji clearly represent two distinctive lineages, each of which
can be diagnosed with respect to the other by a unique com-
bination of morphological apomorphies, as reviewed below.
Though I propose that these two groups are indeed sister taxa,
I separate them into separate genera below. This taxonomic
change better reflects these morphologically and genetically
divergent groups? different evolutionary and biogeographic
histories, and more accurately portrays phylogenetic relation-
ships among the six species of monkey-faced bats.
1 Parnaby?s (2002b) sample included only two actual specimens of P.
anceps: AM M6283, which, as he noted, has the dental features of
P. anceps as diagnosed by Andersen (1912) and AM M6497, which is
an old adult with cheekteeth extensively worn, obscuring their occlusal
features. All of his other specimens from Bougainville were P. flanneryi
sp. nov. (compare with the type series, below). On account of his limited
sample size, Parnaby did not grasp the significance of variation in pelage
and dental traits within the sample.
Systematics
Monophyly of monkey-faced bats
Together, themonkey-faced bats (Pteralopex andMirimiri gen.
nov.) can be immediately distinguished from all other ptero-
podid bats by the following combination of distinctive traits:
1. Skull very robust relative to overall size, with thick, heavy
zygomata and an unusually high sagittal crest.
2. Mandible with well-developed angular process, promin-
ent masseteric ridge and a very high coronoid process
(especially relative to the alveolar margin of the lower
toothrow).
3. Upper canines relatively very large and bicuspidate (bear-
ing a large posterior accessory cusp).
4. Dental formula I2/2, C1/1, PM3/3, M2/3, with PM1 and
I1 excessively reduced.
5. PM4 and molars bearing a complex pattern of cusps.
6. Iris of eye red/orange in life (verified to date for pulchra,
taki and acrodonta).
7. Colour of mantle not markedly contrasting with colour of
mid-back.
8. Tail absent.
9. Wings attached to back near the spine, rather than on the
sides of the body.
10. Second digit of the wing bearing a claw.
11. Body size medium to large for Pteropodidae (forearm
length 117 to 169).
12. Sexually monomorphic (or very weakly dimorphic) in
body size.
Of these character states, numbers 1, 2, 3, 4, 5, 6, 8
and 11 are derived relative to the primitive condition in the
Pteropodidae (according to my understanding of the distribu-
tion of these traits in living and fossil pteropodids). Relative
to flying-foxes (Pteropus and Acerodon), traits 1?6 are more
highly derived in monkey-faced bats, traits 8?12 are shared
with some or all species of flying-foxes, and 7 is less derived.
As Pteralopex and Mirimiri share a considerable number of
apomorphic traits relative to other larger-bodied pteropodid
genera, I strongly suspect that they are sister taxa, a point
as yet unconfimed by molecular phylogenetics (cf. Colgan &
da Costa, 2002). The more remote phylogenetic relationships
of Pteralopex and Mirimiri are less certain, but probably lie
with Pteropus, Acerodon, and/or other pteropodine bats such
as Dobsonia and Aproteles (Andersen, 1909a, 1912; Hill &
Beckon, 1978; Colgan & Flannery, 1995; Colgan & da Costa,
2002).
Hill & Beckon (1978) reviewed in still-useful detail sev-
eral other features that jointly characterize the monkey-faced
bat genera.
Genus Pteralopex Thomas, 1888
Revised diagnosis
The genusPteralopex can be distinguished from all pteropodid
genera but Mirimiri gen. nov. by the distinctive combination of
morphological traits listed above. In addition, each of the fol-
lowing eight characters is perfectly diagnostic for Pteralopex
Paci?c monkey-faced bats 437
with respect to Mirimiri gen. nov.:
1. Postorbital processes well-developed and annectent to the
zygomata in adults.
2. Interorbital region conspicuously narrowed.
3. M2 strongly reduced relative toM1 (less than half themass).
4. Labial margins of PM4 and M1 united in a single elevated
cusp.
5. Length of D4P2 conspicuously longer than D4P1.
6. Six divided median ridges in the soft palate.
7. Three serrated supplementary posterior ridges in the soft
palate.
8. Sexes identical in coloration.
Of these character states, numbers 1, 2, 3, 5, and 6 are
apomorphic with respect toMirimiri.Pteralopex can further be
consideredmore highly derived thanMirimiri in that three of its
five species (Pteralopex anceps, P. flanneryi and P. atrata) are
much larger in body size than the small-bodied M. acrodonta.
Content
The genus Pteralopex comprises five species, all endemic to
the Solomon Islands: P. anceps Andersen, 1909 (Bougainville,
Choiseul); P. atrata Thomas, 1888 (Guadalcanal, New
Georgia); P. flanneryi sp. nov. (Bougainville, Buka, Choiseul,
Ysabel); P. pulchra Flannery, 1991 (Guadalcanal); and
P. taki Parnaby, 2002 (New Georgia, Vangunu, and formerly
Kulumbangra).
Remarks
The traits of P. atrata and P. anceps have previously been
confused; as a result, the taxonomy of these species is reviewed
in detail below in the context of describing a new species,
P. flanneryi. Systematics and basic attributes of the remaining,
small-bodied species (P. pulchra and P. taki) were recently
reviewed adequately by Parnaby (2002b) on the basis of all
available museum material; thus these taxa are not treated
further here.
Pteralopex ?anneryi sp. nov.
(Figs 2, 3 lower, 5 upper)
HOLOTYPE. AM M6282, adult male, skin (Figs. 2, 3) and
skull (Fig. 5), fromBuinDistrict (06?50?S, 155?44?E), 10miles
inland from the coast, altitude 50m, Bougainville (Northern
Solomons Province, Papua New Guinea), collected March
1937 by J.B. Poncelet.
PARATYPES. AM M6498, young adult female, skin and
skull, from the type locality, collected September 1937 by
J.B. Poncelet. AM M6346, adult male, whole specimen in al-
cohol, and AM M6347, adult female, skin and skull, from Iula
(06?50?S, 155?45?E), 15 miles inland from the coast, altitude
150m, Bougainville, collected 3 March 1937 by J.B. Poncelet.
USNM 276973, adult female, skin, skull, and skeleton, from
Cape Torokina (06?15?S, 155?02?E), at sea level, Bougainville,
collected 5 August 1944 by D.H. Johnson. USNM 276974,
adult male, skull, from Puruata Island opposite Cape Torokina,
Bougainville, collected 30 July 1944 by W.L. Necker. USNM
277112, adult male, skin and skull, from Cape Torokina,
Bougainville, collected 22 July 1944 by D.H. Johnson. BBM-
NG61066, adult female, skin and skull, fromPokapa (05?37?S,
154?58?E), altitude 200m, Bougainville, collected 26 February
1968 by A.B. Mirza.
REFERRED SPECIMENS. FMNH 31561, subadult female,
from Tunnabuli (08?24?S, 159?48?E), Ysabel, collected 17
April 1929 by F.C. Wonder (see Sanborn, 1931). BBM-BSIP
23682, adult male, skin and skull, from Malangona (07?01?S,
156?59?E), Choiseul, collected 17 March 1964 by P. Temple
(see Phillips, 1968; Parnaby [2002b] erroneously referred
to this specimen as USNM 23682). AM M19822, subadult
male, skin and skull, fromMalasang-Kakalis Hamlet (05?21?S,
154?41?E), Buka, collected September 1987 by S. Wickler. A
specimen of Pteralopex collected by Leary and Pita (in litt.) on
Barora Fa (a small island adjacent to Isabel=Barola, 07?30?S,
158?20?E) is also referred to P. flanneryi on the basis of its
reported measurements. All specimens but the last have been
examined by the present author.
Diagnosis
Largest-bodied species of Pteralopex, with adult condylobasal
length > 71 and forearm greater than or equal to 159; pelage
relatively short, with ear exposed, fur of the mid-back and the
dorsal surface of the pes adpressed, and the tibia and forearm
naked or very sparsely furred; overall colour black dorsally and
ventrally with occasional light (yellowish or whitish) tipping
to the hairs on the breast (but lacking the expansive white
or yellow chest-patch seen in sympatric P. anceps). Skull
absolutely larger than that of P. anceps, with narrower brain-
case, relatively smaller incisors, larger cheekteeth, and PM4,
M1, and M2 with lengthened, elevated lingual ridges.
Distribution
Recorded from lowland forest at sea level to 200m (see
specimens above) on the north-eastern Solomon Islands of
Bougainville (including the immediately adjacent small island
of Puruata), Buka, Choiseul, and Ysabel (including the im-
mediately adjacent small island of Barora Fa). These islands
represent land-bridge fragments of a single large Late Pleis-
tocene island (a landmass generally called ?Greater Bukida?;
Mayr & Diamond, 2001). Pteralopex flanneryi is sympatric
with P. anceps on Bougainville and Choiseul (Figure 1; see
discussion below).
Etymology
For Professor Tim Flannery, current Director of the South
Australian Museum in Adelaide, in recognition of his studies
of Melanesian mammals, including the mammal fauna of the
Solomon Islands. I suggest ?Greater monkey-faced bat? as an
appropriate common name.
