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Roosting ecology and social organization of the spotted-winged fruit bat,

Balionycteris maculata (Chiroptera: Pteropodidae), in a Malaysian lowland
dipterocarp forest

Article  in  Journal of Tropical Ecology · November 2003

DOI: 10.1017/S0266467403006060

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Journal of Tropical Ecology (2003) 19:667–676. Copyright  2003 Cambridge University Press
DOI:10.1017/S0266467403006060 Printed in the United Kingdom

Roosting ecology and social organization of the spotted-winged

fruit bat, Balionycteris maculata (Chiroptera: Pteropodidae), in a
Malaysian lowland dipterocarp forest

Robert Hodgkison*1, Sharon T. Balding*, Zubaid Akbar† and Thomas H. Kunz‡

*Department of Zoology, University of Aberdeen, Aberdeen, AB24 3TZ, UK

†Department of Zoology, Universiti Kebangsaan Malaysia, 43600 UKM Bangi, Selangor, Malaysia
‡Center for Ecology and Conservation Biology, Department of Biology, Boston University, Boston, Massachusetts 02215, USA
(Accepted 2 November 2002)

Abstract: The aims of this study were to investigate the roosting ecology and social organization of the spotted-winged
fruit bat, Balionycteris maculata (Megachiroptera), within an old-growth Malaysian dipterocarp forest, and test the
hypothesis that males spend a significantly greater proportion of the night in the immediate vicinity of their day roosts
than females. Balionycteris maculata roosted in cavities and formed small harem groups that consisted of one adult
male accompanied by up to nine adult females and their dependent young. Both male and female bats foraged locally,
within approximately 1 km of their roost sites. However, in contrast to females, which remained away from the roost
site throughout the night, both harem and solitary adult males divided their nightly activity between foraging and
returning to their roost sites. Individual females were also found roosting sequentially with up to three different males.
Hence, these results suggest that B. maculata has a resource-defence polygynous mating system, in which the roost
cavity may represent a critical and defendable resource by which males recruit and gain improved reproductive access
to females. Roost cavities occupied by B. maculata were found within a number of different forest structures, including
ant nests, termite nests and the root masses of epiphytes. The consistent shape and position of the cavities, along with
a single observation of cavity enlargement, provide the first evidence of roost-cavity excavation, by a megachiropteran
bat, within the nests of social insects. Although the cues by which females select their roost sites are unclear, some
aspect of roost cavity size, or condition, may provide a possible means by which females can assess the reproductive
fitness of potential mates.

Key Words: ant nest, Balionycteris maculata, Chiroptera, epiphyte, polygyny, Pteropodidae, roosting ecology, tent-
making, termite nest

