![Photographs of the ventral views of the skull of a), CG1985-33, a putative syntype of Vespertilio timoriensis Geoffroy, 1806; b), holotype of N. sherrini and c), holotype of N. major. All are males. Scale bar represents 10 mm. [Photographs b and c, courtesy of the Natural History Museum, London.]](https://www.researchgate.net/profile/Harry-Parnaby/publication/284486766/figure/fig4/AS:667929620267009@1536258239483/Photographs-of-the-ventral-views-of-the-skull-of-a-CG1985-33-a-putative-syntype-of_Q320.jpg)
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39
2009 Australian
Zoologist volume 35 (1)
Introduction
The Greater Long-eared Bat Nyctophilus timoriensis
(Geoffroy, 1806), is considered to be the most widely
distributed and largest extant member of the genus.
Until recently, the prevailing concept of N. timoriensis
was of a species extending across the southern half
of the Australian continent, Tasmania, Timor and
Papua New Guinea (Flannery 1995a; Bonaccorso
1998). The nomenclature and taxonomy of this species
has remained confused since its description in 1806,
partly because it was uncertain whether Geoffroy’s
material actually came from Timor, and also because
the whereabouts of his specimens has remained in
doubt for the past century. Timor has been regarded
by many authors since Tomes (1858) as a locality error,
because the genus had not subsequently been recorded
from Timor. However, a single specimen of Nyctophilus
obtained by Kitchener et al. (1991) from Lembata
Island, Indonesia (described as N. heran Kitchener et
al., 1991), reinstated the possibility than the genus also
occurs in Timor.
The mainland Australian populations of the Greater Long-
eared Bat have often been referred to as N. timoriensis
timoriensis, while larger animals from far south-western
Western Australia have variously been referred to as N.
timoriensis timoriensis, N. timoriensis major Gray, 1844 or N.
major. A separate subspecies N. timoriensis sherrini Thomas,
1915 was recognised from Tasmania, while some authors
considered that large Nyctophilus from Tasmania were N.
gouldi Tomes, 1858, not N. timoriensis (Hall and Richards
1979; Richards 1983). The New Guinea records of the species
arose from a small number of large Nyctophilus specimens that
were tentatively assigned to N. timoriensis timoriensis by Hill
and Pratt (1981). The prevailing nomenclature derives from
Iredale and Troughton (1934) and Tate (1941). However,
Iredale and Troughton regarded major as a synonym of
N. timoriensis, and gouldi as a south-eastern Australian
subspecies of N. timoriensis, being unaware of the presence
of large N. timoriensis in eastern Australia. Hall and Richards
(1979) recognised that N. timoriensis was present in eastern
Australia and distinct from N. gouldi.
A taxonomic review of Australian Greater
Long-eared Bats previously known as Nyctophilus
timoriensis (Chiroptera: Vespertilionidae) and
some associated taxa
H. E. Parnaby
Hon. Research Associate, Mammal Section, Australian Museum, 6 College Street, Sydney NSW 2010, Australia.
Email: parn@ozemail.com.au; and Department of Environment, Climate Change and Water NSW, PO Box
1967, Hurstville NSW 2220, Australia; and formerly BEES, University of New South Wales, Sydney NSW 2052.
ABSTRACT
A comparative morphological and morphometric assessment was undertaken of material from
mainland Australia, Tasmania and Papua New Guinea that has previously been referred to as the
Greater Long-eared Bat Nyctophilus timoriensis (Geoffroy, 1806). Five taxa are recognised: N. major Gray,
1844 from south-western Western Australia; N. major tor subsp. nov. from southern Western Australia
east to the Eyre Peninsula, South Australia; N. corbeni sp. nov. from eastern mainland Australia from
eastern South Australia, through Victoria to Queensland; N. sherrini Thomas, 1915 from Tasmania, and
N. shirleyae sp. nov. from Mt Missim, Papua New Guinea. Vespertilio timoriensis Geoffroy is regarded as
nomen dubium due to uncertainty surrounding provenance of the original specimen(s), the lack of a
definite type specimen, and lack of sufficient detail in the original description and illustration to relate
the name to a singular, currently recognised species.
This review required a consideration of two taxa not usually associated with timoriensis: bifax Thomas,
1915 from eastern Australia and New Guinea, and daedalus Thomas, 1915, previously treated as the
western subspecies of bifax, occurring from western Queensland, the northern part of the Northern
Territory, and northern Western Australia. Nyctophilus daedalus is shown to belong to a separate
species group. The implications of removing daedalus from bifax are discussed in relation to N.
arnhemensis Johnson, 1959, which is considered to be a sibling species of N. bifax.
The inter-specific relationships of these taxa are evaluated. A major species group is recognised,
consisting of major Gray, 1844 and N. corbeni sp. nov., while sherrini Thomas, 1915 is placed in a gouldi
group. The relationships of N. shirleyae from Papua New Guinea remain unclear but it is provisionally
placed in a bifax group. The relationships of N. daedalus, which is likely to be a composite species,
remain unclear and it is provisionally placed in the major group. Nyctophilus howensis from Lord Howe
Island differs from all other members of the genus and its generic status needs re-examination.
Key words: Long-eared Bat, Nyctophilus timoriensis, Nyctophilus, bat taxonomy, new species, Australia, Tasmania,
Papua New Guinea, Timor, Microchiroptera.
40 2009
Australian
Zoologist volume 35 (1)
Eleven species of Nyctophilus were recognised prior
to this study (e.g. Simmons 2005). The most widely
accepted synonomy of the 22 names proposed for the
genus is given in Table 1, along with the type locality and
broad distribution. Six species were considered to occur
on mainland Australia (Churchill 1998), four species
(two endemic) on the island of New Guinea (Flannery
1995a; Bonaccorso 1998), one endemic species on New
Caledonia (Flannery 1995b; Parnaby 2002), and two
endemic species known from single specimens: N. heran
from the Indonesian island of Lembata (Kitchener et
al. 1991) and N. howensis McKean, 1975 from Lord
Howe Island, the latter known only from a sub-fossil.
Recent publications (e.g. Reardon 1999; Churchill 2008;
McKenzie 2008; Turbill et al. 2008) recognise additional
taxa within N. timoriensis and N. bifax and draw from
earlier unpublished findings of this study.
Nyctophilus has been recognised for many decades
as a complex genus in need of extensive taxonomic
revision (Wood Jones 1925; Tate 1941, 1952; McKean
and Price 1967; Hamilton-Smith 1974; Koopman 1984;
Parnaby 1991; Reardon 1999). Important taxonomic
studies of the genus are the revisions of Tomes (1858),
Peters (1861), Thomas (1915) and the reviews of Tate
(1941, 1952). Iredale and Troughton (1934) made a
number of nomenclatural changes in their checklist
of Australian mammals, as did Ride (1970), although
each without discussing their taxonomic decisions. The
most recent published taxonomic treatments are the
reviews of the genus by Koopman (1982) for New Guinea
species, Koopman (1984) for Australian species, and the
unpublished morphological revision of Parnaby (1988).
Most species of Nyctophilus remain poorly diagnosed and
inter-specific relationships are in considerable doubt. Past
difficulties in defining species have arisen partly from an
inadequate appreciation of intra-specific variation within
the genus, which were impeded by the small sample sizes
previously available for most taxa except N. geoffroyi. In
particular, taxa from northern Australia and New Guinea
were known from comparatively few specimens, and New
Guinea species such as N. microdon and N. microtis are still
poorly represented in research collections (see Bonaccorso
1998) as are the Australian taxa N. major, N. sherrini and
N. daedalus Thomas, 1915 recognized in this study.
A further difficulty impeding resolution of species
boundaries within Nyctophilus has been the confusing
and seemingly continuous nature of morphological
variation in metric and non-metric characters that have
Parnaby
Table 1. Synonymy of the 22 available names of Nyctophilus, arranged by the 11 species recognized in recent treatments
(in bold), and giving their broad geographic distributions.
Synonymy Type locality
Australian
mainland
Tasmania
New Guinea
Timor
Lembata Is,
Indonesia
New Caledonia
Lord Howe Is.
Nyctophilus timoriensis (Geoffroy, 1806) x x x ?
Vespertilio timoriensis Geoffroy, 1806 ? Timor
Nyctophilus major Gray, 1844 Perth, WA
Nyctophilus sherrini Thomas, 1915 Tasmania
Nyctophilus gouldi Tomes, 1858 Morton Bay, Qld x
Nyctophilus geoffroyi Leach, 1821 Australia x x
Barbastellus pacificus G r ay, 1831 Unknown
Nyctophilus australis Peters, 1861 ? Western Australia
Nyctophilus unicolor Tomes, 1858 Tasmania
N. geoffroyi pallescens Thomas, 1913 Alexandria, NT
Nyctophilus geayi Trouessart, 1915 Nicholson River, Vic.
Nyctophilus bifax Thomas, 1915 Herberton, Qld x x
Nyctophilus daedalus, Thomas 1915 Daly River, NT
Nyctophilus walkeri Thomas 1892 Adelaide River, NT x
Nyctophilus microtis Thomas, 1888 Sogeri, PNG x
Nyctophilus microtis bicolor Thomas, 1915 Aroa River, PNG
Lamingtona lophorhina McKean & Calaby, 1968 Mt Lamington, PNG
Nyctophilus microdon Laurie & Hill, 1954 Welya, PNG x
Nyctophilus arnhemensis Johnson, 1959 Cape Arnhem Peninsula, NT x
Nyctophilus howensis McKean 1975 Lord Howe Island x
Nyctophilus heran Kitchener et al., 1991 Lembata Island, Indonesia x
Nyctophilus nebulosus Parnaby, 2002 Noumea, New Caledonia x
41
2009 Australian
Zoologist volume 35 (1)
been used in species diagnoses. The principal characters
used to define species include general body size, usually
expressed as forearm length; overall body fur colour;
extent of development and morphology of a dorsal rostral
protuberance posterior to the noseleaf; relative ear size;
baculum shape, particularly whether the distal tip forms
a solid point or is bifid, and the extent of bifurcation;
general skull shape, such as relative proportions and
robustness; relative size of the auditory bulla, and relative
size of the teeth, especially the extent of reduction of the
third molars.
In addition to the above-mentioned limitations in
determining species within the genus, or perhaps because
of them, most workers (Thomas 1915 being a notable
exception) have failed to appreciate the significance of
the frequently subtle morphological differences that now
appear to be useful guides to species boundaries within
the genus. The consequent tendency to synonymise
nyctophiline taxa has hindered unraveling species limits
by significantly underestimating species diversity within
the genus.
A review of the taxonomy of timoriensis requires
principal consideration of the following named forms
of Nyctophilus: major, gouldi, sherrini and howensis.
Consideration of N. daedalus Thomas 1915 is also
necessary. This taxon was usually treated as the western
subspecies of N. bifax Thomas, 1915, following Johnson
(1964), who synonymised daedalus with bifax, though
without discussion. The status of daedalus has ranged
from a full species prior to Tate (1941), who suspected
that daedalus and bifax might be subspecifically distinct,
and the contemporary recognition of daedalus as the
western subspecies of N. bifax. However, a number
of authors have suspected that daedalus and bifax
might not be conspecific (Allison 1982, 1983; Parnaby
1987) and Troughton (1941 and subsequent editions)
considered that daedalus might be synonymous with
N. gouldi (as N. timoriensis gouldi). Koopman (1984)
considered daedalus, bifax and gouldi to be subspecifically
distinct but additional material reported by Parnaby
(1987) clearly indicated that N. gouldi and N. bifax
were distinct species with extensive sympatry. During
the course of this study, a number of large, pale-
furred Nyctophilus were examined from north-western
Queensland and northern Northern Territory. It was
initially unclear whether these specimens were large
N. daedalus, a pale northern form of N. timoriensis, or
perhaps a large northern form of N. gouldi.
The primary focus of this paper is to clarify species limits
within the suite of taxa variously associated with the name
timoriensis. This involves consideration of timoriensis itself,
and of major, gouldi, sherrini, daedalus, bifax, howensis, and
New Guinea material previously referred to N. timoriensis.
Methods
Specimen registration prefixes refer to collections held
by the following institutions: AM, Australian Museum,
Sydney; AMNH, American Museum of Natural History,
New York; BBM, Bernice P. Bishop Museum, Honolulu;
NHM, Natural History Museum, London; C, Museum of
Victoria, Melbourne; CG, Muséum National d’Histoire
Naturelle, Paris; CM, CSIRO National Wildlife Collection,
Canberra; J, JM, Queensland Museum, Brisbane; MG,
Museo Civico di Storia Naturale di Genova “Giacomo
Doria”, Genova, Italy; NTM, Northern Territory Museum
and Art Gallery, Darwin; SAM, South Australian Museum,
Adelaide; QV, Queen Victoria Museum and Art Gallery,
Launceston; WAM, Western Australian Museum, Perth.
The manner is which measurements were taken is shown
in Fig. 1 and their abbreviations used in the text are:
CON – Condylobasal Skull Length: from the posterior
surface of the occipital condyles to the most
anterior extension of the premaxilla;
GL – Greatest length of skull: from the most anterior
extension of the premaxilla to the posterior of the
lambdoidal crest;
CM3 – Length of maxillary tooth row: from anterior
cingulum of canine to posterior cingulum of M3;
C1–C1 – Outer breadth across canines from cingula;
ZYG – Zygomatic breadth, maximum breadth across
zygomatic arches;
INT – Least inter-temporal breadth;
M3–M3 – Maximum breadth from left M3 to right M3,
from labial cingula;
BRH – Braincase height: calliper blade positioned along
basioccipital-basisphenoid bones and along the
sagittal crest;
MAS – maximum breadth across mastoids;
BTB – Least inter-bulla distance, least distance between
each bulla;
BUL – Bulla length, from base of eustachian tube when
present;
BAS – Length of basicranial floor: most anterior point
of foramen magnum to most anterior point of
interpterygoid fossa;
M3L – M3 length measured at cingula;
M3B – maximum breadth of M3 measured at cingula;
PAL – Palatal-sinual length, from the most posterior
extension of the anterior palatial emargination to
the most anterior extension of the pterygoid fossa;
MESO – maximum internal breadth of mesopterygoid
fossa level with the hamular processes;
JWL – length of right dentary from anterior cingulum of
canine to posterior of mandibular condyle;
CM3 – length of tooth row from anterior cingulum of
canine to posterior cingulum of M3;
M1-M3 – length of molar row from anterior cingulum of
M1 to posterior cingulum of M3;
Baculum Length – maximum length from most posterior
tip of proximal arms to distal tip, taken perpendicular
to the dorsal surface of the main shaft;
Baculum Breadth – maximum breadth across proximal
arms at their base;
A taxonomic review of Australian Greater Long-eared Bats
42 2009
Australian
Zoologist volume 35 (1)
Baculum Height – maximum height from ventral extent
of proximal arm to distal tip;
Ear Length – taken from the junction of outer ear margin
near the jaw;
FA – forearm length, taken with the wings folded;
D1 – Digit 1 length to base of claw;
D3.1 – Digit 3 metacarpal length, from the anterior
margin of the forearm to the middle of the joint,
taken with the wings half folded;
D3.2 – Length of the first phalanx of third digit;
D3.3 – Length of the second phalanx of third digit;
D5.1 – Digit 5 metacarpal length, from the anterior
margin of the forearm to the middle of the joint,
taken with the wings half folded;
D5.2 – Length of the first phalanx of fifth digit;
D5.3 – Length of the second phalanx of fifth digit;
HL – Hindleg length, taken with the leg bent and pes
bent, note that this is not equivalent to tibia
length.
External measurements were taken with a vernier
dial calliper to the nearest 0.1 mm. Skull and dental
measurements were recorded to the nearest 0.01 mm,
except for BUL, BTB, BAS, M3L and M3B which were
estimated to the nearest 0.01 mm using an eye piece
graticule of a dissecting microscope.
CT scans were made using a Skyscan model 1174
micro CT scanner, using the following software packages:
NRecon (version 1.5.1.5 (C) Skyscan, Belgium 2008) was
used for reconstruction of 3D data sets from RAW CT
x-ray images; 3D surface models used in illustrations were
generated using CTAn Software (version 1.9.2.3 (C),
Skyscan, Belgium 2003-8), and measurements of selected
bacula were made using DataViewer (version 1.4.0.4 (C)
Skyscan Belgium).
Figure 1. The way in which measurements were taken for cranial, dental, external and bacular characters, see text
for abbreviations.
Parnaby
43
2009 Australian
Zoologist volume 35 (1)
Dental nomenclature follows Menu (1985).
Statistical analyses were undertaken using SYSTAT
version 9.0. Canonical Variates Analysis (CVA) and
Principal Components Analysis (PCA) were used to
explore relationships between specimens, based on
untransformed external, skull and dental measurements
using a correlation matrix to remove the influence of
scale in measurements (Lattin et al. 2003). Forearm
measurements were combined with craniodental
measurements in CVA and PC analyses, partly to
increase sample sizes, and because contrasts between
skull dimensions and forearm length were known to be
diagnostic for some Nyctophilus species. An exploratory
data analytic approach was taken which does not entail
tests of statistical significance.
RESULTS
The status of Vespertilio timoriensis Geoffroy,
1806
Geoffroy (1806) established Vespertilio timoriensis, the first
Nyctophilus to be named, on material collected by Peron
and Lesueur during the Baudin Expedition of 1800–1804.
Geoffroy’s description, paraphrased below, is brief:
“Vesp. timoriensis. A new species which we owe to the
work and research of Peron and Lesueur. The ears are
large, of the length of the head, and are united together
by a small membrane, the tragus is shaped like a half
heart. The fur is brownish black above, ashy brown
underneath; the hairs are very bushy, fairly long and
soft; its measurements are: body 70 mm; tail 40 mm and
wingspan 270 mm.”
Plate 47 accompanying his description illustrates the bust
of a long-eared bat that is consistent with a species of
Nyctophilus, particularly in the characteristic tragus shape.
Authors in the decades following Geoffroy (1806) mostly
paraphrased his original account and state that the species
was from Timor (Desmarest 1820, Lesson 1827, Fischer
1829, Geoffroy 1832). Temminck (1840), in what appears
to be a first hand communication from Geoffroy, notes
that it is uncertain whether timoriensis originated from
South Africa, Asia or Australia.
Geoffroy gave no indication of the number of specimens
upon which his description is based although it is
generally assumed to be a single specimen (Tomes 1858,
Thomas 1914, Tate 1941). However, Temminck (1840)
states that, according to Geoffroy, the species was known
from two specimens, a male and female, that closely
resembled each other.
Two specimens in the Muséum National d’Histoire
Naturelle, Paris have been suspected, at different times,
of being Geoffroy’s original material and thus possible
syntypes of timoriensis. Rode (1941) listed no. 884 (now
registered as CG1990-36), a puppet skin with skull
extracted and lost, as the type. Tate (1941) based his
account of N. timoriensis on a then unregistered male in
alcohol, skull extracted and evidently without locality
or details of collector (now registered as CG1985-33). In
1990 I examined both specimens, through the kindness
of Michel Tranier. Neither specimen is likely to be among
the original material upon which Geoffroy based his
description.