Description
Pteralopex flanneryi is the largest species of Pteralopex
(Tables 1, 2). It agrees with P. atrata in dental and pelage
features as described in detail by Andersen (1912) and Hill &
Beckon (1978), including its overall blackish coloration (both
dorsally and ventrally). However, in adults of P. flanneryi there
is occasionally pale tipping to the hairs on the breast (absent in
438 K. M. Helgen
Figure 2 The Greater monkey-faced bat (Pteralopex ?anneryi), in life. Painting by Peter Schouten, rendered from the holotype (Flannery &
Schouten, 2004).
P. atrata), and thewings always lackmottling (variably present
in P. atrata; see Flannery, 1991; Parnaby, 2002b). It differs
further from P. atrata in its absolutely larger skull (Table 1;
Figs. 4, 5), body size (Table 2), and teeth (Table 3), and
(comparing adults) in its proportionately much more elongate
rostrum (ONL/CBL= 0.575?0.632 in P. flanneryi vs. 0.491?
0.553 in P. atrata).
Several images of this species? skull and external appear-
ance have previously been figured in the literature: Flannery
(1995) figured both a skull (p. 450) and the dorsal aspect of a
Paci?c monkey-faced bats 439
P. atrata P. ?anneryi P. anceps
sample n= 13a n= 7b n= 4c
CBL 64.0? 1.46 74.2? 1.42 70.1? 1.08
61.80?66.62 71.52?75.63 68.53?70.96
ONL 19.1? 0.80 23.0? 0.79 20.2? 0.58
17.80?20.38 21.82?23.94 19.50?20.80
ZYG 38.8? 0.91 43.3? 0.95 40.3? 0.67
37.00?40.03 42.21?44.57 39.53?41.12
BBC 23.2? 0.91 25.1? 1.00 26.6? 0.69
21.50?24.57 23.90?27.20 25.90?27.50
MTR 24.6? 0.55 28.6? 0.54 25.7? 0.66
23.7?25.22 27.63?29.18 25.00?26.60
MH 29.8? 1.00 34.7? 1.02 29.9? 0.59
28.35?31.11 33.05?35.77 29.16?30.38
PL 36.2? 1.08 41.9? 1.14 38.6? 0.89
34.74?37.67 39.99?43.23 37.38?39.38
a AM M3373, AM M19220, BMNH 34.7.2.31, BMNH 88.1.5.9, BMNH
89.4.3.1, MVZ 101673, MVZ 101674, SMF 44786, SMF 48306, SMF 48307,
SMF 48308, SMF 48309, SMF 50721.
b AMM6282, AMM6347, AMM6498, BBM-BSIP 23682, USNM276973,
USNM 276974, USNM 277112.
c AM M6283, AM M6497, BBM-NG 61213, BBM-NG 61435.
Table 1 Selected skull measurements for Pteralopex atrata,
P. ?anneryi and P. anceps. Values represent
means?SD, with ranges given below.
P. atrata P. ?anneryi P. anceps
sample n= 13a n= 11b n= 5c
FA 137.4? 5.19 164.0? 4.04 154.5? 9.11
129?147 159?169 141?160
HB 217.1? 12.50 265.1? 12.81 260.4? 7.30
196?237 247?285 252?270
E 17.5? 1.60 22.5? 2.04 23.8? 1.30
15?20 20?26 22?25
WT 472 790 ?
438?506 (n= 2) 790 (n= 1)
a AM M3373, AM M19220, BMNH 34.7.2.31, BMNH 88.1.5.9, BMNH
89.4.3.1, MVZ 101673, MVZ 101674, SMF 44786, SMF 48306, SMF 48307,
SMF 48308, SMF 48309, SMF 50721. Weights (in grams) from AM M19219
and AM M19220.
b AM M6282, AM M6346, AM M6347, AM M6498, BBM-BSIP 23682,
BBM-NG 61066 (subadult with mature external measurements), FMNH
31561 (subadult with mature external measurements), USNM 276973,
USNM 276974, USNM 277112, Leary and Pita (in litt.). Weight (in grams)
from Leary and Pita (in litt.).
c AMM6283, AMM6497, BBM-NG61213, BBM-NG61435, PNGM22824.
Table 2 External measurements for Pteralopex atrata, P. ?anneryi,
and P. anceps. Values represent means ? SD, with ranges
given below.
study skin (p. 239) (as ?P. anceps?) and Parnaby (2002b: 153)
figured occlusal aspects of the dentition (as ?P. anceps?).
Representative wing measurements for the adult male
holotype and an adult female paratype of P. flanneryi (AM
M6347) are as follows, respectively (measured fromdry skins):
pollex 70.6, 66.5;D1M22.3, 22.9;D1P1 39.3, 39.6;D2M90.8,
87.2; D2P1 26.7, 24.8; D2P2 17.4, 22.4; D3M 117.5, 120.1;
D3P1 82.3, 81.0; D3P2 123.2, 123.8; D4M119.2, 118.0; D4P1
64.7, 70.0; D4P2 81.0, 83.6; D5M 126.5, 125.8; D5P1 52.1,
49.4; D5P2 45.1, 47.6; calcar 19.3, 23.5. Representative wing
measurements for a nearly-mature subadult (AM M19822) are
as follows (measured from dry skin): pollex 62.1; D1M 19.1;
D1P1 39.2; D2M 75.1; D2P1 21.3; D2P2 18.9; D3M 109.1;
D3P1 72.8; D3P2 99.6; D4M 104.3; D4P1 65.1; D4P2 72.0;
D5M 112.7; D5P1 49.0; D5P2 39.2; calcar circa 17.
Remarks
In previous literature, the identity of P. flanneryi has been
confused with both P. anceps (with which it is widely sym-
patric) and P. atrata (its allopatric sister species, endemic to
Guadalcanal and New Georgia). For example, Phillips (1968),
Flannery (1995), Bonaccorso (1998), and Parnaby (2002b)
identified some or all museum specimens of P. flanneryi as
?P. anceps?, and Sanborn (1931) identified the Ysabel speci-
men listed above as ?P. atrata.? Due to this past taxonomic
confusion, the morphological traits of P. flanneryi, P. atrata
and P. anceps are contrasted in detail below.
External differences between sympatric Pteralopex
?anneryi and Pteralopex anceps
Pteralopex flanneryi and P. anceps (the two species of Pter-
alopex sympatric on Bougainville and Choiseul) can be easily
identified on the basis of pelage differences. In P. flanneryi the
pelage is relatively short overall, with the fur on the mid-back
and dorsal surface of the pes strongly adpressed, the forearm
and tibia naked or only sparsely furred, and the venter darkwith
occasional pale tipping on the hairs on the breast (all of these
features but the last are shared with P. atrata of Guadalcanal
and New Georgia). In P. anceps the fur is long and luxuri-
ous, the fur of the mid-back and pes are not at all adpressed,
the proximal third of the forearm and the entire tibia are very
heavily haired, and ventrally the breast bears an expansive,
conspicuous white or yellow patch of fur (Fig. 3). (Each of
these traits is an instantly diagnostic feature for P. anceps rel-
ative to P. flanneryi.) The external ear of P. anceps, though not
actually smaller than that of sympatric P. flanneryi (Table 2),
is more hidden in the long fur than in the latter species.
Pteralopex anceps and P. flanneryi are similar in over-
all size, though P. anceps has a significantly (almost abso-
lutely) shorter forearm (Table 2). Skulls of P. flanneryi are
absolutely wider and longer than P. anceps, suggesting that
the former species is heavier than the latter (P. anceps has yet
to be weighed).
Dental differences between sympatric Pteralopex
?anneryi and Pteralopex anceps
Andersen (1912) accurately described and excellently illus-
trated the trenchant dental differences that separate the allo-
patric species P. atrata and P. anceps. These correspond es-
sentially to the respective dental traits of the two Pteralopex
species sympatric in the northern Solomons (flanneryi and
anceps). Selected traits include the larger size of the incisors
and (vertically) more elongate canines but smaller size of the
440 K. M. Helgen
P. anceps
P. atrata P. ?anneryi
n= 12 n= 5 a b c
canine length 5.4? 0.14 6.0? 0.23 6.00 ? ?
5.26?5.65 5.70?6.29
canine width 4.2? 0.16 4.7? 0.34 4.18 ? ?
4.01?4.47 4.17?5.10
canine height 7.6? 0.31 9.3? 0.32 9.76 ? ?