INTRODUCTION roost sites in order to attract and gain improved reproduc-

Food resources exploited by bats vary widely in their
spatio-temporal distributions (Fleming 1982, Hodgkison
2001, McKenzie et al. 1995). However, where bats have
adapted to exploit food resources that are available
locally, over extended periods of time, it has become
energetically feasible for the males of some species to
divide their nightly activity between foraging and
returning to their day roosts (Fleming 1982, 1988; Gopu-
kumar et al. 1999, Kunz & Diaz 1995, Kunz &
McCracken 1996, Kunz et al. 1983, 1998; Marimuthu et
al. 1998, McCracken & Bradbury 1981, Morrison 1979,
Morrison & Morrison 1981, Storz et al. 2000a, b). This
situation may have helped promote the evolution of
harem-based polygynous mating systems, in which com-
peting adult males defend and, in some cases, construct
1
Corresponding author. Email: rhodgkison@hotmail.com
tive access to females (Storz et al. 2001).
Roost sites defended by harem males can take many
forms. While some species compete for a limited number
of pre-existing structures, such as solution cavities and
tree cavities (Kunz 1982, Kunz et al. 1983, McCracken &
Bradbury 1981, Morrison 1978, 1979; Morrison & Mor-
rison 1981), the males of other species modify large
leaves, fruit clusters and other plant parts to form tents
(Balasingh et al. 1995, Bhat & Kunz 1995, Kunz &
McCracken 1996, Kunz et al. 1994, Storz et al. 2000a, b;
Tan et al. 1997). Hence, in addition to providing shelter
and possible protection from predators, the day roosts of
some harem-roosting bats may also provide a means by
which females can assess the reproductive fitness of
potential mates (Balasingh et al. 1995, Kunz &
McCracken 1996, McCracken & Wilkinson 2000, Storz
et al. 2000a, b).
668 ROBERT HODGKISON, SHARON T. BALDING, ZUBAID AKBAR AND THOMAS H. KUNZ
Although the mating systems of many bat species are still
unknown (McCracken & Wilkinson 2000), resource-
defence polygyny and tent-making behaviour appear to be
relatively common among fruit-eating phyllostomid bat
species of the Neotropics (Brooke 1987, 1990; Choe 1994,
Fleming 1982, 1988; Foster & Timm 1976, Kunz &
Lumsden 2003, Kunz & McCracken 1996, Lewis 1992,
McCracken & Wilkinson 2000, Stoner 2000, Timm & Mor-
timer 1976). Fruit-eating pteropodid bat species of the
Palaeotropics, by contrast, tend to be relatively large (up to
1.5 kg), and travel over greater distances to feed. Hence,
resource-defence polygyny is believed to be relatively rare
(Fleming 1988). To date only two species of fruit-eating
pteropodid bat (Cynopterus brachyotis and Cynopterus
sphinx) are known to modify plant parts to create tents
(Balasingh et al. 1995, Bhat & Kunz 1995, Goodwin 1979,
Kunz & Lumsden 2003, Phua & Corlett 1989, Storz &
Kunz 1999, Tan et al. 1997). However, from what is known
about the social organization, foraging distances and recap-
ture rates of several other species (Funakoshi & Zubaid
1997, Heideman & Heaney 1989, Hodgkison 2001,
McKenzie et al. 1995, Tan et al. 1999), it is likely that
resource-defence polygyny, and possibly even tent-making
behaviour, may be more common among pteropodids than
is currently recognized. This is particularly true of certain
small forest-dwelling species, which have short foraging
Figure 1. Map of study site and roosting localities of Balionycteris maculata at Kuala Lompat, Krau Wildlife Reserve, Peninsular Malaysia. Roosts
F1–F12 were located in old-growth forest. Roosts R1–R6 were located in rubber plantation.
distances and relatively stable long-term local popula-
tions – such as Balionycteris maculata, Thomas.
Balionycteris maculata, is one of the smallest species
of pteropodid bat (c. 13.5 g), and is found exclusively in
old-growth and old secondary re-growth forests through-
out Peninsular Malaysia (Medway 1983), northern and
western Borneo (Payne et al. 1985), with scattered records
in southern Thailand (Lekagul & McNeely 1977), and
Sumatra (Danielsen et al. 1997). Although it roosts in
small groups in cave entrances (Medway 1983), palm
leaves (Lim 1966), epiphytic ferns (Medway 1983), and
hollow termite mounds (Francis 1994), the social organ-
ization and mating system of this species are unknown.
However, the high frequency of long-term local recap-
tures and the low wing loading of this species, indicate
that it forages over short distances and has relatively
stable local populations (Hodgkison 2001). Hence, this
species has many characteristics often associated with
year-round resource-defence polygyny. The aims of this
study, therefore, are (1) to characterize the roost sites used
by Balionycteris maculata in a Malaysian rain forest, (2)
to investigate the social organization of this species and
(3) to test the hypothesis that males spend a significantly
greater proportion of the night in the immediate vicinity
of their day roosts than females.
Roosting ecology of the spotted-winged fruit bat
METHODS
Study site
This study was conducted at Kuala Lompat (3°43′N,
102°17′E), within the Krau Wildlife Reserve, Pahang,
Peninsular Malaysia, for 26 mo between October 1995
and November 1999. One roost site was also recorded in
August 2001.
The Krau Wildlife Reserve consists of a large area of