The specimen CG1990–36 has long ears which are joined
in the midline and has a noseleaf that resembles that of
Nyctophilus. However, it differs from any Nyctophilus that
I have examined in that both surfaces of the ears, snout
and nose-leaf are covered by short thick hairs, unlike
the fine hairs of any species of Nyctophilus. Furthermore,
tragus shape of CG1990–36 differs from all Nyctophilus
in being sharply inflected midway along its length. The
wingspan of this specimen is 264mm but as the left
wing tip is missing, wingspan could have been around
270mm as given in the original description of timoriensis;
Tail length is about 40mm and snout-vent length is
53mm, though the snout is slightly bent. Ear length is
19mm though the ear tip is slightly curled. Thus if body
length included ears, then body length would equate to
about 70mm. Although these dimensions correspond
approximately with those given by Geoffroy for Vespertilio
timoriensis, this specimen is unlikely to have formed the
basis of Geoffroy’s description because its tragus shape is
distinctly unlike that shown in his illustration. Overall, I
am not convinced that this specimen is a Nyctophilus and
further examination is warranted to clarify its identity,
including direct comparisons with material of other long-
eared vespertilionid genera. Irrespective of its identity,
there is no certainty that the specimen represents
Geoffroy’s original material, as discussed below.
Specimen CG1985-33, suspected by Tate (1941) to
be a syntype of Vespertilio timoriensis, is identifiable on
morphological criteria as a specimen of N. sherrini from
Tasmania, recognised here as a full species (see below).
In its size (see Table 9), elongate skull and unreduced
third molars, this specimen equates with N. sherrini
but differs from all other large mainland Australian
forms of the genus. Nyctophilus sherrini has the most
distinctive skull morphology of any of the large extant
forms of the genus in its combination of relatively
narrow skull, broad intertemporal region, unreduced third
molars, short interpterygoid fossa, and elongate posterior
extension of the palate. In all of these features, CG1985-
33 differs substantially from the holotype male of N.
major, but resembles the young adult male holotype of N.
sherrini (Fig. 2). Michel Tranier (in lit., 3 May, 1990) has
established that this specimen was collected in Tasmania
and acquired by the Paris Museum in 1840: “probably
from Gould collection, and maybe as material given to
Verreaux”. Tranier further stated that “I have reviewed
our specimens of Nyctophilus: I am sure that the only
type is No. 884…I am afraid that the exact origin of N.
timoriensis will never be elucidated”.
Thomas (1914) considered that the name timoriensis
should be “dropped for the present, as it is impossible to
identify it with certainty among the Australian species,
and it may yet turn up in Timor.” He proposed that N.
major Gray, 1875 be used instead of N. timoriensis with
reference to populations from Western Australia, and
promoted this approach in his generic revision (Thomas
A taxonomic review of Australian Greater Long-eared Bats
44 2009
Australian
Zoologist volume 35 (1)
1915). As detailed above, I also believe that it is not
possible to determine which species formed the basis of
Geoffroy’s original description and illustration. Moreover,
even if the species identity of the skin CG1990-36 (No.
884) could be determined (e.g. through DNA sequence
analysis), it remains uncertain as to whether this specimen
is a syntype, given that tragus shape is inconsistent with
Geoffroy’s illustration.
The principal justification for prior application of
timoriensis to Australian Nyctophilus, the perception that
Nyctophilus does not occur in Timor, and therefore must
have come from an Australian locality (Tomes 1858;
Iredale and Troughton 1934; Goodwin 1979), became
invalid with the description of N. heran from eastern
Indonesia (Kitchener et al. 1991; Corbet and Hill 1992).
Even more immediately, specimens of a Nyctophilus were
recently obtained from Timor by Kristopher Helgen (pers.
comm.).
Stability of nomenclature is a central tenet of the
International Code of Zoological Nomenclature (ICZN
1999). The confused use of timoriensis and major for
Western Australian populations since 1981 has culminated
in the southwestern populations being listed by McKenzie
(2008) as “Nyctophilus sp.” in the influential text of Van
Dyck and Strahan (2008).
For the present, I follow the opinion of Thomas (1914)
and treat Vespertilio timoriensis Geoffroy, 1806 as a nomen
dubium, based on the level of uncertainty surrounding
the whereabouts of Geoffroy’s material, uncertainty over
the type locality, and the improbability of matching
any current taxon to Geoffroy’s description. A more
formal action to stabilize usage of the name V. timoriensis
Geoffroy, 1806 for the recently (re)discovered Timorese
Nyctophilus will be taken in a separate publication.
This will require comparison with N. heran from nearby
Lembata Island, to which the Timorese material bears a
general resemblance.
The nomenclatural history of timoriensis and
associated taxa
In the first revision of Nyctophilus, Tomes (1858)
questioned the accuracy of the type locality of timoriensis
evidently on the grounds that further specimens had not
been obtained from Timor, despite extensive collections
of bats having been made, and because he examined the
“original specimen” of Geoffroy which he considered to
be “absolutely identical” with three collected for John
Gould from southwestern Western Australia. Tomes
did not mention Gray’s (1844-1875) application of the
name N. major to a specimen from southwestern Western
Australia. Gray (1844-1875) stated that he applied the
name N. major to a large specimen of the genus from
Western Australia, because he was unable to allocate it
to any of the four species recognised in the revision of
Tomes (1858). Gray (loc. cit.) referred to a colour plate
of this specimen but did not provide a description or
measurements of the holotype. Peters (1861) noted that
N. major was not mentioned by Tomes (1858) and thus
omitted from the synonomy of N. timoriensis. Thomas
(1914) drew attention to the fact that, because the
colour plate of N. major was not published by Gray until
1875, authorship should be accredited to Peters, 1861
and not Gray, 1875. However, Mahoney and Walton
(1988) established the publication date of Gray’s plate as
1844, on the basis that, although Gray’s plate and text
were published progressively in parts until completion in
1875, Gray had publicly distributed the plate at various
times from 1844.
In his review of Nyctophilus, Dobson (1878) synonymised
the four species recognised by Tomes (1858), i.e. N.
geoffroyi Leach, 1821, N. unicolor Tomes, 1858 and
N. gouldi, with timoriensis. He believed that the slight
differences used by Tomes to differentiate species were
most likely age or geographic differences and in view
of the limited material, felt that recognition of more
than a single species was unjustified. Dobson’s proposal
was rejected by Thomas (1914) who followed the
species arrangement of Tomes. Thomas suggested that
major Peters, 1861 be used for large Nyctophilus from
Figure 2. Photographs of the ventral views of the skull of
a), CG1985-33, a putative syntype of Vespertilio timoriensis
Geoffroy, 1806; b), holotype of N. sherrini and c), holotype
of N. major. All are males. Scale bar represents 10 mm.
[Photographs b and c, courtesy of the Natural History
Museum, London.]
Parnaby
45
2009 Australian
Zoologist volume 35 (1)
southwestern Western Australia and that timoriensis
be dropped due to the uncertainty surrounding the
identity of Geoffroy’s type(s) among specimens in the
Paris Museum as well as the uncertainty about the type
locality. In his revision, Thomas (1915) recognised major
and gouldi as separate species and described as new, N.
sherrini from Tasmania, N. daedalus from the Northern
Territory and N. bifax from Queensland.
Iredale and Troughton (1934) considered Geoffroy’s (1806)
identification of Timor as the type locality of Vespertilio
timoriensis to be erroneous. They regarded the type locality
to be southwestern Western Australia and populations
from that region to represent nominate timoriensis, with
major as a junior synonym. They treated gouldi and sherrini
as subspecies of N. timoriensis, from southeastern mainland
Australia and Tasmania, respectively.
In his generic review, Tate (1941) recognised a
timoriensis group consisting of the forms major, gouldi,
sherrini and timoriensis. Tate based his concept of
timoriensis on an unregistered spirit specimen in the
Paris Museum which he suspected was Geoffroy’s
original specimen (now CG1985-33; argued above to
be a specimen of N. sherrini, probably collected well
after Geoffroy’s publication). Tate noted differences
in the skull and dentition between the holotype of
major and the presumed holotype of N. timoriensis. In
his 1941 treatment, he appears to have regarded gouldi
and sherrini as races of N. timoriensis. However, in a
subsequent work, Tate (1952) listed major and sherrini
as subspecies of N. timoriensis and N. gouldi as a distinct,
albeit closely related, species.
Through much of the second half of the 20th Century, only
two species of Nyctophilus were recognized in the southern
half of Australia: N. geoffroyi and N. timoriensis (e.g.
Troughton 1967; Ride 1970; Corbet and Hill 1980; Allison
and Koopman in Honacki et al. 1982). Hall and Richards
(1979) provided evidence that N. gouldi and N. timoriensis
are distinct species and summarised the distribution of
each species in eastern Australia. Prior to this, Tate (1941)
had been universally followed in treating gouldi as the
southeastern Australian race of timoriensis; apparently, his
revised opinion, that N. gouldi was a separate species (Tate
1952), had been overlooked. The presence of N. gouldi
in far southwestern Western Australia was first noted by
Kitchener and Vicker (1981).
Most authors subsequent to Tate (1952) have treated
major as a synonym of N. timoriensis (Troughton 1967;
Ride 1970; Corbet and Hill 1980; Allison and Koopman
in Honacki et al. 1982; Richards 1983; Parnaby 1995;
Churchill 1998; Simons 2005; ABRS 2008). Kitchener
and Vicker (1981) used N. major for Western Australian
populations, though without comment. Subsequent
confusion has arisen regarding Western Australian
populations which have been variously called N. major
(e.g. McKenzie and Robinson 1987; Hosken 1996; Bailey
and Haythornthwaite 1998; Hobbs et al. 2003), N.
timoriensis major (e.g. Hosken 1997; Menkhorst and
Knight 2004) or N. timoriensis (e.g. How et al. 2001; Bullen
and McKenzie 2004).
The more recent assessments of Nyctophilus offer divergent
arrangements of the timoriensis group. Corbet and Hill
(1980, 1986) recognise N. timoriensis alone. Hill and
Koopman (1981) and Allison (1982) tentatively recognise
N. timoriensis and N. gouldi as full species. Hill and Pratt
(1981) recognised major and timoriensis as separate taxa but
reserved judgement on whether the differences warranted
subspecies or full species rank. They restricted nominate
N. timoriensis to a possible Timorese population. Koopman
(1984) presents the most recent published revision of
Australian Nyctophilus. He treated N. gouldi as a distinct
species and recognised three subspecies of timoriensis in
Australia: major from southwestern Western Australia;
sherrini from Tasmania; and tentatively referred two
specimens from northern Australia to nominate timoriensis.
In summary, all authors except Thomas (1915) have
applied timoriensis to at least some eastern Australian
Nyctophilus populations and have referred Western
Australian populations either to nominate N. timoriensis,
N. major or to N. timoriensis major. Two other taxa have
been treated as subspecies of N. timoriensis by many
authors in the past: gouldi from southeastern Australia
and sherrini from Tasmania, with Thomas (1915) alone
recognising both as full species.
As noted above, Koopman (1984) identified possible
nominate N. timoriensis within the northern Australian
bat fauna. At the same time, he expressed the opinion
that two other northern nyctophiline taxa, bifax and
daedalus, could be subspecies of N. gouldi (Koopman
1984). In this, he partly reflected Troughton’s (1941)
opinion that daedalus from northern Australia might be
synonymous with N. gouldi (as N. timoriensis gouldi) from
south-eastern Australia.
In his original description of the species, Thomas (1915)
compared N. daedalus with N. gouldi and N. bifax.
Diagnostic criteria listed by Thomas for separating N.
daedalus from N. gouldi are the smaller bullae, shorter
ears and a less developed nasal prominence. The only
character cited as distinguishing N. daedalus from N. bifax
is baculum shape: the distal tip forms a solid point in N.
daedalus but a distinct notch in N. bifax. Tate (1941, 1952)
was ambivalent about the relationship between the two
taxa. In 1941 he noted the consistently greater Zygomatic
Breadth and more reduced M3 of N. daedalus. However, he
also noted an overall similarity between daedalus and N.
bifax and concluded that they might be subspecies. After
examining more specimens of N. bifax, Tate (1952) was
able to confirm a consistent difference in baculum shape,
as noted initially by Thomas. His comments suggest that
he now regarded them as possibly distinct species.
Johnson (1964) treated daedalus as a western, allopatric
subspecies of the eastern Australian N. bifax and most
authors over the past four decades have followed this
opinion (e.g. Ride 1970, Hall and Richards 1979,
Corbet and Hill 1986, Allison 1983). However, Johnson
(1964) did not justify his treatment of daedalus, which
is at odds with the views expressed by the majority
of preceding workers (Iredale and Troughton 1934,
Troughton 1941 and subsequent editions, Tate 1941
and 1952, Johnson 1959).
A taxonomic review of Australian Greater Long-eared Bats
46 2009
Australian
Zoologist volume 35 (1)
As noted above, Troughton (1941) speculated that daedalus
might prove to be a subspecies of N. gouldi from southeastern
Australia. This view was extended by Koopman (1984)
who tentatively synonymised both daedalus and bifax with
southeastern Australian N. gouldi, suggesting that daedalus
was in some respects morphologically intermediate between
gouldi and bifax. However, Allison (1982, 1983) noted
the uncertain status of daedalus and speculated that it
could represent a separate species on the basis of baculum
differences, as did Parnaby (1987).
Hill and Pratt (1981) report two large specimens of
Nyctophilus (an adult male and female) from Mt Missim,
northeastern Papua New Guinea which they provisionally
refer to N. timoriensis. They examined the skull of the
female specimen, of which I have examined photographs.
I have obtained a further three adult female specimens
from Mt Missim, which appear to be conspecific with
those reported by Hill and Pratt.
Systematics
Nyctophilus corbeni sp. nov.
Holotype: Australian Museum number M38833 adult
male, field number 6HP04, body fixed in 80% ethanol and
stored in 75% ethanol, skull extracted, penis separated
from the body and stored in 75% ethanol. Captured by
H. Parnaby on 7th May 2006 in a bat trap (harp trap) set
across a road. Field numbers for vials of tissue samples
preserved in 90% ethanol at the Australian Museum
are: liver, 39810; 39856; pectoral muscle: 39729, 39764.
Measurements of the holotype are given in Table 2.
Paratypes: A total of four, all captured by H. Parnaby
in bat traps set across roads on 7th May 2006:
AM registration numbers, field numbers in brackets:
AM38834 (6HP05), adult female captured at the
type locality, field number for liver sample stored in
90% ethanol is 39779; the remaining three paratypes
Table 2. Cranial and external measurements of holotypes of N. major and associated forms. Measurements taken from
Thomas (1915), Tate (1941), J.E. Hill (pers. comm.) and Glenn Hoye (pers. comm.).
N. major N. m. tor subsp.
nov.
N. corbeni sp.
nov.
N. sherrini N. daedalus
Skull and dental measurements
NHM
44.7.9.20
male
WAM63601
male
AM38833
male
NHM
52.1.15.50
male
NHM
97.4.12.8
male
GL 18.0 18.1 19.43 18.5 17.6
CON 16.51 17.75 17.2 16
CM37.3 6.60 6.90 6.9 6.4
C1–C15.9 5.41 6.01 4.7 5.2
ZYG 12 11.42 12.45 11.4 11.4
INT 3.8 3.70 3.83 4.0 3.7
M3-M37.9 7.30 7.97 7.1 7.35
M3L 0.89 0.70 0.75 0.84 0.62
M3B 2.05 1.95 2.15 2.2 1.8
BRH 6.38 7.14 6.35 6.5
MAS 9.80 10.25 8.9 9.2
BUL 4.05 4.3 4.2 3.7
BTB 1.80 2.1
BAS 6.4 7.3
PAL 6.6 6.3 7.1 6.6
MESO 1.95 2.0 2.0 1.8
JWL 11.95 13.3
CM37.25 7.64
M1-M35.1 5.15
External measurements
Ear length 27.1* 25.2*
Forearm length 44 41.5* 45.4* 45 41
Digit 1 length 6.2 6.4
Digit 3 metacarpal length 39.5 44.2
Digit 3 phalanx 1 length 15.6 16.7
Digit 3 phalanx 2 length 14.6 15.2
Digit 3 phalanx 3 length 9.1 10.3
Digit 5 metacarpal length 38.3 41.6
Digit 5 phalanx 1 length 10.8 11.5
Digit 5 phalanx 2 length 9.3 9.7
Hind-leg length 21.0 21.4*
HB 65 51 59* 55 52
Tail 40 42 54* 45 41
Weight 11* 13.5*
* field measurements.
Parnaby
47
2009 Australian
Zoologist volume 35 (1)
were captured at a site on Old Coghill Track, 0.6 km
west of junction with track to main Gilgai Waterhole
(30° 29’ 51”S, 149° 20’ 01”E, altitude approximately
215m), Pilliga East State Forest, New South Wales:
AM38831 (6HP02) adult male, skull extracted, field
numbers for tissue samples stored in 90% ethanol are
Liver (39820, 39868) and pectoral muscle (39811);
AM38832 (6HP03) adult male, field number for liver
sample stored in 90% ethanol is 39846; and AM38835
(6HP06) adult male, field number for liver sample
stored in 90% ethanol is 39840. Bodies of all four
paratypes were fixed in 80% ethanol and stored in 75%
ethanol. Tissue samples for all four specimens are held
at the Australian Museum.
Type Locality: Old Coghill Track, 0.7 km east of
junction with track to main Gilgai Waterhole; formerly
Gilgai Flora Reserve, Pilliga East State Forest, New
South Wales. Approximate altitude 235 m. Coordinates
obtained from a Garmin GPS are 30° 29’ 58”S, 149°
20’ 53”E.
Diagnosis: A large species similar in body and skull size to
nominotypical N. major but differing in: having a relatively
broader and more robust skull: broader braincase; more
expanded and rounded zygomatic arches; a more truncated
rostrum (compare Figs. 3 and 10); palate shorter relative
to skull length (Fig. 4 and Table 4); baculum usually > 4.6
mm (Fig. 5 and 6, Table 3).
It differs from N. sherrini in: it has a relatively much
broader and more robust skull; more massive and
relatively broader rostrum; relatively narrower INT;
relatively shorter palate (Fig. 4); metacone absent on M3
and distinctly more reduced third molars (Fig. 7); and
relatively smaller bullae.
It differs from N. gouldi in: it has a broader, far more
robust skull; more massive rostrum; relatively smaller
bullae; relatively narrower INT; C1–C1 > 5.5 mm (n
= 125); ZYG > 11.1 mm (n = 125); more reduced
third molars, metacone absent and premetacrista nearly
obsolete; a longer baculum (> 3.7 mm) with a more
slender shaft (Fig. 5); and conspicuously larger body size
in sympatry with N. gouldi: FA > 42.0 mm (females) or
41.0 mm (males); C1–C1 > 5.0 mm.
It differs from N. nebulosus in: paler overall fur colour;
larger skull and dental dimensions; more robust skull;
PAL shorter relative to GL; far greater reduction of third
molars, metacone absent and third commissure obsolete
rather than well developed and subequal to second
commissure; baculum larger (> 4.0 mm) with thinner
shaft (compare Fig. 5 with Fig. 4 of Parnaby 2002).
It differs from N. daedalus in: averaging larger for all
external and cranial dimensions (Tables 3 and 4); C1–C1
> 5.6 mm; PAL relatively shorter and BAS relatively
longer, BAS > 6.5 mm; bullae relatively larger and
BUL > 3.9 mm; protocone of M1 and M2 less reduced
resulting in a convex rather than truncated lingual
margin; baculum longer with a relatively narrower base
and more slender shaft (Fig. 5), baculum length > 4.0
mm (Table 3).
Figure 3. X-ray CT scans of the holotype skull of N. corbeni
sp. nov., adult male AM38833. Scale bar represents 10 mm.