7.20?8.09 8.83?9.61
PM3w 4.3? 0.25 4.6? 0.07 4.24 4.01 4.7
3.93?4.63 4.52?4.71
PM4l 5.0? 0.21 5.8? 0.12 5.04 4.82 5.8
4.64?5.23 5.60?5.89
PM4w 4.4? 0.24 5.0? 0.09 4.61 4.08 4.8
4.04?4.80 4.84?5.09
M1l 4.4? 0.28 5.3? 0.14 4.74 4.68 5.0
4.06?5.00 5.02?5.39
M1w 3.9? 0.12 4.5? 0.08 4.01 3.84 4.2
3.69?4.11 4.39?4.57
M2l 3.0? 0.14 3.6? 0.16 3.13 2.83 3.2
2.80?3.20 3.32?3.70
M2w 2.9? 0.13 3.3? 0.15 3.16 2.73 3.0
2.68?3.08 3.13?3.46
PM3w 3.4? 0.17 3.8? 0.23 3.29 2.97 3.6
3.13?3.60 3.50?4.06
PM4w 3.8? 0.13 4.4? 0.16 3.69 3.36 3.9
3.65?4.04 4.27?4.60
M1w 3.8? 0.15 4.5? 0.18 3.62 3.43 3.9
3.55?4.04 4.23?4.64
M2w 3.5? 0.13 4.0? 0.19 3.52 3.47 3.8
3.25?3.70 3.84?4.29
M3w 2.7? 0.15 3.1? 0.06 3.00 2.60 3.0
2.51?3.00 3.00?3.16
Table 3 Selected dental measurements for Pteralopex atrata, P. ?anneryi, and P. anceps. a=AM
M6283; b=AM M6497; c= holotype of P. anceps (from Andersen, 1912: 441). Values
represent means?SD, with ranges given below. w=width; l= length.
premolars and molars in P. anceps relative to P. flanneryi/
P. atrata, a massive P4 very long relative to its width in
P. anceps (more subsquare in P. flanneryi/P. atrata), heavily-
developed anterior cingula on PM4 and M1 in P. flanneryi/
P. atrata (greatly reduced in P. anceps), and an unbroken el-
evated ridge on the lingual edges of PM4, M1, and M2 in
P. flanneryi/P. atrata (interrupted by rising and falling cusps
in P. anceps). Individuals from Bougainville with the respect-
ive pelage traits of P. anceps and P. flanneryi are also those
with the respective dental traits of each species, confirming
by this concordance the presence of two different species of
Pteralopex on the island.
Craniometric differences between sympatric Pteralopex
?anneryi and Pteralopex anceps
A principal component analysis utilizing six cranial and
mandibular variables (CBL, ONL, ZYG, BBC, MTR, MH)
for all adult Pteralopex from Bougainville segregates these in-
dividuals into two well-separated groupings (Fig. 4; Table 1).
These two clusters correspond exactly to the distribution of
discrete pelage and dental traits as described above, again con-
firming the distinctness of the two species.
Though superficially similar, the skulls of P. anceps and
P. flanneryi differ consistently in shape. Although the skull
of P. anceps is absolutely shorter and narrower than that of
sympatric P. flanneryi, it has a more expansive braincase,
both in relative terms (BBC/CBL= 0.375?0.389 in P. anceps
vs. 0.323?0.363 in adult P. flanneryi) and average measure-
ments (e.g. Table 1). In lateral view, the braincase of P.
anceps is more strongly deflected from the horizontal axis
of the palate/toothrow (more pronounced klinorhynchy; see
Fig. 7: the alveolar line passes above the supraoccipital) than
in P. flanneryi (in which the alveolar line intersects the
supraoccipital). The coronoid process of the mandible is
proportionally reduced in P. anceps relative to P. flanneryi
(MH/CBL= 0.410?0.443 in P. anceps vs. 0.447?0.484 in
P. flanneryi), as is the length of the palate (PL/CBL= 0.545?
0.557 in P. anceps vs. 0.558?0.585 in P. flanneryi). In dorsal
Paci?c monkey-faced bats 441
Figure 3 Ventral colouration (seen here in half tone) in sympatric Pteralopex anceps (AM M6283, above) and Pteralopex ?anneryi
(AM M6347, below), both from Bougainville.
view, the zygomata of P. anceps are more nearly parallel
than in other Pteralopex, including P. flanneryi, in which the
zygomata tend to diverge slightly behind the postorbital bar.
Geographic variation in Pteralopex atrata and
Pteralopex ?anneryi
Pteralopex atrata and P. flanneryi are immediately related al-
lopatric species (Fig. 1) that are largely identical in discrete
dental and pelage traits, but differ absolutely in body, dental,
and cranial size and proportions. These size differences are
aptly demonstrated by univariate measurements (Tables 1?
3) and by multivariate analysis. For example, a principal
component analysis utilizing six cranial and mandibular vari-
ables (CBL, ONL, ZYG, BBC, MTR, MH) cleanly separates
P. atrata and P. flanneryi on the basis of overall skull size
(Fig. 6), and (importantly) suggests further that geographic
variation within each species is minimal. A single skull from
New Georgia (see account of P. atrata below) is indistin-
guishable in size and shape from adult P. atrata skulls from
Guadalcanal. Adult skulls of P. flanneryi from Bougainville
and Choiseul group closely together, and subadult skulls from
Ysabel and Buka (entered into the analysis out of geographic
interest) also fall near this cluster but differ slightly in shape
owing to their incomplete maturity (Fig. 6). Subadults of
P. flanneryi can be distinguishedwith confidence even from the
largest adult specimens of P. atrata by their still-larger overall
size and much larger teeth (e.g. Table 3).
Remarks on Pteralopex ?anneryi
Pteralopex flanneryi was last collected on Bougainville in
1968, on Choiseul in 1964, and on Buka in 1987. Field surveys
in the 1990s failed to record P. flanneryi from Bougainville
(Bonaccorso, 1998; S. Hamilton, in litt.), Choiseul
(Bowen-Jones et al., 1997), Buka (Bonaccorso, 1998), and
Ysabel (Parnaby, 2002b: 158). However, Leary and Pita
(in litt.) recently collected an individual on Ysabel (i.e.
442 K. M. Helgen
f
f
f
f
f
f
a
a
a
a
-0.3 -0.2 -0.1 0.0 0.1 0.2 0.3
PC1: 88.11%
-0.12
-0.10
-0.08
-0.06
-0.04
-0.02
0.00
0.02
0.04
0.06
0.08
0.10
P
C
2:
  7
.6
5%
Figure 4 Principal component analysis: adult Pteralopex skulls from Bougainville. a=Pteralopex anceps; f=Pteralopex ?anneryi.
Variable PC1 PC2
CBL ?0.934 ?0.141
ONL ?0.991 0.022
ZYG ?0.931 ?0.132
BBC 0.552 ?0.832
MTR ?0.966 ?0.043
MH ?0.977 ?0.099
Variance explained 88.11% 7.65%
Table 4 Factor loadings for Fig. 4. Principal components extracted
from a covariance matrix of log-transformed craniodental
variables.
Barora Fa) in 2000 (see specimens listed above). Pteralopex
flanneryi is rarely encountered across its range, is restricted to
primary forest, and has apparently declined in recent decades
(Troughton, 1936; Flannery, 1995; Bowen-Jones et al., 1997;
Bonaccorso, 1998; Parnaby, 2002b; S. Hamilton, in litt.). As a
very large-bodied bat it is undoubtedly susceptible to contin-
ued decline via deforestation and hunting pressure in populated
areas. It has previously been classified by the IUCN as ?Critic-
ally Endangered? (Hilton-Taylor, 2000: as ?P. anceps?), a status
it most certainly deserves.
An adult female from Bougainville was lactating when
collected in July (Phillips, 1968), and a subadults (forearm
length 145 to 162) have been collected in February, April,
and September (see specimens above). Little is known of diet
of this species, but one individual was observed eating green
coconuts on Ysabel (Sanborn, 1931; see Discussion below),
and it is undoubtedly largely frugivorous. The large body size
of P. flanneryi relative to P. atrata may be explained in part
by character displacement in the former species relative to
the similarly large-bodied P. anceps (Table 2), with which it
is sympatric on Bougainville and Choiseul (see account of
P. anceps below).
Pteralopex atrata Thomas, 1888
(Fig. 5 lower)
HOLOTYPE. BMNH 1888.1.5.9, adult male, body in alco-
hol with skull extracted and cleaned, from Aola (09?32?S,
160?29?E), presumably at sea level, Guadalcanal (Central
Province, Solomon Islands), collected by C.M. Woodford
(Thomas, 1888; Woodford, 1890; Andersen, 1912).
Material examined
I have examined 20 specimens of P. atrata (which, as far as I
know, constitute all museum material of this species):
1. The type series of P. atrata from Aola (09?32?S, 160?29?E),
Guadalcanal at BMNH (five specimens; see Andersen
(1912) for details), one specimen of which (BMNH
1888.1.5.10) was sent in exchange to Frankfurt (now SMF
44786).
Paci?c monkey-faced bats 443
Figure 5 Skulls of Pteralopex ?anneryi (above: AM M6282, adult male, Bougainville) and Pteralopex atrata (below: AM M3373, adult male,
Guadalcanal).
2. One specimen (BMNH 1934.7.2.31, adult female, skin and
skull), collected on the ?northwest coast? of Guadalcanal in
July?August 1927 by C.E. Hart.
3. A total of seven specimens from ?Lavors? (=Lavoro Plant-
ation; (09?19?S, 159?39?E) and the Poha River Valley,
400m (09?24?S, 159?54?E), Guadalcanal (AM M3373,
3374, 3375, 4218, 4219 [Lavoro]; AM M19219, 19220
[Poha]).
