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405
CHAPTER 14
Conservation Threats to Bats
in the Tropical Pacific Islands
and Insular Southeast Asia
Gary J. Wiles and Anne P. Brooke
Introduction
More than 27,000 islands in 23 countries and territories, spread across millions
of square kilometers in the Indo-Pacific, have given rise to a diverse bat fauna
characterized by high levels of endemism, particularly in the family Ptero-
podidae (Flannery 1995; Pierson and Rainey 1992). A total of 354 bat species,
including 140 pteropodids and 214 microchiropterans in 9 families (appendix
14.1), reside in the mainly tropical geographic region extending from the Cook
Islands in central Polynesia to the subtropical Ryukyu and Ogasawara Islands
of Japan, and westward through the Indonesian archipelago (fig. 14.1). This
represents about 31% of currently recognized bat species and about 74% of all
pteropodids. These inhabit a combined land area of 2,961,600 km2, which is
smaller than the size of India.
Regional bat diversity is greatest on the large islands off Southeast Asia and
in New Guinea, but declines rapidly on the Pacific islands to the east, where
only a handful of species occur (table 14.1; Carvajal and Adler 2005; Hall et al.
2004; Hutson et al. 2001; Mickleburgh et al. 1992; Rainey and Pierson 1992).
This pattern of occurrence has been strongly influenced by the size and geologi-
cal history of the islands. Intermittently low sea levels during the Pleistocene
reduced interisland distances or connected many of the western islands with
the Asian continent on multiple occasions (Voris 2000), promoting the disper-
sal of bat species and, over time, high levels of endemism and greater spe-
cies richness. The mountainous nature of many Australasian islands has also
contributed to speciation. By contrast, the relative isolation and small sizes of
many oceanic islands have greatly hindered their colonization by bats. From
a conservation perspective, the region contains 8 of 39 globally recognized
hot spots of biodiversity (i.e., Sundaland, Wallacea, the Philippines, eastern
Melanesia islands, New Caledonia, Taiwan, Polynesia-Micronesia, and part of
Japan), with bats comprising a significant faunal component in most of these
(Mittermeier et al. 2005).
406 G. J. Wiles and A. P. Brooke
Within Oceania and insular Southeast Asia, knowledge of the status and
natural history of bat populations varies from being fairly good at a few lo-
calities in Polynesia and Micronesia to highly incomplete across much of In-
donesia, the East Malaysian states of Sarawak and Sabah, the Philippines, and
Melanesia. Inadequate information is a serious detriment to preserving bat
populations (Hutson et al. 2001; Mickleburgh et al. 2002). Basic data on distri-
bution and habitat preferences are incomplete for many of the region’s bats
and are entirely lacking for a few taxa. Major taxonomic questions also remain
about the status of some species and further interfere with the establishment
of conservation priorities.
Latent extinction risk among bats and other mammals is considered high
across much of insular Southeast Asia and the tropical Pacific (Cardillo et al.
2006). Regional threats to bats come from a variety of sources, many of which
stem from expanding human populations and their ever-increasing pressure
on natural ecosystems. Currently, 70 of the region’s bat species are recognized
as threatened at some level (appendix 14.1), with 11 species classified as criti-
cally endangered, 22 as endangered, and 37 as vulnerable (IUCN 2008). An-
other 30 species are considered near threatened, 177 are of least concern, 71 are
data deficient, 2 (Pteropus pilosus and P. tokudae) are presumed extinct, and 4
have not been evaluated. Of the 6 families with more than 2 species present in
the region, the family Pteropodidae has by far the highest percent of threatened
members (34.3%, 48 of 140 species), followed by Hipposideridae (12.2%, 5 of
Figure 14.1. Map of the 23 countries, territories, and island groups in the Pacific and insular South-
east Asia covered in this chapter, all of which have tropical or subtropical climates. Temperate
locations such as New Zealand, Lord Howe Island off Australia, and the main islands of Japan are
not discussed, nor are the subtropical Hawaiian Islands.
Conservation Threats to Bats in the Pacific and Southeast Asia 407
41 species), Vespertilionidae (10.6%, 11 of 104 species), Molossidae (10.0%, 2 of
20 species), Rhinolophidae (7.1%, 2 of 28 species), and Emballonuridae (5.9%,
1 of 17 species).
In this chapter, we summarize available information on five major categories
of threats to bat populations in the region: habitat loss and alteration, hunting,
cave disturbance, severe storms, and introduced species. We do not discuss
several additional concerns, which are described elsewhere. These include
global climate change (Rainey 1998), epizootic diseases (Rainey 1998), conflicts
with fruit producers (Fujita and Tuttle 1991), and pesticide use (Tarburton
2002). Species are often threatened by multiple factors, some of which work in
conjunction to drive populations down. For example, severe tropical cyclones
can result in increased hunting of flying foxes (e.g., Pierson et al. 1996) and land
clearing can increase human access to nearby caves occupied by bats.
Habitat Loss and Alteration
Forest loss and degradation through logging, development, and fire are the
principal threats to biodiversity in many tropical Pacific and insular Southeast
Asian countries and territories, with lowland forests and their associated biota
being especially vulnerable ( Jepson et al. 2001; Sodhi and Brook 2006; Wikra-
manayake et al. 2002). The vast majority of the region’s bats rely on forests
completely or to some extent; thus declining forest cover has major impacts
on most taxa by reducing foraging and roosting habitats (Hutson et al. 2001;
Racey and Entwistle 2003).
Tropical Pacific and insular Southeast Asian countries vary considerably in
the amount of forest cover lost over time (table 14.2; FAO 2006). Since 1990,
Indonesia has had one of the highest deforestation rates in the world, with
more than 280,000 km2 of forest permanently lost to human encroachment at
a mean rate of decline of about 1.6% annually. More than 90% of the primary
forest in the Philippines has already been destroyed, giving it one of the small-
est relative amounts of coverage of this habitat for any country in the region.
The Solomon Islands, Timor-Leste, the East Malaysian states of Sarawak and
Sabah, and Papua New Guinea also show relatively high rates of loss. In Japan
there has been extensive clearing for agriculture and other development in the
Ryukyu and Ogasawara Islands. Data are imprecise for many of the smaller
Pacific nations, but ongoing deforestation rates generally appear to be rela-
tively low (table 14.2). However, it should be noted that forest cover statistics
for the region, including those of the FAO (2006), are often misleading because
they commonly incorporate habitats of lower ecological value, such as heavily
disturbed secondary forests, forest monocultures (e.g., rubber trees) that pro-
vide fewer resources for forest bats, and even clear-cut lands left to regenerate
naturally. Hence, existing figures can underestimate actual rates of forest deg-
radation and can substantially overestimate coverage by high-quality forests.
Table 14.1. Numbers of bat species per family for 23 countries, territories, and island groups in the
tropical Pacific and insular Southeast Asia
Country or island group Pteropodidae Rhinolophidae Hipposideridae Megadermatidae Rhinopomatidae
Indonesia 78 20 30 1 1
Sarawak and Sabah
(East Malaysia)
16 10 12 1 —
Papua New Guinea 37 4 13 — —
Philippines 26 10 9 1 —
Brunei 16 6 8 1 —
Solomon Islands 25 — 7 — —
Taiwan 2 2 2 — —
Timor-Leste 11 4 4 — —
Ryukyu Islands
(Nansei Shoto)c
2 3 1 — —
Vanuatu 4 — 2 — —
New Caledonia 4 — — — —
Fiji 4 — — — —
Federated States
of Micronesia
4 — — — —
Samoa 2 — — — —
Guam 2 — — — —
Palau 2 — — — —
American Samoa 2 — — — —
Ogasawara and Iwo Islands 1 — — — —
Commonwealth of the
Northern Mariana Islands
1 — — — —
Tonga 1 — — — —
Wallis and Futuna 1 — — — —
Niue 1 — — — —
Cook Islands 1 — — — —
Table 14.1. (continued)
Emballonuridae Nycteridae Mollossidae Vespertilionidae Total speciesa
Endemic
species
Threatened
speciesbSources
12 2 13 65 222 54 33 3, 4, 14, 15,
21, 23, 24, 25
5 1 3 49 97 2 7 21, 23, 24
10 — 6 24 94 17 7 4, 13, 23, 24
3 — 4 25 78 21 6 6, 7, 8, 11,
16, 23
4 1 3 19 58 — 3 17, 23
4 — 1 6 43 14 12 9, 13, 24
— — 1 29 36 5 — 2, 18, 19, 24
2 — — 10 31 1 2 10, 22, 23, 24
— — 1 6 13 5 3 1, 5, 24
1 — 1 4 12 2 5 9, 12, 24
— — — 5 9 6 6 9
1 — 1 — 6 1 4 20
1 — — — 5 3 3 26
1 — — 1 4 1 1 9
1 — — — 3 1 2 26
1 — — — 3 1 2 26
1 — — — 3 — 1 9
— — — 1 2 2 1 1, 24
1 — — — 2 — 2 26
1 — — — 2 — 1 9
— — — — 1 — — 9
— — — — 1 — — 9
— — — — 1 — — 9
Sources: 1 = Abe et al. 1994; 2 = BAT 2008; 3 = Bates et al. 2007; 4 = Bonaccorso 1998; 5 = BSCGJ 2005; 6 = Esselstyn 2007; 7 =
Esselstyn et al. 2004a; 8 = Esselstyn et al. 2008; 9 = Flannery 1995; 10 = Goodwin 1979; 11 = Heaney et al. 1998; 12 = Helgen
2004; 13 = Helgen 2005; 14 = Helgen 2007; 15 = Helgen and Wilson 2002; 16 = Helgen et al. 2007; 17 = Kofron 2002; 18 = Kuo
et al 2006; 19 = Lin et al. 1997; 20 = Palmeirim et al. 2007; 21 = Payne et al. 1985; 22 = Polhemus and Helgen 2004; 23 = SAMD
2006; 24 = Simmons 2005; 25 = Struebig et al. 2006; 26 = Wiles 2005a.
aIncludes species that have become extirpated or extinct in historic times. Species tallies for the Philippines and Taiwan
include some taxa identified only to genus (see BAT 2008; Heaney et al. 1998).
bIncludes species classified as critically endangered, endangered, or vulnerable by IUCN (2008).
cIncludes all of Japan’s southwestern islands, including the Osumi, Tokara, Amami, Okinawa, Sakishima, Yaeyama, and
Daito island groups.
