J. Zool., Lond. (2004) 264, 117?124 C? 2004 The Zoological Society of London Printed in the United Kingdom DOI:10.1017/S0952836904005667
A new species of bandicoot, Microperoryctes aplini,
from western New Guinea
Kristofer M. Helgen* and Timothy F. Flannery
School of Earth and Environmental Sciences, University of Adelaide, Adelaide SA 5005, Australia, and South Australian Museum, North Terrace,
Adelaide SA 5000, Australia
(Accepted 16 February 2004)
Abstract
A distinctive new striped bandicoot of the genus Microperoryctes (M. aplini new species) is described and compared
with Microperoryctes murina, another little-known bandicoot with which the new species has been confused in
the past. The smallest of all living bandicoots, M. aplini is apparently restricted to high-elevation forests in the
Arfak Mountains of the Vogelkop Peninsula of western New Guinea. The similarly small M. murina is known only
from high-elevation forests on Gunung Sumuri, a peak in the Weyland Range of western New Guinea. The two
species differ markedly in fur colour and ornamentation, pelage texture, external proportions, and quantitative and
qualitative craniodental aspects, and may not be immediately related to one another.
Key words: Microperoryctes, bandicoot, taxonomy, New Guinea
INTRODUCTION
The mouse bandicoots (genus Microperoryctes) are
a group of small-bodied, ground-dwelling marsupials
endemic to forested highlands in the Central Ranges of
New Guinea. Most species are light brown to reddish-
brown, attractively ornamented with a black dorsal stripe,
and, in some species, with facial and rump stripes. One
species of the genus, M. murina, differs from all others
in its inornate, smoky-grey pelage, which is soft and
woolly like that of many shrews and moles. This unusual
bandicoot is the smallest of all described bandicoots,
with a total length of <300 mm. It is a rare species,
known only by the type series of three specimens from the
Weyland range of western New Guinea. Four additional
specimens of another very small, striped bandicoot (in
museums on four different continents) were studied, all of
which originated from the Arfak Mountains of western
New Guinea. Although these latter specimens have
been tentatively identified in museum collections and in
literature asM.murina (almost entirely on the basis of their
very small size), it has been recognized for some time that
they differ substantially from the type series of that species
(see collective discussion by Ziegler, 1977; George &
Maynes, 1990; Groves & Flannery, 1990; Menzies, 1991;
Flannery, 1995; Aplin, 1998; Aplin, Pasveer & Boles,
1999; Aplin explicitly referred to the Arfak pygmy bandi-
coot as ?a potentially distinct species?). In this report the
*All correspondence to: K. M. Helgen.
E-mail: kristofer.helgen@adelaide.edu.au
taxonomic content of the genus Microperoryctes is briefly
reviewed before describing the Arfak pygmy bandicoot as
a new species.
MATERIALS AND METHODS
Specimens discussed here are deposited in the collections
of the Bernice P. Bishop Museum in Honolulu (BBM),
the Museum fu?r Naturkunde in Berlin (ZMB), the
Museum Zoologicum Bogoriensis in Cibinong, Indonesia
(MZB), the Papua New Guinea National Museum and
Art Gallery in Port Moresby (PNGM), the American
Museum of Natural History in New York (AMNH),
and the Australian Museum in Sydney (AM). External
measurements were recorded by the original collectors
in the field. Craniodental variables were measured by
the first author with hand-held callipers to the nearest
0.01 mm (rounded here to 0.1 mm). External variables
are abbreviated as follows: head and body length (HB),
tail length (TV), hindfoot length (HF), ear length (E).
SYSTEMATICS
Order Peramelemorphia Kirsch, 1968
Family Peramelidae Gray, 1825
Subfamily Echymiperinae McKenna & Bell, 1997
Comments: Familial and subfamilial classification follows
Groves (in press).
118 K. M. HELGEN AND T. F. FLANNERY
Fig. 1. Map of New Guinea, with important localities mentioned in text. ?, Hatam, Arfak Mountains, type locality of Microperoryctes
longicauda; 
, Lake Anggi Giji, Arfak Mountains, type locality of Microperoryctes aplini; , Gunung Sumuri, Weyland Range, type
locality of Microperoryctes murina; , Doormanpad-bivak, Mamberamo Basin, type locality of M. l. dorsalis; , Star Mountains region,
site of apparent overlap between M. longicauda and M. ornata; H17009, mouth of Aroa River, near type locality of Microperoryctes ornata;
, Mount Mura, Owen Stanley Range, type locality of Microperoryctes papuensis;, Mount Simpson, Owen Stanley Range, type locality
of M. ornata magna.