A taxonomic review of Australian Greater Long-eared Bats
48 2009
Australian
Zoologist volume 35 (1)
Easily distinguished from N. howensis which has a
much larger skull: GL, 23.09 vs 20.82; ZYG 13.88 vs
13.2; C1–C1 (outer alveoli), 6.7 vs 6.5 (outer cingula);
BRH 7.4 vs 7.45; INT 4.23 vs 4.04; CM3 8.1 vs 7.4;
M3-M3 (outer alveoli) 8.71 vs 8.5 (outer cingula); MAS
estimated at 11.3 vs 11.0; PAL, 9.46 vs 7.1; BAS 7.57
vs 7.71. N. howensis has a relatively much narrower
and more gracile skull (compare Figs 3 and 29):
relatively narrower ZYG and rostrum, and narrower
more elongate braincase; relatively narrower anterior
palatal emargination, rostral sulcus and mesopterygoid
fossa; relatively longer PAL; and M3 less reduced: the
metacone is clearly present.
It differs from N. heran, which has a better developed
postnasal bump, more pronounced membrane uniting the
distal median sides of the paired post-nasal prominences,
and in being larger for most measurements (comparisons
are for adult males): e.g. GL > 18.0 mm vs < 17.0 mm;
C1–C1 > 5.7 mm vs 4.5 mm; in having relatively much
smaller bullae; baculum larger, baculum length > 4.0
mm, with relatively smaller basal arms and main shaft
tapers less to distal point (compare Fig. 5 with Fig. 5 of
Kitchener et al. 1991).
Easily distinguished from N. geoffroyi in: having a simpler
post-nasal elevation which has a simple median vertical
grove, rather than an more developed pair of mounds
joining in the distal mid-line by an elastic membrane which
forms a distinctive “Y”-shaped structure; by larger size, e.g.
compared to sympatric N. geoffroyi, GL > 18.0 mm vs ≤ 16.7
mm (n = 126, sexes combined for mainland N. geoffroyi);
C1–C1 > 5.6 mm vs ≤ 4.8 mm (n = 117) relatively smaller
bullae; skull far more robust; more reduced M1 protocone
such that lingual margin is truncated rather than convex;
M3 more reduced with more rudimentary third commissure
and metacone not present; distal tip of glans penis blunt and
rounded rather than forming an elongate “beak”, lacking a
Figure 4. Plot of PAL vs GL for N. major complex.
a) adult females: 1, specimen from
Mundra billa (WAM22953); 2, Katanning
(AMNH197281); 3, Woodanilling (AM37642);
b), adult males, 1, specimen from Balladonia (AM39802);
2, specimen from Madura (WAM28398); X = unallocated
adults from Dr yandra Woodlands. Species symbols are:
N. corbeni sp. nov. (), N. major major (○) , N. m. tor subsp.
nov. (□), N. sherrini (∇) , N. daedalus (◊) and N. shirleyae
sp. nov. (▼).
a.
b.
Table 3. Summary statistics for bacula of various Nyctophilus
species.
Mean s.d. Range N
Baculum Length
N. corbeni sp. nov. 4.97 0.372 4.54–5.73 11
N. major 4.46 0.124 4.30–4.57 4
N. m. tor subsp. nov. 4.38 0.136 4.18–4.59 12
N. sherrini 4.32 0.296 4.00–4.51 3
N. gouldi 3.26 0.159 2.99–3.69 26
N. daedalus 3.50 0.237 3.20–3.85 6
N. nebulosus 2.95 1
N. bifax 3.49 0.129 3.28–3.73 10
Baculum Breadth
N. corbeni sp. nov. 1.24 0.194 0.78–1.46 10
N. major 1.27 0.039 1.23–1.31 3
N. m. tor subsp. nov. 1.18 0.064 1.07–1.31 11
N. sherrini 1.28 0.089 1.18–1.35 3
N. gouldi 1.09 0.090 0.90–1.27 26
N. daedalus 1.16 0.074 1.07–1.23 5
N. nebulosus 1.11 1
N. bifax 1.21 0.105 1.07–1.39 9
Baculum Height
N. corbeni sp. nov. 1.69 0.218 1.44–2.09 11
N. major 1.35 0.294 1.02–1.60 3
N. m. tor subsp. nov. 1.42 0.106 1.23–1.60 11
N. sherrini 1.48 0.142 1.37–1.64 3
N. gouldi 1.06 0.119 0.86–1.35 26
N. daedalus 1.24 0.061 1.15–1.31 5
N. nebulosus –
N. bifax 1.24 0.111 1.07–1.43 10
Parnaby
49
2009 Australian
Zoologist volume 35 (1)
Table 4. Summary statistics for 11 external and 15 skull and dental dimensions of adult specimens examined of N. corbeni
sp. nov., N. major major, N. major tor subsp. nov. and N. daedalus.
N. corbeni sp. nov.
Female Male
Mean s.d. Min Max N CV Mean s.d. Min Max N CV
EAR 26.40 1.794 24.3 29.3 10 6.8 26.43 1.069 24.4 28.3 26 4.0
D1 7.25 0.446 6.4 7.6 6 6.2 6.48 0.459 5.6 7.2 16 7.1
FA 46.65 1.238 44.7 48.9 14 2.7 44.72 1.669 41.3 49.4 35 3.7
D31 45.96 1.261 44.2 47.6 13 2.7 43.59 1.748 41.2 49.5 28 4.0
D32 18.50 1.072 16.8 20.4 10 5.8 17.03 0.727 16.0 18.3 21 4.3
D33 16.00 0.865 14.6 17.1 8 5.4 15.16 0.654 13.9 16.2 21 4.3
D51 44.95 1.284 43.4 47.1 13 2.9 42.44 1.684 40.1 48.3 29 4.0
D52 12.75 0.845 11.4 14.1 10 6.6 11.50 0.375 10.7 12.1 21 3.3
D53 10.55 0.750 9.3 11.3 6 7.1 9.57 0.875 7.8 11.6 21 9.1
HL 21.96 0.903 20.5 23.6 13 4.1 21.28 0.799 20.0 23.0 29 3.8
WT 16.22 2.055 14.3 20.0 6 12.7 13.52 0.996 11.2 15.5 20 7.4
CON 18.20 0.340 17.80 18.76 10 1.9 17.54 0.420 16.70 18.30 23 2.4
GL 20.11 0.447 19.50 20.82 10 2.2 19.20 0.508 18.00 20.20 23 2.6
CM37.25 0.159 7.00 7.44 10 2.2 6.98 0.211 6.55 7.40 23 3.0
C1-C16.27 0.155 5.90 6.50 10 2.5 6.06 0.172 5.70 6.50 23 2.8
ZYG 12.77 0.261 12.45 13.20 10 2.0 12.35 0.278 11.90 12.80 22 2.3
INT 3.75 0.145 3.60 4.00 10 3.9 3.74 0.160 3.45 4.04 23 4.3
M3–M38.22 0.256 7.77 8.50 10 3.1 7.92 0.224 7.50 8.30 23 2.8
BRH 7.16 0.167 6.95 7.45 10 2.3 6.93 0.189 6.60 7.30 23 2.7
MAS 10.59 0.326 9.90 11.00 10 3.1 10.18 0.188 9.90 10.50 23 1.8
BTB 2.26 0.165 1.97 2.50 10 7.3 2.14 0.103 1.97 2.30 21 4.8
BUL 4.42 0.126 4.18 4.60 10 2.9 4.27 0.113 4.10 4.50 21 2.6
BAS 7.42 0.287 6.97 7.71 10 3.9 7.12 0.236 6.70 7.71 21 3.3
M3L0.87 0.039 0.82 0.90 4 4.5 0.85 0.055 0.78 0.98 10 6.5
M3B2.41 0.039 2.38 2.46 4 1.6 2.23 0.087 2.09 2.42 10 3.9
PAL 6.74 0.237 6.40 7.10 7 3.5 6.45 0.220 6.10 6.90 15 3.4
N. major major
Female Male
Mean s.d. Min Max N CV Mean s.d. Min Max N CV
EAR 26.22 1.426 24.4 28.6 13 5.4 26.30 0.957 24.7 27.5 6 3.6
D1 6.69 0.553 6.0 7.6 9 8.3 6.93 0.153 6.8 7.1 3 2.2
FA 45.91 1.354 43.5 48.4 19 2.9 44.55 1.584 42.5 47.5 8 3.6
D31 43.84 1.250 42.2 45.9 14 2.9 42.29 1.716 40.05 44.9 6 4.1
D32 17.11 0.651 16.1 17.9 10 3.8 16.52 0.746 15.5 16.6 5 4.5
D33 15.27 0.650 14.2 16.1 10 4.3 15.13 0.643 14.4 15.6 3 4.2
D51 42.68 1.188 40.8 44.7 14 2.8 41.56 1.540 39.8 44.0 6 3.7
D52 11.91 0.415 11.4 12.6 10 3.5 11.46 0.844 10.7 12.5 5 7.4
D53 9.69 0.536 9.1 10.7 10 5.5 9.57 1.007 8.5 10.5 3
HL 21.96 0.649 21.1 23.2 13 3.0 20.80 0.860 20.2 22.3 5 4.1
WT 15.0 16.5 2 12.0 13.5 2
CON 18.37 0.394 17.70 19.00 12 2.1 17.87 0.560 17.18 18.89 8 3.1
GL 19.95 0.477 19.10 20.70 12 2.4 19.36 0.523 18.83 20.32 8 2.7
CM37.46 0.230 7.15 7.80 13 3.1 7.27 0.159 7.04 7.55 8 2.2
C1-C16.05 0.213 5.70 6.35 13 3.5 5.90 0.265 5.70 6.52 8 4.5
ZYG 12.28 0.404 11.70 13.30 13 3.3 11.99 0.384 11.60 12.50 7 3.2
INT 3.83 0.156 3.50 4.00 13 4.1 3.88 0.124 3.71 4.10 8 3.2
M3–M38.09 0.311 7.70 8.60 13 3.8 7.88 0.216 7.60 8.10 8 2.7
BRH 6.93 0.293 6.50 7.50 12 4.2 6.92 0.093 6.80 7.06 7 1.3
MAS 10.20 0.326 9.70 10.70 12 3.2 10.02 0.300 9.70 10.64 7 3.0
BTB 2.02 0.165 1.89 2.30 8 8.2 1.97 0.083 1.89 2.05 5 4.2
BUL 4.34 0.121 4.18 4.51 10 2.8 4.25 0.071 4.15 4.35 5 1.7
BAS 6.96 0.293 6.60 7.38 9 4.2 6.66 0.351 6.40 7.22 5 5.3
M3L0.86 0.054 0.78 0.94 6 6.2 0.87 0.047 0.82 0.90 3 5.4
M3B2.19 0.104 2.05 2.34 6 4.7 2.20 0.085 2.13 2.30 3 3.9
PAL 7.30 0.243 7.00 7.75 11 3.3 7.14 0.222 6.70 7.35 8 3.1
A taxonomic review of Australian Greater Long-eared Bats
50 2009
Australian
Zoologist volume 35 (1)
Table 4. continued
N. m. tor subsp. nov.
Female Male
Mean s.d. Min Max N CV Mean s.d. Min Max N CV
EAR 24.23 1.638 21.5 27.5 19 6.8 25.02 1.452 21.3 27.3 43 5.8
D1 6.28 0.364 5.6 6.8 16 5.8 6.16 0.376 5.3 7.0 32 6.1
FA 41.34 1.188 39.3 44.6 26 2.9 40.94 1.347 37.6 43.3 53 3.3
D31 39.84 1.354 37.7 43.0 25 3.4 39.32 1.292 36.8 42.0 53 3.3
D32 15.73 0.760 14.2 17.0 16 4.8 15.46 0.687 13.9 17.0 37 4.4
D33 13.87 0.732 12.7 15.4 16 5.3 13.89 1.256 7.8 15.4 35 9.0
D51 39.11 1.141 36.9 41.6 25 2.9 38.44 1.183 36.4 40.7 52 3.1
D52 11.08 0.672 10.1 12.6 16 6.1 10.87 0.463 9.9 11.7 35 4.3
D53 9.09 1.137 6.1 10.6 16 12.5 9.31 0.605 8.1 10.6 36 6.5
HL 20.68 0.890 19.2 22.5 17 4.3 19.79 0.720 18.4 21.5 43 3.6
WT 11.0 11.5 2 10.82 1.361 8.0 12.3 18 12.6
CON 16.68 0.238 16.20 17.00 12 1.4 16.61 0.314 15.90 17.20 30 1.9
GL 18.06 0.276 17.50 18.30 12 1.5 18.04 0.347 17.20 18.75 31 1.9
CM36.68 0.156 6.40 7.00 12 2.3 6.76 0.163 6.50 7.10 31 2.4
C1-C15.34 0.218 5.00 5.60 12 4.1 5.39 0.187 5.00 5.70 31 3.5
ZYG 11.25 0.271 10.70 11.60 11 2.4 11.12 0.281 10.50 11.62 31 2.5
INT 3.59 0.089 3.50 3.80 12 2.5 3.65 0.120 3.49 4.00 31 3.3
M3–M37.36 0.226 7.00 7.80 12 3.1 7.31 0.185 6.90 7.70 31 2.5
BRH 6.31 0.135 6.10 6.60 12 2.1 6.46 0.177 6.10 6.80 31 2.7
MAS 9.48 0.286 9.00 9.80 12 3.0 9.42 0.223 9.00 9.90 31 2.4
BTB 1.81 0.090 1.64 1.97 10 5.0 1.84 0.135 1.56 2.13 28 7.3
BUL 4.06 0.116 3.94 4.26 11 2.9 4.02 0.109 3.77 4.20 28 2.7
BAS 6.20 0.174 5.82 6.48 10 2.8 6.23 0.208 5.82 6.60 24 3.3
M3L0.81 0.057 0.74 0.90 7 7.0 0.81 0.047 0.74 0.90 20 5.7
M3B2.10 0.145 1.80 2.26 7 6.9 2.07 0.090 1.93 2.26 19 4.3
PAL 6.65 0.191 6.50 6.90 4 2.9 6.56 0.229 6.10 7.00 23 3.5
N. daedalus
Female Male
Mean s.d. Min Max N CV Mean s.d. Min Max N CV
EAR 23.52 1.355 20.5 25.1 13 5.8 23.75 1.083 21.9 25.8 17 4.6
D1 6.61 0.500 6.0 7.7 12 7.6 6.34 0.280 6.0 6.8 10 4.4
FA 43.30 1.597 40.2 45.8 13 3.7 40.52 1.561 38.3 43.7 18 3.9
D31 41.39 1.720 38.3 43.4 13 4.2 38.61 1.630 36.8 42.6 18 4.2
D32 16.08 0.728 15.1 17.3 12 4.5 15.20 0.503 14.5 15.8 10 3.3
D33 14.30 0.858 13.0 16.2 12 6.0 13.67 0.397 12.9 14.1 10 2.9
D51 40.93 1.213 38.9 42.8 13 3.0 38.73 1.087 37.3 41.8 18 2.8
D52 10.93 0.602 9.8 11.7 12 5.5 10.30 0.488 9.5 11.1 10 4.7
D53 9.57 0.631 8.7 10.8 12 6.6 8.74 0.690 7.7 9.7 10 7.9
HL 20.74 0.906 19.6 22.0 11 4.4 19.37 0.798 18.2 21.5 17 4.1
WT 12.07 1.629 10.2 13.2 3 13.5 7.88 0.954 6.5 9.0 10 12.1
CON 16.12 0.342 15.60 16.60 9 2.1 15.55 0.262 15.20 16.10 15 1.7
GL 17.68 0.465 17.00 18.30 8 2.6 17.09 0.318 16.72 17.70 15 1.9
CM36.60 0.200 6.30 6.90 9 3.0 6.30 0.184 5.90 6.60 15 2.9
C1-C15.17 0.218 4.90 5.50 9 4.2 4.96 0.145 4.70 5.20 15 2.9
ZYG 11.23 0.466 10.50 11.90 9 4.2 10.72 0.353 10.20 11.30 15 3.3
INT 3.69 0.169 3.40 4.00 9 4.6 3.59 0.108 3.50 3.90 15 3.0
M3–M37.24 0.235 6.90 7.50 9 3.2 6.98 0.191 6.60 7.30 15 2.7
BRH 6.46 0.260 6.10 6.90 9 4.0 6.30 0.191 6.00 6.60 15 3.0
MAS 9.40 0.387 8.80 9.90 9 4.1 9.11 0.283 8.80 9.70 15 3.1
BTB 2.14 0.215 1.80 2.46 9 10.0 2.02 0.170 1.72 2.38 15 8.4
BUL 3.71 0.085 3.53 3.77 8 2.3 3.69 0.128 3.44 3.85 15 3.5
BAS 5.92 0.249 5.58 6.31 9 4.2 5.74 0.248 5.41 6.30 13 4.3
M3L0.82 0.054 0.74 0.86 9 6.6 0.78 0.054 0.70 0.86 9 7.0
M3B2.05 0.100 1.85 2.21 9 4.9 1.96 0.074 1.88 2.13 9 3.8
PAL 6.47 0.197 6.20 6.80 6 3.0 6.24 0.217 5.90 6.65 13 3.5
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2009 Australian
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distal median dorsal serrated ridge; distal tip of baculum not
fully ossified, occasionally with very weak notch compared
to solid point; baculum length > 4.6 mm vs 2.4 - 2.9 mm (n
= 13 for mainland and Tasmanian N. geoffroyi).
Etymology: Named in honour of Christopher John Corben,
bat researcher, frog expert, ornithologist and technophile,
in recognition of his contribution to Australian zoology
from his largely unfunded pioneering development and
ceaseless refinement of technology and software for
detection, storage and analysis of bat echolocation calls
which has revolutionised bat research and inventory in
Australia and on other continents.
Figure 6. Plot of Baculum Height vs Baculum Length for
N. corbeni sp. nov. (), N. major major (•), N. m. tor subsp.
nov. (□), N. sherrini (▼) and N. daedalus (◊).
Figure 7. Scanning electron micrographs of left M1 (left), M2
and M3 (right) of: a), N. corbeni sp. nov. (AM3909, female);
b), N. major major (AM6319, male) c), N. m. tor subsp. nov.
(AM35884, male); d), N. sherrini (AM37936, male) and e),
N. gouldi (AM3912, male). Scale bar represents 1 mm.
Figure 5. X-ray CT scans of the baculum showing lateral
(left), and dorsal (right) views of: a) N. major major
(AM39797); b), N. m. tor subsp. nov. (WAM63601); c) N.
corbeni sp. nov. (AM38833); d), N. sherrini (AM34455); e),
N. gouldi (AM38841); f), N. daedalus (AM34451); and g), N.
bifax (CM11628). Scale bar represents 2 mm.
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Distribution: Drier areas of Queensland, New South
Wales and South Australia (see Fig. 8). Most records
are from inland of the Great Dividing Range. The most
northerly record is from Yebna Station, 80 km west of
Taroom, Queensland; Danggali Conservation Park, South
Australia is the most westerly locality for which I have
examined specimens; however, specimens from Canegrass,
South Australia (33o 35’ 37” S, 140 o
03’ 06”E; SAM
17320-21) identified from gene sequencing (B. Appleton,
T. Reardon, et. al. in progress) represents the most western
record of the species. The western distribution of this
species appears to be truncated by the Flinders Ranges in
South Australia. A comprehensive review of field records
of this species is presented by Turbill and Ellis (2006).
This species is sympatric with N. geoffroyi throughout its
entire range; in the southern and eastern parts of its range
it shows extensive sympatry with N. gouldi.
Specimens examined: A total of 64, see Appendix.
Remarks: The presence of this large distinctive species
in eastern Australia was evidently overlooked until Hall
and Richards (1979) drew attention to its distinction from
N. gouldi and summarised distribution data for the few
specimens available (as N. timoriensis).
Larger examples of N. gouldi from higher rainfall areas
are of the same general size as N. corbeni sp. nov.