Table 14.2. Forest coverage and amount of change for 23 countries, territories, and island groups in the tropical Pacific and insular Southeast Asia
Country or island group
Total land
area (km2)
Total forest
cover in 2005
(km2)a
Total forest cover
in 2005 as % of
total land areaa
Primary forest
cover in 2005
(km2)
Mean % annual
change in forest
cover, 1990–2005
Total forest
loss, 1990–2005
(km2)
Indonesia 1,811,570 884,950 48.8 487,020 –1.61 280,720
Papua New Guinea 452,860 294,370 65.0 252,110 –0.44 20,860
Philippines 298,170 71,620 24.0 8,290 –2.15 34,120
Sarawak and Sabah (East Malaysia) 198,070 125,040b63.1bn.a.c–0.67b19,340b
Solomon Islands 27,990 21,720 77.6 n.a. –1.44 5,960
Timor-Leste 14,870 7,980 53.7 n.a. –1.16 1,680
Brunei 5,270 2,780 52.8 2,780 –0.75 350
Locations where forest
cover trends are less certain
Total land
area (km2)
Total forest cover
in 2005 (km2)
Total forest cover
in 2005 as % of
total land area Remarks on continuing forest loss
Taiwan 35,970 21,000 58.5 Forest cover is stable or slightly increasing.d
New Caledonia 18,280 7,170 39.2 Minor loss continues from logging, mining,
and other causes.e
Fiji 18,270 10,000 54.7 Minor overall loss since 1990, but extensive
conversion to forests of poorer quality;f see
Ash 1992 for additional remarks.
Vanuatu 12,190 4,400 36.1 Extensive logging of lowland forests is ongoing.e
Ryukyu Islands (Nansei Shoto)g4,500 n.a. n.a. Some deforestation continues.e
Samoa 2,830 1,710 60.4 Minor overall loss since 1990, but extensive
conversion to forests of poorer quality.f
Tonga 720 n.a. n.a. Unknown.
Federated States of Micronesia 700 630 90.6 Minor loss on most islands since 1990, but
extensive on Pohnpei (see text).h, j
Guam 550 260 47.1 Minor loss since 1990.h
Palau 460 400 87.6 Minor loss since 1990.h
Commonwealth of the
Northern Mariana Islands
460 330 72.4 Little or no loss on most islands since 1990,
except Anatahan, where loss is nearly
complete (see text).h, j
Wallis and Futuna 274 n.a. n.a. Some ongoing loss since the 1980s.e
Niue 260 140 53.8 Mean annual loss of forest cover was 1.2%
from 1990 to 2005.k
Cook Islands 230 160 69.6 Mean annual increase of forest cover was
0.4% from 1990 to 2005.k
American Samoa 200 180 89.4 Mean annual loss of forest cover was 0.2%
from 1985 to 2001.l
Ogasawara and Iwo Islands 100 n.a. n.a. Little forest cover remains, thus additional
loss is probably minor.e
Source: Statistical data originate from FAO 2006 unless otherwise noted and are for the period from 1990 to 2005.
aIncludes primary and secondary forests, mangroves, and monoculture tree plantations (including nonnative species) used for forest products or protective purpose. Habitats
must be greater than 0.5 ha in size and exceed tree heights of 5 m and a canopy cover of 10%, or be capable of reaching these thresholds to qualify as forest. It is unclear whether
coconut plantations are consistently included in the data, but other three crops such as oil palm and fruit trees are excluded.
bI. E. Henson, pers. comm. Data are for the period from 1980 to 2000. Additional background appears in Jomo et al. 2004.
cN.a. = data are not available or, in a few cases, are considered unreliable as presented in FAO 2006.
dTsai 1999.
eStattersfield et al. 1998.
fJ. Atherton, pers. comm.
gIncludes all of Japan’s southwestern islands, including the Osumi, Tokara, Amami, Okinawa, Sakishima, Yaeyama, and Daito island groups.
hG. J. Wiles, pers. obs.
iMerlin and Raynor 2005.
jC. C. Kessler, pers. obs.
kFAO 2006.
lDonnegan et al. 2004.
412 G. J. Wiles and A. P. Brooke
Techniques for measuring forest cover also frequently differ among studies,
further complicating comparisons over time and among countries.
Fragmentation is a significant component of forest disruption and threat-
ens populations of forest-dwelling bats through increased isolation, related
stochastic factors, and reductions in microhabitat quality. Size of fragments,
degree of isolation, level of matrix contrast, and species vagility are among
the factors that affect persistence of bat assemblages in fragmented landscapes
(Struebig et al. 2008). Forest fragmentation is considered most severe in the
Philippines and the Greater Sunda Islands of Indonesia, whereas compara-
tively intact tracts of forest persist in New Guinea, Melanesia, and Wallacea
(Wikramanayake et al. 2002).
Few analyses of the effects of timber harvest and land conversion on bat
communities have been published for insular Southeast Asia and the tropical
Pacific; thus it is instructive to look at investigations from neighboring areas to
gain a better understanding of impacts. Singapore has lost more than 95% of its
original forest cover since the early 1800s and has seen bat species diversity fall
by as much as 69–75% for microchiropterans and about 60% for megachirop-
terans (Lane et al. 2006). Projected declines in species richness are particularly
apparent among hipposiderids, rhinolophids, members of the vespertilionid
subfamilies Murininae and Kerivoulinae, and other forest-dependent taxa. Sur-
viving species tend to be microchiropterans that prefer open and edge habitats
and megachiropterans that are widespread and select agricultural and second-
ary habitats or that can travel sizable distances (Lane et al. 2006). These dire
findings are made worse by the small population sizes for many of Singapore’s
remaining bats, suggesting that additional extinctions are likely. Research from
peninsular Malaysia shows that forest-interior microchiropterans are especially
vulnerable to changes in forest structure associated with human disturbance
(Kingston et al. 2003; Struebig et al. 2008; Zubaid 1993). Many such species are
characterized by wing morphologies and echolocation calls that are adapted for
foraging in dense forest understories, and hence are unable to detect prey ef-
ficiently in more open environments (Kingston et al. 2003; Meijaard et al. 2005).
Additionally, loss of large trees with hollows or exfoliating bark eliminates the
preferred roosting sites for some species.
Forest disturbance, clearance, and fires probably represent the most impor-
tant threats to many bat species in Indonesia and East Malaysia. Preliminary
data from Sumatra indicate that logging and conversion of forests to planta-
tions of oil palm and rubber can reduce species richness by 50–88% through
the loss of both microchiropterans and pteropodids (Danielsen and Heegaard
1995). In and around the Sangkulirang limestone karst formations in eastern
Kalimantan, deforestation has reduced habitat availability for the area’s di-
verse microchiropteran fauna, with forest-roosting species underrepresented
in surveys of sites where large mature trees have been lost to fires (Suyanto
and Struebig 2007; M. J. Struebig, pers. comm.). In the oligotrophic forests of
Conservation Threats to Bats in the Pacific and Southeast Asia 413
southern Kalimantan, populations of species using tree hollows as roosts ap-
pear more limited in disturbed locations than in undisturbed ones (Struebig et
al. 2006). Elsewhere on Borneo, habitat destruction has probably contributed
to the decline of Cheiromeles torquatus by reducing opportunities for foraging
and roosting in tree cavities (Hutson et al. 2001). Other species reliant on tree
hollows (e.g., Rhinolophus sedulus, Megaderma spasma, Nycteris tragata, and Ke-
rivoula papillosa) are also considered at risk on this island (Meijaard et al. 2005).
Logging, land clearing, and plantation establishment have produced impover-
ished pteropodid communities in northern Sulawesi, the Sangihe Islands, and
islands off Irian Jaya (Meinig 2002; Riley 2002b). Forest loss is also a major
threat to Pteropus vampyrus on Java (Bergmans 2001).
The greatest threat to Philippine bats is habitat loss (Hutson et al. 2001).
At least 18 of 26 pteropodids occur entirely or primarily in forests and have
experienced some level of population decline due to land clearing and continu-
ing modification of mature and secondary forests (Esselstyn 2007; Heaney et
al. 1998; Mickleburgh et al. 1992; Utzurrum 1992). Deforestation of lowland
areas, where pteropodid diversity is greatest (Utzurrum 1998), is one of the
chief reasons for the declines of many taxa, especially Acerodon jubatus, Dob-
sonia chapmani, Nyctimene rabori, and Pteropus leucopterus (Heaney and Heide-
man 1987; Heaney et al. 1998; Heaney et al. 1999; Stier and Mildenstein 2005).
Populations of species inhabiting middle or higher elevation forests, such as
Haplonycteris fischeri, Harpyionycteris whiteheadi, and Otopteropus cartilagonodus,
are thought to be more stable, but remain vulnerable to changes in extent and
quality of habitat (Heaney et al. 1998). Observations by Paalan et al. (2004)
suggest that a number of Philippine pteropodids, including some threatened
endemics, are somewhat tolerant of moderate forest fragmentation. At least 29
of the country’s 52 microchiropteran species also inhabit forest, and many are
undoubtedly affected by habitat loss. Heaney et al. (1998) reported that the cut-
ting of large hollow trees during logging has caused substantial harm to some
rhinolophids and hipposiderids, particularly those inhabiting lowland diptero-
carp forests. Populations of cave-dwelling species (e.g., Hipposideros bicolor, H.
pygmaeus, and Miniopterus schreibersii) may be decimated by the elimination of
forested foraging habitat near roost caves (Heaney et al. 1999).
Deforestation is a known or potential concern for at least 8 of 37 megachi-
ropterans and 1 microchiropteran in Papua New Guinea (Bonaccorso 1998).
Intensive logging on New Britain and New Ireland threatens several species
with relatively small geographic distributions, including Dobsonia praedatrix,
Pteropus capistratus, and Kerivoula myrella (Bonaccorso 1998). In the Solomon
Island chain, habitat disturbance has been implicated in the possible extinction
of Nyctimene sanctacrucis (Mickleburgh et al. 1992), and timber harvest threat-
ens several rare species of Pteralopex that are heavily reliant on primary forests
(Bowen-Jones et al. 1997; Fisher and Tasker 1997; Helgen 2005). Pteralopex taki
roosts in the hollows of large-diameter trees and is therefore vulnerable to
414 G. J. Wiles and A. P. Brooke
selective timber cutting, as evidenced by its apparent extirpation from the
island of Kolombangara following extensive logging operations in the 1970s
(Fisher and Tasker 1997; Flannery 1995).