4. Two specimens (MVZ 101673 and 101674, skins and
skulls) from the Nalimbiu River (09?24?S, 160?09?E),
1mile inland, 15m,Guadalcanal, collected inOctober 1944
by L. Adams.
5. Four specimens at SMF (skins and skulls) collected on
Guadalcanal in 1974 by H.L. Bregulla: SMF 48306 (adult
female) and SMF 48307 (adult female) from Vilu; SMF
48308 (adult female) from ?Lakalaka and Komambulu?
(=Komovaolo? 09?55?S, 160?34?E); SMF 48309 (adult
male) from Verawaulu (09?47?S, 159?57?E).
6. A single specimen (SMF 50721, adult female, skull,
skin now missing) from ?Jericho and Munda Village?
(Munda= 08?21?S, 157?13?E), New Georgia (Western
Province, Solomon Islands), collected April-May 1975 by
Bregulla. This is the first record of P. atrata from outside of
Guadalcanal. The skull of this individual does not appear
to differ morphologically or morphometrically from skulls
of P. atrata from Guadalcanal (e.g. Fig. 6). However, it
bears an annotation suggesting that it may represent a new
species of Pteralopex (D. Kock, in litt.), which possibly
suggests that the skin (now missing) is distinctive relative
to Guadalcanal P. atrata. Until the skin is rediscovered or
additional specimens are collected, I can conclude nothing
more about the systematic status of this specimen. Previ-
ously, only P. taki has been known from New Georgia.
Diagnosis
Pelage variable in length but usually relatively short, with
ear exposed, fur of the mid-back and the dorsal surface of
the pes adpressed, and the tibia and forearm naked or very
sparsely furred; overall colour black dorsally and ventrally
(with wings occasionally mottled). Size medium (adult con-
dylobasal length< 67; forearm< 150), but absolutely larger
than sympatric P. pulchra and P. taki (Parnaby, 2002b), with
subsquare PM4, relatively heavy cheekteeth, and PM4, M1,
and M2 with lengthened, elevated lingual ridges. Other diag-
nostic dental traits reviewed by Andersen (1912).
444 K. M. Helgen
g
g
g
g
g
g g g
g
g
g
g
n
b
b
b
b
b
b
c
y
bu
-0.5 -0.4 -0.3 -0.2 -0.1 0.0 0.1 0.2 0.3 0.4 0.5
PC1: 89.11%
-0.15
-0.10
-0.05
0.00
0.05
0.10
0.15
0.20
P
C
2:
  6
.0
6%
Figure 6 Principal component analysis of skulls of Pteralopex atrata (g=Guadalcanal; n=New Georgia) and Pteralopex ?anneryi.
b=Bougainville; c= Choiseul; y= Ysabel (subadult); bu=Buka (subadult).
Variable PC1 PC2
CBL ?0.982 ?0.130
ONL ?0.960 ?0.228
ZYG ?0.949 0.003
BBC ?0.776 0.595
MTR ?0.957 0.184
MH ?0.961 ?0.004
Variance explained 89.11% 6.06%
Table 5 Factor loadings for Fig. 6. Principal components extracted
from a covariance matrix of log-transformed craniodental
variables.
Distribution
Recorded from primary lowland forest on Guadalcanal (from
sea level to 400m, but possibly up to 1000?1200 m; Flannery,
1991) and from New Georgia (see above). Most specimens
originate from at or near sea level; the mean elevation of all
sites where this bat has been collected is 52m (n= 8; range
0?400m).
Remarks
Pteralopex atrata was recently reviewed by Flannery (1995)
and Parnaby (2002b). Andersen (1912) and Hill & Beckon
(1978) reviewed aspects of dental morphology, Thomas (1888)
provided a colour illustration of the head of this species (re-
figured by Flannery, 1995), and Flannery (1995) figured a
freshly-killed specimen. Andersen (1912), Flannery (1995),
and Parnaby (2002b) figured skulls, and Andersen (1912)
provided representative wing measurements.
Apart from brief notes presented by Flannery (1991,
1995), little has been recorded of the basic biology of P. atrata.
Judging from sight records (Flannery, 1995) and museum spe-
cimens, it may be more common than other Pteralopex, and
is apparently distributed in primary lowland forest across the
island of Guadalcanal (Fig. 1). However, its IUCN classifica-
tion of ?Critically Endangered? (Hilton-Taylor, 2000; Parnaby,
2002b) probably remains appropriate (Parnaby, 2002b).
P. atrata is regionally sympatric with P. pulchra (i.e.
both occur on Guadalcanal) but the two species have not been
collected syntopically and are probably altitudinally strati-
fied, with P. atrata occupying primary lowland forest and P.
pulchra endemic to cloud forest situated at 1200m and higher
(Flannery, 1991). Interestingly, certain morphometric relation-
ships between these two Guadalcanal species are analogous
to those between P. flanneryi and P. anceps in the north-
eastern Solomons. In both cases, the higher-elevation species
(P. pulchra and P. anceps) are smaller-bodied (FA 118 in the
only known specimen of pulchra), with longer pelage, smaller
teeth, proportionally wider braincases (BBC/CBL= 39.8% in
pulchra vs. 33.6?38.8% in atrata), and reduced coronoid pro-
cesses relative to their respective sympatric lowland congeners.
Paci?c monkey-faced bats 445
Figure 7 Skull of Pteralopex anceps (AM M6283, young adult male, Bougainville). A photograph of the skull of this species has not previously
been published.
Both P. atrata and P. taki are now recorded from New
Georgia, where they are likely to occur syntopically in primary
lowland forest (see Discussion below).
Maa (1966) correctly recorded the batfly Cyclopodia
macracantha macracantha Theodor, 1959 as an ectoparasite
of this species on Guadalcanal. He also recorded Cyclopodia
solomonarum Theodor, 1959 from this species (listed from
a wide array of localities in the Solomon Islands), but
these records appear to be mistaken, based instead on bat-
flies taken from various flying-foxes (Pteropus spp.), which
C. solomonarum certainly parasitizes (Theodor, 1959; Maa,
1966).
Pteralopex anceps Andersen, 1909
(Figs 3 upper, 7)
HOLOTYPE. BMNH 1908.11.16.7, subadult female, skin and
skull, from Bougainville (Northern Solomons Province, Papua
New Guinea), presumably in coastal lowland forest, collected
April 1904 by A.S. Meek.
Other material examined
AM M6283 (adult male, skin and skull) and AM M6497
(adult male, skin and skull), from Buin District (ca. 06?50?S,
155?44?E), 10 miles inland, 150 meters, Bougainville, collec-
ted March 1937 and June 1938, respectively, by J.B. Poncelet.
BBM-NG 61435 (adult male, skin and skull), from Mt.
Balbi (05?55?S, 154?59?E), 1900meters (?moss forest? on skin
label), Bougainville, collected 15 April 1968 by A.B. Mirza.
PNGM 22824, adult female, skin and skull, from
Mutahi (=Muguai, 06?43?S, 155?45?E), 1100 meters (?rain-
forest? on skin label), Bougainville, collected 10 March 1968
by A.B. Mirza. BBM-NG 61213 (adult female, skin and skull)
and BBM-NG 61214 (subadult female, skin and skull), from
Mutahi, 1100m (?rainforest? on skin label), Bougainville, col-
lected 12 March 1968 by A.B. Mirza. Habitat details (in quo-
tations) taken from specimen tags.
Diagnosis
Forearm less than or equal to 160 (vs. 159?169 in sympatric
P. flanneryi); pelage very long and thick, with ear unreduced
but inconspicuous in the long pelage, the fur of the mid-back
not adpressed, and the tibia, dorsal surface of the pes, and
proximal third of the forearm very densely furred; overall col-
our blackish brown, but venter with a expansive yellow or
white fur patch on the breast. Skull smaller than sympatric
P. flanneryi (Table 1; Fig. 4), with wider and more horizont-
ally deflected braincase, reduced coronoid process, and less
elongate palate; compared to P. flanneryi, incisors relatively
larger and canines (vertically) longer, but cheekteeth relat-
ively smaller except for a massive, rectangular PM4 (Table 3);
PM4, M1, and M2 without unbroken, elevated lingual ridges
along their lengths. Other, lesser diagnostic dental traits were
reviewed by Andersen (1912).
Distribution
Pteralopex anceps is known only from Bougainville and
Choiseul, where it has a wide recorded altitudinal range. On
Bougainville it has been collected in coastal and inland forests
at and near sea level, in mid-elevation tall-canopy rainforest
at 1100m, and in mossy montane forest at 1900m on Mt.
Balbi. Notably, Pteralopex anceps and P. flanneryi have been
collected syntopically at 150m near Buin (series at AM).
This species is recorded from Choiseul on the basis of
an unambiguous sight record reported by Bowen-Jones et al.
(1997) from primary forest at 850m in the vicinity of Mt.