old-growth forest (Clark 1996), which ranges in elevation
from 50 m, at Kuala Lompat, to over 2000 m at the
summit of Gunung Benom. At Kuala Lompat, the habitat
can be classified as lowland evergreen mixed-dipterocarp
forest. However, because it is relatively poor in diptero-
carps, and unusually rich in large Leguminosae, the site
is not typical of most dipterocarp forests in Peninsular
Malaysia (Raemaekers et al. 1980).
Although the study site has previously been described
in detail by Raemaekers et al. (1980), the area sur-
rounding the reserve has undergone considerable changes
in recent years. At the time of study, the forest bordered
directly onto gardens and plantations, without a buffer
zone (Figure 1). Although most of the field work for this
project was conducted within the lowland dipterocarp
forest of the reserve, some observations of roosting beha-
viour were also made in the adjoining rubber plantation.
Bat capture and roost location
Balionycteris maculata was sampled over an area of
approximately 1.5 km2 of old-growth forest using 6 × 3-m
mist nets (Avinet Inc., USA). These nets were set singly,
across trails, or stacked together in groups, to create a
continuous wall of net that extended from ground level to
approximately 28 m (see Hodgkison 2001, Hodgkison et
al. 2002). The target species was identified following
Medway (1983) and Payne et al. (1985), and notes were
made on the sex, age and reproductive condition of each
captured bat (Racey 1988), along with measurements of
body mass and forearm length. Standard methods for
marking bats (Kunz 1996) proved to be problematic with
this species. Initial trials, using stainless steel ball-chain
necklaces (Ball-Chain Manufacturing Company, New
Jersey, USA), were quickly abandoned, due to severe
injury to the bats. Trials were then conducted with flanged
monel bands (Lambournes Ltd., UK), which were clasped
around the forearm through a small incision made in the
propatagium. Tolerance to these bands was mixed. Whilst
many individuals appeared to show no adverse effects,
others suffered infections where the bands contacted the
skin. Therefore, concurrent with the banding, a third trial
was conducted, which explored the potential of using the
naturally occurring wing markings to aid identification.
This method was based on the assumption that each indi-
669
vidual bat has its own unique pattern of wing markings.
The method of recording wing markings involved map-
ping the individual spots on the right wing of each indi-
vidual, onto a photocopied line-drawing wing template.
These individually numbered wing-spot diagrams were
then coded, according to the number of spots on each
wing-section and digit, and filed for future reference. All
captured individuals could then be verified against
existing spot-codes and diagrams, and recaptures could be
recognized by both the number, and relative positions, of
their wing-spots. After several months of trials (during
which time the validity of the method was tested, using
wing bands as a control), this method proved to be a very
reliable technique for recognizing individual bats. There-
fore, in the welfare interest of the study species, wing-spot
diagrams became the method of choice and banding was
discontinued.
The roost sites of B. maculata were identified using
small radio-transmitters (0.6 g Holohill, Canada, model
BD-2A). These were attached to the bats on the upper
back, between the shoulder blades, using a latex-based
skin-bond glue. A small patch of reflective tape, glued to
the top of each transmitter, increased the visibility of the
bats in their roosts, and aided in the recovery of the trans-
mitters after use. Radio-transmitters weighed less than 5%
of the body mass of the study species (after Aldridge &
Brigham 1988) and remained attached to the bats for up to
10 d. Recovered radio-transmitters were re-used on other
individuals.
The roost sites of radio-tagged bats were located, the
following day, using a portable radio-receiver (Custom
Electronics Inc., USA) equipped with a five-element Yagi
antenna. The detection range of the signal was limited
within the forest to approximately 300 m. Because the
signal was often blocked and reflected by trees, and par-
ticularly by standing water within the swamps, the search
was made more efficient by measuring the direction of the
signal from triangulation points. To minimize disturbance
to the bats, once a roost was located, the final approach
was performed with extreme caution. Where possible the
presence of fruit bats was confirmed visually, with the aid
of a torch, or sunlight directed by a small mirror. With
sufficient light, it was usually possible to record the
number of individuals present.
Social organization and roost fidelity
Initial attempts to capture bats, using a long-handled hoop
net, resulted in some individuals deserting the roosts.
Because one of the primary objectives of a parallel project
was to study the diet of B. maculata, most roosts were left
undisturbed upon discovery. However, towards the end
of the study, entire roosting groups from several of the
remaining roost sites were captured using a hoop net, to
670 ROBERT HODGKISON, SHARON T. BALDING, ZUBAID AKBAR AND THOMAS H. KUNZ