Although total forest coverage has changed little in Fiji and Samoa since
1990, there has been a significant qualitative shift toward increasing amounts
of secondary forest and disturbed forest dominated by introduced species
( J. Atherton, pers. comm.). Habitat concerns are a conservation issue for all six
bat species present in the two island groups (Palmeirim et al. 2005; Palmeirim
et al. 2007; Wilson and Engbring 1992). Pteropus samoensis is especially depen-
dent on tracts of native forest to meet its foraging and roosting needs, but even
P. tonganus, which feeds more extensively in disturbed areas, has lost habitat
with the conversion of lands to grasslands, sugarcane plantations, and other
open sites (Banack 1998; Palmeirim et al. 2005; Palmeirim et al. 2007; Wilson
and Engbring 1992). Mirimiri acrodonta is restricted to a small area of montane
forest on Taveuni, Fiji, most of which is secure within Bouma National Heritage
Park. However, further forest clearing on unprotected lower slopes in the fu-
ture could result in more foraging by P. tonganus at higher elevations, thereby
increasing competition for M. acrodonta (Palmeirim et al. 2005). On a localized
scale, Palmeirim et al. (2005, 2007) noted that the removal of large overstory
trees outside the mouths of caves used by Emballonura semicaudata can promote
the growth of shrubby vegetation, thereby blocking entrances and preventing
bats from entering.
In Micronesia the largest anthropogenic forest loss on any island during
the past several decades has occurred on Pohnpei, where more than 70% of
the remaining upland native forest has been destroyed or heavily degraded
since 1975 by intensified cultivation of the shrub Piper methysticum (Merlin
and Raynor 2005), which is used to produce the mildly narcotic drink known
as kava or sakau. About 120 km2 of upland forest was lost by 2002, represent-
ing a decline from 42% to 15% of the island’s land cover. The impacts of such
loss on Pohnpei’s two bats, Pteropus molossinus and Emballonura semicaudata,
are unknown but may be moderately severe. On Aguiguan in the Common-
wealth of the Northern Mariana Islands, the occurrence of E. semicaudata is
probably closely linked to the island’s remaining forest cover (Esselstyn et al.
2004b). Nonnative ungulates have had an important role in damaging forests
and reducing forest cover on a number of the Marianas (Kessler 2002; Wiles
et al. 1999; Worthington et al. 2001).
Although large-scale deforestation is certainly harmful to most bat species
in insular Southeast Asia and Oceania, many taxa are in fact tolerant of lim-
ited anthropogenic habitat modification. This probably results from the long
history of human disturbance to native forests in much of the region (e.g.,
Bayliss-Smith et al. 2003; Mercado 2003) and, for megachiropterans, to the
often-shared fruit preferences among bats and people (Marshall 1983; Wiles
and Fujita 1992). Near human settlements, sizable areas of forest have long been
Conservation Threats to Bats in the Pacific and Southeast Asia 415
converted to agroforest, where tree crops such as breadfruit, coconuts, man-
gos, avocados, bananas, and numerous other species are interspersed among
native trees. Coconut plantations, another food source for pteropodids, have
also been widely established for commercial purposes. More than half of the
96 pteropodid species occurring in New Guinea, the Moluccas, and Oceania
regularly enter younger secondary forests, gardens, and plantations to feed
(Bonaccorso 1998; Flannery 1995; Helgen 2007). This number is lower in the
Philippines, where only 7 of 26 megachiropterans visit these types of heavily
altered habitats (Heaney et al. 1998). Several widespread taxa (e.g., Cynopterus
brachyotis, Eonycteris spelaea, Macroglossus minimus, Pteropus hypomelanus, and
Rousettus amplexicaudatus) are particularly common in agricultural lands, ur-
banized areas, and disturbed forests, indicating that they have benefited from
large-scale habitat change (Abdullah et al. 1997; Hall et al. 2004; Heaney et al.
1998; Helgen 2007; Mickleburgh et al. 1992; Mohd-Azlan et al. 2003). Examples
of species known to forage extensively in agroforest include Melonycteris mela-
nops (Bonaccorso 1998; Flannery 1995), Macroglossus minimus (Bonaccorso 1998;
Flannery 1995), Pteropus tonganus (Banack 1998; Banack and Grant 2003; Nelson
2003; Palmeirim et al. 2005), and P. mariannus yapensis (Falanruw 1988).
By virtue of their many caves and fissured cliffs, limestone karst landscapes
are of inherent importance to bat populations across much of the region (e.g.,
Alcala et al. 2004; Lee et al. 2007; Suyanto and Struebig 2007). However, eco-
nomic expansion has brought increasing destruction of these ecologically sensi-
tive environments (Clements et al. 2006; Whitten 2002). Karsts are most threat-
ened by quarrying for limestone, which is used in the manufacturing of cement
and other products. Wildfires and deforestation are additional concerns.
A number of nations in the region, especially Indonesia, the Philippines,
Papua New Guinea, the Solomons, Vanuatu, Japan, and the Northern Mari-
ana Islands, are positioned along the edges of tectonic plates and experience
regular volcanic activity. The impacts of volcanism on bat populations in
these countries have been rarely described, but major eruptions resulting in
the destruction of forested habitats can eliminate or reduce populations on a
local scale (also see Pedersen et al., chapter 11, this volume). For example, in
the Northern Marianas, recurring volcanic activity since 2003 has eliminated
nearly all forest and flying foxes on the island of Anatahan (C. C. Kessler,
pers. comm.). This island, which is 32 km2 in area, was considered one of the
most important remaining sites for P. mariannus in the 1980s and 1990s (Wiles
et al. 1989; Worthington et al. 2001). Eruptions on other Mariana Islands have
reduced the amount of forest available for bats during the past few centuries
(Wiles et al. 1989). Tidemann et al. (1990) described the gradual recolonization
of the Krakatau Archipelago and nearby parts of Java by at least 11 species of
pteropodids and 20 microchiropterans following the cataclysmic eruption in
1883 (also see Shilton and Whittaker, chapter 7, this volume). Long Island off
eastern New Guinea lost its biota in a similar destructive eruption in about
416 G. J. Wiles and A. P. Brooke
1645 and has since been reinhabited by six pteropodids and a single microchi-
ropteran (Thornton et al. 2001).
Hunting
Flying foxes and other bats continue to be a traditional food and delicacy on
many islands, although not on all. Historically, flying foxes were taken with a
variety of tools and techniques, such as stone projectiles, sticks, long-handled
hoop nets, tree platforms, bows and arrows, blowguns, and thorny vines as
the bats came to feed at fruiting or flowering trees or flew near the ground at
certain locations along flyways (e.g., Chambers and Esrom 1988; Fritz 1904;
Kubary 1885; Loeb 1926; MacGillivray 1860). Cave-dwelling pteropodids (e.g.,
Dobsonia moluccensis and Notopteris macdonaldi) were sometimes caught by
blocking cave entrances with vegetative barricades or fire, and then capturing
the animals trapped inside by various means (Dwyer 1968; Palmeirim et al.
2005). Harvesting of bats sometimes involved considerable ritual (Falanruw
1988). As with many birds, aboriginal hunting pressure probably contributed
to the extinctions of some megachiropteran species or populations, although
habitat loss and ecological changes associated with human settlement were also
likely involved (Koopman and Steadman 1995; Steadman 1995, 2006; Weisler
et al. 2006).
Within the past 50 years, improved road and boat access to remote areas;
the wider availability of firearms, air rifles, and nets; and fewer cultural re-
strictions have enabled hunters to take increased numbers of flying foxes and
other bats relatively easily. In some locations, growing human populations
and increased access to markets have also produced greater demand for bats.
Overhunting for both subsistence and commercial purposes is considered the
greatest threat to larger pteropodids on many islands, including some with few
or no people (Brooke and Tschapka 2002; Cousins and Compton 2005; Craig et
al. 1994a; Heaney and Heideman 1987; Pierson et al. 1996; Riley 2002b; Stinson
et al. 1992; Wiles 1992; Wiles et al. 1989; Wiles et al. 1997). Unfortunately, in
many areas where these bats were historically abundant, there are no data
on population size to evaluate the impacts of hunting. To our knowledge,
the hunting of flying foxes and other pteropodids is not managed sustainably
anywhere in the region.
Detailed information on the extent of hunting is rarely available for any
population, and no study has monitored long-term harvest levels. Most data
have been collected via interviews or questionnaires of hunters and nonhunt-
ers. One of the best documented cases is for Niue, where an annual hunting
season for Pteropus tonganus lasts 2–4 months. In 1998, field surveys estimated
the flying fox population at between 2,000–4,000 animals. Interviews after the
hunting season estimated that a quarter to half of the bat population was killed
(Brooke and Tschapka 2002). Based on the population’s size and the reproduc-
Conservation Threats to Bats in the Pacific and Southeast Asia 417
tive potential of the species, the number of bats removed was clearly unsus-
tainable. Widespread misperceptions regarding the actual abundance of bats
played a significant role in overhunting. Older hunters remembered a much
larger bat population in the past and supported limits on hunting, but many
others believed an infinite number of bats roosted in areas protected from
hunting by a taboo and consequently thought the hunt could continue with
no impact on the bat population.
Annual harvests of Pteropus tonganus and P. samoensis on Tutuila, American
Samoa, were estimated to total 500–1,600 animals under normal conditions
during the early 1990s (Craig et al. 1994a). However, the increased vulnerability
of bats after Cyclone Ofa resulted in an estimated 3,100 bats being killed within
six months of the storm.
In Palau, Wiles et al. (1997) estimated roughly that up to 2,000–5,000 P. mari-
annus pelewensis were killed annually and that as many as 40–50% of Palauans
occasionally ate bats. Harvest levels of P. tonganus are also high in the Cook
Islands, although Cousins and Compton (2005) did not estimate total take.
Based on questionnaire results, they determined that 8% and 20% of adult re-
spondents on Rarotonga and Mangaia, respectively, hunted flying foxes more
than twice per year. The majority of people had little idea of the threat that
hunting posed to local bat populations, but most supported restrictions if de-
clining numbers of animals could be demonstrated.
In parts of northern Sulawesi, Indonesia, bats are hunted as a source of
meat for family and neighbors and perhaps to limit crop depredation (Lee
2000). Bats, presumably mostly pteropodids, were the second most commonly
harvested taxonomic group after rodents, comprising 19–25% of the total wild
animals caught in and around two protected areas. Thousands were taken
annually and greatly exceeded ungulates, primates, and cuscuses in total bio-
mass. Subsistence hunting of pteropodids also occurs in Java (Fujita and Tuttle
1991), the Sangihe and Talaud Islands (Riley 2002b), Kalimantan (Mohd-
Azlan et al. 2003), northern Sumatra (G. Fredriksson, pers. comm.), and
probably many other parts of the country.