Genus Microperoryctes Stein, 1932
(type species Microperoryctes murina Stein, 1932)
Synonyms
Ornoryctes Tate & Archbold, 1937 (type species Per-
oryctes longicauda Peters & Doria, 1876 [=Micro-
peroryctes longicauda]); considered here a valid subgenus
(see below).
Diagnosis
The genus Microperoryctes was originally erected by
Stein (1932) to reflect the diminutive size and unusual
pelage characteristics of a then newly discovered
bandicoot Microperoryctes murina from the Weyland
range of the western Central Cordillera of New Guinea.
Microperoryctes murina is unique among bandicoots in
its very small size and its dark and woolly pelage, and
most subsequent workers followed Stein and maintained
Microperoryctes as a monotypic genus (Tate & Archbold,
1937; Tate, 1948; Laurie & Hill, 1954; George & Maynes,
1990). More recently, based on cladistic considerations,
Groves & Flannery (1990) united the striped bandicoots of
the subgenus Ornoryctes under the genus Microperoryctes
rather than Peroryctes (where they have traditionally been
placed) and provided revised diagnoses of the resultant
redefined genera. According to Groves & Flannery
(1990), among the bandicoots, species of Microperoryctes
(including Ornoryctes) can be diagnosed by a unique
combination of derived characters: a broadened braincase,
a deeply wavy coronal suture, two pairs of palatal
vacuities, with the posterior palatal vacuities extending
from P3 to M2 or M3 (dental terminology following
Luckett, 1993), and straight (rather than bracket-shaped)
posterior palatal margins. This revised diagnosis is
accepted here. The species of Microperoryctes also share a
relatively small body size, a relatively long tail (generally
>60% of head and body length), small ears and granular
foot pads.
Distribution
All species of the genus Microperoryctes are endemic to
higher altitude forests (above 1000 m) in the mountain
ranges along the Central Cordillera of New Guinea, from
Mt Simpson in the far south-eastern Owen Stanley Range
(Milne Bay Province, Papua New Guinea) to the mountain
ranges of the Vogelkop Peninsula in the north-east (Fig. 1;
Flannery, 1995). Aplin, Pasveer & Boles (1999) suggested
that M. longicauda may occur at elevations lower than
1000 m in the Vogelkop Peninsula based on its presence
in upper levels excavated from Kria Cave in the Ayamaru
Lakes region, a lowland site. The genus is conspicuously
absent from the north coast ranges of northern New
Guinea (e.g. the Cyclops, Torricelli and Bewani ranges)
and is also unrecorded from the Huon Peninsula, although
hunters interviewed by the first author in the Finisterre
Range on the Huon universally claim familiarity with an
animal similar to Microperoryctes ornata.
Each species of Microperoryctes is (or is probably)
sympatric with one or more congeners in all or some part
of its range (see below).
New species of bandicoot 119
Table 1. Selected external attributes of the species of Microperoryctes. Measurements (mm) for M. longicauda
and M. ornata are taken from series at BBM and AMNH. Measurements for M. papuensis are taken from
Flannery (1995)
murina aplini longicauda ornata papuensis
Dorsal stripe Absent Present Variable Present Present
Lateral rump stripes Absent Absent Absent/faint Present Present
White tail-tip Absent Present Present Present Present
Head?body length 152?174 142?160 250?290 239?303 175?205
Tail length 105?111 115?120 160?205 161?258 142?158
n for measurements 2 3 30 73 7
Content
There are five valid species of Microperoryctes: M.murina
Stein, 1932; M. longicauda (Peters & Doria, 1876);
M. ornata (Thomas, 1904); M. papuensis (Laurie, 1952);
M. aplini new species (Fig. 1, Table 1).
Until now, only three species have generally been
recognized within the genus Microperoryctes (Groves,
1993; Flannery, 1995). This traditional scheme is modified
here by describing M. aplini as a new species distinct
from M. murina, and by recognizing both M. longicauda
and M. ornata, previously considered conspecific, as two
separate biological species. Microperoryctes longicauda
has traditionally been divided into three or four sub-
species that are thought to form a stepped cline in
coloration from west to east (see Tate, 1948; Laurie, 1952;
George & Maynes, 1990; Flannery, 1995; Westerman,
Springer & Krajewski, 2001; Groves, 2005). These are
M. l. longicauda of the Vogelkop Peninsula, which lacks
a dorsal stripe and rump stripes; M. l. dorsalis of the
western Central Cordillera (from the Weyland Range to the
Strickland River), which generally possesses a moderately
pronounced dorsal stripe but lacks rump stripes; and
M. l. ornata and M. l. magna of east-central and south-
eastern New Guinea, respectively, which are boldly
patterned with a dorsal stripe and lateral rump stripes.