investigate the social organization of this species. Groups
that consisted of one adult male, accompanied by at least
one adult female and their dependent young, were
described as harems. Data on group size are presented as
means ± 1 SD.
Records of roost fidelity were recorded (1) when bats
switched roost sites during the period of radio attachment,
(2) when re-captured bats were repeatedly radio-tagged
on separate occasions or (3) when previously radio-tagged
bats were re-captured at roost sites, using a hoop net.
Roost descriptions
The nature of the roost structure, its height above ground,
and, where possible, the dimensions of the roost cavity
(i.e. width, breadth and depth) were recorded for all roost
sites located within the study area. These measurements
were made at night, to minimize disturbance to the bats.
Where possible, the duration of roost occupancy, and the
minimum longevity of each roost structure, were also
recorded to the nearest month. Plant nomenclature follows
Holttum (1950, 1974). The abandoned termite nest was
identified after Abe (1978). Data on roost dimensions are
presented as means ± 1 SD.
Foraging movements
The nightly movements of male and female B. maculata,
relative to their roost sites, were compared using radio-
telemetry. Using a pre-existing system of trails and
markers (Figure 1), selected bats were tracked continu-
ously within the forest for a period of 12 h (from dusk to
dawn), for between one and three nights. Individuals
selected for radiotracking included one solitary and two
harem males (all of which were adults) and three adult
females. The compass bearings of each radio-tagged bat
were recorded at 15-min intervals, from at least two
marked positions within the forest. The trail and marker
system was then mapped, over an area of approximately
1.5 km2, and the bearings of each 15-min signal pairing
triangulated to estimate the position and movements of
each bat, relative to its day roost, throughout the night.
All paired signal bearings triangulated at an angle of
greater than 20°.
RESULTS
During 26 mo of field work, including 10 242 (6 × 3 m)
mist net hours of sampling, 113 B. maculata individuals
were captured and marked, or otherwise recorded in the
study area. The male to female sex ratio of the population
was significantly different from unity for adults (1:2.15;
Gadj. = 8.78, df = 1, P < 0.01, n = 78), but not for juveniles
(1:1.50; Gadj. = 1.39, df = 1, P > 0.05, n = 35). Adult males
were significantly smaller than females in relation to fore-
arm length (t = −7.64, df = 69, P < 0.001) but not body
mass (t = 0.69, df = 55, P > 0.05). Forearm lengths of
adult males and females (mean ± 1 SD) were 40.9 ± 0.7
mm and 41.9 ± 1.1 mm, respectively. Body mass measure-
ments of adult males and non-pregnant adult females were
13.3 ± 0.8 g and 13.5 ± 1.0 g, respectively.
Radio-transmitters were applied to 25 bats on 31 occa-
sions. Three bats were radio-tagged on more than one
occasion. Twenty-five of the 31 radio-transmitters were
relocated to reveal 18 roost sites. Twelve roost sites were
located in the old-growth forest (Roosts F1–F12), six were
located in the adjoining rubber plantation (Roosts R1–
R6). Six bats (three males and three females) were tracked
continuously from dusk to dawn, for a total of 11 nights.
Individuals selected for all-night tracking included one
solitary adult male (n = 3 nights), two adult harem males
(n = 3 nights) and three adult females (n = 5 nights).
Roost sites
Balionycteris maculata used at least five different types
of structure for roosting in the old-growth forest at Kuala
Lompat (Roosts F1–F12). These included two epiphytic
plant species, Asplenium nidus Linn. (Aspleniaceae)
(n = 3 roosts; 25%) and Hedychium longicornutum Baker
(Zingiberaceae) (n = 1 roost; 8%), active arboreal nests of
the ant Crematogaster ebinina Forel (n = 5 roost; 42%),
an abandoned arboreal termite Bulbitermes sp. nest
(n = 1; 8%) and the hollow base of a large detached
branch (n = 1 roost; 8%). Only one roost site, used by at
least two radio-tagged bats, was unidentified in the forest
(roost F12, Figure 1). But judging by the weak and highly
directional signals, received from both radio-tagged bats,
it is possible that this roost site may have been located
within a tree cavity. Dense vegetation cover in the sur-
rounding area, however, precluded visual confirmation.
All roost sites occupied by B. maculata in the rubber
plantation were located within epiphytic Asplenium nidus
ferns (Roosts R1–R6).
Roost description
All but two roost sites (roosts R4 and R5 – Figure 1)
located during this study provided B. maculata with a
semi-enclosed sheltered space, in the form of a smooth-
surfaced, bell-shaped cavity (Figure 2). These cavities
were positioned at the base of the roost structure, and
were unobstructed by vegetation. All roost structures were
comprised of relatively soft materials, and neither species
of ant or termite were notably aggressive. Cavity widths
ranged from 8.0–20.0 cm in diameter (mean = 13.7 ± 4.0
cm, n = 9), and were generally as broad as the individual
resource permitted (mean = 11.9 ± 3.0 cm, n = 9). Cavity
depths ranged from 7.0–22.0 cm (mean = 12.3 ± 5.3 cm,
Roosting ecology of the spotted-winged fruit bat 671