Pteropodids, including Eonycteris spelaea and Acerodon leucotis, were
among the more commonly caught animals by an impoverished commu-
nity of shifting cultivators in southern Palawan, Philippines (Shively 1997).
Bats and other small game species were exploited on a subsistence basis
and were probably targeted because they provided greater hunting success
in comparison to larger mammals. Small pteropodids were hunted by 14%
of households, with members of those households making an average of
20 trips per year to seek bats and catching a mean of 11.7 bats per trip.
Among the 9% of households that hunted Acerodon leucotis, a mean of 15
trips per household was made annually to obtain this species, with 3.9 bats
caught per trip on average. Blowguns and poles armed with thorns or fish-
hooks were the primary capture methods. In a second Philippine study,
418 G. J. Wiles and A. P. Brooke
bats comprised 11 of the 72 bird and mammal species killed by hunters on
Negros (Cariño et al. 2006). Three threatened species (A. jubatus, Dobsonia
chapmani, and Pteropus pumilus) were among those captured. Although
harvest rates were not calculated, bats collectively were regularly hunted.
Most hunters were subsistence farmers who tended to be poorly educated,
earned low incomes, and killed bats and other wildlife primarily for home
consumption. Remaining hunters were mostly better-educated professional
or skilled workers who hunted for recreation. The main capture methods
in this study were nylon lines with hooks and air guns.
Megachiropterans are widely hunted in both the Philippines and Papua
New Guinea, where at least 14 of 26 species and 12 of 37 species are caught,
respectively (Bonaccorso 1998; Esselstyn 2007; Flannery 1990; Heaney et al.
1998; Mickleburgh et al. 1992; Shively 1997; Utzurrum 1992). Even small spe-
cies such as Syconycteris australis are sometimes taken (Craven 1988). Hunting
in these countries seriously threatens at least three species, Acerodon jubatus,
Aproteles bulmerae, and Dobsonia chapmani (Alcala et al. 2004; Cariño 2004; Flan-
nery and Seri 1993; Heaney and Heideman 1987; Heaney et al. 1998; Paguntalan
et al. 2004), and possibly a fourth, Styloctenium mindorensis (Esselstyn 2007).
Although most species in Papua New Guinea are harvested for their meat,
Dobsonia inermis and Melonycteris woodfordi are collected for their canine teeth,
which are used in ornamental necklaces (Bonaccorso 1998).
In Vanuatu, Chambers and Esrom (1988) reported that flying foxes were a
common part of the diet and that most people ate them at least occasionally.
Recent evidence from Fiji suggests that hunting pressure differs among spe-
cies. Palmeirim et al. (2005) reported that hunting of Pteropus tonganus and P.
samoensis may be declining because of greater access to alternative forms of pro-
tein through modern commerce. However, exploitation remains a concern for
Notopteris macdonaldi and may be causing some populations in Fiji to decline. In
the Mariana Islands, hunting of P. mariannus continues on most islands despite
legal protection for the species (Lemke 1992; Wiles et al. 1989; Worthington et
al. 2001; G. J. Wiles, pers. obs.). Bats are regularly killed on islands with larger
numbers (i.e., Rota and islands in the northern half of the island chain) and
then smuggled to the main inhabited islands of Saipan, Tinian, and Guam.
In 2006 and 2007, bats sold for US$40 on Rota (P. Wenninger, pers. comm.)
to as much as $100 per animal on Tinian and Saipan. Illegal hunting is also
widespread for P. ornatus in New Caledonia (Flannery 1995). Hunting has
probably caused the near extinction of the subspecies P. dasymallus formosus,
which survives only on Lutao Island (also known as Green or Kashoto Island)
off Taiwan (Rainey 1998).
Hunting pressure has not been well-documented in the Solomon Islands,
but at least one species, Pteropus rayneri, is hunted heavily (Flannery 1995).
However, other reports indicate that most people apparently consume ptero-
podids only on an infrequent basis (Bowen-Jones et al. 1997; Fisher and Tasker
Conservation Threats to Bats in the Pacific and Southeast Asia 419
1997; Whewell 1992). In Tonga, harvests of P. tonganus appear to be relatively
small and of no threat to populations (K. R. McConkey, pers. comm.), and on
Ulithi Atoll in the Carolines, only small numbers of people hunt and eat flying
foxes (Wiles et al. 1991). On some islands, such as Chuuk, Pohnpei, and Kosrae,
bats are not hunted for local use (Rainey 1990).
Cultural factors are sometimes involved in the hunting of pteropodids on
islands where people have retained more of their traditional values. For ex-
ample, on Yap, where flying foxes are considered less desirable than marine
foods, only people of certain social strata with limited or no access to the coast
hunt flying foxes (Falanruw 1988; M. V. C. Falanruw, pers. comm.). Religious
and cultural beliefs prevent some people from eating or catching flying foxes in
Vanuatu (Bani 1992; Chambers and Esrom 1988). Localized superstitions also
afford some protection to several pteropodids in Papua New Guinea (Flannery
and Seri 1990). Traditional hunting methods continue to be employed in some
remote locations (Chambers and Esrom 1988; Palmeirim et al. 2005).
Sizable commercial markets for Pacific flying foxes became established on
Guam during the 1960s and in the neighboring Northern Mariana Islands prob-
ably sometime between the late 1970s and 1985 (Stinson et al. 1992; Wiles 1992;
Wiles and Payne 1986; Wiles et al. 1997). This trade resulted in the importation
of between 7,600 and 29,500 bats annually (mean = 13,960) to Guam from 1975
to 1994, although estimates exceeding 19,000 animals per year from 1976 to
1980 are very likely inflated. Smaller numbers of flying foxes were shipped to
the Northern Marianas, ranging from 750 to 8,600 bats annually (mean = 4,682)
from 1986 to 1994. Flying foxes from 11 islands or island groups dominated
the trade, with those in Palau, Yap, Pohnpei, Chuuk, Samoa, American Samoa,
and perhaps Kosrae seriously depleted for several years or longer (Rainey
1990; Wiles 1992; Wiles et al. 1997). During the 1970s and early 1980s, bats were
sent to Guam mainly from nearby islands (i.e., Palau, Yap, Rota, Saipan, and
Tinian). However, depletion of many of these sources and changing business
factors led to greater exploitation of bat populations on more distant islands
(i.e., Samoa, American Samoa, Pohnpei, Tonga, Papua New Guinea, and the
Philippines) from about 1982 to 1990 (Wiles 1992). This international trade
was greatly restricted in 1989 after seven of the species involved were added
to appendix I of the Convention on International Trade in Endangered Spe-
cies of Wild Fauna and Flora (CITES), and all remaining Pteropus and Acerodon
were placed on appendix II. However, trade continued until 1994 because of
a loophole that allowed the importation of Palauan bats (Wiles 1994). Little
international smuggling of bats has occurred since then, and it appears that
CITES restrictions have been highly successful in terminating this trade (G. J.
Wiles, pers. obs.). Several important factors facilitated the existence of this
trade, including the expansion of interisland airline traffic, greater availability
of refrigeration, and the greater affluence of Chamorro consumers in Guam
and the Northern Marianas (Wiles and Payne 1986). Elsewhere in the Pacific,
420 G. J. Wiles and A. P. Brooke
small-scale shipments of Pteropus from Vanuatu to New Caledonia were docu-
mented in the 1990s (Rainey 1998).
Intensive hunting to supply market demand is a major threat to flying
foxes and other pteropodids in parts of Indonesia, especially northern Su-
lawesi and Kalimantan (Bergmans and Rozendaal 1988; Clayton and Milner-
Gulland 2000; Fujita and Tuttle 1991; Lee et al. 2005; Struebig et al. 2007). Five
species, Pteropus vampyrus, P. hypomelanus, P. alecto, Acerodon celebensis, and
A. humilis, have been exploited in alarmingly high numbers, resulting in large
population declines (Fujita and Tuttle 1991; Lee et al. 2005; Struebig et al. 2007).
Muslim religious beliefs protect many wildlife species from being hunted, but
in areas with non-Muslim populations, flying foxes have been intensively
hunted as a delicacy and a medicinal. Bat meat and liver are valued as a cure
for kidney ailments, general malaise, and respiratory problems, particularly
by ethnic Chinese (Fujita and Tuttle 1991). North Sulawesi is a major center
of commercial bat harvest, with at least 13 species of 11 pteropodid genera
sold (Bergmans and Rozendaal 1988). Huntable populations of bats report-
edly have been eliminated over much of the peninsula (Argeloo 2001 cited in
Bergmans 2001; Lee et al. 2005), and hunting to supply demand occurs increas-
ingly farther from Manado and other cities (Bergmans 2001; Lee et al. 2005;
Riley 2002a, 2002b). Market surveys indicate that although large numbers of
bats are sold, they typically comprise just a small proportion of the total rev-
enue earned by wild meat dealers (Clayton and Milner-Gulland 2000). Riley
(2002b) described aspects of the trade in the Sangihe and Talaud Islands. In
portions of Kalimantan, demand for flying foxes has expanded greatly since
the mid-1990s, especially in the city of Palangkaraya (Struebig et al. 2007).
In peat swamp forests near the city, hunters captured over 4,500 P. vampy-
rus during a single month in 2003, when animals became seasonally common
(Struebig et al. 2007). Hunters observed noticeable declines in catches between
the early to mid-1990s and 2005, and extended their harvest activities to more
distant locations. Significant numbers of Pteropus are also marketed in Jakarta
(Fujita and Tuttle 1991), and commercial harvesting of Pteropus and Rouset-
tus has been recently observed in northern Sumatra (G. Fredriksson, pers.
comm.).
In Sarawak, hunting is thought to be an important cause in the decline
of Pteropus vampyrus (M. Gumal, pers. comm. in Hall et al. 2002), with large
numbers being sold in the capitol of Kuching during the late 1980s (Fujita and
Tuttle 1991). Heavy market hunting may have eliminated a colony of about
12,000 Eonycteris spelaea from Niah Cave during the 1990s (Hall et al. 2002). In
this case, trapping occurred inside the cave, a designated national park. This
species is caught at other caves in Sarawak as well.