Maetambe (07?05?S, 157?01?E) (?identification of the bats was
enabled . . . by the absence of large protruding ears . . . the chest
was conspicuously white in colour?). Phillips? (1968) earlier
record of this species from Choiseul is based on a misidenti-
fied specimen of P. flanneryi (see above). Pteralopex anceps
has not been recorded from Buka (contra Flannery [1995],
Bonaccorso [1998], and Parnaby [2002b]), or Ysabel (contra
Parnaby [2002b]); these distributional records are instead also
based on misidentified specimens of P. flanneryi (see above).
However, I suspect that this species does occur onYsabel, even
though there no valid records to date (see Discussion below).
Remarks
The correct identity and unique traits of P. anceps have gener-
ally been obscured or ignored in the literature since Andersen?s
446 K. M. Helgen
P. anceps P. atrata and P. ?anneryi
Tibia heavily furred naked or sparsely furred
Fur of mid-back not adpressed adpressed
Fur of breast with white or yellow chest patch without chest patch
Braincase wider relative to skull length narrower relative to skull length
I1 and I2 larger, generally broader than long smaller, generally longer than broad
PM4 and M1 anterior cingula weak anterior cingula heavy
PM4 much longer than broad subsquare
PM4, M1, and M2 without unbroken lingual ridges with unbroken lingual ridges
Table 6 Selected qualitative distinctions between Pteralopex anceps and P. atrata/P. ?anneryi.
(1909b, 1912) papers; however Hill & Beckon (1978) cor-
rectly described characteristic dental and pelage traits of this
species on the basis of the holotype. As noted above, Laurie &
Hill (1954) arranged P. anceps as a subspecies of P. atrata
without supporting evidence, an arrangement later criticized
on morphological grounds by Hill & Beckon (1978). Taxo-
nomic concepts of P. anceps employed by Phillips (1968),
Koopman (1994), Flannery (1995), Bonaccorso (1998), and
Parnaby (2002b) all failed to distinguish between P. anceps
and P. flanneryi, which are sympatric on Bougainville and
Choiseul, and thus refer to both of these species. How-
ever, a drawing figured by Bonaccorso (1998) correctly de-
picts the appearance and external diagnostic traits of this
species.
Representative wing measurements for an adult male
(AM M6497) are as follows (measured from dry skins): pollex
64.4; D1M 19.9; D1P1 41.7; D2M 83.6; D2P1 20.3; D2P2
19.7; D3M 118.9; D3P1 72.4; D3P2 107.7; D4M 112.1;
D4P1 64.3; D4P2 74.5; D5M 117.7; D5P1 48.4; D5P2 39.3.
Andersen (1912: 440) presented wing measurements for the
immature holotype.
Four of the seven known museum specimens of
P. anceps from Bougainville were collected in high eleva-
tion rainforest or mossy montane forest (i.e. cloud forest)
at elevations of 1100m or higher, where the larger-bodied,
shorter-furred P. flanneryi has never been recorded and may
not occur. OnChoiseul,P. anceps has been observed only in in-
land primary forest at 850m (Bowen-Jones et al., 1997), and,
on the basis of local interviews, Bowen-Jones et al. (1997)
suggested that it does not occur in coastal forest there (the
mountains of Choiseul rise to 1060 m; Table 8). These obser-
vations potentially highlight differences in habitat preference
between P. anceps and P. flanneryi. The long, thick pelage of
P. anceps (recalling that of Pteralopex pulchra, a moss forest
endemic from montane Guadalcanal) lends additional sup-
port to this supposition. The mean altitude of all sites where
P. anceps has been collected or observed is 800m (n= 5, range
0 to 1900m); the mean for P. flanneryi is 50m (n= 8, range 0
to 200m).
Pteralopex anceps has been observed roosting solitar-
ily and in groups (Bowen-Jones et al., 1997: 212). Subadults
(forearm 137 to 138.5) have been collected in March and April
on Bougainville (see specimens above). Little is known of the
diet of this species, but its large incisors and small molars re-
lative to sympatric P. flanneryi undoubtedly denote trenchant
differences in feeding ecology between the two species; in
particular, P. anceps may engage in tree-gouging for exud-
ate feeding (see Discussion below). Judging from museum
specimens, toothwear in adults is extensive, suggesting hard,
abrasive fruits also form part of the diet.
Maa (1968) recorded the batfly Cyclopodia macracantha
longiseta (taken from AM M6497) from this species on
Bougainville.
Anatomically, P. anceps is in manyways the most highly-
derived species of monkey-faced bat. It differs from other
Pteralopex in its considerably deflected braincase, expanded
incisors, reduced molars, and lack of lingual ridges on the
lower first and secondmolars. These autapomorphies, together
with the very close genetic relationship assumed here between
P. flanneryi and P. atrata, and that demonstrated between
P. taki and P. pulchra (Ingleby & Colgan, 2003), may sug-
gest that P. anceps is the most basal species of Pteralopex
(sensu stricto). Alternately, the blackish fur and large size of
P. anceps may betray a sister relationship with P. flanneryi and
P. atrata. No genetic study has yet included samples of all
six monkey-faced bat species, but this remains an important
priority for resolving the pattern, timing, and geography of
diversification in the group.
Museum specimens demonstrate that P. anceps has
not been collected on Bougainville since 1968, despite
some survey efforts (e.g. Flannery, 1995; Bonaccorso, 1998;
S. Hamilton, in litt.). The species was recorded most recently
fromChoiseul by Bowen-Jones et al. (1997), who observed the
species only twice during six months? fieldwork on Choiseul
in 1995. Given its limited distribution, apparent rarity, and
restriction to old-growth forests, the current IUCN designa-
tion of ?Critically Endangered? for P. anceps (Hilton-Taylor,
2000; Parnaby, 2002b) is still appropriate, despite the present
changes in the species? taxonomic definition.
Mirimiri gen. nov.
Diagnosis
The genus Mirimiri can be distinguished from all pteropo-
did genera but Pteralopex by the distinctive combination of
morphological traits listed above (?Monophyly of monkey-
faced bats?). In addition, each of the following eight
characters is perfectly diagnostic for Mirimiri with respect to
Paci?c monkey-faced bats 447
Taxonomic and geographic arrangement of Pteralopex: Parnaby (2002b)
Pteralopex atrata: Guadalcanal
Pteralopex anceps: Bougainville, Buka, Choiseul, Ysabel
Pteralopex pulchra: Montane Guadalcanal (1200 m and higher)
Pteralopex taki: New Georgia and Vangunu, formerly Kulumbangra
Pteralopex acrodonta: Montane Taveuni (1000 m and higher), possibly Vanua Levu
Taxonomic and geographic arrangement of Pteralopex (and Mirimiri): this study
Pteralopex atrata: Guadalcanal, New Georgia
Pteralopex ?anneryi: Bougainville, Buka, Ysabel, Choiseul
Pteralopex anceps: Bougainville, Choiseul, possibly Ysabel
Pteralopex pulchra: Montane Guadalcanal (1200 m and higher)
Pteralopex taki: New Georgia and Vangunu, formerly Kulumbangra
Mirimiri acrodonta: Montane Taveuni (1000 m and higher), possibly Vanua Levu
Table 7 Changes to Pteralopex taxonomy and distribution.
Pteralopex:
1. Postorbital processes not annectent to the zygomata in
adults.
2. Interorbital region of skull not conspicuously narrowed.
3. M2 not strongly reduced relative to M1 (greater than half
the mass).
4. Labial margins of PM4 and M1 divided into three distinct
cusps.
5. Length of D4P2 subequal to D4P1.
6. Five divided median ridges in the soft palate.
7. Two serrated supplementary posterior ridges in the soft
palate.
8. Slight sexual dichromatism in pelage, with the back and
rump more strikingly golden in males than females (more
khaki-coloured in the latter).
Of these eight traits that distinguish the monkey-faced
bat genera, traits 4, 7, and 8 represent character states more
highly derived in Mirimiri than Pteralopex, while ( judging
from their distribution among the Pteropodidae) the other states
are probably more primitive. Parnaby (2002b: 150) illustrated
differences in traits 2 and 5 between Mirimiri and Pteralopex
graphically. The palatal ridges (traits 6 and 7) differ from
Figure 8 Skull of Mirimiri acrodonta (AM M24030, adult male, Taveuni).
Pteralopex not only in number, but also in details of struc-
ture (Parnaby, 2002b). The cheekteeth of Mirimiri differ from
Pteralopex not only in details of cuspidation along the labial
margins of PM4 and M1 (as described above), but also in other
features as reviewed in further detail by Hill & Beckon (1978).
CONTENT. Monotypic; comprising only M. acrodonta of
montane Taveuni (Fiji).
ETYMOLOGY. Fijian, ?mist? (Capell, 1968), in allusion to the
cloud forests this monkey-faced bat inhabits, and appropriate
for such an elusive and endangered animal.
Mirimiri acrodonta (Hill and Beckon, 1978)
(Figs 8, 9)
HOLOTYPE. BMNH 1977.3097, adult male, skin and
skull, from Mt. Koroturanga (=Des Voeux Peak, 16?50.5?S,
179?58?W), 1170m, Taveuni, Fiji, collected 3 May 1977 by
W.N. Beckon.