Figure 2. (a) Root mass of a bird’s nest fern (Asplenium nidus) excavated and used as a roost site by Balionycteris maculata; (b) small harem group
of B. maculata occupying a cavity in an excavated bird’s nest fern; (c) root mass of an epiphytic ginger (Hedychium longicornutum) excavated and
used as a roost site by B. maculata; (d) small harem group of B. maculata occupying a cavity in the root mass of an epiphytic ginger; (e) arboreal
ant nest (Crematogaster ebinina) excavated and used as a roost site by B. maculata; (f) small harem group of B. maculata occupying the interior of
an excavated arboreal ant nest. Arrows indicate openings to roost cavities from below.
672 ROBERT HODGKISON, SHARON T. BALDING, ZUBAID AKBAR AND THOMAS H. KUNZ

90
n = 9) and each roost structure was between 2.30–6.20 m 80 (a)
above ground (mean = 3.39 ± 1.32 m, n = 17). 70
60

% NRP
Social organization and roost fidelity 50
40
Balionycteris maculata roosted in small groups of 1–14 30

bats (mean = 5.6 ± 4.2, n = 19). Each group consisted of 20

10
one adult male, accompanied by up to nine females
0
(mean = 2.5 ± 3.0, n = 15) and their dependent young. < 51 51 - 100 101 - 250 251 - 500 501 - 1000
Multiple linear regression revealed no statistically signi- Distance from the roost (m)
ficant association between group size and any combina-
Females Males
tion of roost cavity dimension (r2 = 0.311, SE = 3.55,
F = 2.35, df = 4, 8; P = 0.141). Individuals roosted in
tight clusters, and were often in physical contact. 120
(b)
Individual females were recorded within up to three dif- 100
ferent roost sites. In contrast, no cavity-dwelling male was

HRP
80

%HRP
ever found to have moved to an alternative roost site 60
within the study area, whilst his old roost site was still
%
40
intact. Indeed, adult males that had disappeared from
20
intact roost sites were never captured again within the
study area, despite a continuous mist-netting effort. 0
20h00

21h00

22h00

23h00

00h00

01h00

02h00

03h00

04h00

05h00

06h00

07h00
Only two adult males were directly observed to occupy
more than one roost site within the study area. The first Time
Time
individual (a solitary male) displayed transient fidelity to Females Males
two epiphytic bird’s nest ferns (Roosts R4 and R5 –
Figure 1), both of which were within the rubber planta- Figure 3. Nightly foraging movements of male and female Balionycteris
tion, and neither of which possessed a cavity. The second maculata in relation to their day roosts, showing (a) the mean percentage
individual (a harem male), by contrast, only moved to an of nightly radio-telemetry positions (NRP) recorded at different dis-
alternative roost site after his existing roost site had been tances from the day roost, and (b) the mean percentage of hourly radio-
destroyed by a tree-fall. This male was first discovered in telemetry positions (HRP) recorded within 50 m of the day roost. All
radio-telemetry positions were recorded at 15-min intervals from dusk
April 1997, roosting in a Crematogaster ebinina nest (19h00) to dawn (07h00). Bars indicate standard deviation.
(Roost F4), along with three adult females. The tree-fall
that destroyed the roost occurred in May 1998. When cap-
tured and radio-tagged on a second occasion, almost 1 y
togaster ebinina nests. One roost cavity (Roost F7 – see
later (April 1999), the same bat was discovered roosting
above), which contained a solitary male, was 5 cm deep
alone in a second newly constructed Crematogaster ebin-
when measured in April 1999. When re-measured in Nov-
ina nest (Roost F7), within 10 m of the original observa-
ember of the same year, the depth of the cavity had
tion (Figure 1). In August 2001, this second roost site,
increased to 10 cm, by which time the male was accom-
which was last observed in November 1999, had also been
panied by two other individuals, both of which were
destroyed. Inspection of the surrounding area, however,
females.
revealed a third new ant nest, which was located within 1
m of the original observation (Roost F8 – Figure 1). A
cavity within this nest contained the same adult male, Duration of roost occupancy and longevity of roost sites
along with eight females and their dependent young.
Balionycteris maculata occupied roost sites for between 1
Hence, one adult male had occupied three different ant
d to at least 13 mo. Three forest roost sites (roosts F4, F5
nests, within the same small patch of forest, over a period
and F10) were successively occupied by more than one
of more than 4 y. All three roost sites had deep cavities,
solitary or harem-roosting adult male. At least one of
and, because of their close proximity, all three nests were
these roost sites (roost F4) was unoccupied for several
also likely to have been created by the same colony of
months before being appropriated by another bat. Some
ants. Measurements of cavity depth, in the second roost
roost sites remained structurally sound for at least 28 mo.
site (Roost F7) are detailed below.