Bats are occasionally sold in local markets throughout the Philippines (T. L.
Mildenstein, pers. comm.; L. R. Heaney, pers. comm.), but the extent of this
trade is poorly documented. Numbers are probably much smaller than the
Conservation Threats to Bats in the Pacific and Southeast Asia 421
number killed for personal consumption. Acerodon jubatus and Pteropus vampy-
rus are among the species sold (Cariño 2004; Cayunda et al. 2004).
Smaller commercial markets catering to local demand exist elsewhere. For
example, in Palau, a significant portion of the Pteropus mariannus pelewensis
killed are sold in town rather than being eaten by the hunter’s family (G. J.
Wiles, pers. obs.), whereas in Vanuatu, only small numbers are available in
markets (Chambers and Esrom 1988). In both island groups, a few restaurants
serve small numbers of flying foxes to tourists (Chambers and Esrom 1988;
Wiles et al. 1997), and one Vanuatu hotel advertises bat hunting on its web site.
At least two species of megachiropterans, P. neohibernicus and Dobsonia inermis,
are sold locally in Papua New Guinea (Bonaccorso 1998). There is minor de-
mand for P. tonganus in Fiji, with hunters occasionally selling animals directly
to customers (Palmeirim et al. 2005; Palmeirim et al. 2007).
The extent that microchiropteran populations in the region are hunted for
food or for medicinal purposes is poorly understood, but a number of species
are probably caught. Miniopterus species and other cave-dwelling insectivorous
bats are collected from accessible roost sites by traditional peoples in New
Guinea (Craven 1988; Flannery and Seri 1990). Bergmans and Rozendaal (1988)
occasionally found microchiropterans, including Cheiromeles parvidens, being
sold in markets in northern Sulawesi. Hunting is considered a contributing
factor in the declines of Cheiromeles torquatus in Borneo (Hall et al. 2002; Hutson
et al. 2001) and Chaerephon bregullae in Fiji (Palmeirim et al. 2005; Palmeirim et
al. 2007). Species roosting in bamboo culms (possibly Tylonycteris) and furled
banana leaves (Myotis muricola) are eaten in Kalimantan (Mohd-Azlan et al.
2003). Hutson et al. (2001) remarked on the heavy harvest of insectivorous
bats in the Solomon Islands, but did not provide details. Scotophilus kuhlii is
sometimes harvested in the Philippines, as documented by the presence of 64
animals in a shipment mistakenly sent to Guam in 1987 during the flying fox
trade (G. J. Wiles, unpublished data).
Cave Disturbance
Natural and artificial caves provide permanent shelter for numerous bat spe-
cies in the region. Many locations serve as traditional day roosts and mater-
nity sites, and may be inhabited by more than one species. Sites can support
anywhere from one or a few individuals to colonies of thousands. Suitable
caves are frequently limited in availability; thus animals often display strong
attachment to particular sites. As commonly noted elsewhere in the world
(Hutson et al. 2001), sustained disturbance at caves can result in severe long-
term impacts on the viability of resident bat populations. In insular Southeast
Asia and the tropical Pacific, cave-dwelling bats are vulnerable to many forms
of human disturbance, such as guano mining, the collection of edible swiftlet
(Aerodramus spp.) nests, hunting within caves, rock quarrying, and visitation
422 G. J. Wiles and A. P. Brooke
by tourists, cavers, and vandals (Hutson et al. 2001). Rates of cave disturbance
are considered especially high in the Philippines (Cariño 2004; Hutson et al.
2001), but this is true of other areas as well. Indonesia and Malaysia are by far
the largest sources of swiftlet nests for the international nest trade (Lau and
Melville 1994), with many caves in both countries experiencing heavy harvest
pressure. In the past, nest gathering at some localities (e.g., parts of Sarawak)
was traditionally restricted to two or three relatively short periods per year.
However, expansion of the trade in recent decades (e.g., nest imports in Hong
Kong increased threefold between 1959 and 1988) has led to greater levels of
visitation by collectors, many of who now work year-round (Lau and Melville
1994). This has undoubtedly resulted in significantly more disturbance of bats
sharing the same caves.
The region’s cave-dwelling fruit bats include Aproteles bulmerae, Penthetor
lucasi, and species of Rousettus, Dobsonia, Eonycteris, Notopteris, Ptenochirus,
and Cynopterus. Most species roost in the dimly lit portions of caves or be-
neath overhangs at cave entrances, but Rousettus are able to inhabit the deeper
reaches of caves because of their ability to echolocate. To date, human activity
at caves has been identified as a serious threat to A. bulmerae, D. anderseni,
D. chapmani, E. spelaea, N. macdonaldi, N. neocaledonica, R. amplexicaudatus, and
possibly D. inermis (Bonaccorso 1998; Heaney et al. 1998; Heaney et al. 1999;
Mickleburgh et al. 1992; Palmeirim et al. 2007). Two of the primary factors
for the rarity of D. chapmani are cave disturbance from guano mining and the
harvesting of animals inside caves (Heaney and Heideman 1987; Paguntalan
et al. 2004). These same problems, plus limestone mining at caves, threaten E.
spelaea in the Lesser Sundas of Indonesia (Mickleburgh et al. 1992). The remain-
ing populations of N. macdonaldi in Fiji and N. neocaledonica in New Caledonia
are both known from small numbers of caves and are therefore particularly
vulnerable to disturbance (Flannery 1995; Mickleburgh et al. 1992; Palmeirim
et al. 2005; Palmeirim et al. 2007). Dwyer (1968) described the disruptive activi-
ties that traditional hunters can have at caves occupied by megachiropterans,
in this case D. moluccensis.
Only anecdotal accounts seem to exist describing the effects of human cave
visitation on the region’s microchiropterans. In the Philippines, populations
of many rhinolophid species and the molossid Chaerephon plicatus have been
harmed by cave disturbance, with extirpation of some colonies occurring for C.
plicata (Heaney et al. 1998). Heaney et al. (1999) reported that few if any caves
in the vicinity of Mount Isarog, Luzon, have escaped frequent disturbance
from guano miners and bat hunters. Recreational caving is also considered a
growing problem in the country (Hutson et al. 2001). Swiftlet nest collecting
is a major disruptive factor for cave bats on the Sangkulirang peninsula of
eastern Kalimantan (Suyanto and Struebig 2007) and is partially responsible
for the large declines of Cheiromeles torquatus in Sarawak (Hutson et al. 2001).
Hsu (1997) identified the closure of caves by farmers during the expansion of
Conservation Threats to Bats in the Pacific and Southeast Asia 423
croplands as the main threat to bats in Kenting National Park on Taiwan. On
Iriomote island in the Ryukyu Islands, publicity on the threatened status of Hip-
posideros turpis produced the undesired outcome of greater public visitation of
the caves occupied by the bats (Hutson et al. 2001). During World War II, major
cave disturbances and closures must have negatively impacted Emballonura
semicaudata and other insectivorous species in Micronesia and other parts of
the Pacific war theater. However, cave disturbance by people is not currently
considered an important threat to E. semicaudata (Grant et al. 1994; Palmeirim
et al. 2005; G. J. Wiles, unpublished data). Although not directly related to hu-
man visitation, forest fires of probable anthropogenic origin have burned into
ground-level caves and apparently eliminated resident bat colonies in eastern
Kalimantan (Suyanto and Struebig 2007).
Severe Storms
Severe tropical cyclonic storms, known variously as typhoons, cyclones, or
hurricanes, are regular occurrences on many Pacific islands, the Philippines,
and Taiwan. Most storms produce relatively minor localized damage to natural
ecosystems on affected islands and have few if any impacts on bat populations.
However, exceptionally strong storms occasionally cause far more damage to
forests, which may not recover for several years or longer. The effects of such
storms on bat populations can be disastrous, as evidenced by the substantial
reductions in flying fox abundance at various locations in both the Pacific and
Indian Oceans (Pierson and Rainey 1992). This is especially true in areas of
extensive deforestation, where storms may leave no refugia for bats (Pierson
et al. 1996). In American Samoa and Samoa, Pteropus tonganus and P. samoensis
decreased by an estimated 80–99% following Cyclone Ofa in 1990 and Val in
1991 (Craig et al. 1994b; Pierson et al. 1996). A similar overall decline of about
80% in P. tonganus occurred in the Vava’u archipelago of Tonga following
Cyclone Waka in 2001 (McConkey et al. 2004). On the island of Rota in the
Northern Marianas, numbers of P. mariannus were reduced by an estimated
57% after Typhoon Roy in 1988 (Stinson et al. 1992) and 70% after Typhoon
Pongsona in 2002 (Esselstyn et al. 2006). Storm-related losses have been re-
ported, but not quantified, from other locations, including Niue (P. tonganus,
Cyclone Heta in 2004, Anonymous 2005), the Solomon Islands (P. rayneri and
P. tonganus, Cyclone Namu in 1986, Flannery 1989; Pteropus sp., Cyclone Ida
in 1972, Bowen-Jones et al. 1997), Vanuatu (Pteropus sp., Chambers and Esrom
1988), Fiji (P. tonganus and P. samoensis, Palmeirim et al. 2005; Palmeirim et al.
2007), and Guam (P. mariannus, Wiles 1987b).
Accounts indicate that flying foxes usually suffer much greater mortality
during the several months following tropical cyclones than from high winds
and flying debris during storm passage. Poststorm mortality typically results
from starvation, dehydration, overhunting for food and recreation, and
424 G. J. Wiles and A. P. Brooke
predation. Extensive defoliation of forests during severe storms not only
greatly reduces food availability, but also increases vulnerability to hunting
by removing protective cover. Starvation can drive bats to forage diurnally in
places they would otherwise avoid, such as on or near the ground and in areas
of human activity (e.g., near villages and in plantations). This further exposes
animals to human hunting as well as predation. The impacts of cyclone-caused
resource scarcity on flying fox populations are probably density-dependent,
with smaller (i.e., less dense) populations less likely to be affected. Pierson et al.
(1996) observed that P. tonganus in Samoa and American Samoa displayed far
fewer overt signs of starvation after Cyclone Val than when populations were
much larger two years earlier following Cyclone Ofa. Guam’s small population
of P. mariannus showed no evidence of food stress after several major typhoon
strikes during the 1990s (G. J. Wiles, pers. obs.) and another in 2002 (Esselstyn et
al. 2006). This was also true on Rota, where P. mariannus occurs below carrying
capacity as well, after two of three large typhoons since 1988 (Esselstyn et al.
