INTRODUCTION
Chaerophon plicatus (family Molossidae, sub-
family Molossinae) is a widely distributed bat
throughout lowland South Asia and Southeast Asia,
from India east to the Philippines and south to
Lombok (Heaney et al., 1998; Molur et al., 2002).
The species is a facultative cave-dweller, and in suit-
able locations forms very large cave colonies. In
northern Borneo, spectacularly large colonies are
well-known from Simud Hitam Cave (Gomantong
Forest Reserve, Sabah — Fig. 1) and Deer Cave
(Mulu National Park, Sarawak). A formerly very
large colony, now much reduced, occupies Niah
Cave (Niah National Park, Sarawak — Medway,
1958; Struebig et al., 2010). Foraging areas have
not been explicitly measured for this species, but
by analogy to the North American member of the
same subfamily — Tadarida brasiliensis (Ammer-
man et al., 2012) which has measured nightly
foraging ranges averaging 41 km around the Carls-
bad Caverns (New Mexico) colony (Best and
Geluso, 2003).
It can be presumed that major C. plicatus
colonies are top insect predators over very large
expanses of lowland rainforest and agricultural
mosaics. These colonies feed on some nine kg of
insect prey per 1,000 bats per night (based on
mass scaled feeding rates of T. brasiliensis from
Kunz et al., 1995; C. plicatus mean body mass of
15.5 g from Leelapaibul et al., 2005 versus 12.3 g
for T. brasiliensis from Wilkins, 1989), and their
accumulated guano supports rich subterranean
ecosystems (Chapman, 1983) including a number
of endemic species (Ng, 1991; McFarlane et al.,
2009).
Acta Chiropterologica, 17(1): 171–177, 2015
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/15081109ACC2015.17.1.014
Estimating colony size of the wrinkle-lipped bat, Chaerephon plicatus
(Chiroptera: Molossidae) at Gomantong, Sabah, by quantitative image analysis
DONALD A. MCF ARLANE
1, 5
, GUY V AN RENTERGEM
2
, ANNEMIEKE RUINA
1
, JOYCE LUNDBERG
3
,
and KEITH CHRISTENSON
4
1
Keck Science Center, The Claremont Colleges, Claremont, CA 91711, USA
2
NA c9 3803919, Koningin Astridstraat 57, 9800 Deinze, Belgium
3
Department of Geography and Environmental Studies, Carleton University, Ottawa, ON K1S 5B6, Canada
4
2026 Peach Orchard Drive Apt. 23, Falls Church, VA 22043, USA
5
Corresponding author: E-mail: dmcfarlane@kecksci.claremont.edu
The Gomantong cave system, Kinabatangan, Sabah (Malaysia) hosts one of the largest bat colonies known from north Borneo. The
nightly exodus of Chaerophon plicatus from this site is an economically valuable tourist attraction, and must impose significant
controls on the regional ecology. Monitoring ecosystem health requires monitoring bat population size, but no quantitative
assessments for the Gomantong colony are available in the literature. Traditional censusing techniques (based on packing density
and roost area or on roost exodus counts) yield notoriously unreliable results. Here we have applied innovative image analysis and
counting techniques to videographic, photographic and terrestrial laser scanning data obtained in July 2012. The colony exits in
a consistent stream along a narrow exit trajectory. The laser scanning of the large cave entrance allowed precise measurement of bat
position. Video data provided 0.02-second time resolution. Average flight speed was calculated (10.38 ± 0.85 m/sec), bat exit rate
was estimated at three minute intervals (peaking at 14,000 bats/minute) and then summed over the 40 minutes of bat flight. The
resultant colony size estimate of 275,625–276,939 (95% confidence interval) individuals of C. plicatus in Gomantong Caves in 2012
demonstrates that the technique is viable, provides a realistic basis for ecosystem management, and can be repeated to monitor
ecosystem change. The estimated insect consumption by the colony of this size is 927 ± 191 metric tons of insects per year over an
estimated 270 km
2
area, a very important component in maintaining ecosystem stability in the Gomantong Forest Reserve and the
Kinabatangan Wildlife Sanctuary, as well as pest control over the large agricultural tracts of the region.
Key words: Borneo, Kinabatangan, bats, caves, colony size, image analysis, terrestrial laser scanning