Overall external differences are quite striking between
the boldly patterned eastern ?races? (ornata and magna)
on the one hand and the weakly patterned western ?races?
(longicauda and dorsalis) on the other, and here we note
that the two overlap without apparent introgression in the
Star Mountains of central New Guinea (Fig. 1), as shown
by extensive series of Microperoryctes at BBM collected
by A. B. Mirza (a point preliminarily noted by Flannery,
1995: 113?114). The BBM series demonstrates that in the
Star Mountains region, ornata occurs in mid-elevation
rainforest and moss forest from c. 1400 to 2600 m (see
also Flannery & Seri, 1990: 179), while dorsalis occurs in
higher-elevation moss forests and in subalpine grassland at
3100 m and above (elsewhere in New Guinea where these
two forms do not co-exist, each has an elevational range
from roughly 1000 to 4000 m; see Tate, 1951: 328?329).
The two do not generally differ in overall size and standard
external proportions across most of their largely allopatric
ranges (see Table 1). However, in the Star Mountains
(their area of overlap), series of dorsalis have relatively
Table 2. External measurements (mm) for series of Microper-
oryctes ornata and M. longicauda dorsalis from the Star Mountains
(specimens at AM and BBM). HB, head?body length; TV, tail
length; E, ear length; HF, hindfoot length
ornata dorsalis
HB 253.8 ? 15.29 266.8 ? 11.13
232?276 (n = 8) 250?290 (n = 11)
TV 183.4 ? 7.80 167.7 ? 8.62
174?198 (n = 8) 156?183 (n = 11)
E 25.1 ? 1.81 30.4 ? 0.67
22?27 (n = 8) 29?31 (n = 11)
HF 58.2 ? 3.27 60.1 ? 2.39
53?64 (n = 8) 57?65 (n = 11)
much shorter tails and longer ears compared to series of
ornata from lower elevations (Table 2), as well as more
elongate rostra, thicker pelage, darker dorsal coloration, a
less well-defined dorsal stripe, no lateral rump stripes, and
grey-based rather than pure white belly fur. (Interestingly,
the Seltaman people of Western Province, Papua New
Guinea, may have different names for these taxa; see
Whitehead (2000).) Thus ornata and dorsalis apparently
displace one another altitudinally but do not introgress
where their ranges overlap along the New Guinean
Central Cordillera. This sympatric (if not syntopic)
occurrence suggests that two biological species should
be recognized within the previous taxonomic boundaries
of Microperoryctes longicauda. These are M. longicauda
(Peters & Doria, 1876) (withM. l. dorsalis [Thomas, 1922]
as a subspecies), occurring on the Vogelkop Peninsula
(M. l. longicauda) and along the Central Cordillera from
the Weyland range to the Star Mountains (M. l. dorsalis),
and M. ornata (Thomas, 1904) (with magna [Laurie,
1952] as a subspecies), occurring along most of the eastern
Central Cordillera (M. l. ornata) and in the Agaun area and
on Mt Simpson in far south-eastern Papua New Guinea
(M. l. magna). However, phylogeographic patterns within
M. ornata are probably more complicated than current
taxonomy suggests (Laurie, 1952; Westerman et al., 2001:
99; K. M. Helgen, pers. obs.).
Only the two pygmy species of the genus are reviewed
in further detail below, in the context of describing a new
pygmy species of Microperoryctes.
120 K. M. HELGEN AND T. F. FLANNERY
Microperoryctes murina Stein, 1932
Type material and type locality
The holotype of murina is ZMB 44229, adult male,
skin and skull, from Gunung Sumuri, 2500 m, Weyland
Mountains, west-central New Guinea (Papua Province,
Indonesia), collected 31 August 1931 by G. Stein. There
are only two other known specimens, both collected by
Stein at the type locality: AMNH 103262, young adult,
probably female (the sex of this specimen is given as
female by Stein (1932) and as possibly male by Tate &
Archbold (1937)), collected 27 August 1931; and ZMB
91462, immature female, skin and skull, collected
3 September 1931.
Diagnosis
Microperoryctesmurina can be immediately distinguished
from all other species of bandicoots by its combination of
extremely small body size (head and body length 152?