In particular, FA measurements (used extensively in
field identifications of Nyctophilus) show broad overlap
between each species for each sex. However, N. corbeni
sp. nov. has a noticeably broader head and snout, as
reflected by C1-C1: for males, N. gouldi maximum = 5.2
mm (n=85) vs minimum for N. corbeni = 5.7 (n=26);
for females, maximum for N. gouldi = 5.4 (n=58)
compared with minimum of 5.9 (n=15) for N. corbeni.
The two species are broadly sympatric inland of the Great
Dividing Range in northern and northwestern Victoria,
New South Wales and Queensland and both species have
been captured in the same trap on the same night. Inland
N. gouldi are generally smaller than those of montane or
subcoastal regions (Parnaby 1987; Lumsden 1994; Young
and Ford 2000) and are readily distinguished from N.
corbeni sp. nov., as seen in a plot of FA vs C1-C1 (Fig. 9).
Nyctophilus major Gray, 1844
Holotype: NHM no. 44.7.9.20, male skin and skull.
Collected by J. Gilbert 20 March 1843 (Mahoney and
Walton 1988). Gilbert’s field number 23 (J. Mahoney
pers. comm. 1984).
Figure 8. Distribution of specimens examined of N. corbeni sp. nov. (), N. major major (○) N. m. tor subsp. nov. (•), N.
daedalus (■), and N. sherrini ().
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2009 Australian
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Type locality: Perth, Western Australia.
Re-diagnosis (of nominotypical form): Nyctophilus major
major differs from N. corbeni sp. nov. in: its relatively
narrower skull (Figs 3 and 10; Table 4); relatively narrower
and less rounded zygomatic arches; narrower braincase;
squarer more nearly parallel-sided rostrum; proportionally
longer palate (Fig. 4); usually smaller, more slender
baculum; relatively more reduced protocone on M1
resulting in more truncated lingual margin (Fig. 7).
It differs from N. gouldi in: its more reduced third
molars; baculum longer, > 4.0 mm with more slender
shaft. It is further distinguished from southwestern
Australian populations of N. gouldi in: its considerably
larger size; more massive, narrower skull; larger rostrum;
conspicuously more reduced protocone on M1 and larger,
more slender baculum.
It differs from N. sherrini in: its relatively broader, more
massive skull; relatively broader rostrum, C1–C1 > 5.6
mm; relatively smaller bullae; relatively more reduced
protocone on M1 and M2 resulting in a more truncated
lingual margin (Fig. 7); reduced third molars with metacone
absent and third commissure obsolete rather than well
developed; relatively narrower INT; and relatively shorter
proximal end on the baculum.
It differs from N. daedalus in: its larger skull size (Table 4);
larger bullae; darker fur colour; larger baculum, baculum
length > 4.0 mm (Table 3, Figs 5 and 6) with a more
slender shaft.
Skull readily distinguished from that of N. howensis by
conspicuously smaller skull dimensions, more reduced M3
(see also re-diagnosis of that species).
Figure 9. Plot of C1–C1 vs FA showing separation of adult
N. gouldi (□) and N. corbeni sp. nov. (○), sold symbols
are males. a), N. gouldi from all localities in southeastern
Australia; b), N. gouldi from inland of the Great Dividing
Range from Victoria to Queensland.
Figure 10. Photographs of the skull and dentary of a young
adult male N. major major (AM5477) from Tambellup,
Western Australia. Scale bar represents 10 mm.
a.
b.
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Distribution: Southern Australia from the southwest
corner, east to Eyre Peninsula of South Australia (Figs. 8
& 11). Two subspecies are distinguished (see below), the
nominotypical form in southwestern Western Australia,
and a new subspecies from the wheatbelt of Western
Australia, east to Eyre Peninsula.
Material examined: A total of 43 specimens of the
nominotypical form, listed in the Appendix. Six specimens
are referred to an inland subspecies, described below.
Black and white photographs of the holotype skull and
dentary of N. major major.
Remarks: Nominotypical N. major and N. corbeni sp.
nov. are morphologically very close, with considerable
overlap in both external and craniodental metric variation.
They are distinguished by proportional differences in the
cranium, as noted above, and by the development of the
protocone on M1, which is usually larger in N. corbeni sp.
nov., resulting in a more rounded lingual margin (Fig. 7).
On external criteria, D3.1 length is usually shorter relative
to FA in N. major major.
Size variation within N. major
Two size morphs are evident within Western Australian
samples of N. major. Individuals from lower rainfall,
typically inland, localities across southern Western
Australia are generally smaller in body and skull size
than N. major from the higher rainfall areas of far
south-western Western Australia. Greater variation in
body and skull size is evident amongst specimens from
districts of intermediate rainfall, viz. the Dryandra
woodlands of the wheatbelt, as well as the subcoastal
districts of elevated rainfall around Balladonia and
the Roe Plain. The majority of specimens examined
from these districts are relatively small but there are
a small number of animals that are as large as those
from high rainfall districts, as well as some animals
of intermediate size. The following assessment of
size variation in N. major focuses on specimens from
Western Australian localities.
The variation within N. major is evident in a plot of
ZYG against GL for females (Fig. 12a). In the wheatbelt
region, two adult females from the Katanning district fall
into each of the large and small morphs, as do the two
females from the Woodanilling area, 18 km northwest of
Katanning. In the Roe Plains area south of the Nullarbor
Plain in eastern Western Australia, one of two females
from different localities south of Madura falls within the
small morph and the other is intermediate but closest to
the small morph, while a female from Kuthala Pass on the
edge of the Hampton Tableland at Mundrabilla, clearly
falls within the large morph.
A similar, though less clear, trend is evident in a plot of
ZYG against GL for males from all Western Australian
localities (Fig. 12b). This includes specimens from
the three districts of apparent sympatry between size
morphs. The seven animals from Dryandra Woodlands
(wheatbelt district) include two that fall within the
small morph, one that falls within the large morph,
and four that are intermediate. Similarly, four animals
from the Balladonia district include two that fall
within the small morph, one in the large morph, and
one intermediate. Five animals from Kuthala Pass,
Figure 11. Distribution of N. major major (□) and N. m. tor subsp. nov. (•) based on specimens examined. Districts of near
sympatry are: 1, Dr yandra woodlands; 2, Woodanilling; 3, Katanning; 4, Balladonia, 5, Madura and Roe Plain, and 6, Kuthala
Pass, Mundrabilla – 7, represents the type locality of N. m. tor subsp. nov.
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2009 Australian
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Mundrabilla, on the edge of the Hampton Tableland
fall within the small morph, while one from the
Madura district of the adjoining Roe Plain, falls within
the large morph.
The size contrast between males (few female specimens
are available) from inland and the far south-western areas
(home of nominotypical N. major) is clearest in a plot of
ZYG against GL from a more restricted region, localities
west of longitude 122oE in Western Australia (Fig. 12c).
Of the seven males from Dryandra woodlands, two fall
within the smaller inland morph, two fall within or close
to the large morph, and the remaining three could be
considered to be intermediate.
The small magnitude of the differences that separate the
two morphs on the basis of individual measurements (often
less than 1 mm) is deceptive. For example, measurements
for two adult females representing both morphs from the
Katanning district are respectively, GL 19.4 mm vs 18.5
mm, ZYG 12.2 vs 11.2 mm, and MAS 10.0 vs 9.5 mm, yet
the size difference is clearly evident from direct comparison
of skulls (Fig. 13). This probably reflects the inadequate
extent to which standard skull measurements, considered
individually, capture overall size and shape differences that
are apparent from direct visual comparisons. I know of at
least three bat researchers who captured live examples
of the small morph and all independently tentatively
identified them as N. gouldi, i.e. it was recognised as being
distinct from larger N. major. This indicates that the
differences apparent from comparative museum studies
are evident in live animals.
Size variation within N. major was examined using
a Principal Components Analysis (PCA) based on a
correlation matrix of FA and nine skull and dental
measurements of 67 adult specimens. Measurements were
selected for the analysis in order to maximise sample size.
Figure 12. Plot of ZYG vs GL for N. major major (○)
and N. major tor subsp. nov. (□). a), adult females from
WA and SA; b), males from WA localities; c), males from
WA localities west of longitude 122o E. Localities are: B,
Balladonia; D, Dwellingup; K, Katanning, M, Mundrabilla;
Ma, Madura; W, Wodanilling; X= males from Dr yandra
Woodlands, wheatbelt, Western Australia. H represents
holotype of N. major tor subsp. nov.
Figure 13. Photographs illustrating differences in skull size
between, left, N. major major (WAM6375); and right, N. m.
tor subsp. nov. (AMNH197281): both are adult females
from the Katanning district, Western Australia. Scale bar
represents 10 mm.
a.
b.
c.
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Standardised coefficients for each measurement on the
first PC axis suggest that this axis, which accounts for
79% of the total measurement variance (Table 5), reflects
overall skull and FA size because all coefficients are of the
same approximate magnitude and sign. Coefficients on
the second axis reflect an inverse relationship between
INT and BRH, on the one hand, against the remaining
measurements. Principal Component scores for specimens
on the first two PC axes fall into two broad groups (Fig.
14) which correspond to the two size morphs recognised
here. PC scores for most of the 7 specimens (all male)
from Dryandra woodlands are intermediate between each
group. A partial separation by sex is evident for both
morphs on PC 2. A PCA based on the external characters
EAR, FA, D31, D51 and HL showed extensive overlap
between size morphs and sex (not shown).
Geographic variation in size was examined further by
plotting scores for specimens on PC 1, used as an
indicator of general size, against longitude (Fig. 15).
Three points are evident: a) the greater size of specimens
from far south-western Western Australia, including some
individuals from the wheatbelt; b), the intermediate size
of some individuals from Dryandra; and c) the fact that
several individuals from localities south of the Nullarbor
Plain fall within the size range for the large size morph, as
do two adult males from the Balladonia district.
The integrity of the two size morphs and the relationships
of intermediate specimens were examined through a
Canonical Variates Analysis (CVA), using the same ten
characters and 67 specimens used in the PCA. Specimens
were assigned to groups based on size morph and sex, while
ten specimens were entered in the analysis ungrouped. The
latter series included seven specimens from Dryandra, the
large adult male and female from the Mundrabilla region,
and a female from the wheatbelt. The first two CV axes
captured 98.4% of the variance (Table 6) and a plot of
scores for individuals on the first two CV axes (Fig. 16)
shows a similar separation of size morphs as on the first
axes in the PCA. The jack-knifed classification function
assigned all females to the correct size morph, although
a substantial proportion was allocated to the wrong sex
Table 6. Standardised character coefficients, eigenvalues
and percentage of total variance for the first three CV
axes of a CVA of FA and 9 skull and dental dimensions of
N. major major and N. m. tor subsp. nov.
CV 1 CV 2 CV 3
FA 0.360 0.706 0.089
CON 0.255 -0.010 -1.277
GL 0.293 0.384 1.570
CM30.173 -0.886 0.819
C1–C1-0.044 -0.519 -0.194
ZYG 0.016 0.367 -0.071
INT 0.237 -0.166 -0.454
M3-M30.206 0.943 -0.701
BRH 0.185 -0.597 -0.230
MAS -0.255 -0.404 0.012
Eigenvalues 5.386 0.710 0.096
% variance 87.0 11.4 1.6
Table 5. Standardised character coefficients and variance
for each axis of a PCA based on a correlation matrix of
FA and 9 cranial and dental dimensions of N. major major
and N. m. tor subsp. nov..
PC 1 PC 2
FA 0.847 0.153
CON 0.971 0.102
GL 0.969 0.073
CM30.930 0.085
C1–C10.914 0.072
ZYG 0.929 0.090
INT 0.732 -0.591
M3-M30.899 0.196
BRH 0.790 -0.426
MAS 0.885 0.072
% variance 79.13 6.34
Figure 14. Plot of specimen scores on the first two PC
axes for adult N. major major (○) and N. m. tor subsp.
nov. (□) based on a PCA of FA and 9 cranial and dental
measurements. Females are solid symbols, males are open
symbols. Locality designations for ten specimens that were
not allocated to a group in the accompanying canonical
variates analysis are: seven males from Dryandra (X), a
female from Mundrabilla (M), a male from Madura district
(N), and a female from Woodanilling (W).
Parnaby
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2009 Australian
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(Table 7). Of the 10 specimens not allocated to a group
prior to the analysis, 3 of the 7 male Dryandra specimens
were allocated to males of the small morph, 3 to males
of the large morph and 1 to the female small morph.
The remaining three specimens were allocated to the
respective sex and morph expected on the basis of size, viz.
the female from Kuthala Pass and the male from Madura
to the large morph, the small female from Woodanilling
to the small morph. (Several key female specimens from
the wheatbelt were excluded from the CVA due to
missing measurements). The intermediate nature of these
specimens is confirmed by the PCA and CVA. However,
the classification of intermediate specimens in the CVA
should be interpreted with caution due the small sample
sizes. Further, CVA performs poorly when allocating
specimens that form a gradation compared with discrete
groups because it maximises between-group differences
relative to within-group variation.
Three scenarios can be invoked to explain size variation
within N. major, none of which were unequivocally
rejected by this study. The simplest is that size variation is
a response to environmental factors, such as moisture or
temperature gradients. It is not surprising that animals from
lower rainfall areas are, on average, smaller. Alternatively,
two or more crypticspecies might exist in the region,
Figure 15. Plot of PC 1 scores vs longitude for adult N. major major (○) and N. m. tor subsp. nov. (□) from a PC analysis
using FA and 9 cranial and dental measurements. Solid symbols represent females, open symbols are males. The ten
specimens entered in the CVA as ungrouped are indicated as per the caption for Fig. 14.
Figure 16. Plot of specimen scores on first two CV axes
for adult N. major major (○) and N. m. tor subsp. nov.
(□) based on an analysis using FA and 9 skull and dental
characters. Females Solid symbols represent females, open
symbols represent males; (+) indicates the group centroid
for each sex. The ten specimens entered in the CVA as
ungrouped are indicated as per the caption for Fig. 14.
Table 7. Results of jackknife classification function of CVA of FA and 9 skull and dental dimensions of N. major major and
N. m. tor subsp. nov., showing number of misclassified specimens per group, and allocation of 10 specimens entered in
the CVA as ungrouped.
Female
N. m. tor subsp. nov.
Female
N. m. major
Male
N. m. tor subsp. nov.
Male
N. m. major % correct
Female 9 0 3 0 75
Female N. m. major 0 5 0 5 50
Male N. m. tor subsp. nov. 5 0 22 1 79
Male N. m. major 0 3 1 3 43
Total 14 8 26 9 68
Ungrouped a priori 2 1 3 4
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either with complete reproductive isolation but a degree
of morphometric overlap, or with hybridisation and/or
introgression. If the latter scenario is correct, the likely
zone of interaction is in the wheatbelt region of south-
western Western Australia, and at locations of elevated
rainfall along the southern, near coastal areas of eastern
Western Australia. Irrespective of the significance of the
size morphs, it is clear that they co-occur; in at least one
instance, both were captured on the same night at the
same site. An adult lactating female from Kuthala Pass,
near Mundrabilla Hotel (WAM22953) falls within the
large morph (e.g. Fig. 12a) and groups with the large
morph in the PCA (Fig. 14) and CVA (Fig. 16). Eight
adult males of the small morph were evidently captured
in the same trap with this specimen. The other instances
of apparent sympatry or parapatry between size morphs
occur in several districts, as noted previously. However,
sympatry or close parapatry, perhaps due to habitat
separation, cannot be established due to imprecise locality
data. This is problematic, given that steep gradients in
rainfall and vegetation changes occur over comparatively
short distances in these areas.
On balance, I suspect that two cryptic species are present,
which are broadly sympatric in the wheatbelt and in
southern subcoastal areas of eastern Western Australia.
However, although the data are suggestive of two species,
I am unable to refute the simpler hypothesis of a variable
species with environmentally induced size variation, for
which infra-specific variation is inadequately defined in
this study due to the limited number of specimens available
from strategic locations. Resolution of this complex
problem will depend on further, targeted collecting and
detailed genetic investigations using multiple markers
to document the contemporary pattern of gene flow
between populations. In the interim, one option is to
treat N. major as a single, highly variable taxon. Another
is to recognise the small size morph as a distinct taxon,
but at subspecific level within N. major. While this
action might be unpopular at a time when the subspecies
category is treated by many taxonomists as an essentially
meaningless entity, it is taken in this instance for several
reasons: 1) it enables a refinement in diagnoses and
identification of other southern Australian Nyctophilus; 2)
formal recognition of the small morph will reduce the risk
of future confusion with N. gouldi; and 3) providing the
small morph of N. major with a formal identity should lead
to greater likelihood that it will attract the further work
that is needed to determine its true status.
Nyctophilus major tor subsp. nov.
Holotype: WAM63601 (previously AM39782), field
number 7HP51, adult male body fixed in 80% ethanol
and stored in 75% ethanol, skull extracted and cleaned.
Penis stored separately in 75% ethanol. Liver sample
stored in 95% ethanol (field number 48159) held at
the Australian Museum and liver sample (field number
48120) stored in liquid nitrogen held at the South
Australian Museum. Captured in a bat trap (harp trap)
set next to Johnnies Dam by T. Reardon, A. Reside, A.
Scanlon and H. Parnaby, 2 December, 2007. Dimensions
of the holotype are given in Table 2.
Paratypes: A total of five, field number in brackets: AM
M39815 (7HP50) adult male, skull extracted, body fixed in
80% ethanol and stored in 75% ethanol, captured in a mist
net by T. Reardon, A. Reside, A. Scanlon and H. Parnaby,
2 December, 2007 at Johnnies Dam, Jaurdi Station, 30o
46’S 120o 07’ E. Liver sample stored in 95% ethanol and
held at the Australian Museum, liver sample stored in
liquid nitrogen held at the South Australian Museum; AM
M39801 (7HP46), adult female body fixed in 80% ethanol
and stored in 75% ethanol, skull extracted, captured in
a mist net by T. Reardon, A. Reside, A. Scanlon and H.
Parnaby, 29 November, 2007 at Eagle Rock, approximately
105 km NW of Southern Cross, Goldfields district, WA
30o 26’ 17”S, 118o 40’ 31”E. Liver sample (field number
48170) stored in 95% ethanol held at the Australian
Museum, liver sample stored in liquid nitrogen stored at
the South Australian Museum; AM38843 (WA08), adult
male with skull extracted, AM38844 (WA09), adult male
with skull extracted, and AM38845 (WA10) adult male,
with skull extracted - bodies of all three were fixed in 10%
formalin and stored in 75% ethanol and all three captured
in mist nets by M. Pennay, T. Reardon, A. Reside, and A.
Scanlon, 13 November, 2007, Goongarrie Station, WA,
29o 59.528’S, 121o 03.464’ E. Liver samples of the latter
three specimens are stored in liquid nitrogen held at the
South Australian Museum.
Type locality: Johnnies Dam, Jaurdi Station, 30o 46’
22”S, 120 o 07’ 55”E, 125 km west of Kalgoorlie, Western
Australia. Altitude approximately 435 m.
Diagnosis: It differs from nominotypical N. major in:
smaller average size, e.g. FA for adult females typically
< 44 mm, adult males typically <42 mm; GL < 18.8
mm; CM3 mostly < 7.1 mm; C1–C1 usually < 5.7 mm;
relatively longer ears, and in relatively longer baculum
(Fig. 6). Means of all external, skull and dental dimensions
are smaller, see Table 4. The protocone of M1 and M2 is
often more reduced in N. m. tor subsp. nov., resulting in
a more truncated lingual margin (Fig. 7) and M3 is often
slightly more reduced.