Other material examined
Three specimens from the type locality, collected in May 1990
by Pavel German (AM M24030, adult male, 1190 m; AM
448 K. M. Helgen
M24031, adult female, 1010 m; AM M24579, adult female,
1010m). Apart from these three specimens and the holo-
type, this species is known only by one additional specimen
(the paratype, 1190m) and an individual that was netted but
escaped (1170m); see Hill & Beckon (1978).
Diagnosis
As for the genus.
Distribution
Mirimiri acrodonta is recorded only from mossy montane
forest (i.e. cloud forest) on Mt. Koroturanga (=Des Voeux
Peak) on the volcanic Fijian island of Taveuni, at altitudes
between 1000 and 1200m (Hill & Beckon, 1978; Flannery,
1995). Aspects of the vegetation and microclimate of this
unique forest (Fig. 10) were reviewed in useful detail by Ash
(1987) and Mueller-Dombois & Fosberg (1998). This species
probably occurs also in highmontane forest on the larger, adja-
cent island of Vanua Levu (especially Delaikoro Peak), where
it is recorded by unvouchered observations (Hill & Beckon,
1978; P. German, pers. comm.; Flannery, 1995), but this awaits
confirmation. Themean elevation of the four sites onMt. Koro-
turanga where this bat has been mistnetted (Hill & Beckon,
1978; Flannery, 1995) is 1140m (range 1010?1190m).
Remarks
All that is known of the biology of the Fijian monkey-faced
bat has previously been reviewed by Hill & Beckon (1978),
Flannery (1995), and Parnaby (2002b). Hill & Beckon (1978)
provided an excellent descriptive morphological review;
Parnaby (2002b) discussed additional museum specimens
and figured the species? skull, and Flannery (1995) provided
brief biological notes and figured a photograph of the species
in life. Using allozymes, Ingleby & Colgan (2003) studied
genetic differences between Mirimiri and Pteralopex, con-
cluding that ?the level of separation betweenPt. acrodonta and
its two congeners in the Solomon Islands [fixed differences at
38% of loci] . . . is about the level found in two other Pacific
megachiropteran genera for which electrophoretic data are
available [Nyctimene and Paranyctimene].?
Mirimiri joins a number of other terrestrial vertebrate
genera endemic to Fiji, including five living avian genera
(Prosopeia, Phigys, Trichocichla, Lamprolia, and Xanthotis),
two extinct avian genera (Natunaornis, Megavitiornis), one
living snake genus (Ogmodon), an extinct iguanid genus (Lapi-
tiguana), and the extinct mekosuchine crocodylian genusVolia
(Stattersfield et al., 1998; Worthy, 2000, 2001; Molnar et al.,
2002; Pregill & Worthy, 2003). The iguanid genus Brachylo-
phus is another near-endemic genus, extending only to Tonga
(Gibbons, 1981; Pregill & Worthy, 2003).
Mirimiri acrodonta forms part of a relatively depau-
perate Fijian bat fauna which also includes two flying-foxes
(Pteropus samoensis and P. tonganus), the blossum-bat No-
topteris macdonaldi, the sheathtail-bat Emballonura semi-
caudata, the mastiff-bat Chaerephon bregullae, and possibly
a species of the vespertilionid genus Nyctophilus (Flannery,
1995; Helgen & Flannery, 2002; Parnaby, 2002a; Ingleby &
Colgan, 2003). Of these, M. acrodonta is the most ancient ele-
ment, and the only endemic taxon in the Fijian bat fauna apart
from the flying-fox subspeciesPteropus samoensis nawaiensis
Gray, 1870.
Mirimiri acrodonta is currently classified as ?Critically
Endangered? (Parnaby, 2002b). Though little remains known
of its general biology, this is a status it most certainly deserves.
Whether this species extends to Vanua Levu or not, its total
geographic range is undoubtedly minuscule. As Ash (1987)
noted:
The extent of cloud-forest in Fiji is very limited, occupying only
the ridges on the highest peaks [above about 900m altitude]. In
Taveuni cloud-forest occurs only along about 10 km of the main
range and in the other islands it is restricted to 1?2 km of ridges
around isolated peaks which are separated by distances of 30?
100m.
Discussion
Ecomorphological differences between sympatric
Pteralopex species
Until relatively recently (Hill & Beckon, 1978) all species of
Pteralopex were thought to be allopatric, but four islands in
the Solomons are now known to support two sympatric species
ofPteralopex. Flannery (1991) reported the first examplewhen
he documented the presence of both P. atrata and P. pulchra on
Guadalcanal. There the two species appear to be elevationally
stratified, with P. pulchra endemic to mossy montane forests
at 1200m and above, and P. atrata occupying primary forest at
lower elevations (Flannery, 1991). Among many other traits,
P. pulchra differs from P. atrata in its absolutely smaller body
size, and longer pelage (Flannery, 1991, 1995).
On New Georgia, both P. taki and P. atrata have been re-
corded (this paper), although the latter species is known from
that island by a single specimen (collected in 1975) without
accompanying elevational data. Pteralopex taki is thus far
known only from lowland forest on NewGeorgia and Vangunu
(Flannery, 1995; Fisher&Tasker, 1997; Parnaby, 2002b). I sus-
pect that P. atrata also occurs in lowland forest on New Geor-
gia, as elsewhere it is recorded only from lowlandGuadalcanal.
Hence Pteralopex taki and P. atrata may occur syntopically in
lowland forest on New Georgia. If still present there, P. atrata
is no doubt much the rarer species, given that it was never
encountered by Fisher & Tasker (1997) during their survey
of pteropodids on New Georgia and adjacent islands. Though
ecomorphological distinctions between these taxa remain es-
sentially unstudied, their sympatric occurrence is no doubt fa-
cilitated in part by the absolute size difference between the two
species: P. taki has a forearm length of 112.8 to 122.8mm and
an adult body weight of 225 to 351 grams (Parnaby, 2002b),
while P. atrata has a forearm length of 129 to 147mm and an
adult body weight of 438 to 506 grams (Table 2).
On Bougainville and Choiseul, where P. anceps and
P. flanneryi both occur, the two species do not simply segre-
gate vertically, as they have been collected together in lowland
forest. Nor is the contrast in body size between P. flanneryi
and P. anceps as striking as that between P. atrata and
Paci?c monkey-faced bats 449
Figure 9 The Fijian monkey-faced bat (Mirimiri acrodonta), from Taveuni, in life. Photograph by Pavel German.
P. pulchra on Guadalcanal or between P. atrata and
P. taki on New Georgia. Instead, Pteralopex anceps and
P. flanneryi presumably segregate ecologically, preferentially
utilizing both different habitats (see account of anceps, above)
and different food sources (at least in part; see below). The
latter is suggested by their very different dentitions, with, for
example, P. anceps having relatively very large incisors and
smaller molars andP. flanneryiwith smaller incisors and larger
molars.
Nothing about the diet of P. anceps has been recorded in
the literature except that individuals have been observed roost-
ing in fig (Ficus) trees (Bowen-Jones et al., 1997), and very
450 K. M. Helgen
Figure 10 Cloud forest on Mt. Des Vouex, Taveuni, Fiji, 1100 m: habitat of Mirimiri acrodonta. Photograph by Tim Flannery.
little is known of the diet of P. flanneryi. (Feeding ecology of
only P. taki has been studied in detail: Fisher & Tasker, 1997.)
Only two observations relevant to the diet of P. flanneryi are
available: Sanborn (1931) observed a subadult from Ysabel
eating green coconuts, and A.B. Mirza noted on a museum tag
that one adult (BBM-NG 61066) was shot at night in a coconut
tree on Bougainville. In light of these observations, it is inter-
esting to note Parnaby?s (2002b: 158) following comments:
hunters in the Hograno district of Isabel . . . assured [me] that the
Pteralopex in their region did not feed on coconuts, young or old.
A number of people reported that the species chewed bark on
the trunk of selected trees. The expectorated pulp of Pteralopex
could evidently be readily recognized from that of Pteropus sp. by
shape and greater size. Individuals were said to be equally likely
to be encountered in coastal forests as on the higher elevations
of the nearby mountains, which reach some 1000m, the highest
mountains on Isabel.
Although only P. flanneryi has been recorded to date
from Ysabel, given that that species is indeed known to eat
coconuts and has never been collected above 200m, it seems
quite likely that Parnaby?s local informants were actually de-
scribing the behaviour of P. anceps. I suspect that the larger,
more robust molars of P. flanneryi are utilized to break open
coconuts and hard fruits with powerful lateral bites (Dumont &
Herrell, 2003; Marshall, 1985), while the more strongly-
developed incisors, anteriorly expanded braincase, and greater
cranial klinorhynchy of P. anceps (see above) may instead
facilitate tree-gouging (as described by Parnaby?s informants
above), perhaps for exudate feeding ? an ecological strategy
not uncommon among various small and medium-sized mam-
malswith similar adaptations in other tropical forests (Dumont,
1997; Vinyard et al., 2003). However, the extensively worn
cheekteeth of some older specimens of P. anceps suggests that
harder foods also factor significantly in the diet.