2006). However, when combined with restricted ranges or forest availability,
even small populations can become vulnerable to extirpation following the
worst storms (Palmeirim et al. 2005; Robertson 1992).
Starvation and ground foraging after tropical cyclones have been described
by a number of authors (Anonymous 2005; Craig et al. 1994b; Daschbach 1990;
Esselstyn et al. 2006; Flannery 1989; McConkey et al. 2004; Palmeirim et al. 2005;
Pierson et al. 1996), and flying foxes “blown to the ground” in Vanuatu (Cham-
bers and Esrom 1988) were probably in fact animals searching for food. This
type of feeding behavior can result in considerable predation by cats, dogs, and
domestic pigs, and even mortality from vehicles on roadways (Anonymous
2005; Palmeirim et al. 2007; Pierson et al. 1996).
Intensive harvest of food-stressed flying foxes has been documented on
several islands after major tropical cyclones (Pierson and Rainey 1992). Hunt-
ing of P. tonganus and P. samoensis was severe in Samoa and American Samoa
after Cyclone Ofa (Pierson et al. 1996), with data suggesting that it was respon-
sible for about half of all mortality in American Samoa (Craig et al. 1994b).
Many bats were apparently killed by boys for recreation rather than for food
(Daschbach 1990). Stinson et al. (1992) reported that hunting accounted for
roughly two-thirds of the decline in P. mariannus on Rota following Typhoon
Roy, with hunting-related emigration to neighboring islands responsible for
the remaining losses. Intensified poaching also occurred on Rota after Typhoon
Pongsona and again caused the majority of the population’s decrease (Essel-
styn et al. 2006). The capture of large numbers of ground-foraging flying foxes
after cyclones has been noted in Vanuatu (Chambers and Esrom 1988) and the
Solomons (Flannery 1989). Hunting was also detected in Tonga after Cyclone
Waka (McConkey et al. 2004). On Niue, Cyclone Heta struck during the annual
hunting season for P. tonganus and apparently exacerbated harvest mortality
(Brooke 2004).
Conservation Threats to Bats in the Pacific and Southeast Asia 425
Direct storm mortality has been documented in only a few instances. S.
Campbell (pers. comm.) observed 40–50 dead P. tonganus washing ashore in
a small bay following Cyclone Waka in Tonga, indicating that animals were
blown into the sea and drowned (McConkey et al. [2004] reported this as “large
numbers”). Wiles (1987b) presented secondhand evidence of dead P. mariannus
found under roost trees on Guam.
Flowers and fruits from plants that are sturdy enough to sustain relatively
less damage from high winds have been identified as important foods for fly-
ing foxes in the first few months following severe tropical cyclones. Examples
include coconut flowers (McConkey et al. 2004; Pierson et al. 1996), flowers
and fleshy bracts from the woody liana Freycinetia reineckei (Pierson et al. 1996),
and Pandanus fruits (Stinson et al. 1992). Emerging leaves and petioles can also
be important soon after storms (Nelson et al. 2000b; Pierson and Rainey 1992).
Although forest recovery usually proceeds fairly rapidly, it can lag substan-
tially after the worst of storms (Elmqvist et al. 1994). In Tonga, McConkey et
al. (2004) observed that food resources for flying foxes were still reduced by
85% or more six months after the passage of Cyclone Waka.
Damage from severe tropical cyclones can be highly disruptive to normal
foraging and roosting behavior in flying foxes. Food-stressed individuals may
respond by becoming more active for longer periods while searching for food
and increasing their daytime foraging effort (Esselstyn et al. 2006; Grant et al.
1997; Pierson et al. 1996; Stinson et al. 1992). Dramatic changes in roosting pat-
terns may result from damage to roost trees, the sudden loss of food sources,
and hunting and other human activity, or a combination thereof. Brooke et
al. (2000) reported that colonies of P. tonganus broke into smaller groups and
moved frequently among alternate sites after cyclones in American Samoa.
This was probably caused initially by heavy damage to primary roost trees,
but hunting and greater scarcity of food resources also likely contributed. Colo-
nies coalesced and returned to prestorm roost sites on steep mountain slopes
and cliff faces within four years. Stinson et al. (1992) observed a substantial
increase in the number of solitary P. mariannus after Typhoon Roy on Rota
in the Northern Marianas and attributed this to the greater effort needed to
find food. However, repeated hunting incidents at the main roosts also kept
colonies broken apart. In contrast, the single small colony of P. mariannus on
Guam temporarily doubled in size the month after Typhoon Pongsona hit the
island in 2002, most likely because of immigration from the neighboring island
of Rota (Esselstyn et al. 2006).
Impacts of tropical cyclones on microchiropterans are poorly described. In
American Samoa, Grant et al. (1994) reported that storm-generated waves from
Cyclone Ofa may have destroyed nearly all of the bats in one of the few re-
maining colonies of Emballonura semicaudata, which resided in a sea cave. These
authors also speculated that intense stormy weather during the four days of
Cyclone Val’s passage possibly prohibited foraging and perhaps caused some
426 G. J. Wiles and A. P. Brooke
animals to starve. Tarburton (2002) attributed the loss of one of the few known
remaining colonies of E. semicaudata in Samoa to these same cyclones.
Tropical cyclones in the Pacific basin have grown in destructive power,
but not frequency, since the mid-1970s in response to global climate change
(Emanuel 2005). This trend, if it continues, will undoubtedly have major effects
on many of the region’s bat populations (Rainey 1998).
Introduced Species
Although the negative impacts of invasive species are commonly observed
among island ecosystems (Courchamp et al. 2003; D’Antonio and Dudley 1995;
Sherley 2000; Veitch and Clout 2002), relatively few harmful interactions be-
tween exotics and tropical Pacific bats have been described thus far. Probably
the greatest effect occurs indirectly through perturbations to native habitats,
thereby altering food availability for bats. A number of exotic species estab-
lished in the region have produced large-scale changes in forest composition
or reduced the abundance of particular plant species. Ungulates have been in-
troduced to relatively few islands with pteropodids but have caused significant
damage to ecosystems on those where they have become well established. On
some islands in the Marianas, high densities of feral pigs (Sus scrofa), feral goats
(Capra hircus), and Philippine deer (Rusa marianna [formerly Cervus mariannus])
have been linked to reduced plant diversity in forests and declines of various
food species (e.g., Artocarpus mariannensis, Pandanus tectorius, Premna obtusifo-
lia, Pipturus argenteus, Dendrocnide latifolia, and probably many others; Kessler
2002; Wiles 2005b; Wiles et al. 1999; Worthington et al. 2001; G. J. Wiles, pers.
obs.). Ungulate damage to forests has also been reported in New Caledonia
(Bouchet et al. 1995; de Garine-Wichatitsky et al. 2005) and Fiji (Ash 1992). Seed
predation by introduced rats (Rattus spp.), which are common throughout the
tropical Pacific, can substantially reduce the recruitment of some plant species
(McConkey et al. 2003), thereby altering habitats. Ecosystem-wide changes
such as these likely affect microchiropteran bats as well because of changes in
the availability of invertebrate prey. Rats also feed on some fruits (e.g., Termi-
nalia spp.) in situ and therefore may directly compete with frugivorous bats
for food (Weisler et al. 2006; D. R. Drake, pers. comm.). Introduced insects also
have the potential to eliminate some foods regularly eaten by flying foxes.
For example, on Guam, a large scarab beetle (Protaetia orientalis) introduced
in about 1972 commonly swarms on and consumes the freshly ripened fruit
of seeded breadfruit (Artocarpus mariannensis; G. J. Wiles, pers. obs.), which is
eaten by Pteropus mariannus. Another insect, the cycad aulacaspis scale (Au-
lacaspis yasumatsui), arrived on the island in 2002. It attacks Cycas micronesica
and threatens to seriously reduce or entirely eliminate the species from Guam
(Terry and Marler 2005). The erythrina gall wasp (Quadrastichus erythrinae)
is now widespread on Tutuila, American Samoa, where it is destroying
Conservation Threats to Bats in the Pacific and Southeast Asia 427
Erythrina variegata, and is newly detected on Guam. These species are food
sources for Pteropus mariannus, P. tonganus, and P. samoensis.
Only a few cases of introduced predators killing bats have been reported
in the region, but these illustrate the vulnerability of island bats and show
that the threat should not be minimized. On Guam, brown tree snakes (Boiga
irregularis) have been implicated in the decline of Pteropus mariannus. Snakes
were accidentally brought to the island in the aftermath of World War II, most
likely in military cargo shipped from the Admiralty Islands. They are nocturnal
and arboreal; individuals may grow as long as 3 m and are able to consume
prey up to 70% of their body weight (Rodda et al. 1997; Rodda et al. 1999). Data
collected at Guam’s main colony of flying foxes from 1982 to 2006 have indi-
cated a largely consistent pattern wherein small pups are routinely recorded
with their mothers, but medium-sized young are much rarer, and large young
are virtually absent (Wiles 1987b; Wiles et al. 1995; G. J. Wiles, unpublished
data; D. S. Janeke, pers. comm.). Most observations were made in areas where
native forest bird populations had collapsed, indicating the establishment of
high densities of tree snakes. However, for several months in 1983 after the
colony moved to a new location where native birds persisted, sizable numbers
of larger pups were detected. It is unlikely that the snake played a role in the
decline and loss of Emballonura semicaudata on the island (Fritts and Rodda
1998; G. J. Wiles, pers. obs.).
Predation on bats by nonnative mammalian predators has apparently been
documented only in Samoa and Fiji, where domestic cats, dogs, and pigs have
been observed killing Pteropus forced to forage on the ground near human habi-
tations after the passage of cyclones (Palmeirim et al. 2007; Pierson et al. 1996).
Feral cats have also been suggested as a potentially important predator of E.
semicaudata in Fiji (Palmeirim et al. 2005; Palmeirim et al. 2007), but supportive
data are lacking. Nevertheless, predation by exotic mammals, especially rats
and cats, is probably more frequent than indicated by these few records. Sup-
port for this contention comes from several islands bordering the geographic
area discussed in this chapter. Predation by feral cats, rats, and other mammals
has been linked to declines in Chalinolobus tuberculatus and Mystacina tuberculata
in New Zealand (Lloyd 2005; Pryde et al. 2005) and is suspected as a contrib-
uting factor in the possible extinction of Nyctophilus howensis on Lord Howe
Island (Richards and Hall 1998). Feral cats regularly catch Pteropus melanotus
feeding in shrubs and small trees on Christmas Island in the Indian Ocean
(Tidemann et al. 1994), suggesting that other pteropodids may be vulnerable
to this predator when circumstances lead to their foraging near the ground.