It differs from N. corbeni sp. nov. in: its smaller overall body
and skull size; e.g. adult male mean FA 40.94 mm vs 44.72
mm, mean GL 18.04 mm vs 19.20 mm; skull relatively
narrower and conspicuously less robust (Fig. 3 and Fig. 17,
Fig. 18): ZYG < 11.7 mm vs > 12.2 mm (females), < 11.6
mm vs > 11.9 mm (males); PAL relatively longer (Fig. 4,
Table 4); mean baculum length shorter, 4.38 mm vs 4.97
mm and ≤ 4.6 mm, with proportionately broader base:
mean Baculum Breadth 1.18 mm vs 1.24 mm.
It differs from N. sherrini in: smaller size; skull relatively
broader, with broader zygomatic arches and broader
rostrum; PAL relatively shorter (Fig. 4); INT relatively
narrower; third molars far more reduced: third commissure
of M3 rudimentary and metacone absent (Fig. 7); protocone
on M1 and M2 more reduced resulting in far more
truncated lingual margin (Fig. 7); baculum of equivalent
length but with more slender main shaft.
It differs from N. daedalus in: its darker fur colour;
generally larger size; longer baculum (> 4.0 mm),
narrower skull; relatively larger bullae, narrower
mesopterygoid fossa; basisphenoid pits shallow or
absent; less reduced third molars.
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2009 Australian
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It differs from N. bifax in: grey-brown dorsal fur colour
rather than tawny brown: postnasal elevation relatively
higher rather than a low rounded bump; proportionately
larger skull, GL larger for equivalent FA; relatively larger
bullae; third molars far more reduced, third commissure
of M3 rudimentary rather than being subequal to second
commissure (Fig. 7 and Fig. 21); distal tip of baculum a
simple point or with a weak notch compared to strong
distal bifurcation, baculum length > 4.1 mm vs < 3.9
mm; glans penis with relatively much larger urethral
lappets, and in which the distal tip is a simple rounded
point, rather than being enlarged into a sub-spherical
protrusion as in N. bifax.
It differs from eastern Australian N. gouldi in: relatively
more reduced protocone on M1 and M2 resulting in more
truncated lingual margin (Fig. 7); more reduced third
molars, metacone absent; generally more robust skull;
and a larger more slender baculum shaft; baculum length
> 3.8 mm. Although few specimens of N. gouldi were
available from south-western Western Australia, this
population is distinguished from N. m. tor in: its smaller
overall size; less massive skull; relatively larger bullae;
unreduced third molars in which the metacone is well
Figure 17. X-ray CT scans of the holotype skull of N.
major tor subsp. nov., WAM63601 adult male. Scale bar
represents 10 mm.
Figure 18. Photographs of the skulls of N. corbeni sp. nov.
(C3240; left) and N. m. tor subsp. nov. (WAM22973; right)
showing relatively broader and more robust skull of N.
corbeni. Both are adult males. Scale bar represents 10 mm.
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60 2009
Australian
Zoologist volume 35 (1)
developed. Although ranges of FA for equivalent sex for
adults overlap between both taxa, N. gouldi is clearly a
smaller animal, as reflected by C1–C1: adult females mean
= 4.72 mm (4.6–4.8, n = 4) compared with mean = 5.43,
> 5.0; adult males mean = 4.46 mm (4.3–4.63, n = 5)
compared to mean = 5.38, > 5.0 mm.
It differs from N. nebulosus in: relatively narrower INT;
relatively longer CM3; bullae relatively smaller and set
further apart; more reduced third molars: greater reduction
of third commissure of M3 and metacone absent; baculum
with relatively more slender shaft and longer, > 4.1 mm
vs < 3.0 mm.
It differs from N. heran in having far less developed post-
nasal elevation, which is a rounded mound consisting of
a pair of mounds separated medially by a thin vertical
groove compared with paired mounds joined medially by
a conspicuous membrane that expands distally to form
a “Y” shape; C1–C1 > 5.0 mm vs 4.5 mm; main shaft of
baculum thicker.
It differs from N. geoffroyi in: having a simpler post-nasal
elevation which has a simple median vertical grove, rather
than an more developed pair of mounds joining in the
distal mid-line by an elastic membrane which forms a
distinctive “Y”-shaped structure; by larger average size,
e.g. compared to South Australian and southern Western
Australian N. geoffroyi, adult female mean FA 41.34 mm
vs 36.32 mm (33.6–39.6, n = 48), males 40.94 mm vs
34.87 mm (32.3–37.7, n = 28); having GL > 16.7 mm;
C1–C1 > 4.8 mm, CM3 > 6.1 mm; relatively smaller
bullae; more reduced M1 protocone such that lingual
margin is truncated rather than convex; M3 more reduced
with more rudimentary third commissure and metacone
not present; baculum > 3.8 mm; and distal tip of glans
penis blunt and rounded rather than forming an elongate
“beak”, lacking a distal median dorsal serated ridge; distal
tip of baculum not fully ossified, with very weak notch
compared to solid point; baculum length > 4.1 mm
vs < 2.9 mm (n = 13 for mainland and Tasmanian N.
geoffroyi).
Skull readily distinguished from N. howensis by
conspicuously smaller skull dimensions, more reduced
M3 which lacks a metacone, and as indicated in the
re-diagnosis of that species.
Etymology: a random combination of letters, selected for
brevity.
Distribution: Throughout Western Australia south of the
Hamersley Range and across South Australia as far east as
the Eyre Peninsula (Fig. 8). It appears to be absent from far
south-western Western Australia. In addition to extensive
sympatry with N. geoffroyi, this species is closely parapatric
with N. daedalus in the Hamersley Range of north-western
Western Australia.
Specimens examined: A total of 92, see Appendix.
Remarks: Formal recognition of the smaller morph as a
subspecies of N. major represents a further step toward
clarification of the taxonomy of this group but it is a
compromise, pending a more detailed assessment using
an integrated morphometric and genetic approach. The
relatively small sample available for N. major major has
hindered an assessment of individual variation in that
taxon. Field workers in Western Australia should be alert
to the possibility that the small morph could occur in the
higher rainfall areas of the far south-west.
The type locality of N. major is given as “Perth” (Thomas
1915) and Mahoney and Walton (1988) note that the
collection date on the holotype label is three days after
the collector, Gilbert, returned to Fremantle from the
Houtman Abrolhos. Whittell (1942) notes that little is
known of the collecting itinerary during the time Gilbert
left Perth on a trip overland to Albany, but that the route
went via the settlements of Williams (30 km south-west
of Narrogin) and Kojonup. It is therefore possible that the
holotype of major was collected during the overland trip,
within the geographic range of N. m. tor subsp. nov., as
both taxa occur on the western edge of the wheatbelt. I
have examined high quality photographs of the holotype
skull of N. major but have not had the opportunity to
examine the holotype. The available dimensions of the
holotype of N. major (Table 2) exceed the maximum
recorded dimensions of N. m. tor subsp. nov. for a number
of characters: ZYG of 12 mm exceeds the upper range of
11.6 mm, although the zygomatic arches are broken on
one side, measurements taken from the scaled photograph
suggest that it was at least 12 mm; CM3 of 7.3 mm exceeds
the upper range of 7.1 mm for N. m. tor subsp. nov.
Other available measurements of the holotype of N. major
fall within the overlap of ranges of males between each
taxon for FA, M3-M3 and INT. GL as given in Table 2 falls
within the range for N. m. tor subsp. nov; however, this
measurement is incomplete because the posterior of the
braincase is missing in the holotype (Fig. 2). The holotype
groups within the nominotypical N. major cluster in a plot
of FA vs CM3 (Fig. 19) and also in a plot of FA vs ZYG
(not shown). Consequently, the holotype of N. major is
unlikely to be an example of the small morph, herein
designated as N. major tor subsp. nov.
Figure 19. Plot of CM3 vs FA for adult male N. major major (○)
and N. m. tor subsp. nov. (□) from localities west of longitude
122oE, showing the size of the holotype skull of N. major (H)
relative to seven adult males from Dryandra Woodlands (X)
and the holotype of N. m. tor subsp. nov. (■).
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2009 Australian
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Nyctophilus daedalus Thomas, 1915
Holotype: NHM No. 97.4.12.8, adult male in alcohol
collected by Knut Dahl.
Type Locality: Daly River, Northern Territory.
Re-diagnosis: A moderate to large species, closely
resembling N. major tor subsp. nov. but differing in: paler
fur colour; generally smaller; relatively broader skull;
moderate to deep basisphenoid pits; baculum length <
4.0 mm with a relatively larger proximal end (Fig. 5);
and smaller bullae which are relatively further apart, as
indicated by a plot of BTB against CON (Fig. 20).
It differs from N. bifax in: having a relatively broader
skull; relatively smaller and more reduced third molars
(Fig. 21); the presence of a slight notch on the distal
tip of the baculum which is never deeply bifurcate
as in N. bifax; and a pronounced difference in the
external morphology of the glans penis which has
relatively much larger urethral lappets and lacks the
large rounded distal protuberance present in N. bifax
(Fig. 22).
It differs from N. gouldi in: generally relatively smaller
postnasal prominence; a generally broader and more
robust skull; more reduced protocone on M1 and M2
resulting in truncated rather than strongly convex lingual
margin; far more reduced third molars, metacone absent
and third commissure obsolescent rather than subequal to
second commissure; bullae that average smaller and are
set further apart: the bullae are closer together in N. gouldi
of equivalent BUL (Fig. 23); and distal tip of baculum is
partially ossified rather than a solid ossified point.
Figure 20. Plots of BTB vs CON for N. daedalus (◊) and
N. m. tor subsp. nov. (□), showing the greater BTB in N.
daedalus for equivalent CON. a), females, solid symbols
are females from northwestern Queensland; b), males,
solid symbol is specimen from Mt Bruce.
Figure 21. Scanning electron micrographs of M3 of left, N.
shirleyae sp. nov. (holotype female); centre, N. bifax (AM
M17299, male); and right, N. daedalus (AM M9411, male).
Scale bar represents 0.4 mm.
Figure 22. Scanning electron micrographs showing fronto-
lateral views of the glans penis of a), N. bifax (AM13249);
and b), N. daedalus (AM M34450), showing the much
larger urethral lappets of N. daedalus (indicated by X)
and the sub-spherical distal knob of N. bifax. Scale bar
represents 0.5 mm.
a.
b.
A taxonomic review of Australian Greater Long-eared Bats
62 2009
Australian
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It differs from N. nebulosus in: having paler fur colour;
shorter ears for equivalent FA; skull usually relatively
broader; narrower INT relative to GL; more reduced
protocone on M1 and M2 resulting in truncated rather
than strongly convex lingual margin; far more reduced
third molars, metacone absent from M3 and third
commissure nearly obsolete rather than being well
developed and subequal to second commissure; and
baculum length > 3.1 mm.
It differs from N. arnhemensis in: lighter fur; and other
features as outlined for N. bifax. Nyctophilus daedalus of
the same sex are significantly larger than N. arnhemensis
for most external and cranial dimensions.
It differs from N. heran in: having a less developed post-
nasal prominence; relatively smaller bullae; and main
shaft of baculum thicker distally.
Readily distinguished from N. howensis in skull shape
and smaller cranial dimensions (e.g. GL < 18.3 mm
vs 23.1 mm), and as outlined in the rediagnosis of
that species.
Distribution: Extends from the Hamersley Range
region of Western Australia across northern Northern
Territory to north-western Queensland (Fig. 8).
Distributional limits are Weeli Wolli Springs in the west
and Lawn Hill in the east. Most records are within 300
km of the coast.
Nyctophilus daedalus is evidently sympatric with N. major
tor subsp. nov. in the Hamersley Range in Western
Australia: a single record of N. major tor subsp. nov.
from Mt Bruce is some 70 km west of specimens of N.
daedalus collected at Cadgeput Springs. In northwestern
Queensland, N. daedalus is parapatric with N. bifax. The
most western records of N. bifax are from Cloncurry
(AM2547 and a specimen reported by Thomas 1915)
which is 300 km southeast of Lawn Hill.
Specimens examined: A total of 33, see Appendix. Black
and white photographs of the holotype skull and dentary.
Morphological Variation
Considerable variation exists in overall body size, relative
ear length, degree of development of the post-nasal
swelling, and skull morphology. This variation occurs both
within regions and across the range of the taxon; a more
detailed evaluation will be presented elsewhere.
A trend of increasing body size from the Pilbara region
through to western Queensland is illustrated by a plot
of CON vs longitude (Fig. 24) and FA shows a similar
pattern. However the variation is not a simple size cline,
as demonstrated in a plot of GL vs FA (Fig. 25). The
configuration of specimens in Fig. 25 could be interpreted
in terms of two sexually dimorphic forms. However, group
membership suggested for some specimens in Fig. 25 do not
hold when other characters are examined, although there
is general agreement with the morphs described below. A
reverse trend occurs of decreasing relative ear size (Fig. 26).
Figure 23. Plot of BTB vs BUL showing separation of N.
daedalus (□) from N. gouldi (○). Solid symbols represent
males, open symbols are females.
Figure 25. Plot of GL vs FA for N. daedalus grouped by
sex and putative morph. Localities are P, Pilbara region; K,
Kimberley region; N, Northern Territory, and Q, western
Queensland. Capital letters represent females, lower case
represents male specimens. 1, holotype of N. daedalus, 2,
N. daedalus (AM9411) from the type locality.
Figure 24. Plot of CON vs longitude for N. daedalus
showing trend of increasing size of specimens from the
Pilbara to western Queensland. Adult females, solid
symbols, adult males open symbols.
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2009 Australian
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Broadly concordant patterns of variation occur between
the external characters noted above and the extent of
reduction of M3, relative skull breadth, relative bullar
size and distance between bullae. In broad terms, at least
two forms are evident within N. daedalus, although some
individuals are difficult to allocate. These are:
a. larger bifax-like animals with relatively short ears, a
rudimentary post-nasal swelling, and relatively broader,
more robust skulls with reduced third molars and
relatively smaller bullae that are clearly set further apart
on account of the relatively broader skulls.
b. smaller animals that externally superficially resemble
smaller N. gouldi in the relatively long ears and more
developed post-nasal swelling and less robust skulls.
Although most are from the Pilbara, there is a small
number of specimens from the Kimberley region and
the Northern Territory.
c. a small number of large-bodied animals from the
Kimberley region, the Northern Territory and north-
western Queensland; these are of equivalent size to
larger southern Australian Nyctophilus.
The status of several large female specimens from
western Queensland (Lawn Hill) requires further
clarification and is currently being reviewed. These
specimens differ in several skull and dental features
from specimens from the Northern Territory and it
is unclear whether they represent larger examples of
N. daedalus or a northern variant of a larger southern
taxon such as N. m. tor subsp. nov. The two specimens
from Lawn Hill resemble a pale-furred version of N. m.
tor subsp. nov. in external appearance and fall within
the size range of that taxon for several dimensions, e.g.
C1–C1 (Fig. 27) but they have smaller bullae than N. m.
tor subsp. nov. of equivalent GL. Several large-bodied
female specimens from localities in the Kimberley region
of Western Australia and the Northern Territory also
require investigation but I have not yet examined their
skulls. Koopman (1984) tentatively assigned an adult
female from Port Essington (Northern Territory) to N.
timoriensis timoriensis, believing it to be distinct from
daedalus which he regarded to be a subspecies of N.
gouldi. Measurements of the Port Essington specimen
(NHM 47.7.2.1.1) provided by Koopman (pers. comm.,
1988) for FA (46 mm), condylobasal length (16.6 mm)
and CM3 (6.5 mm) are comparable to those of the Lawn
Hill specimens.
Remarks: There is no doubt that Thomas (1915) was
correct in distinguishing N. daedalus and N. bifax as full
species; indeed, as will be suggested below, it is likely that
each belongs to a separate major clade within the genus.
Pronounced differences exist between the glans penis of
these two species: the urethral lappets are much larger in
N. daedalus, in which there is no trace of the conspicuous
spherical distal swelling of N. bifax (Fig. 22).
Figure 27. Plot of C1–C1 vs FA for N. daedalus (◊) and N.
major tor subsp. nov. (□). (a), adult females, solid symbols
are three specimens from western Queensland; (b), adult
males, solid symbol is subadult from Mt Bruce, Pilbara
region.
Figure 26. Plot of EAR/FA ratio vs longitude for N.
daedalus showing trend of decreasing relative ear size
between specimens from the Pilbara compared to those
of Queensland. Open symbols represent females, solid
symbols, males.
a.
b.
A taxonomic review of Australian Greater Long-eared Bats
64 2009
Australian
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Thomas (1915) listed the simple distal point of the baculum
and relative ear length as the main feature differentiating
N. daedalus from N. bifax, which has relatively longer ears
and an obvious notch in the distal tip of the baculum.
The ten bacula of N. bifax examined in the present study
all have a prominent distal folk. Thomas stated that the
distal tip of the baculum of N. daedalus forms a simple
point as in N. gouldi. In the specimens examined here, the
distal tip is only partly ossified and a small cartilaginous
groove is visible which is similar to some specimens of
N. major, although this is likely to be overlooked in dried
bacula. Although ear length is relatively shorter than N.
bifax in most specimens of N. daedalus from the Northern
Territory, specimens from the Pilbara and the Kimberley
region have relatively long ears, similar to N. bifax and
N. gouldi.
When specimens are pooled from throughout their
geographic range, mensural ranges for all external and
cranial dimensions overlap for each sex between N.
daedalus and N. bifax (Tables 4 and 8). Skulls of N.
daedalus from the Northern Territory and north-western
Queensland differ consistently from N. bifax in the more
reduced M3 and in being generally more robust. Thus
the zygoma and braincase are relatively wider with a
prominent lambdoidal crest. The M3 of N. daedalus, as
well as being smaller relative to M2, has the metacone
and premetacristae reduced to a small ridge, and in this
respect, is similar to that of N. m. tor subsp. nov. (Figs 7
and 21).
A specimen of N. major tor from Mt Bruce indicates
close parapatry between this taxon and N. daedalus
in the Hamersely Range. The closest records of N.
daedalus are from Cadjeput Springs and Weeli Wolli
Springs, both some 70 km to the east. The Mt Bruce
specimen closely resembles an adult male N. daedalus
from Weeli Wooli Springs (WAM18976) in skull shape
but differs in its larger size and in having very shallow
compared to deep basioccipital pits. Though not fully
mature, this specimen is significantly larger in skull
dimensions than four skulls of N. daedalus of both sexes
from adjoining localities (e.g. GL 18.2 mm vs 16.9 -
17.4 mm) and is well within the overall range of male
N. major tor subsp. nov. (17.2 - 18.75 mm, n = 31).
These differences are also evident in a bivariate plot
Table 8. Summary statistics for 11 external and 15 skull and dental dimensions of adult specimens examined of
Australian N. bifax.