Geography of monkey-faced bats and future
survey priorities
Throughout the south-west Pacific, flying-foxes (species of
Pteropus) are common inhabitants of islands of all sizes, in-
cluding very small and extremely remote islands.Many aspects
of their ecology and physiology are suited to survival on small
islands (McNab, 1994; McNab & Bonaccorso, 2001), includ-
ing predilection for or tolerance of disturbed habitats, such as
secondary forest, native gardens, or cyclone-damaged land-
scapes (Flannery, 1990; Flannery, 1995; Grant et al., 1997). In
this regard, basic ecological requirements of flying-foxes con-
trast strongly with those of monkey-faced bats despite their
resemblance in size and external appearance. Across their
range, monkey-faced bats (both Pteralopex and Mirimiri) oc-
cur only in old-growth forest ? probably necessary both for
suitable roosting sites and favoured food plants (Fisher &
Tasker, 1997). In the case of P. atrata, P. flanneryi, and
P. taki this is primary lowland forest (below about 1000m),
for P. pulchra and M. acrodonta this is montane forest above
1200 and 1000m respectively, and for P. anceps this is both
primary lowland and montane forest (though forests above
about 800m may be most important). Further, monkey-faced
bats are not known from any island less than 400 km2 in area
apart fromBarora Fa and Puruata, small islands situated imme-
diately along the respective coasts of Ysabel and Bougainville
Paci?c monkey-faced bats 451
Island km2 m Species recorded
Bougainville 8591 2591 2 (?, an)
Guadalcanal 5281 2448 2 (at, pu)
Malaita 4307 1280 0
Ysabel 4095 1250 1 (?)
Makira 3090 1040 0
Choiseul 2966 1060 2 (?, an)
New Georgia 2044 1006 2 (at, ta)
Kulambangra 704 1768 1 (ta: extinct)
Vella Lavella 640 793 0
Buka 611 402 1 (?)
Vangunu 544 1124 1 (ta)
Table 8 The eleven Solomon Islands (excluding the outlying island
of Rennell) greater than 400 km2 in area, with area (km2),
maximum elevation (m), and number of recorded
Pteralopex species. an=P. anceps; at=P. atrata; ?=P.
?anneryi; pu=P. pulchra; ta=P. taki. Areas and
elevations are from Mayr and Diamond (2001: 60), except
for highest elevation for Choiseul (1060 m), taken from
Bowen-Jones et al. (1997).
(Table 8). This suggests that monkey-faced bats require re-
latively large areas of old-growth forest to sustain resident
populations (although the montane species may be restricted
to quite small upland areas on their respective islands). Indeed,
P. taki has apparently been extirpated on Kulumbangra follow-
ing extensive deforestation (Flannery, 1995; Fisher & Tasker,
1997), which has claimed much of the lowland forest of that
island over the last 40 years (Whitmore, 1989).
Of the eleven ?core? islands of the Solomons greater than
400 km2 in area (i.e. excluding the outlying island of Rennell),
eight are known to support (or to have until recently suppor-
ted) at least one Pteralopex species (Table 8). The three excep-
tions are the large islands of Malaita, Makira (San Cristobal),
and Vella Lavella. Malaita and Makira are large, mountainous
islands that have been very incompletely surveyed for mam-
mals, and I suspect future surveys will show these islands to
support species of Pteralopex (cf. Parnaby, 2002b). Surveys
are also needed to determine whether the distribution of P. taki
extends to Vella Lavella, a poorly-surveyed island seemingly
large enough to support a species of Pteralopex. In addition, as
noted above, I suspect future fieldwork will demonstrate that
P. anceps (as well as P. flanneryi) occurs on the large island of
Ysabel.
At 435 km2, Taveuni in Fiji is the smallest island that
supports a monkey-faced bat (M. acrodonta). As noted by
Flannery (1995) and Parnaby (2002b), surveys are necessary
to establish whether this species extends to cloud forest on
the adjacent island of Vanua Levu, which is much larger
(5538 km2) but has lower mountains (maximum elevation
ca. 1030m, vs. 1241m on Mt. Uluingalau on Taveuni [Ash,
1987]).
Finally, because monkey-faced bat genera occur in the
Solomon Islands (Pteralopex) and Fiji (Mirimiri), related spe-
cies might be expected to occur in the geographically inter-
vening archipelago of Vanuatu (the New Hebrides). Lowland
forests throughout Vanuatu have now been relatively well-
surveyed for mammals (Medway & Marshall, 1975; Flannery,
1995; Helgen, unpubl. data), and Pteralopex does not occur
there. However, mammal faunas from the higher peaks in the
western part of the largest island of Vanuatu ? Espiritu Santo ?
remain poorly known (Medway & Marshall, 1975). Several
of these mountains rise higher than 1000m (the tallest be-
ing Mt. Tabwemasana at 1879m) and mossy upper montane
forests on these peaks (at and above about 1000m) support
an avifauna quite distinct from that of the lowlands, which
includes the endemic montane starling Aplonis santovestris
and the warbler Cichlornis whitneyi whitneyi (Bregulla, 1992).
If monkey-faced bats are found to occur in Vanuatu, it will
probably be in these mountains.
Acknowledgements
For access to specimens and other assistance during museum visits,
I thank Sandy Ingleby and Tish Ennis (AM), Paula Jenkins, Richard
Harbord, and Daphne Hills (BMNH), Carla Kishanami, Kathleen Ig-
eta, Allen Allison, and Neal Evanhuis (BBM), Dieter Kock, Gerhard
Storch, andKatrin Krohmann (SMF), DonWilson, LindaGordon, and
Craig Ludwig (USNM), Jim Patton and Chris Conroy (MVZ), and
Bulisa Iova (PNGM). My research has been supported by grants and
fellowships from the U.S. National Science Foundation (Graduate Re-
search Fellowship), the Australian-American Fulbright Commission,
the American Society of Mammalogists (Fellowship in Mammalogy),
the University of Adelaide, and travel grants from the Smithsonian
Institution and the South Australian Museum.
I thank Harry Parnaby, Norberto Giannini, Paula Jenkins, Tim
Flannery, and Tanya Leary for insightful reviews that improved this
manuscript. I am additionally grateful to Tanya Leary and John Pita
for sharing unpublished data, Don Wilson and Russell Baudinette
for various assistances, Lauren Johnston for producing the maps, and
Pavel German, Elizabeth Tasker, Steve Hamilton, Sandy Ingleby, Don
Colgan and Steve Richards for helpful discussion.
References
ANDERSEN, K. 1909a. On the characters and affinities of ?Des-
malopex? and Pteralopex. Annals and Magazine of Natural His-
tory (series 8) 3, 213?222.
ANDERSEN,K. 1909b. Twonewbats from theSolomon Islands.Annals
and Magazine of Natural History (series 8) 3, 266?270.
ANDERSEN, K. 1912. Catalogue of the Chiroptera in the collection of
the British Museum. Volume I. Megachiroptera. Second edition.
London: British Museum (Natural History).
ASH, J. 1987. Stunted cloud forest in Taveuni, Fiji. Pacific Science 41,
191?199.
BONACCORSO, F.J. 1998. Bats of Papua New Guinea. Washington,
D.C.: Conservation International.
BOWEN-JONES, E., ABRUTAT, D., MARKHAM, B. & BOWE, S. 1997.
Flying-foxes on Choiseul (Solomon Islands) ? the need for con-
servation action. Oryx 31, 209?217.
BREGULLA, H.L. 1992. Birds of Vanuatu. Shropshire, UK: Anthony
Nelson.
CAPELL, A. 1968. A New Fijian Dictionary. Suva, Fiji: Government
of Fiji.
COLGAN, D.&DACOSTA, P. 2002.Megachiropteran evolution studied
with 12S rDNAand c-mosDNAsequences. Journal ofMammalian
Evolution 9, 3?22.
COLGAN, D. & FLANNERY, T.F. 1995. A phylogeny of Indo-West Pa-
cificMegachiroptera based on ribosomalDNA. Systematic Biology
44, 209?220.
452 K. M. Helgen
DUMONT, E.R. 1997. Cranial shape in fruit, nectar, and exudate
feeders: implications for interpreting the fossil record. American
Journal of Physical Anthropology 102, 187?202.
DUMONT, E.R. & HERRELL, A. 2003. The effect of gape angle and
bite point on bite force in bats. Journal of Experimental Biology
206, 2117?2123.
FISHER, D. & TASKER, E. 1997. Natural history of the New Georgia
monkey-faced bat Pteralopex sp. nov. from the Solomon Islands.
Pacific Conservation Biology 3, 134?142.
FLANNERY, T.F. 1990. Flying foxes in Melanesia: populations at risk.
Bats 7, 5?7.
FLANNERY, T.F. 1991. A new species ofPteralopex (Chiroptera: Ptero-
podidae) from montane Guadalcanal, Solomon Islands. Records of
the Australian Museum 43, 123?130.
FLANNERY, T.F. 1995. Mammals of the South-West Pacific and Moluc-
can Islands. Ithaca, NY: Cornell University Press.
FLANNERY, T.F. & SCHOUTEN, P. 2004. Astonishing Animals: Ex-
traordinary Creatures and the Fantastic Worlds they Inhabit.