Other assorted interactions between bats and exotic species have been noted.
Biting red ants became established on Choiseul in the Solomons in 1987 and
may have altered the roosting patterns of Pteropus rayneri by discouraging the
use of coconut trees (Bowen-Jones et al. 1997). In the Ryukyus, P. dasymallus
occasionally die after becoming entangled in the leaf fibers of two nonnative
428 G. J. Wiles and A. P. Brooke
palms (Roystonea regia and Washingtonia robusta; K. Kinjo, unpublished data).
Spennemann and Wiles (2002) described the deliberate introduction of a dis-
ease, avian cholera, to Upolu, Samoa, by European planters in the 1890s to
control flying foxes in an effort to limit their damage to fruit crops. Replace-
ment of native forest with introduced trees, especially for the establishment
of monocultures for timber, can be detrimental to bats by reducing food avail-
ability and roosting opportunities (A. P. Brooke, pers. obs.). Exotic vines and
other plants can grow densely at cave entrances (G. J. Wiles, pers. obs.), thereby
blocking access for bats.
In contrast to the negative interactions described above, Pacific flying foxes
have benefited from the establishment of a number of widely introduced plants
that serve as food sources. These include a mix of Paleotropical and Neotropi-
cal species, with some of the more commonly recorded being Annona spp.,
Artocarpus altilis, Cananga odorata, Carica papaya, Ceiba pentandra, Eugenia javan-
ica, E. malaccensis, Mangifera indica, Musa spp. Passiflora spp., Persea americana,
Psidium guajava, Spondias dulcis, S. pinnata, and Syzygium spp. (Banack 1998;
Nakamoto et al. 2007; Stier and Mildenstein 2005; Wiles and Fujita 1992; Wiles
et al. 1997).
Species Accounts
Few populations of bats are regularly monitored in Oceania and insular South-
east Asia. Here we briefly review the status and biology of four species from
the Pacific whose populations are fairly well known on some islands.
Pteropus mariannus
IUCN endangered; threatened status for the United States; protected in the
Commonwealth of the Northern Mariana Islands and on Guam.
Distribution and Genetics
Pteropus mariannus occurs in western and central Micronesia. Koopman (1993)
recognized seven subspecies, as follows: P. m. mariannus in the southern Mari-
anas, including Guam; P. m. paganensis in the northern Marianas; P. m. ulthiensis
at Ulithi Atoll in the western Carolines; P. m. pelewensis at Palau; P. m. yapensis
at Yap; P. m. ualanus at Kosrae; and P. m. loochoensis at Okinawa. Flannery (1995)
classified P. m. pelewensis, P. m. yapensis, and P. m. ualanus as separate species
without explanation, and Yoshiyuki (1989) split P. m. loochoensis as also distinct.
Flannery’s classification (1995) was provisionally followed by Simmons (2005),
but we prefer to retain P. m. pelewensis, P. m. yapensis, and P. m. ualanus as part
of the species until justification for separation is provided. Furthermore, P. m.
paganensis is probably an invalid subspecies (Wiles et al. 1989). Preliminary
analyses suggest that bats within the Marianas are genetically similar, but dis-
tinct from those in Palau (G. McCracken, unpublished data).
Conservation Threats to Bats in the Pacific and Southeast Asia 429
Biology
P. mariannus occupies a variety of habitat types, including native forest, coastal
strand, mangroves, agroforest, and isolated trees in open areas (Falanruw 1988;
Stinson et al. 1992; Wiles and Johnson 2004; Wiles et al. 1997; Worthington et al.
2001). The species is generally colonial, with roosts often containing hundreds
of bats and rarely reaching as many as 2,000 animals (Falanruw 1988; Stinson
et al. 1992; Wiles et al. 1989; Worthington et al. 2001). In several populations,
however, most aggregations hold fewer than 75 bats (Wiles and Johnson 2004;
Wiles et al. 1991; Wiles et al. 1997). Solitary animals are present in all popu-
lations and sometimes comprise a sizable portion of total numbers. Roosts
form at locations seldom visited by people, such as near remote cliffs, at other
isolated upland sites, and in mangroves (Falanruw 1988; Stinson et al. 1992;
Wiles 1987a; Wiles et al. 1997). P. mariannus feeds on the fruits, flowers, and
leaves of at least 78 plants (Wiles and Fujita 1992; Wiles et al. 1997). Young
are born throughout the year in P. m. mariannus and P. m. yapensis (Falanruw
1988; Wiles 1987a).
Population Status
Wiles et al. (1989) estimated a total minimum population of 8,700–9,000 P. m.
mariannus and P. m. paganensis for the entire Marianas chain in 1983 and 1984.
Abundance has generally declined since then. Numbers on Guam decreased
steadily through the 1990s (Utzurrum et al. 2003) to fewer than 100 animals
in 2006 (D. S. Janeke, pers. comm.). Rota’s population has varied from about
2,600 to 1,000 bats since the mid-1980s, with hunting mortality being the major
influence on abundance (Esselstyn et al. 2006; Stinson et al. 1992; Utzurrum et
al. 2003). Numbers stood at about 1,000 in 2003. Populations have remained
low, generally fewer than 25 to 200 bats each, on Saipan, Tinian, and Aguiguan
since the 1970s (Utzurrum et al. 2003). The population on Anatahan fell from
a minimum of 3,000 bats in 1983 (Wiles et al. 1989) to about 2,000 bats in 1995
(Worthington et al. 2001). Intermittent volcanic eruptions since 2003 have cov-
ered much of the island with volcanic ash and further reduced bat numbers to
about 110 animals in 2006 (C. C. Kessler, pers. comm.). Sarigan has generally
maintained about 125–235 bats since the 1980s, although its population is some-
times supplemented by additional animals from neighboring islands (Wiles
and Johnson 2004). The northernmost islands of Guguan, Alamagan, Pagan,
Agrihan, Asuncion, and Maug have not been fully surveyed since 1983–1987,
when minimum populations were estimated to total about 4,300 bats (Utzur-
rum et al. 2003; Wiles et al. 1989). Brief visits in 2000 and 2001 to some islands
were inconclusive.
Population information is much sparser for the other subspecies. Surveys of
P. m. pelewensis in Palau in 1991 and 2005 did not yield population estimates,
but found the subspecies to be fairly common in 1991 and somewhat more
abundant in 2005 (Wiles et al. 1997; G. J. Wiles, unpublished data). At Yap,
430 G. J. Wiles and A. P. Brooke
P. m. yapensis increased from about 1,000 bats in 1981 to about 2,500–5,000 bats
in 1986 (Mickleburgh et al. 1992), but no estimates have been made since then.
Numbers of P. m. ulthiensis at Ulithi Atoll have not been surveyed since 1986,
when Wiles et al. (1991) estimated a total of 895–1,060 bats. No valid population
estimates exist for P. m. ualanus.
Pteropus tonganus
IUCN least concern; protected in Samoa and American Samoa; partially pro-
tected in Tonga; five-year ban on hunting in Niue beginning in 2004.
Distribution and Genetics
Pteropus tonganus occurs widely across the southern Pacific (Miller and Wilson
1997). Genetic studies using allozyme electrophoresis and mtDNA found no
evident geographic pattern in P. t. tonganus, which appears to form a single
wide-ranging population from Fiji to the Cooks (Ingelby and Colgan 2003;
Utzurrum et al. 2000). Distribution of this subspecies formerly extended east-
ward to at least Rurutu in the Austral Islands of French Polynesia (Weisler et
al. 2006). Two other subspecies include P. t. geddiei at New Caledonia, Vanuatu,
and the Solomons, and P. t. basilicus on Karkar and Koil islands off northern
New Guinea (Flannery 1995).
Biology
Colonies range in size from a few individuals to several thousand and prefer
tall emergent trees for roosting. Where bats are hunted, colonies seek sites that
are difficult for people to reach: cliffs, volcanic craters, steep hills, mangrove
swamps, and small uninhabited islands (Brooke et al. 2000; Cousins and Comp-
ton 2005; Palmeirim et al. 2005). When not hunted, roosts may occur closer to
human habitation, such as throughout the village of Kolovai, Tonga. Colonies
are commonly found on small islets in Tonga and Fiji, where animals commute
nightly to larger neighboring islands up to 10 km away (McConkey and Drake
2007; Palmeirim et al. 2005). The diet includes numerous fruits and flowers
(Banack 1998; Miller and Wilson 1997). Mating and births occur throughout
the year (Grant and Banack 1999).
Population Status
Population estimates of P. t. tonganus exist for several locations. Regular is-
landwide roost counts on Tutuila, America Samoa, have been conducted since
1992. Early surveys in 1987–1989 estimated 12,750–28,000 individuals, but
unregulated hunting following two severe cyclones in 1990 and 1991 caused
the population to decline to about 1,500–2,500 animals (Craig et al. 1994b).
After hunting was banned in 1992, colonies regrouped in inaccessible areas
and increased to about 6,300 bats by 2000 (Brooke et al. 2000; Utzurrum et
al. 2003). Surveys in Tonga during 1995 found a robust population of about
6,000 bats on 14 islands in the Vava’u group (Grant 1998), but abundance fell
Conservation Threats to Bats in the Pacific and Southeast Asia 431
by more than 80% after Cyclone Waka hit the area in 2001 (McConkey et al.
2004). Niue’s population ranged from an estimated 2,040 to 4,080 bats in 1998
(Brooke and Tschapka 2002), but numbers have declined greatly since then
because of uncontrolled hunting and damage from Cyclone Heta in January
2004. Surveys in 2004 failed to detect any colonies, with only 60 bats counted
in 27 surveys, a decrease of 95% from those conducted in 1998 at the same
locations (Brooke 2004). A five-year ban on hunting enacted in late 2004 may
help to restore this population. In the Cook Islands in 2002, abundance was
estimated at about 1,730 bats on Rarotonga and about 80 bats on Mangaia, with
probably an overall declining trend (Cousins and Compton 2005). Insufficient
habitat appears to be the critical factor affecting abundance on Mangaia. In
addition, bats are hunted on both islands without restriction. More than half
of the global population of P. t. tonganus is believed to occur in Fiji (Palmeirim
et al. 2005; Palmeirim et al. 2007). Although population estimates do not exist
for this island group, animals are plentiful and widespread on large and small
islands alike. Some hunting, mostly for personal consumption, occurs here
(Palmeirim et al. 2005).