N. bifax
Female Male
Mean s.d. Min Max N CV Mean s.d. Min Max N CV
EAR 24.34 1.385 20.8 27.1 55 5.7 23.62 1.544 19.2 26.7 51 6.5
D1 6.58 0.520 5.1 7.8 50 7.9 6.31 0.615 4.4 8.1 48 9.7
FA 42.74 1.282 39.5 46.8 61 3.0 40.93 1.252 37.5 42.8 61 3.1
D31 39.62 1.110 36.4 42.3 54 2.8 38.64 1.212 36.1 40.8 57 3.1
D32 16.01 0.922 11.1 17.3 48 5.8 15.59 0.618 13.6 16.9 49 4.0
D33 14.96 0.725 13.3 16.6 48 4.8 14.74 0.691 13.3 16.3 49 4.7
D51 39.47 1.111 36.4 41.7 51 2.8 38.33 1.151 36.0 40.7 51 3.0
D52 11.03 0.458 9.9 12.3 49 4.2 10.66 0.432 9.6 11.6 48 4.1
D53 10.16 0.829 8.0 11.5 49 8.2 9.78 1.014 7.3 11.5 49 10.4
HL 21.12 0.934 18.9 22.7 50 4.4 20.57 0.943 18.9 22.3 49 4.6
WT 9.24 1.011 7.7 12.0 38 10.9 8.08 1.122 5.0 11.8 37 13.9
CON 15.71 0.402 14.80 16.50 25 2.6 15.30 0.360 14.60 16.20 43 2.3
GL 17.07 0.410 16.30 17.70 25 2.4 16.78 0.374 16.10 17.70 43 2.2
CM36.43 0.184 6.10 6.80 25 2.9 6.25 0.147 6.00 6.60 43 2.4
C1-C14.92 0.191 4.50 5.30 25 3.9 4.80 0.174 4.4 5.20 43 3.6
ZYG 10.76 0.257 10.30 11.37 25 2.4 10.59 0.223 10.20 11.00 43 2.1
INT 3.63 0.159 3.30 4.00 25 4.4 3.58 0.154 3.20 4.00 43 4.3
M3–M37.05 0.236 6.50 7.45 25 3.3 6.87 0.172 6.50 7.20 43 2.5
BRH 6.39 0.184 6.00 6.70 25 2.9 6.34 0.224 6.00 6.80 43 3.5
MAS 9.05 0.235 8.50 9.40 25 2.6 8.88 0.216 8.40 9.50 43 2.4
BTB 2.20 0.110 1.97 2.46 23 5.0 2.16 0.118 1.89 2.46 38 5.5
BUL 3.59 0.095 3.36 3.77 23 2.6 3.52 0.128 3.28 3.85 38 3.6
BAS 5.82 0.227 5.33 6.15 24 3.9 5.66 0.189 5.33 6.15 38 3.3
M3L0.89 0.062 0.83 0.97 6 7.0 0.83 0.022 0.80 0.85 5 2.6
M3B1.95 0.097 1.78 2.05 6 5.0 1.93 0.096 1.82 2.05 5 5.0
PAL 6.55 0.174 6.33 6.86 11 2.7 6.24 0.287 5.70 6.66 9 4.6
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2009 Australian
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of C1–C1 vs FA (Fig. 27). The considerably worn teeth
of some of these specimens of N. daedalus suggests that
their smaller size is not due to age differences. In its
larger size, darker fur colour and shallow basisphenoid
pits the specimen from Mt Bruce contrasts with N.
daedalus from adjoining localities yet resembles N.
major tor subsp. nov.
The recognition of N. daedalus and N. bifax as distinct
species raises the issue of the status of N. arnhemensis.
This taxon closely resembles N. bifax in external, cranial,
penile and bacular morphology but is smaller overall.
However, morphological evidence to be presented
elsewhere suggests that N. arnhemensis is distinct from
N. bifax, and also supports the view of Koopman (1984)
that N. arnhemensis and N. microtis from New Guinea
are separate species. Nyctophilus arnhemensis differs from
N. daedalus in having darker fur colour, often relatively
longer ears, overall smaller size for equivalent sex, and
smaller skull size (GL < 16.5 mm). The glans penis
and baculum of N. arnhemensis resemble those of N.
bifax in the pronounced distal spherical protruberance,
small urethral lappets and conspicuous distal notch in
the baculum.
The presence of a large Nyctophilus species in northern
Australia, comparable in body size to N. timoriensis from
southern Australia, has been overlooked, apart from
Koopman’s tentative identification of a Northern Territory
specimen as N. timoriensis (Koopman 1984).
A clearer diagnosis of N. daedalus will rest on clarification
of the status and relationships of smaller individuals,
particularly those from the Pilbara region.
Nyctophilus sherrini Thomas, 1915
Holotype: NHM no. 52.1.15.50, adult male in alcohol,
collected by Ronald Gunn (Thomas 1915).
Type locality: “Tasmania” (Thomas 1915).
Re-diagnosis: Distinguished from all other members of the
genus by the combination of: large size (compare tables 4,
8-10); unreduced third molars (see Figs 7, 21 and 29);
Table 9. Summary statistics for 11 external and 15 skull and dental dimensions of adult specimens examined of N. sherrini.
W T are field weights taken from Taylor et al. (1987).
Female Male
Mean s.d. Min Max N CV Mean s.d. Min Max N CV CG1985-33
male
N. sherrini
NHM
52.1.15.50
holotype male
EAR 29.00 29.0 1 28.45 1.392 27.2 29.8 4 4.9
D1 6.70 6.7 1 7.20 0.392 6.7 7.6 4 5.4
FA 45.20 45.2 1 45.54 0.940 44.30 46.4 5 2.1 46.3 45
D31 42.80 42.8 1 43.36 1.450 41.3 44.9 5 3.3
D32 16.00 16.0 1 16.28 0.705 15.5 17.0 5 4.3
D33 13.90 13.9 1 14.14 0.513 13.3 14.7 5 3.6
D51 41.40 41.4 1 41.54 1.201 40.1 43.2 5 2.9
D52 11.40 11.4 1 11.36 0.555 10.9 12.1 5 4.9
D53 9.90 9.9 1 9.86 1.178 8.6 11.0 5 11.9
HL 20.00 20.0 1 20.64 0.462 20.1 21.1 5 2.2
WT 13.1 1.5 9.8 14.9 10 12.7 2.3 9.9 18.9 13
CON 17.28 0.263 16.90 17.50 4 1.5 17.12 0.148 16.90 17.30 5 0.9 - 17.2
GL 18.85 0.404 18.30 19.20 4 2.1 18.85 0.152 18.60 19.00 6 0.8 19.0 18.5
CM36.98 0.126 6.80 7.10 4 1.8 6.84 0.184 6.45 7.00 7 2.7 6.95 6.9
C1-C15.35 0.129 5.20 5.50 4 2.4 5.31 0.184 4.95 5.50 7 3.5 5.48 4.7
ZYG 11.13 0.299 10.70 11.40 4 2.7 11.10 0.237 10.70 11.40 6 2.1 11.4 11.4
INT 4.13 0.050 4.10 4.20 4 1.2 4.12 0.117 3.90 4.20 6 2.8 4.2 4.0
M3–M37.55 0.252 7.20 7.80 4 3.3 7.51 0.143 7.30 7.70 6 1.9 7.55 7.1
BRH 6.68 0.126 6.50 6.80 4 1.9 6.68 0.164 6.40 6.80 5 2.5 - 6.35
MAS 9.83 0.171 9.60 10.00 4 1.7 9.82 0.148 9.60 10.00 5 1.5 8.9
BTB 1.83 0.047 1.80 1.89 3 2.6 1.85 0.195 1.56 1.97 4 10.6
BUL 4.16 0.041 4.10 4.18 4 1.0 4.12 0.037 4.10 4.18 5 0.9 4.2
BAS 6.15 0.116 6.07 6.31 4 1.9 6.00 0.107 5.90 6.15 5 1.8
M3L0.95 0.021 0.94 0.98 4 2.2 0.96 0.024 0.94 0.98 3 2.5 0.84
M3B2.14 0.021 2.13 2.17 4 1.0 2.14 0.052 2.09 2.21 4 2.4 2.2
PAL 7.42 0.284 7.20 7.80 4 3.8 7.32 0.117 7.10 7.40 6 1.6 - 7.1
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a relatively narrow skull with unexpanded zygoma and
narrow rostrum, yet with relatively broad temporal region
(see Figs 2 and 28); inflated braincase; and comparatively
large bullae (Fig. 28).
It differs from N. major major and N. m. tor subsp. nov.
in: relatively larger third molars; a proportionately
narrower skull, i.e. relatively less expanded zygomatic
arches and relatively broader intertemporal region; and
greater lateral inflated of the anterior of the braincase.
The baculum shaft is stouter and proximal arms are
relatively shorter. It differs further from N. m. major and
N. corbeni sp. nov. in a relatively much narrower rostrum
and far less robust skull.
It differs from N. gouldi in: larger skull size for equivalent
sex; slightly broader skull with braincase relatively more
expanded; and longer baculum (> 4.0 mm). It is similar
in external appearance and size to larger examples of
southeastern Australian N. gouldi. The skull differs from that
species in: a relatively more inflated braincase; slightly less
expanded zygomatic arches; generally wider interpterygoid
fossa; and relatively greater INT. The baculum resembles that
of N. gouldi but is larger (baculum length > 4.0 mm, n= 3).
It differs from N. nebulosus in: larger in most skull and
dental measurements except BTB, e.g. GL > 18.0 mm;
relatively narrower skull; relatively larger bullae that are set
closer together; and longer baculum, > 4.0 mm (Table 3).
Table 10. Summar y statistics for 13 external and 19 skull and dental dimensions of adult specimens of N. shirleyae sp.
nov.
Mean s.d. Min Max N CV
AM37711
8005 f
holotype
AM37710
8004 f
AM37712
8025 f
NHM
80.498 f**
EAR 25.33 0.058 25.3 25.4 3 0.2 25.3* 25.4* 25.3* 23.9
D1 8.13 0.153 8.0 8.3 3 1.9 8.3 8.0 8.1
FA 46.77 1.252 45.5 48.5 4 2.7 46.6* 45.5* 46.5* 48.5
D31 43.63 0.586 43.2 44.3 3 1.3 43.4 43.2 44.3 -
D32 17.70 0.458 17.3 18.2 3 2.6 17.6 17.3 18.2 -
D33 16.93 0.493 16.6 17.5 3 2.9 16.6 17.5 16.7 -
D51 43.87 0.850 42.9 44.5 3 1.9 44.2 42.9 44.5 -
D52 12.60 0.624 12.1 13.3 3 5.0 12.1 13.3 12.4 -
D53 11.13 0.513 10.7 11.7 3 4.6 10.7 11.0 11.7 -
HL 22.97 0.451 22.6 23.5 3 2.1 22.6* 22.8* 23.5* -
HB 57.67 6.028 52 64 3 10.5 57* 52* 64* -
TL 49.67 2.08 48 52 3 4.2 49* 48* 52* -
WT 12.33 0.764 11.5 13.0 3 6.2 12.5* 11.5* 13.0* -
CON 17.15 0.172 16.9 17.3 4 1.0 16.9 17.2 17.3 17.2
GL 19.03 0.126 18.9 19.2 4 0.7 19.0 19.0 18.9 19.2
CM37.06 0.136 6.9 7.2 4 1.9 7.0 7.1 6.95 7.2
C1-C15.64 0.079 5.5 5.7 4 1.4 5.7 5.5 5.6 5.7
ZYG 11.58 0.153 11.4 11.7 4 1.3 11.4 11.7 11.5 11.7
INT 3.92 0.147 3.8 4.1 4 3.7 3.8 4.0 4.1 3.8
M3–M37.64 0.249 7.4 7.9 4 3.3 7.5 7.4 7.8 7.9
BRH 7.13 0.153 7.0 7.3 3 2.1 7.0 7.1 7.3 -
MAS 9.76 0.304 9.4 10.1 4 3.1 10.1 9.65 9.9 9.4
BTB 2.62 0.153 2.45 2.75 3 5.8 2.65 2.45 2.75 -
BUL 3.79 0.131 3.6 3.9 4 3.5 3.6 3.8 3.85 3.9
BAS 6.23 0.208 6.0 6.4 3 3.3 6.0 6.4 6.3 -
M3L0.87 0.029 0.85 0.9 3 3.3 0.9 0.85 0.85 -
M3B2.08 0.029 2.05 2.1 3 1.4 2.05 2.1 2.1 -
PAL 6.55 0.129 6.4 6.7 4 2.0 6.6 6.7 6.4 6.5
MESO 2.08 0.065 2.0 2.15 4 3.1 2.05 2.0 2.15 2.1
JWL 12.84 0.387 12.45 13.3 4 3.0 12.45 13.0 12.6 -
CM37.59 0.207 7.34 7.8 4 2.7 7.52 7.72 7.34 7.8
M1-M35.07 0.094 5.0 5.18 3 1.9 5.0 5.18 5.04 -
* field measurements; ** measurements from Hill and Pratt (1981).
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It differs from N. geoffroyi in: its larger overall size; reduced
postnasal prominence; the distinctive shape of the glans
penis; larger baculum; narrower skull; and relatively
smaller bullae.
Easily distinguished from N. howensis, which has a much
more thick-set skull which is larger (GL > 20 mm, CM3
> 7.5 mm) and flatter, and has a proportionately much
larger rostrum.
Distribution: Restricted to Tasmania (Fig. 8) where it
is widely distributed, including in the coastal southwest
of the State (Schulz and Kristensen 1996), though with
relatively few records. Taylor et al. (1987) provide a
distribution map for this species (as N. timoriensis), based
on their field work.
Specimens examined: A total of 17, see Appendix. I have
examined black and white photographs of the holotype
skull and dentaries.
Remarks: In the past most authors have associated
N. sherrini with Australian mainland populations of N.
timoriensis, presumably due to its large size. However,
of the mainland Australian species, N. sherrini most
resembles N. gouldi, as implied by Hall and Richards
(1979) and Richards (1983). Although a relatively small
number of specimens of N. sherrini were available for this
study, it is clear that N. sherrini and N. gouldi are distinct
species. Larger adult examples of N. gouldi from Victoria
overlap in FA and C1–C1 with N. sherrini of equivalent
sex. Field workers in Tasmania should consider the
possibility that N. gouldi might also occur in that State.
If so, it is not clear at present how these taxa might be
distinguished using external criteria, though this might
be more evident in live animals than voucher specimens.
Ranges of body weights of Victorian N. gouldi taken in
the field overlap with those of N. sherrini given by Taylor
et al. (1987). Externally N. sherrini is also similar to
Figure 28. Photographs of the skulls of male, left, N. sherrini (AM M34456) from Fortesque Forest, Tasmania; and right,
N. gouldi (C26051) from Mt Eccles, Victoria, showing the more inflated braincase and broader INT of N. sherrini. Scale
bar represents 10 mm.
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68 2009
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larger southeastern Australian N. gouldi. The overall size
of the skull of N. sherrini is larger, with a more inflated
braincase and the interpterygoid fossa is usually relatively
wider. Thus the skull of an adult male N. sherrini (AM
M34456) from Fortesque Bay, while only slightly larger
in most dimensions than a male N. gouldi (MV C26051)
from Mt Eccles, western Victoria (GL 18.8 vs 18.4 mm;
CM3 6.9 vs 6.8 mm; C1–C1 both 5.3 mm; ZYG 11.1 vs
10.5 mm; INT 4.2 vs 4.1 mm; MAS 9.8 vs 9.7 mm;
BRH 6.8 vs 6.5 mm) has a much larger braincase which
is clearly more expanded anteriorly (Fig. 28). In most
cases, the posterior extension of the pterygoids is slightly
greater in N. sherrini. While the braincase is relatively
larger and wider in N. sherrini, the zygomatic arches are
relatively less expanded, resulting in a generally slightly
narrower skull than N. gouldi. INT is relatively broader
in N. sherrini.
Baculum shape is similar in N. sherrini and N. gouldi,
although the proximal end tends to be relatively higher
in N. sherrini (Figs 5 and 6, Table 3) and the baculum is
considerably larger than in N. gouldi (length 4.0-4.5 mm, n
= 3 vs mean = 3.26, 3.0 - 3.7, n = 26). The glans penis of
N. sherrini is far narrower than that of N. gouldi, being more
compressed laterally in the three specimens examined.
Nyctophilus howensis McKean, 1975
Holotype: ANWC CM4724, cranium with periotic bones
and dentaries missing, collected by G. F. van Tets. The
skull was found on a rock ledge on the cave wall, but
post-cranial material was not found with the skull (G. F.
van Tets, pers. comm.).
Type locality: Lord Howe Island, “cave at north end of
Island, north east of North Bay Beach” (McKean 1975).
The skull was found on a mezzanine ledge in Goosebury
Cave (Van Tets, quoted in Richards and Hall 1999). A
label associated with the type skull notes “cave entrance
in vine-covered opening in forest”.
Re-diagnosis: Evidently a large bat, as judged by cranial
dimensions (see Tables 4, 8-10; Fig. 29). Skull is largest
recorded for the genus, compared with maximum
measurements of the next largest species, N. major and N.
corbeni sp. nov.: GL 23.1 mm vs 20.8; ZYG 13.9 mm vs
13.3; CM3 8.1 mm vs 7.8; C1–C1 6.7 (from alveoli) vs 6.5
(from cingula); PAL 9.4 vs 7.7. Lateral profile of skull is
low, unlike any other large member of the genus.
It differs from other large species of the genus, viz, N.
corbeni sp. nov., N. major major, and N. sherrini in:
ant-orbital foramina being relatively much narrower
and smaller; relatively much smaller anterior palatal
emargination and narrower rostral sulcus; interdental
palate relatively broader and shallower; and interpterygoid
fossa width similar in absolute size but relatively much
narrower due to larger skull size.
It further differs from N. m. major, N. corbeni sp. nov., N.
m. tor subsp. nov. and N. daedalus in its less reduced M3,
and further differs from N. corbeni and N. daedalus by a
relatively much longer palate.
It further differs from N. sherrini by a greater reduction of
M3 and a relatively broader rostrum.
Distribution: known only from the holotype skull from
Lord Howe Island.
Measurements of holotype (mm): GL, 23.09;
CON, 21.26; ZYG, 13.88; INT, 4.23; Anterior palatal
emargination, diameter, 1.9; Ant-orbital breadth, 6.93;
Incisor socket, diameter, 1.27; Canine socket diameter,
1.78; Outer width C1–C1 (alv.), 6.71; Inner width C1–C1
(alv.), 3.56; CM3 right side, alv., 8.1; Outer width PM4-
PM4 alv., 7.59; Inner width PM4-PM4 (alv.), 4.48; Outer
breadth M3-M3 (alv.), 8.71; Inner breadth M3-M3 (alv.),
4.34; Glenoid fossa breadth, 2.91; BRH, 7.40; Braincase
breadth, 9.76; MAS (incomplete, estimated), 11.3; right
side socket for auditory capsule, maximum length, 5.2;
maximum width, 4.76; Minimum width of basi-sphenoid
between sockets of auditory capsules, 1.5; BAS, 7.55;
PAL, 9.46; Mesopterygoid fossa width at root of hamular
process, 2.72; Mesopterygoid fossa width at posterior
base, 2.51; Foramen magnum breadth, 4.24. McKean
Figure 29. Photographs of the holotype skull of N.
howensis (ANWC CM4724), sex unknown. Scale bar
represents 10 mm.
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2009 Australian
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(1975) gives other measurements for dentition. (NB:
foramen magnum breadth, braincase breadth, MAS
and auditory capsule socket length and breadth were
taken in 1991. The right occipital condyle and an
adjoining section of the cranial vault, including that
forming the border of the auditory capsule socket, is
now missing.)
Material examined: the holotype skull.
Remarks: No further material of this species appears to
have been reported since its description. This species
is clearly not conspecific with any known species of
the genus. The general size of the holotype skull is
much larger than the largest specimens examined of
N. corbeni and N. major - the largest of the extant
species of Nyctophilus. Although GL of the holotype of
N. howensis is only a few mm greater than the largest
skull of N. corbeni sp. nov. (20.8 mm), the skull of the
latter is considerably smaller in overall appearance
than N. howensis. The general form of the skull is more
gracile than in N. corbeni, and superficially resembles
that of N. sherrini; no close relationship with the latter
taxon is suggested.
The overall morphology of the holotype skull superficially
resembles that of large species of Nyctophilus. A single
large upper incisor socket, and no trace of a socket in the
narrow gap between it and the canine alveoli indicates that
the specimen has a single upper incisor, as indicated in the
original description. Compared to large Nyctophilus, the
skull of N. howensis has a longer palate (Fig. 30a) as noted
by McKean, but a comparatively short tooth row (Fig. 30b).