174 mm) and unpatterned, smoky-grey, very soft pelage
(reminiscent of many shrews). The scrotum is furry and
greyish-white.
Distribution
Microperoryctes murina has been recorded to date only
from the type locality (Fig. 1).
Discussion
Almost nothing is known about the basic biology of
M. murina, although Tate & Archbold (1937) suggested
that it is probably ?highly fossorial?. Its woolly, soft,
shrew-like pelage is unique among bandicoots and
suggests a lifestyle convergent on insectivorous or
vermivorous shrews and shrew-mice, small mammals
that occupy highland forests elsewhere in south-east
Asia and Melanesia (cf. Musser, 1982; Flannery, 1995).
(Little is known of the feeding habits of other species of
Microperoryctes, but most bandicoots are thought to be
omnivorous; see Flannery, 1995; Hume, 1999: 82.) Given
its extremely limited known distribution (montane forest
on Gunung Sumuri), M. murina may be vulnerable to
extinction.
The Weyland Range has been well-surveyed for
mammals thanks to efforts by collectors during the middle
decades of the 20th century. Especially notable collections
from these mountains were amassed in the 1930s by
the collectors G. Stein (series at ZMB and AMNH) and
F. Shaw-Mayer (series at the Natural History Museum,
London). In addition to Microperoryctes murina, several
additional mammal species are apparently endemic to
the Weyland range. These include the distinctive murine
Macruromys elegans Stein, 1933 and a mosaic-tailed
rat, Paramelomys steini (Ru?mmler, 1935). Macruromys
elegans (the lesser small-toothed rat) is known only by
the type series of four individuals collected between
1400 and 1800 m on Gunung Kunupi in October 1931
(at ZMB and AMNH, see Tate, 1951; Flannery, 1995),
and Paramelomys steini (Stein?s paramelomys) is known
only by the type series collected from 2000 to 2600 m on
Gunung Sumuri (see Menzies, 1996; Musser & Carleton,
in press). Interestingly, all three endemic mammal species
of the Weylands are restricted to Mts Kunupi and Sumuri,
and do not extend to the Gebroeders region, another well-
surveyed area of the range.
Microperoryctes aplini new species
Holotype
BBM 22457, young adult male, study skin (figured
by Flannery, 1995: 116) and skull (Fig. 2), from
Kampong Sururai, Lake Anggi Giji (01?23?S, 133?58?E),
6200 ft (= 1890 m), Arfak Mountains, eastern Vogelkop
Peninsula, western New Guinea (Papua Province,
Indonesia), collected 2 March 1963 by M. C. Thompson.
Paratypes
ZMB 40293, unsexed subadult skull, from Koffo (01?25?S,
133?52?E), Lake Anggi Giji, 2000 m, Arfak Mountains,
collected 13 June 1928 by E. Mayr. PNGM 25375, adult
male, study skin, skull, and body in alcohol, from ?Arfak
Mountains?, 2200 m, collected 21 September 1986 by
I. Craven.
Referred specimen
MZB 2951, adult male, skin only, collected in 1928 by
E. Mayr in ?Netherlands New Guinea? (i.e. Papua
Province, Indonesia). The locality written on the tag
in script seems to be ?Liwi? or ?Siwi?; probably Siwi,
01?29?S, 134?02?E.
Diagnosis
Microperoryctes aplini can be immediately distinguished
from all other species of bandicoots by its combination
of extremely small body size (head and body length
<160 mm; total length 300 mm) and sleek brown
dorsal fur with a well-pronounced dorsal stripe. It is prob-
ably the smallest known species of bandicoot, similar in
size only to M. murina of the Weyland range (see above).
Distribution
Microperoryctes aplini has been recorded to date only
from the Arfak Mountains of the Vogelkop Peninsula of
western New Guinea in a narrow elevational band between
1890 and 2200 m (Fig. 1). Remains of M. aplini have not
New species of bandicoot 121
Fig. 2. Skull of BBM 22457, holotype of Microperoryctes aplini.
been recovered in extensive Quaternary mammal deposits
from Kria and Toe Caves in the Ayamaru Lakes area of
the central Vogelkop (Aplin et al., 1999). Likewise, the
species is not represented in large lots of trophy material
collected in a hunting camp situated in moss forest at
2500 m in the Mokwam area in the Arfaks by T. Flannery
in 1992 (specimens at AM) or in the notable collection of
mammals from the Arfaks made by F. Shaw-Mayer in the
1920s, deposited in the Natural History Museum, London.