Cavity enlargement Foraging movements

Fragments of broken nest material were consistently The nightly movements of male and female B. maculata,
found below the cavities of B. maculata roosts in Crema- relative to their day roosts, are summarized in Figure 3.
Roosting ecology of the spotted-winged fruit bat
Although insufficient data were available for a detailed
analysis of home range, the results show that all indi-
viduals foraged locally within approximately 1 km of their
roost sites. Indeed, only two individuals, one male and
one female, were recorded more than 500 m from their
day roosts throughout the radio-tracking period of this
study.
Males spent a significantly greater proportion of the
night within 50 m of their day roosts than females (Gadj. =
59.5, df = 1, P < 0.001). Two males that travelled less
than 500 m to forage (Roosts F3 and F5) returned to their
day roosts periodically throughout the night, whereas one
male that travelled over 500 m to feed (Roost F2) returned
to his roost at 00h00, after a continuous absence of
approximately 4 h. Observations of both radio-tagged and
untagged individuals indicate that males use regular long-
term feeding perches, close to their day roosts. Females,
by contrast, stayed away from the roost throughout the
night, irrespective of the foraging distance they travelled.
The presence of both male and female bats could be
easily detected within the forest by their social calls,
which were distinctive bird-like ‘peeps’. Males also pro-
duced a distinctive ‘twittering’ type variation of this call.
This often appeared to be triggered by human presence,
particularly within the vicinity of the roost.
Apart from one male, which was observed tugging at
the antenna of his radio-transmitter, all radio-tagged
animals were out of visual contact throughout the tracking
period of this study. Grooming and antenna tugging
seemed to affect the modulation of the radio signal. Thus,
from a distance, it was not always possible to determine
whether the bat was stationary or in flight.
Although this species was evidently aware of, and irri-
tated by the application of the radio-transmitter, apart
from a small amount of temporary hair loss, none of the
bats appeared to suffer any injury as a result of this treat-
ment.
DISCUSSION
Social organization and mating system
Balionycteris maculata roosted in cavities, and formed
small harem groups that consisted of a single adult male,
accompanied by up to nine adult females and their
dependent young. Both male and female bats foraged
locally, within approximately 1 km of their roost sites.
However, in contrast to females, which remained away
from the roost site throughout the night, both harem and
solitary adult males divided their nightly activity time
between foraging and returning to their day roosts. Indi-
vidual females were also found roosting sequentially with
up to three different males. Hence, these results suggest
that B. maculata is likely to have a resource-defence poly-
gynous mating system (Clutton-Brock 1989, Emlen &
673
Oring 1977), in which the roost site, or perhaps more
specifically the roost cavity, may represent a critical and
defendable resource for the recruitment of females. There-
fore, in addition to providing shelter from wind and rain,
and concealment from potential predators, the cavities
used by B. maculata males could also provide a possible
means by which females can assess the reproductive fit-
ness of potential mates (Balasingh et al. 1995, Kunz &
McCracken 1996, Storz et al. 2000a).
Roost cavities and cavity formation
Roost cavities occupied by B. maculata were located
within a number of different forest structures, including
ant nests, termite nests and the root masses of epiphytes.
The use of insect nests by bats has also been described in
the Neotropics, where several species of phyllostomids,
including the round-eared bats (Tonatia silvicola, T.
minuta, T. carrikeri) and the greater spear-nosed bat
(Phyllostomus hastatus), have all been observed roosting
in cavities formed within the active and abandoned arbor-
eal nests of termites (Allen 1911, Goodwin & Greenhall
1961, Kalko et al. 1999, McCarthy et al. 1983, 1992;
Tuttle 1970). Although the origins of these cavities are
unknown, based on observations of T. silvicola in Panama
and Venezuela, Kalko et al. (1999) suggested that Tonatia
may exploit abandoned nest cavities, created within termi-
taria by birds such as trogons (Trogon sp.) or parakeets
(Brotogeris sp.).
Several species of bird are also known to nest in the
arboreal domatia of ants and termites in the Old World
tropics (Hindwood 1959). Indeed, abandoned birds’ nests
were found in several ant and termite nests within the
study area, including one within a Crematogaster ebinina
ant nest, of the same type used by B. maculata. However,
in contrast to B. maculata roost cavities, which were basal
and semi-enclosed, the nest cavities created by birds
entered the ant nest from the side, through a narrow open-
ing of approximately 5 cm diameter, and hence were
almost completely enclosed. Thus, birds were unlikely to
have produced the roost cavities described in this study.
It may be possible, in the case of the ant and termite
nests, that the roost cavities occupied by B. maculata were
partially formed as a result of insect-eating birds and/or
mammals, breaking open the nests from below during
feeding. But, the consistent shape and position of the cav-
ities, and the fact that they were also found within the
root masses of epiphytes, suggests that this is unlikely
to explain the formation of the completed structure. An
alternative hypothesis, therefore, which is supported by an
observation of roost cavity enlargement by a single adult
male, is that B. maculata excavates its own roost sites.
The roost-making behaviour of two other closely
related pteropodids (Hill 1992), lends further support to
this hypothesis. The short-nosed fruit bats Cynopterus
674 ROBERT HODGKISON, SHARON T. BALDING, ZUBAID AKBAR AND THOMAS H. KUNZ