The skull is narrow (Fig. 30c) and remarkably flat (Fig. 30d).
McKean stated that the palate is much broader than any
species of Nyctophilus. The rostral sulcus is smaller than in
other species of Nyctophilus and terminates less posteriorly,
as does the anterior palatal emargination, which is also
narrower and has an evenly rounded posterior margin.
The morphology of the premolars and molars broadly
resembles that of other species of Nyctophilus. The
shape of M2 differs from that of N. major and N. corbeni
sp. nov. in having both the anterior and posterior sides
of the tooth straight. M3 is reduced, but the second
and third commissures are present and subequal, and
although nearly worn flat, it is evident that a reduced
metacone is present.
Figure 30. Plots of selected measurements against GL of the holotype skull of N. howensis and large species of Nyctophilus
of either sex, illustrating that N. howensis has: a), a long palate; b), a short tooth row; c), a narrow skull; and d), a relatively
low braincase. Species symbols are: N. howensis (♦), N. corbeni sp. nov. (), N. major major (●), N. m. tor subsp. nov. (■),
N. sherrini (∇), and N. shirleyae sp. nov. (▼).
a.
c.
b.
d.
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The generic status of howensis warrants a more detailed
reassessment that is possible here. McKean expressed
reservation about placing the taxon in Nyctophilus, and
I concur. There appears to be no specific reason for
assigning the holotype to Nyctophilus, other than its
superficial resemblance in dental and cranial structure
compared to any other genera in the Australian region.
The genus Nyctophilus is defined by a combination of an
abruptly truncated snout with a low horse-shoe shaped
narial nose-leaf, a variably developed fleshy postnasal
mound, one upper incisor and premolar, ears joined in the
midline (except for N. microtis) and a relatively slender
baculum (Miller 1907; Hill and Harrison 1987). Of these
criteria, only the single upper incisor can be confirmed
for N. howensis. McKean noted the presence of well
developed basioccipital depressions which he considered
to be characteristic of Nyctophilus and the allied genus
Pharotis. Basioccipital pits occur in a range of vespertilionid
genera (DeBaeremaker and Fenton 2003). There appears
to be no convincing evidence that the holotype had
either a nose-leaf or large ears. McKean interpreted
the presence of a rostral depression in the holotype as
indicative of a moderately developed nose-leaf which he
speculated was possibly of similar development to that of
Nyctophilus timoriensis. This would have been a reference
to the secondary nose-leaf which forms part of a postnasal
mound in some species of Nyctophilus. However, a wide
range of vespertilionid genera contain species that lack
any form of nose-leaf or postnasal mound, yet have similar
or more developed rostral depressions. Both auditory
capsules are missing from the holotype and there appears
to be no means of establishing ear size of the holotype.
In conclusion, there is little evidence that howensis was a
long-eared bat on the basis of skull morphology.
Nyctophilus shirleyae sp. nov.
Holotype: Australian Museum number M37711 (field
number 8005), adult female, body fixed in 10% formalin
and stored in 75% ethanol, skull extracted. Field numbers
for tissue samples stored in liquid nitrogen at the Australian
Museum are: liver (8005L), kidney (8005K) and heart
(8005H). Collected by H. Parnaby in a mist net on Mt
Missim, 8 July 1988. Measurements of the holotype are
given in Table 10.
Paratypes: Australian Museum number M37710 (field
number 8004) adult female, body fixed in 10% formalin
and stored in 75% ethanol, skull extracted and in good
condition. Captured in a mist net by H. Parnaby on 8 July
1988, at the type locality on Mt Missim. Field numbers for
tissue samples stored in liquid nitrogen at the Australian
Museum are: liver (8004L), kidney (8004K) and heart
(8004H). Australian Museum number M37712 (field
number 8025), adult female with regressed teats, body
fixed in 10% formalin and stored in 75% ethanol, skull
extracted. Captured in a mist net by H. Parnaby on 11
July 1988 on the southwestern slopes of Mt Missim, Kuper
Range, PNG: the site (17o 15’ S, 146o 47’ E) was of higher
altitude than the type locality.
Referred specimen: Natural History Museum number
80.498, adult female in alcohol, skull separate, Mt
Missim.
Type locality: Southwestern slopes of Mt Missim, Kuper
Range, Morobe Province, Papua New Guinea, 17o 16’
S, 146o 46’ E. The holotype and paratype females were
captured in mist nets in mature montane rainforest at
an approximate altitude of 1600-1800 m. The exact
location and altitude of the type locality and the location
of the higher altitude collection site for the paratype
M37712 could not be determined but are within about
a km radius of the co-ordinates given above. The higher
altitude site was close to the main trail leading up the
southwestern slopes to the summit of Mt Missim, but the
holotype and paratype M37710 were collected on a spur
northwest of the main trail.
Diagnosis: Distinguished from all other Nyctophilus by
the combination of: reduced postnasal prominence (Fig.
31); large skull size (GL for females ≥ 18.9 mm) (Fig. 32
and 33); moderately reduced third molars (Fig. 21); bullae
relatively small and set comparatively far apart (BTB >
2.5 mm); and bullae more reduced relative to periotic
bone exposing a larger proportion of periotic bone.
It differs from N. major major, N. m. tor subsp. nov. and
N. corbeni sp. nov. in: less reduced third molars in which
the metacone is clearly present; smaller bullae (BUL of
adult females < 4.0 mm) which are relatively further apart
(BTB > 2.5 mm). It further differs from N. major major
and N. m. tor in relatively shorter palate (Fig. 4).
It differs from N. sherrini in: relatively broader and more
massive rostrum with broader zygomatic archers; relatively
smaller INT; posterior extension of the palate relatively
shorter; mesopterygoid fossa relatively broader; tympanic
bulla relatively much smaller and also absolutely smaller,
BUL < 4.0 mm (Tables 9 and 10); and M3 substantially
more reduced (compare Figs 7 and 21).
It differs from N. daedalus in: relatively narrower skull;
narrower ZYG, C1–C1, INT and MAS relative to skull
length; a relatively shorter palate (Fig. 4); and considerably
less reduced third molars (Fig. 21).
It differs from N. nebulosus in: relatively lower post-nasal
elevation; in being larger: FA > 45 mm (n = 4) vs < 44 mm
(n = 3); larger skull: GL ≥ 18.9 mm; C1–C1 > 5.5 mm vs
4.9–5.0 (n = 2); GL much larger relative to FA; a relatively
shorter palate; mesopterygoid fossa relatively narrower; INT
relatively much narrower; bullae relatively smaller; and in
having substantially greater reduction of third molars.
It differs from N. gouldi in: relatively lower postnasal
prominence; relatively broader skull with a proportionately
larger braincase, e.g. BRH for females 7.0 mm and
greater, vs mean = 6.19, 5.9–6.6 (n = 42 for the largest
populations of female N. gouldi which occur in montane
New South Wales and Victoria); proportionately much
smaller bullae which are set further apart e.g. BTB ≥ 2.45
mm vs mean = 1.71, 1.4–2.0 (n = 37).
Easily distinguished from N. geoffroyi which is smaller,
e.g. maximum FA for female N. geoffroyi from population
of largest individuals (Tasmania, n = 13) 41.7 mm vs
minimum of 45.6 mm for N. shirleyae sp. nov.; N. geoffroyi
has a more developed postnasal elevation with a well
developed median membrane joining each prominence
that is most developed distally; grey-white tips to ventral
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2009 Australian
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fur; relatively much larger bullae, and is smaller than N.
shirleyae sp. nov. in cranial dimensions except BUL: e.g.
compared to maximum dimensions of populations of
the largest female N. geoffroyi (from Tasmania, n = 14):
maximum GL 17.1 mm vs minimum of 18.9 mm; CM3 6.0
mm vs 6.9 mm; C1–C1 4.8 mm vs 5.5 mm.
Figure 31. Photographs of the adult female paratype
(AM37710) of N. shirleyae sp. nov. from Mt Missim, Papua
New Guinea, taken by H. Parnaby, 8 July, 1988.
Figure 32. X-ray CT scans of the adult female holotype
skull and dentary (AM37711) of N. shirleyae sp. nov. from
Mt Missim, Papua New Guinea. Scale bar represents 10
mm.
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72 2009
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Easily distinguished from N. heran in: much darker
ventral fur; far less developed post-nasal elevation, which
is a low rounded mound consisting of a pair of mounds
separated medially by a thin vertical groove compared
with paired mounds joined medially by a conspicuous
membrane that expands distally to form a “Y” shape;
being much larger for most dimensions (see Table 10);
e.g. FA > 45 mm vs 39.3 mm, GL > 18.9 mm vs 16.7
mm (n = 1), C1–C1 > 5.5 mm vs 4.5 mm; and relatively
much smaller bullae – BUL = 3.6–3.9 mm v 3.9 mm.
It differs from Australian populations of N. bifax in:
larger size for most skull dimensions (Tables 8 and 10;
Fig. 33), e.g. adult females have GL > 18.0 mm, C1–C1
> 5.4 mm; palate shorter relative to GL (PAL/GL <
0.355); GL larger relative to FA (Fig. 34) third molars
more reduced: M3 more reduced than N. bifax, second
and third commissures shorter relative to first (Fig. 21);
and bullae relatively smaller and less developed than
N. bifax such that a much greater proportion of periotic
bone is exposed.
Readily distinguished from N. microtis by external
features: ears relatively longer and joined at their base
by an obvious median membrane, compared with N.
microtis in which the median membrane is either absent
or scarcely visible above the fur; anterior margin of
tragus strongly convex rather than straight or weakly
convex; larger overall size, e.g. field body weights
of adult females > 11 gm vs 9 gm or less (n = 6);
dimensions of adult females: FA > 44 mm compared
to mean 39.91 mm (38.5–42.7, n = 10), Ear Length >
24 mm, compared to 17.50 (14.9–19.5, n = 7); skull
larger: GL > 18.9 mm, compared to mean 15.34 mm
(14.5–16.3, n = 8); C1–C1 > 5.4 mm compared to 5.0
mm or less (n = 8); CM3 > 6.9 mm vs mean 5.78 mm
(5.4–6.1, n = 8).
Readily distinguished from N. walkeri by much larger
overall size, e.g. FA > 37 mm; adult female WT > 10.0
gm; GL > 14.0 mm; C1–C1 > 4.5 mm; Ear Length >
16.5 mm and ears relatively much larger; anterior margin
of tragus convex as is typical of the genus, rather than
straight or weakly concave as in N. walkeri.
Easily distinguished in the field from N. microdon which
has smaller body size: e.g. FA > 45 mm vs < 42 mm,
C1–C1 > 5.4 mm vs < 4.4 mm; postnasal mound low
and rounded compared to two well developed mounds
joined in the midline by an obvious elastic membrane; a
much smaller tragus relative to ear size; tragus relatively
narrower and distal end of tragus rounded rather than
truncate; dorsal and ventral body fur grey-brown rather
than the rich red-brown of N. microdon.
Readily distinguished from N. howensis by skull shape and
size (the far less inflated cranium of N. howensis results in a
nearly linear lateral skull profile); a relatively much larger
rostrum; skull relatively more elongate; with relatively
longer palate and is larger, e.g. GL 23.1 mm compared
to < maximum of 19.2; CM3 is 8.1 mm compared to a
maximum of 7.2 mm.
Etymology: I name this long-eared bat after my mother,
Shirley Jean Parnaby (nee Slade), a great admirer of
the people of the Papua New Guinea nation and its
biodiversity, and who encouraged my childhood interest
in mammals.
Remarks: This is the largest of the four species of
Nyctophilus known from New Guinea. It is immediately
distinguished in the field from N. microtis, which has
relatively short ears which lack an obvious membrane
that connects the base of the ears; has a narrower tragus,
the anterior margin of which is either straight or slightly
convex in the midline, rather than being strongly convex;
and is conspicuously smaller. It is also easily recognized
from each of N. microdon and Pharotis imogene which are
smaller in body size, e.g. FA < 42 mm vs > 45 mm; C1–C1
< 4.4 mm vs > 5.4 mm, and both of which have very
large ears and large tragi, relative to head size.
Figure 33. Photographs of the dorsal view of the skulls of
left, adult female N. shirleyae sp. nov. (AM37710), showing
the larger size and braincase compared to a large female
N. bifax (CM4508) from Atherton Tableland, Queensland.
Scale bar represents 10 mm.
Figure 34. Plot of GL vs FA for adult female specimens of
N. shirleyae sp. nov. (●) and N. bifax (■). 1, adult female
cf. N. bifax (BBM-NG 60073) from Port Moresby, Papua
New Guinea; 2, adult female N. bifax (CM4508) from
Atherton Tableland, Queensland.
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2009 Australian
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In general size and external appearance N. shirleyae
sp. nov. resembles a large version of N. bifax. In body
size, this species is about the same size as the largest
Australian female N. bifax that I have examined, but has
a relatively much larger skull. Thus although FA lengths
overlap, the two species clearly separate on skull size, as
illustrated in a plot of GL vs FA (Fig. 34).
Several specimens from New Guinea have been referred
to N. bifax. Thomas (1922) considered that an adult
female (which I have not examined), unfortunately
without adequate locality data, compared well with N.
bifax from Queensland. Tate (1952) regarded a specimen
(AMNH 152462) from Idenburg River, northeast West
Papua, to be indistinguishable from a series of N. bifax
from Cape York Peninsula, Australia. Koopman (1982)
considered this specimen and another from the Fly River
to be quite similar to N. bifax from north Queensland,
noting that they are at the larger end of the size range.
Although the Idenburg River specimen resembles N.
bifax in skull shape and size (e.g. GL 17.2 mm vs
16.1–17.7 for 43 males) the ears are decidedly smaller
than any Australian N. bifax that I have examined. The
taxonomic status of this specimen is unclear. An adult
female (BBM-NG 60073, skin and skull) from Brown
River Forestry Station, Central Province, Papua New
Guinea resembles N. bifax from northern Australia in
the relatively long ears and in general skull shape but
the skull is larger (GL 18.1 mm vs 16.3–17.7 for 25
adult females). The latter two specimens could well be
at least subspecifically distinct from Australian N. bifax.
Neither appears to represent N. shirleyae sp. nov.. Other
material is assigned to N. bifax by Flannery (1995a) and
Bonaccorso (1998).
Other than for its large size, N. shirleyae would appear
to have little in common with the N. major complex
or N. sherrini. It could be most closely related either to
the bifax or microtis species groups, as defined below.
The morphology of the glans penis and baculum has
not been reported, but these are likely to be highly
informative regarding its interspecific relationships.
I have not located the adult male reported by Hill
and Pratt (1981). Thane Pratt (pers. comm. 2005) has
suggested that this specimen is likely to be lodged either
with the Wau Ecology Institute, PNG, or in the Papua
New Guinea National Museum. If the relationships
of N. shirleyae sp. nov. lie with N. bifax or N. microtis,
as suggested by skull and dental morphology, it is
likely that penile morphology would consist of a pair
of relatively small, narrow urethral lappets and a
subspherical distal nob, which is the broad shape for
both N. bifax and N. microtis.
Although currently only known from Mt Missim, it is
likely that N. shirleyae has a wider distribution within
Papua New Guinea, particulary given that bat surveys
have not been undertaken in many area. Preliminary
examination of specimens recently obtained from the
low elevations of the Fly River region, Western Province
by Steve Hamilton (pers. comm., University of New
South Wales) indicates a close resemblance with N.
shirleyae and will be reported elsewhere.
Interspecific relationships within
Nyctophilus
The primary assessment of interspecific relationships
within Nyctophilus is that of Tate (1941) who recognised
four species groups:
a. timoriensis group, including major, sherrini and gouldi;
b. bifax group with bifax and daedalus;
c. geoffroyi group with australis, pacificus, unicolor
and pallescens characterized by a highly developed
postnasal elevation;
d. microtis group, including microtis, bicolor and walkeri.
Tate did not specifically define the characters for each
group. He noted the distinctiveness of N. walkeri but
tentatively placed it within his microtis group which he
considered to be the most primitive species group. Tate
reserved judgement about the taxonomic status of most
taxa, many of which were known from few specimens,
thus preventing any useful evaluation of within-species
variation.
Interspecific relationships of the taxa examined in
this study require more detailed examination than is
possible here. However, I propose the following tentative
arrangement based on an extensive unpublished
examination of external features, skull and dentition,
and external morphology of the glans penis:
a. A major group, consisting of major, m. tor subsp.
nov., corbeni sp. nov. and possibly daedalus. This
group has the most extreme reduction of the third
molars, and broadly similar external morphology
of the glans penis, i.e. comparatively large urethral
lappets, the distal portion is simple and lacks any
protrusions. The relationships of N. daedalus are
unclear; it is provisionally included in this group
though in some respects it resembles the gouldi
group. Part of the difficulty could be due to daedalus
being a composite species.
b. gouldi group consisting of three geographic forms
of gouldi: far south-western Western Australia,
inland southeastern Australia and montane and
subcoastal eastern Australia; sherrini and nebulosus
are also tentatively included in this group. All have
unreduced third molars, and a thick baculum shaft
with a solid distal point, (baculum morphology of
the south-western Western Australian form of gouldi
has not been examined in this study) and unadorned
glans penis morphology similar to the major group.
The status of smaller animals from the Pilbara that
are currently included with daedalus requires further
consideration, particularly in relation to eastern
Australian inland form of N. gouldi;
c. howensis, which does not form part of the major
complex, and differs from all other members of the
genus in cranial characters;
d. microtis group, with the distinctive walkeri tentatively
associated. Both have unreduced third molars, small
bullae, relatively very short ears and a linear or only
A taxonomic review of Australian Greater Long-eared Bats
74 2009
Australian
Zoologist volume 35 (1)
slightly convex anterior tragus margin; and a distinct
distal notch in the baculum;
e. bifax group, which includes arnhemensis – both taxa
share small bullae, unreduced third molars and distal
baculum bifurcation, and similar penile morphology
with the microtis group, but have a less specialized
postnasal prominence than microtis and walkeri and
the ears are joined medially by a distinct membrane.
The relationships of N. shirleyae sp. nov. remain
unresolved but it is provisionally placed with bifax
which it most closely resembles. The bifax group
might belong with the microtis group.
f. microdon – this highly distinctive species differs
from all other described species of the genus in the
enlarged tragus, distinctive morphology of the glans
penis, baculum, and in a number of cranial and dental
features. There is no support for the suggestion of
Koopman (1984) that microdon is closely related to,
but more primitive than, N. geoffroyi;
g. geoffroyi group: previous authors have synonymised
australis, pacificus, unicolor and pallescens but I have
not attempted an evaluation of the status of these
forms. Differs in the unique serrated longitudinal
dorsal ridge on the distal portion of the glans
penis, highly developed snout mound posterior to
the noseleaf, and relatively inflated bullae. The
affinities of N. heran require clarification, athough
it is clearly a distinct species from any of the named
forms of geoffroyi. Kitchener et al. (1991) compared
N. heran with N. geoffroyi, and I have tentatively
placed it with this group though it differs in penile
morphology, which more closely resembles the gouldi
group and N. daedalus.
I have examined external morphology of the glans penis
of all currently recognized species except N. shirleyae
sp. nov. and N. heran (described and illustrated by
Kitchener et al. 1991), and N. microtis bicolor which is
known only from the holotype from Papua New Guinea.
My detailed observations on penile morphology will
be published separately; however, they suggest the
presence of three main groups within Nyctophilus:
a. group with large paired urethral lappets, and which
lack a pronounced terminal subspherical structure.