Both museum specimens of M. aplini bearing a definite
locality originated from the vicinity of Lake Anggi Giji;
thus the distribution of this species may be restricted to
the vicinity of this highland lake. A label on the holotype
notes that it was collected in ?forest?, and the collector
of the holotype also secured specimens of Phascolosorex
dorsalis and Melomys cf. mollis in the immediate vicinity
of the type locality.
Etymology
Named for the zoologist Ken Aplin, to recognize his stud-
ies of Australasian vertebrates in general and Vogelkop
biogeography in particular. We recommend ?Arfak pygmy
bandicoot? as an appropriate common name.
Description
Microperoryctes aplini is a very small bandicoot with
sleek, light-brown dorsal fur with grey bases, a relatively
wide and moderately pronounced dark stripe extending
from crown to rump along the mid-dorsum, and a relatively
long tail with a short but distinct white tail-tip (about
one-sixth the length of the tail). The tail is dorsoventrally
bicoloured (brown above and white below), the venter is
grey-brown with a mid-ventral creamy-white stripe, and
the claws are long and slender. Superficially M. aplini
resembles a dwarf version of the other small species
of Microperoryctes with a prominent dorsal stripe
(M. papuensis of south-eastern New Guinea) except that
it lacks the distinctive lateral rump stripes of that species.
In M. aplini, the length of the head and body averages
shorter than in the similarly tiny bandicoot M. murina
(Table 3). However, the tail of M. aplini is longer than
that of M. murina, both absolutely (Table 3) and relative
to head and body length (74?83% of HB in M. aplini vs
64?69% in M.murina). No weights are available for either
M. aplini or M. murina, but both probably weigh <100 g
as adults (see Flannery, 1995: 105).
Table 3. External dimensions (mm) of the pygmy bandicoots
M. aplini and M. murina. Values for MZB 2951 are measured
from the study skin by the first author and should be considered
approximate. HB, head?body length; TV, tail length; E, ear length;
HF, hindfoot length
HB TV HF E
Microperoryctes aplini
UPNG 25375 Young adult ? 142 115 33 20
BBM 22457 Young adult ? 145 120 38 22
MZB 2951 Adult ? 160 118 33 ?
Microperoryctes murina
ZMB 44229 Adult ? 174 111 34 23
AMNH 103262 Young adult ? 152 105 31 21
122 K. M. HELGEN AND T. F. FLANNERY
Table 4. Selected skull measurements (mm) for Microperoryces
aplini and M. murina
aplini murina
UPNG BBM ZMB AMNH
25375 22457 44229 103262
Greatest length of skull 43.2 41.7 45.6 42.4
Condylobasal length 41.1 39.5 44.0 40.2
Zygomatic width 16.8 15.3 16.4 15.6
Nasal length 17.5 17.1 19.2 17.8
Coronoid height of mandible 8.9 7.9 8.8 8.0
Greatest length of mandible 32.3 30.3 34.3 31.1
Although sample sizes are small for both species, cur-
rently available series suggest that while M. aplini and
M. murina do not differ from one another in overall skull
length, M. aplini has shorter nasals (Table 4) and slightly
larger molars than M. murina (Fig. 3).
In addition to differences in coloration, pelage texture,
and external and craniodental proportions, M. aplini
differs from M. murina in some notable qualitative cranial
features. For example, the position and size of the incisive
and anterior palatal foramina differ characteristically
between the two species (Fig. 3). In M. aplini, the incisive
foramina extend posteriorly to or just beyond the posterior
alveolus of I5; in M. murina they extend well behind this
tooth. In M. aplini the anterior palatal foramina are more
expansive than in M. murina, extending anteriorly only as
far as P1 in M. murina but well in front of this tooth in
M. aplini. In fact, in M. aplini the anterior palatal
foramina are relatively more expansive and elongate than
in any other member of the genus Microperoryctes. In
dorsal view the occiput of the cranium of both M. aplini
and M. murina appears rounded and convex rather than
Fig. 3. Conformation of palatal foramina in Microperoryctes aplini (left, ZMB 40293, unsexed sub-adult) and M. murina (right, ZMB
91462, sub-adult female). Specimens of similar dental development.
concave with the lateral margins protruding, as in most
bandicoots.
In other qualitative aspects M. aplini agrees closely
with the other four species of Microperoryctes (as revised
and reviewed by Groves & Flannery, 1990), which, despite
their occurrence in overlapping sympatry across the island
of New Guinea, are very similar to each other in most
qualitative craniodental characters (Tate, 1948; Laurie,
1952; Groves & Flannery, 1990).