brachyotis and Cynopterus sphinx are both known to ACKNOWLEDGEMENTS

modify leaves, root masses, stems and flower and fruit
We would like to express our thanks to the Department of
clusters to create roost spaces, or tents – some of which
Wildlife and National Parks (Malaysia), for their generous
show a striking similarity in both shape and position to
support throughout this project, and for providing us with
the roost cavities occupied by B. maculata (Balasingh et
the free use of the field station facilities at Kuala Lompat.
al. 1995, Bhat & Kunz 1995, Tan et al. 1997). To con-
This study would not have been possible without the per-
struct roost cavities within the nests of social insects
mission of the Economic Planning Unit of Malaysia.
would presumably require a similar level of ingenuity, and
Valuable assistance and encouragement in the field was
could also be achieved in the same manner – through the
given by Ahmad Bin Dagu, Tigga Kingston and Zamiza
use of teeth and jaws. Hence, the results of this study
Bin Zainal. Constructive comments on the manuscript
appear to provide the first evidence of roost cavity excava-
were offered by P. A. Racey, M. B. Fenton, Colin O’Don-
tion, by a megachiropteran bat, within the nests of social
nell, Kerryn Parry-Jones and an anonymous reviewer. The
insects.
ant specimens were kindly identified by Min-Sheng
Wang. Alison Sandison helped with the preparation of the
map. This study was supported by a grant from the Lubee
Roost appropriation and cavity enlargement
Foundation Inc., Florida, to T. H. Kunz. Additional spon-
Several roost sites observed during this study were aban- sorship, to R. Hodgkison, was also provided by the Carne-
doned by B. maculata, without re-occupation. The causes gie Trust for the Universities of Scotland, the Royal Soci-
of desertion were not always clear, but human disturbance ety South East Asia Rain Forest Regeneration and
was probably the primary cause within the rubber planta- Recovery Programme (RS-112) and Bat Conservation
tion. Three of the 12 roost cavities located within the International.
forest, however, were sequentially occupied by more than
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