This includes the major and gouldi groups defined
above and N. daedalus, and N. heran. It is likely that
N. geoffroyi also belongs within this clade, though
this species complex has a distinctive modification
unique in the genus;
b. group in which the paired urethral lappets are
relatively much smaller and more elongate than the
above clade, and in which a subspherical distal nob
is usually present. This includes the microtis and bifax
groups and N. walkeri is tentatively placed in this
group; and
c. group which has very small, elongate urethral lappets
and an entirely different distal structure to either of
the above clades. The two species of this group are
N. microdon and an unnamed species from Papua
New Guinea.
A cladistic analysis based on morphological characters
is hindered by inadequate knowledge of intraspecific
variation, poorly defined species boundaries in some
groups (e.g. the geoffroyi and gouldi groups and N.
daedalus) and uncertainty over appropriate outgroup
comparsions. While it is acknowledged that the species
groups recognized here are primarly phenetic and
may be based as much on shared primitiveness as
on synapomorphy, they are considered a useful step
such for a confused and poorly understood genus.
A collaborative study with a team led by Belinda
Appleton (University of Melbourne) is in progress,
in which comparative morphological work will be
integrated with a molecular phylogeny of the genus.
Discussion
The central aim of this paper is clarification of the
taxa that comprise what has hitherto been referred
to as N. timoriensis. What has long been regarded as a
single widespread species, N. timoriensis, is here shown
to represent five taxa, at least four of which are full
species: N. major (including N. m. tor subsp. nov.), N.
corbeni sp. nov., N. sherrini, and N. shirleyae sp. nov. In
order to clarify the status of timoriensis, it was necessary
to evaluate variation within a further six taxa, viz. N.
gouldi, N. daedalus, N. bifax, N. arnhemensis, N. heran,
and N. howensis.
The extent of variation within some taxa is considerable,
particularly in N. daedalus and N. gouldi. A more
refined diagnosis of all of these taxa must await a more
thorough evaluation of morphological variation, which
will be greatly assisted by further collection of material
from strategic geographic regions, and the application
of molecular analyses. Selection of reliable criteria for
field identification is currently hindered by a lack of
understanding of within-taxon variation.
Considerable variation within N. gouldi was discernable
during the course of this study, which will be reported
elsewhere. The small size of the inland N. gouldi in
eastern Australia has been previously recognized (e.g.
Parnaby 1987; Lumsden 1994) and small individuals
from Queensland were recognized as being different
from montane southeastern Australian N. gouldi by
Churchill et al. (1984), who regarded it as a separate
unnamed species. Clarification of the status of these
forms will greatly facilitate the diagnosis of N. gouldi
from N. corbeni and N. sherrini, and the reassessment
of N. daedalus.
The considerable morphological variation within N.
daedalus suggests that this is a composite of two, and
possibly three distinct forms. Furthermore, if more than
one taxon is currently included within N. daedalus,
they are most likely to be broadly sympatric throughout
the current range of N. daedalus – a critical issue for
field workers trying to identify Nyctophilus in northern
Australia. A morphological assessment of variation
in N. daedalus is in progress. Resolution of variation
within N. daedalus is further necessary to clarify its
diagnosis relative to N. gouldi, N. bifax, N. m. tor, N.
nebulosus and N. heran.
Parnaby
75
2009 Australian
Zoologist volume 35 (1)
Description of the new species and subspecies taxa
in this paper represents a step towards resolving
the number of species and their diagnosis within
Nyctophilus. However, the determination of species
within this complex genus should be considered a work
in progress. A substantial number of issues require
resolution before it can be confidently assumed that
the majority of taxa have been recognized, let alone
adequately diagnosed.
Geographic regions for further strategic
collecting
A number of geographic areas can be identified in which
further strategic collecting is required to further clarify
species limits of the taxa covered in this paper:
1. Additional material is required to define the extent
of infra-specific variation within N. major from
the far south-west of Western Australia, as it is
known from relatively few voucher specimens and
basic data such as body weights are not available.
In particular, further work is needed in areas of
potential sympatry between N. major major and N.
m. tor, such as the wheatbelt region of south-western
Western Australia (e.g. the Katanning and Narrogin
districts), the Roe Plain and Madura districts and
surrounding region south of the Nullarbor and the
Balladonia district.
2. Given the considerable variation within populations
currently referred to N. daedalus, it is important to
target the entire range of that taxon: the Pilbara,
Kimberley, the northern Northern Territory, and
northern inland Queensland. Field workers active
across that entire region should be alert to any
Nyctophilus that is not obviously N. walkeri or
N. geoffroyi – I also anticipate difficulties with
remaining species including N. arnhemensis.
3. A transition zone between the smaller inland form
of N. gouldi and the larger montane and subcoastal
form of N. gouldi should be examined to determine
the relationships of these morphologically distinct
populations; the smaller inland form extends from
northern Victoria to northern Queensland.
4. Further survey work is required in Tasmania, where
forest environments are currently undergoing
accelerating and already severe degradation from
clear-cut logging operations. This is necessary, both
to obtain more material of N. sherrini, which is very
poorly represented in world research collections,
and to determine whether N. gouldi also occurs in
Tasmania. If the latter species does occur there,
it is likely have been confused with N. sherrini
in the past, particularly because identification of
Nyctophilus species would most likely have been
based on the presence or absence of a distinctive
Y-shaped groove on the post-nasal bump, which is
characteristic of N. geoffroyi, and general body size.
5. Extensive survey work is needed in Papua New
Guinea and West Papua, and more widely in
eastern Indonesia, both in rainforest and in eucalypt
savannahs. Few specimens exist of N. shirleyae and
the status of the small number of specimens assigned
to N. bifax from those regions needs clarification.
6. Efforts should be made to obtain further material of
N. howensis, including post-cranial material, from
cave deposits on Lord Howe Island.
Field workers in all regions should anticipate ongoing
difficulties in identifying Nyctophilus – reliable field
criteria cannot be derived until species diagnoses are
refined, which in turn requires further collecting of the
many poorly represented taxa to assess intraspecific
morphological variation. Unfortunately, recognition of
the imperative to retain voucher specimens has declined
in Australia over the past decade or so. There has
been a significant increase in field work but a decrease
in the number of voucher specimens being lodged in
museums. This has probably arisen from a combination
of factors, one being a failure to appreciate that species
taxonomy is unrefined for many taxa. It is generally
assumed that taxonomic confusion is confined to a
minority of taxa, whereas the reality is that many genera
of Australian mammals (and other vertebrates) remain
poorly resolved.
Management implications
The results of this study provide yet another reminder
of the imperative of a refined understanding of species
taxonomy and the implications of species taxonomy
for effective conservation management strategies –
a recurrent theme in the literature of Australian
mammals. For example, one of the most intensively
studied Australian mammals, the small dasyurid
Antechinus stuartii, was shown to consist of four
largely allopatric species with restricted distributions
(Dickman et al.1998; Van Dyck and Crowther 2000).
Many other examples could be cited for Australian
mammals and many more can be anticipated in the
coming decade.
Despite the implications of unrecognized species for
effective conservation management, and consequently
the obvious relevance of taxonomic studies, species
taxonomy still appears to be perceived as either an
irrelevancy, a low priority by managers and funding
bodies alike, or as the domain of academic research
rather than management, i.e. someone else’s problem.
Paradoxically, academia itself tends to view taxonomy
as applied science, at best, and unworthy of pursuit
or reward. In my opinion, reasons for the neglect by
managers of something as fundamental as species
taxonomy for a high profile group like mammals, should
be sought in the ideological, political and social arenas.
Consequently, the taxonomic impediment of confused
species limits of Australian Microchiroptera discussed,
for example, by Wood Jones (1925), Frith (1973),
Hamilton-Smith (1974), Parnaby (1991) Richards
and Hall (1998) and Reardon (1999), still remain
substantially unresolved.
A taxonomic review of Australian Greater Long-eared Bats
76 2009
Australian
Zoologist volume 35 (1)
Acknowledgements
The initial stages of this research were undertaken as
a Ph.D under the patient guidance of Mike Archer,
School of Biological Sciences, University of New
South Wales. I am grateful for the support from staff
at the Australian Museum over the long duration
of this study: especially to Sandy Ingleby, Mammal
Section for endless assistance, Tish Ennis for French
translations; Carl Bento (Photography) for skull photos
and guidance with PhotoShop; Sue Lindsay SEM
Unit, for electron microscopy, and David McAlpine,
Entomology Department, for advice on nomenclature.
Angela Frost kindly assisted with photography. Thanks
to: Sue Hand who took notes on specimens in Paris,
Pat Woolley for inspiring discussions and providing
her notes on Mt Missim, and Thane Pratt for advice on
the location of specimens. Glenn Hoye (Fly by Night
Consultancies) shared his measurements and photographs
of type specimens in the British Museum. Thanks to the
substantial time commitment from Fred Ford (Sustainable
Ecosystems, CSIRO) for the CT scans, and 3D models of
skulls and bacula. The manuscript was greatly improved by
input from Ken Aplin (Sustainable Ecosystems, CSIRO),
Kris Helgen (Smithsonian Institute, Washington) and
Dan Lunney and Mathew Crowther (Department of
Environment, Climate Change and Water, Hurstville) and
Norm McKenzie (DEC, Wanneroo): an arduous task, much
appreciated. This study was partly funded through a grant
to the author from the Estate of Winifred Violet Scott.
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APPENDIX 1
Registration number NLocality State Lat. Long.
Nyctophilus corbeni sp. nov. (n=64)
AM5282, AM5515 2“Calumet” 26 ml N of Binnaway NSW 31 17 149 44
AM36753–55 3Arakoola Nature Reser ve NSW 29 18 150 48
AM36635 1Arthurs Seat State Forest NSW 29 21 150 59
AM33176 1Attunga State Forest NSW 30 56 150 54
AM36636 1Baldwin Range, Manilla NSW 30 38 150 35
AM36883, 36734 2Bebo State Forest NSW 28 50 150 55
ANWC31 1Buddigower Nature Reserve NSW 34 05 147 05
ANWC24479 1Bullock Creek, W of Echucha Vic 36 13 144 12
AM11160 1Cocopara NP NSW 34 15 146 15
AM7946 1Copeton NSW 29 55 151 01
SAM11329-30, SAM11765-11771,
AM35880, AM37585 11 Dangali Nature Reser ve SA 33 12 140 39
C5195 1Deniliquin, NSW NSW 35 32 144 57
AM36715 1Dubbo City centre NSW 32 14 148 36
AM25355-56 2Dunsandle Station NSW 29 08 146 24
AM34675 1Goonoo State Forest NSW 32 04 148 54
AM36880 1Hell Hole Creek NSW 30 05 150 19
ANWC4910 1Lake Cowal NSW 33 42 147 21
C28470 1Lake Mournpall, via Hattah Vic 34 43 142 21
AM3909 1Millmerran Qld 27 53 151 16
AM37639 1Moonbi NSW 31 01 151 04
C3240 1Mopoke Tank, 30 ml W of Hattah Vic 34 46 141 46
ANWC4385-86 2Mt Pluto Qld 25 00 147 05
AM35881, AM37586 2Mungo National Park NSW 33 15 145 00
AM36759 1Horton River private proper ty NSW 30 21 150 19
AM37645 1Pilliga East State Forest - Clay Dam NSW 30 35 149 26
AM37644 1Pilliga East SF - Delwood Dam NSW 30 47 149 42
AM37717, AM37721-22, AM37732-33 5Pilliga Nature Reser ve, Borah Ck NSW 30 58 149 32
AM38831-35 5Pilliga East: Gilgai Flora Reser ve NSW 31 00 149 21
AM36878-79 1Plagyan State Forest NSW 30 27 150 17
AM36877 1 Plagyan State Forest NSW 30 27 150 13
SAM10003 1Sandfords Dam SA 33 20 140 54
AM36934 1South Warialda State Forest NSW 29 43 150 36
SAM9777 1Tipperary Dam, Morgan Vale SA 33 14 140 43
AM32038 1Top Hut Homestead, 8.9 km W NSW 33 32 142 54
AM36082 1Warrabah NP -Mt Kapitar NSW 30 33 151 00
AM36761 1Woodsreef TSR NSW 30 21 150 46
SAM490 1Yarrock Vic 36 17 141 12
AM23540-23541 2Yathong Nature Reserve NSW 32 38 145 33
Nyctophilus major major (n=43)
WAM18829 12 km from Boddington WA 32 47 116 28
AM37643 170 km SE of Perth WA 32 20 116 15
WAM2955, 2958 2Albany WA 35 00 117 52
WAM28057 1Albany, Kalgan River WA 34 31 117 43
WAM6715 1Broadwater WA 34 29 115 41
WAM1268 1Chorkerup Siding =?Chorkerrys WA 34 50 117 41
AMNH197280 1Contine WA 32 50 116 50
WAM16850, 16853, 24018 3Dwellingup WA 32 38 116 03
A taxonomic review of Australian Greater Long-eared Bats
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Australian
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Registration number NLocality State Lat. Long.
Nyctophilus major major continued.
WAM24862-63 2Forest Grove WA 34 04 115 06
WAM6094 1Mundaring WA 31 57 116 08
SAM522 1Greenbushes WA 33 51 116 03
WAM6375 1Katanning WA 33 41 117 33
WAM24547 1Ludlow WA 33 37 115 29
MG, unregistered 1Perth WA 31 57 115 51
AM4573, AM5473-79, AM5774, AM6319-20 11 Tambellup WA 34 02 117 38
WAM7666 1Vasse WA 33 40 115 15
WAM1247 1Wonnerup WA 33 38 115 26
AM M39797-98, M39808 4Nor thcliffe WA 34 47 116 04
AM M39800 1Dwellingup WA 32 37 116 01
AM39814 1Waroona WA 32 48 116 01
WAM6335-6, WAM6363, 6367 4Woodanilling WA 33 34 117 33
WAM22953 1Kuthala Pass via Mundrabilla WA 31 49 128 13
WAM28398 1Nullabor - Madura Quad 2 WA 33 07 127 21
Nyctophilus major tor subsp. nov. (n=92)
Registration number NLocality State Lat. Long.
WAM28400 1Nullabor WA ? ?
AM35884 1Nullabor, Eyre Highway, (car grill) WA/SA ? ?
WAM17431 1Jibberding area: White Well WA 29 50 116 56
AM37933-37934, M39774-75, M39779,
M39792, M39804-05. 8Dryandra woodlands WA 32 53 116 58
AM37642 1Woodanilling WA 33 34 117 33
AMNH197281 1Katanning WA 33 41 117 33
WAM14847 1Kodjkodjin WA 31 26 117 46
WAM15163 1Mt Bruce WA 22 37 118 08
WAM9916 1Dragon Rocks Reserve WA 32 49 119 05
WAM17725 1Marda WA 30 13 119 16
WAM20697 1Die Hardy Range WA 29 57 119 26
WAM20696 1Mt Manning Range WA 30 00 119 36
WAM17763-64 2Bungalbin Hill WA 30 14 119 49
WAM20166 1Woodline area WA 31 50 122 19
WAM23364 1Queen Victoria Springs WA 29 53 123 30
AM4976 1Booanya via Balladomia WA 32 46 123 36
WAM28399 1Nullabor - Balladonia Quad 3 WA 32 04 124 03
WAM8735 1Madura, 12 ml S WA 32 05 127 04
WAM28402 1Nullabor -Madura Quad 2 WA 33 07 127 21
WAM28401 1Nullabor - Madura Quad 4 WA 32 13 127 26
unreg, field nu. 112-012 1Great Victoria Desert WA 28 20 127 56
WAM22944, 22949, 22951-52, 22954,
22962, 22967-68, 22973 9Kuthala Pass via Mundrabilla WA 31 49 128 13
SAM11296 1Red Gate Tank, 3km S SA 31 23 131 16
SAM14285 1Maralinga, 8.5 km SW SA 30 13 131 31
SAM14286-87 2Maralinga, 12 km SSW SA 30 16 131 33
SAM9325, 9327-9329, 9331-9332, 9334-
9342, AM21170-21172 18 Maralinga SA 30 10 131 35
SAM11288 1Nanwoora Well, 6 km S SA 31 25 131 36
SAM14288 1Maralinga, airstrip SA 30 09 131 37
SAM14280-14284 5Ooldea Siding SA 30 28 131 59
SAM14279 1Immarna Siding SA 30 33 132 08
SAM14292 1Mount Christie Siding SA 30 55 133 16
APPENDIX 1
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2009 Australian
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Registration number NLocality State Lat. Long.
Nyctophilus major tor subsp. nov. continued.
SAM14289 1Yumbarra Conservation Park SA 31 47 133 25
SAM14291 1Mt Finke, 2 km E SA 30 55 134 02
SAM14290 1Mt Finke, 11 km NE SA 30 52 134 06
SAM13337 1Calpatanna Water Hole SA 33 01 134 21
SAM15020 1Karcultaby SA 32 46 134 58
SAM12396 1Gawler Range, Yandinga Wells SA 32 33 135 19
SAM11363 1Hambidge Conservation Park SA 33 26 135 47
SAM10315-10320 6Lake Gillies SA 32 58 136 45
AM39782, M39815 2Jaurdi Station WA 30 46 120 07
AM38843-38845 3Goongarrie WA 29 59 121 03
AM35879 1Cowell SA 33 41 136 55
AM39801 1Eagle Rock, Goldfields district WA 30 26 118 40
AM38842 1Balladonia WA 32 15 123 25
AM39802 1Balladonia WA 32 18 123 32
SAM13004 1Iron Baron SA 32 58 137 07
Nyctophilus daedalus (n=33)
WAM15898 1Beverley Springs Homestead WA 16 43 125 28
AM34453 1Blackfellow Creek NT 13 45 130 52
WAM22356-58 3Cadgeput Springs WA 22 46 119 08
WAM22557 1Cocky Well WA 16 40 122 45
AM22126-28 3Corktree Bore, Pilbara region WA 22 47 119 18
AM9411 1Daly River NT 13 45 130 41
AM34450 1Darwin, 35 k S NT 12 34 131 05
ANWC7592 1Deaf Adder Creek Valley NT 13 06 133 00
SAM6815 1Doomadge Mission Qld 17 56 138 45
WAM552 1Drysdale River WA 14 07 126 43
WAM14097 1Drysdale River National Park WA 15 02 126 55
JM5246 1Lawn Hill NT 18 42 138 29
JM5260 1Louie Ck, Lawn Hill Qld 18 48 138 30
WAM22558 1Martins Well WA 16 34 122 51
WAM19631-33 3Millstream Station WA 21 34 117 03
WAM30586-87 2Millstream Station WA 21 35 117 04
WAM21578 1Mitchell Plateau WA 14 53 125 45
SAM489 1Palmerston (near Darwin) NT 12 14 131 18
ANWC4824 1Rookery Plains NT 12 32 132 23
AM13351 1Roper River, Mataranka NT 14 56 133 07
AMNH216686, WAM18976, unreg. field
numbers WA15, WA16 4Weeli Wolli Spring WA 22 54 119 13
AM34451-52 2West Alligator R., junction Highway NT 12 47 132 10
Nyctophilus sherrini (n=17)
AM34456-57 2Fortesque Forest Tas 43 10 147 50
AM34458 1Scottsdale, 12 km W Tas 41 10 147 31
AM37942-43 2Brittons Swamp Tas 40 59 144 57
QV1984.1.10 1Dip Falls Tas 41 07 145 22
AM34454-55 2Mole Creek Tas 41 34 146 24
AM37935 1Fortesque Tas ? ?
AM37936-37941 6Fortesque Bay Tas 43 09 147 56
CG1985-33 1Tasmania Tas ? ?
AM37944 1Grassy Forest, near Lake Leake Tas 42 10 147 57
APPENDIX 1
A taxonomic review of Australian Greater Long-eared Bats