Discussion
Almost nothing is yet known about the basic biology of
M. aplini, although Ziegler (1977) suggested that it
is subfossorial. Given its extremely limited known
distribution (the vicinity of Lake Anggi Giji), it should
certainly be considered vulnerable to extinction.
The Vogelkop Peninsula (or ?Bird?s Head? of New
Guinea) is a notable centre of endemism for vertebrates
(see Aplin, 1998). In addition to Microperoryctes aplini,
several other mammal species are endemic to the Arfaks,
including the pseudocheirids Pseudochirops coronatus
and Pseudochirulus schlegeli (Flannery, 1994, 1995) the
murine Stenomys arfakiensis (see Musser & Carleton,
2004), and undescribed species of the murine genera Mal-
lomys and Leptomys (K. M. Helgen, pers. obs.). A number
of other mammals are endemic to the greater Vogelkop
region (some including the Arfaks); these include the
petaurid Dactylopsila kambuayai and the pseudocheirid
Petauroides ayamaruensis (both of which remain known
only by Holocene subfossil remains from the Ayamaru
Lakes area; see Aplin et al., 1999), the tree-kangaroo
Dendrolagus ursinus, and a number of distinctive endemic
subspecies, including a dasyure Phascolosorex dorsalis
dorsalis, a wallaby Dorcopsis muelleri muelleri, and two
New species of bandicoot 123
ringtails Pseudochirops albertisii albertisii and Pseudo-
chirulus canescens bernsteini.
In addition to Microperoryctes aplini, a larger congener,
M. longicauda, also occurs on the Vogelkop Peninsula,
including the Arfak Mountains (Peters & Doria, 1876;
Flannery, 1995), where the two species probably occur
syntopically. Judging from modern and fossil collec-
tion localities (Flannery, 1995; Aplin et al., 1999),
M. longicauda is both more common and more wide-
spread on the Vogelkop than M. aplini. Species of Micro-
peroryctes also occur in sympatry in other mountain
ranges of New Guinea, such as throughout south-eastern
New Guinea (whereM. ornata andM. papuensis co-occur;
see Aplin & Woolley, 1993) and in the Star Mountains of
western Papua New Guinea (where both M. ornata and
M. longicauda apparently occur, see above). Microper-
oryctes longicauda and M. murina both occur in the
Weyland Range of western New Guinea, but have yet to
be collected at the same locality.
Microperoryctes aplini shares several derived traits with
M. longicauda, M. ornata, and M. papuensis, including a
relatively long tail (generally >70% of head and body
length), a white tail tip, and a dorsal stripe (the last
of which is variably pronounced in M. longicauda).
These features are absent in both M. murina and in the
Melanesian and Wallacean bandicoot genera Echymipera,
Rhynchomeles and Peroryctes, all or some of which are
closely related to Microperoryctes (cf. Groves & Flannery,
1990; Westerman et al., 2001), as well as in all or most
of the remaining, less closely related bandicoots (the dry-
country, mostly Australian genera Isoodon and Perameles
as well as the more distantly related genera Macrotis
and Chaeropus). Based on this suite of synapomorphous
external characters, and despite its previous classification
within M. murina, M. aplini is probably best arranged
with M. longicauda, M. ornata, and M. papuensis in the
subgenus Ornoryctes (see above), while M. murina can be
retained alone in the subgenus Microperoryctes to reflect
its more plesiomorphic external features relative to its
congeners.
Acknowledgements
For access to specimens and other assistance during
museum visits, we thank Carla Kishinami (BBM), Peter
Giere, Irene Thomas, and Nils Hoff (ZMB), Bulisa Iova
(PNGM), Sandy Ingleby and Tish Ennis (AM), Darrin
Lunde and Rob Voss (AMNH), and Ibnu Maryanto and
M. H. Sinaga (MZB). KMH also thanks N. Evenhuis,
A. Chien, and C. Dren for notable kindnesses during his
travels to Honolulu and New York, Lauren Johnston for
producing the map, Steve Richards for helpful discussion,
and Don Wilson for ongoing support. KMH is supported
by fellowships from the Australian-American Fulbright
Commission, the University of Adelaide, and the U.S.
National Science Foundation. Funding for our visits to
overseas museums was provided by grants from the South
Australian Museum and the Smithsonian Institution (to
KMH) and from the Winifred Violet Scott Foundation (to
TFF).
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