Boston: Atlantic Monthly Press.
GIBBONS, J.R.H. 1981. The biogeography of Brachylophus including
the description of a new species, B. vitiensis, from Fiji. Journal of
Herpetology 15, 255?273.
GRANT, G.S., CRAIG, P. & TRAIL, P. 1997. Cyclone-induced shift in
foraging behavior in flying foxes in American Samoa. Biotropica
29, 224?228.
GRAY, J.E. 1870. Catalogue of Monkeys, Lemurs, and Fruit-eating
Bats in the Collection of the British Museum. London: Trustees of
the British Museum (Natural History).
HELGEN, K.M. & FLANNERY, T.F. 2002. Distribution of the en-
dangered Pacific sheathtail bat Emballonura semicaudata. Aus-
tralian Mammalogy 24, 209?212.
HILL, J.E. & BECKON, W.N. 1978. A new species of Pteralopex
Thomas, 1888 (Chiroptera: Pteropodidae) from the Fiji Islands.
Bulletin of the BritishMuseum (Natural History) Zoological Series
34, 65?82.
HILTON-TAYLOR, C. 2000. 2000 IUCNRed List of Threatened Species.
Gland, Switzerland: IUCN.
INGLEBY, S. & COLGAN, D. 2003. Electrophoretic studies of the sys-
tematic and biogeographic relationships of the Fijian bat gen-
era Pteropus, Pteralopex, Chaerephon, and Notopteris. Australian
Mammalogy 25, 13?29.
KOOPMAN, K.F. 1994. Chiroptera: systematics. Handbuch der Zoolo-
gie 8(60), 1?217.
LAURIE, E.M.O. & HILL, J.E. 1954. List of land mammals of New
Guinea, Celebes, and adjacent islands, 1758?1952. London:
British Museum (Natural History).
MAA, T.C. 1966. Partial revision of the Cyclopodiinae (Diptera:
Nycteribiidae). Pacific Insects 8, 648?685.
MAA, T.C. 1968. Additions to the Cyclopodiinae. Part I. (Diptera:
Nycteribiidae). Pacific Insects 10, 1?23.
MARSHALL, A.G. 1985. Old World phytophagous bats (Megachirop-
tera) and their food plants: a survey. Zoological Journal of the
Linnean Society 83, 351?369.
MATSCHIE, P. 1899. Die Megachiroptera des Berliner Museums fu?r
Naturkunde. Berlin: George Reimer.
MAYR, E. & DIAMOND, J. 2001. The Birds of Northern Melanesia:
Speciation, Ecology, and Biogeography. Oxford: Oxford Univer-
sity Press.
MCNAB, B.K. 1994. Resource use and the survival of land and fresh-
water vertebrates on oceanic islands. American Naturalist 144,
643?660.
MCNAB, B.K. & BONACCORSO, F.J. 2001. The metabolism of New
Guinean pteropodid bats. Journal of Comparative Physiology B
171, 201?214.
MEDWAY, L. & MARSHALL, A.G. 1975. Terrestrial vertebrates of
the New Hebrides: origin and distribution. Philosophical Trans-
actions of the Royal Society of London Series B 272, 423?
465.
MOLNAR, R.E.,WORTHY, T. &WILLIS, P.M.A. 2002. An extinct Pleis-
tocene endemic mekosuchine crocodylian from Fiji. Journal of
Vertebrate Paleontology 22, 612?628.
MUELLER-DOMBOIS, D. & FOSBERG, F.R. 1997. Vegetation of the
Tropical Pacific Islands. New York: Springer-Verlag.
PARNABY, H.E. 2002a. A new species of long-eared bat (Nyctophilus:
Verspertilionidae) from New Caledonia. Australian Mammalogy
23, 115?124.
PARNABY, H.E. 2002b. A taxonomic review of the genus Pter-
alopex (Chiroptera: Pteropodidae), the monkey-faced bats of
the south-western Pacific. Australian Mammalogy 23, 145?
162.
PHILLIPS, C.J. 1968. Systematics of megachiropteran bats in the So-
lomon Islands.University of Kansas Publications, Museum of Nat-
ural History 16, 779?837.
PREGILL, G.K. & WORTHY, T.H. 2003. A new iguanid lizard
(Squamata, Iguanidae) from the Late Quaternary of Fiji, south-
west Pacific. Herpetologica 59, 57?67.
SANBORN, C.C. 1931. Bats from Polynesia, Melanesia, and Malaysia.
Publications of the Field Museum of Natural History, Zoological
Series 18, 7?29.
SIMPSON, G.G. 1945. The principles of classification and a classific-
ation of mammals. Bulletin of the American Museum of Natural
History 85, 1?350.
STATTERSFIELD, A.J., CROSBY, M.J., LONG, A.J. & WEGE, D.C. 1998.
Endemic Bird Areas of the World: Priorities for Biodiversity Con-
servation. Cambridge, UK: BirdLife International.
THEODOR, O. 1959. A revision of the genus Cyclopodia
(Nycteribiidae, Diptera). Parasitology 49, 242?308.
THOMAS, O. 1888. Diagnoses of six new mammals from the Solomon
Islands. Annals and Magazine of Natural History (series 6) 2,
155?158.
TROUGHTON, E. LE G. 1936. The mammalian fauna of Bougainville
Island, Solomons group. Records of the Australian Museum 19,
341?353.
VINYARD, C.J., WALL, C.E., WILLIAMS, S.H. & HYLANDER W.L.
2003.Comparative functional analysis of skullmorphology of tree-
gouging primates. American Journal of Physical Anthropology
120, 153?170.
WHITMORE, T.C. 1989. Changes over twenty-one years in the
Kolombangara rain forests. Journal of Ecology 77, 469?
483.
WOODFORD, C.M. 1890. A naturalist among the head-hunters, being
an account of three visits to the Solomon Islands in the years
1886, 1887, and 1888. Melbourne, Australia: E.A. Petherick and
Co.
WORTHY, T.H. 2000. The fossil megapodes (Aves: Megapodiidae)
of Fiji with descriptions of a new genus and two new spe-
cies. Journal of the Royal Society of New Zealand 30, 337?
364.
WORTHY, T.H. 2001. A giant flightless pigeon gen. et sp. nov. and
a new species of Ducula (Aves: Columbidae), from Quaternary
deposits in Fiji. Journal of the Royal Society of New Zealand 31,
763?794.
Paci?c monkey-faced bats 453
Appendix
Key to species of monkey-faced bats
(Pteralopex and Mirimiri) (modi?ed
from Parnaby, 2002b: 161)
1a. Body size relatively small (FA < 130; CBL < 60); post-
orbital processes not annectent to the zygomata in adults;
M2 not strongly reduced relative to M1 (greater than half
the mass); labial margins of PM4 and M1 divided into
three distinct cusps; length of D4P2 subequal to D4P1;
five divided median ridges in the soft palate; two serrated
supplementary posterior ridges in the soft palate . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Mirimiri acrodonta
1b. Body size small to large (FA 112?170; CBL 52?76); post-
orbital processes annectent to the zygomata in adults; M2
strongly reduced relative to M1 (less than half the mass);
labial margins of PM4 and M1 united in an elevated cusp;
length of D4P2 conspicuously longer than D4P1; six di-
vided median ridges in the soft palate; three serrated sup-
plementary posterior ridges in the soft palate . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . genus Pteralopex (2)
2a. Body size smaller (FA< 125; CBL < 56) . . . . . . . . . . . . . 3
2b. Body size larger (FA> 128; CBL > 61) . . . . . . . . . . . . . . 4
3a. Dorsal fur dense and long, black; ventral fur dark with
yellowish tips; teeth smaller (MTR< 20); second lower
incisors separated from each other by at least half the
breadth of each tooth . . . . . . . . . . . . . . .Pteralopex pulchra
3b. Dorsal fur generally sparse, short, overall fur colour light
brown,without yellowish venter; teeth larger (MTR > 20);
second lower incisors nearly in contact or separated by less
than half the breadth of each tooth . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Pteralopex taki
4a. Body size large (FA 141?160); pelage very long and thick,
fur of mid-back not adpressed; tibia, dorsal surface of
the pes, and proximal third of the forearm very densely
furred; overall colour blackish brown, but venter with a
expansive yellow or white fur patch on the breast; incisors
and canines relatively very large but cheekteeth relatively
small except for PM4; PM4,M1, andM2 without unbroken,
elevated lingual ridges along their lengths . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Pteralopex anceps
4b. Body size medium to very large (FA 129?169); pelage
generally short; fur of mid-back adpressed; tibia, dorsal
surface of the pes, and proximal third of the forearm very
sparsely furred; overall colour blackish brown above and
below; cheekteeth relatively heavy; PM4,M1, andM2 with
unbroken, elevated lingual ridges along their lengths . . . 5
5a. Size smaller (FA< 150); rostrum less elongate (ONL/
CBL= 0.491?0.553) . . . . . . . . . . . . . . . .Pteralopex atrata
5b. Size larger (FA> 158); rostrum more elongate (ONL/
CBL= 0.575?0.632) . . . . . . . . . . . . . Pteralopex flanneryi