P. t. basilicus was highly visible in 1992 on Karkar Island, Papua New Guinea,
although its lowland habitat had been largely converted to plantation forests
(Bonaccorso 1998). No recent information is available on population status for
Koil Island. Hunting probably threatens populations on both islands (Bonac-
corso 1998). Relatively little information is available on P. t. geddiei, which is
found in New Caledonia, the Solomon Islands, and Vanuatu.
Pteropus samoensis
IUCN near threatened; protected in Samoa and American Samoa.
Distribution and Genetics
Two subspecies exist: Pteropus samoensis samoensis in the Samoan archipelago
and P. s. nawaiensis in Fiji (Banack 2001). Fossil remains predating Polynesian
settlement have been found in Tonga (Koopman and Steadman 1995). Ingelby
and Colgan (2003) did not observe notable differences in allele frequencies from
three Fijian islands (Vanua Levu, Viti Levu, and Taveuni), suggesting few bar-
riers to gene flow. In contrast, Utzurrum et al. (2000) found haplotypes differed
greatly within and among island populations, suggesting that P. samoensis once
had a larger distribution and has undergone a dramatic decline with substan-
tial structuring among islands. P. samoensis appears to be most closely related
to P. nitendiensis from the Solomon Islands (Ingelby and Colgan 2003).
Biology
P. samoensis is most common in or near native forest and forages on a variety
of leaves, flowers, and fruits of both native and agricultural plants (Banack
1998; Mickleburgh et al. 1992; Nelson et al. 2000a, 2000b; Palmeirim et al. 2005;
Palmeirim et al. 2007). Bats are most active at night but are seen throughout the
432 G. J. Wiles and A. P. Brooke
day, especially near dawn and dusk, when thermal updrafts are used for soar-
ing (Brooke 2001; Norberg et al. 2000; Thomson et al. 1998, 2004). The species
typically roosts singly or in small groups usually comprising either a female
and young of the year or a mated pair (Brooke 2001).
Population Status
Surveys of P. s. samoensis were initiated in American Samoa in 1986, with an
estimated 700 bats present on Tutuila prior to Cyclone Ofa in 1990 and 200–400
bats present after Cyclone Val in 1991 (Utzurrum et al. 2003). From 1995 to
2000, the population remained stable at about 900 animals, whereas numbers
in the Manu’a Islands remained low at about 100 individuals. The low numbers
recorded in Samoa after Cyclone Val had not appreciably increased by 1996
(Brooke 1997; Wilson and Engbring 1992).
P. s. nawaiensis is moderately common in some lowland areas of Viti Levu
and Vanua Levu, the two largest islands in Fiji (Palmeirim et al. 2005; Pal-
meirim et al. 2007). It also occurs on some medium-sized islands, but usually
avoids smaller islands.
Emballonura semicaudata
IUCN endangered; candidate status for the United States; protected in Ameri-
can Samoa, Guam, and the Commonwealth of the Northern Mariana Islands
(Hutson et al. 2001).
Distribution
Four subspecies occur as follows: Emballonura semicaudata semicaudata in Vanu-
atu, Fiji, Tonga, Samoa, and American Samoa; E. s. sulcata in Chuuk and Pohn-
pei in the central Caroline Islands; E. s. palauensis in Palau; and E. s. rotensis from
Guam to Saipan in the southern Marianas (Koopman 1997; Simmons 2005).
Biology
E. semicaudata is most common on limestone islands with caves and rock over-
hangs, but also inhabits volcanic islands. Roosts occur in small to large caves,
lava tubes, rock depressions, and hollow trees. Illumination at roosts can vary
from twilight to dark sections of caves and includes well-lit rock depressions
(Palmeirim et al. 2005; Wiles et al. 1997; G. J. Wiles, unpublished data). Zooar-
chaeological excavations on Guam found skeletal remains common under the
overhangs of limestone cliffs (D. W. Steadman, pers. comm.). Most colonies
range in size from a few to several hundred bats, but some roosts in Fiji may
have once harbored thousands of individuals (Palmeirim et al. 2005; Sawyer
and Andrews 1901 cited in Palmeirim et al. 2007; Wiles et al. 1997).
E. semicaudata feeds on aerial insects. The last surviving population in the
Marianas, located on the island of Aguiguan, forages preferentially in native
forest (Esselstyn et al. 2004b), but elsewhere, animals can forage in a variety of
Conservation Threats to Bats in the Pacific and Southeast Asia 433
habitats, including urban areas (Palmeirim et al. 2005; Wiles et al. 1997). Large
numbers of individuals have been observed transiting distances of 5 km or
more to reach feeding locations in Palau (Wiles et al. 1997).
Population Status
Many populations of E. semicaudata have undergone dramatic decline during
the past 50 years for reasons that are unclear, but which may involve forest
loss, insecticide use, severe tropical cyclones, introduced predators, or human
disturbance of caves (Hutson et al. 2001; Palmeirim et al. 2005; Palmeirim et
al. 2007). Relatively healthy populations remain only in Palau, Pohnpei, and
Chuuk, and on some of Fiji’s smaller islands. In Palau, an estimated 5,000–
10,000 bats were counted departing several roosting islands, with bats being
widespread and regularly detected elsewhere in the island group in 1991 and
again in 2005 (Wiles et al. 1997; G. J. Wiles, unpublished data). The species ap-
peared fairly common on Pohnpei in 1999 and Chuuk in 1989 (G. J. Wiles, pers.
obs.), but surveys have not been conducted at either location.
Populations elsewhere contain few remaining bats or have been extirpated.
In the Marianas, E. semicaudata survives only on Aguiguan, where an estimated
450–600 bats occurred in 2008 (G. J. Wiles, unpublished data). The species has
experienced tremendous decline in American Samoa and Samoa since the 1970s
or earlier (Grant et al. 1994; Hutson et al. 2001; Tarburton 2002) and may now
be gone from one or both locations. Populations on Fiji’s two largest islands
were widespread and common into the 1970s, but surveys made in 2000 and
2001 found only a single colony (Palmeirim et al. 2005; Palmeirim et al. 2007).
Fijian populations now persist primarily on smaller limestone islands. Status is
poorly known for Tonga (Helgen and Flannery 2002; Koopman and Steadman
1995), and no records exist for Vanuatu since 1929 (Helgen 2004).
Conservation Needs
The threats described in this chapter, combined with the many socioeconomic
problems inherent to the region such as large human populations, poverty,
ineffective governments, and corruption (e.g., MacKinnon 2006), portray a pes-
simistic future for bat populations in many parts of insular Southeast Asia and
the tropical Pacific. We nevertheless believe that some successes are achievable
in conserving bats and concur with the general needs and solutions discussed
by Sodhi and Brook (2006), such as improving public awareness, empower-
ing citizens, and increasing resource protection whenever feasible. Two of the
greatest challenges in protecting bats throughout the region are preserving
adequate amounts of habitat (including roosting sites) and reducing overex-
ploitation of populations. This can only be done by convincing local people
that they have a stake in preserving natural resources. There are glimmers of
progress, one being the expansion of conservation-oriented nongovernmental
434 G. J. Wiles and A. P. Brooke
organizations working at the national and local levels. Some groups, such as
the Bat Association of Taiwan, work specifically for the benefit of bats, while
others like the Foundation for the Philippine Environment and the Palau Con-
servation Society have sponsored valuable bat-related projects. We are also
heartened by the increasing participation of resident biologists working with
bats in Taiwan, the Philippines, East Malaysia, and Indonesia.
Greater implementation of improved logging practices and timber conces-
sion management, as summarized in Meijaard and Sheil 2008 and Dennis et al.
2008, has begun in the region and offers promise for the conservation of forest-
dwelling bats and other biota in production forests. However, for sustainable
forest management to succeed on a large scale, greater enforcement and ac-
countability are required as well as increased demand for certified timber in
international markets (Meijaard et al. 2005).
Many of the conservation activities recommended in two action plans for
bats (Hutson et al. 2001; Mickleburgh et al. 1992) remain pertinent. These in-
clude (1) improving public and government support for bats so that meaningful
protection of populations and habitats is achieved and (2) conducting invento-
ries, population monitoring, and ecological studies of species. One particular
challenge in the region is preventing the overharvesting of bats as food, espe-
cially pteropodids. Banning commercial sales of bats may be one useful mea-
sure in some locations for reducing harvest pressure (Bennett et al. 2000). Other
widely needed efforts are greater enforcement of existing laws and amend-
ing conservation laws so that they include bats (Corlett 2007). Several studies
have reported that flying foxes have a poor capacity for increase (McIlwee and
Martin 2002; Pierson and Rainey 1992), but our experiences suggest that at
least some populations of Pteropus (e.g., P. mariannus pelewensis in Palau [Wiles
et al. 1997], P. m. yapensis in Yap [Mickleburgh et al. 1992], and P. tonganus in
American Samoa [Brooke et al. 2000; Utzurrum et al. 2003]) readily respond to
reduced hunting pressure and can grow fairly rapidly. This may bode well for
the recovery of other pteropodids if harvest levels can be controlled.
Conclusions
Islands in the tropical Pacific and insular Southeast Asia hold more than 350 bat
species, or about 31% of the world’s total, making the region a major center of
bat diversity, especially for pteropodids. Many of these species are thought to
be declining, with 28% (100 species) designated as threatened or near threat-
ened by the IUCN. This chapter discusses five important conservation concerns
facing bat populations in the region: habitat destruction and alteration, hunt-
ing, cave disturbance, severe tropical cyclones, and exotic species.
Deforestation is a major threat for large numbers of forest-dependent bats,
especially in Indonesia, the Philippines, East Malaysia, the Solomon Islands,
Conservation Threats to Bats in the Pacific and Southeast Asia 435
and Papua New Guinea, which feature some of the world’s highest rates of
forest loss and alteration. Studies from mainland Southeast Asia reveal the
presence of impoverished bat communities after forest disturbance and de-
struction, and suggest that similar outcomes can be expected in insular South-
east Asia and the Pacific.
Bats are a traditional food source in most of the region and continue to
be harvested extensively for both subsistence and commercial purposes. Fly-
ing foxes and other pteropodids are targeted because of their larger sizes, but
microchiropterans are also caught in some areas. Commercial markets are ap-
parently most active in Indonesia. A large international trade in Pteropus that
was formerly centered on Guam and the Northern Mariana Islands ended in
1994 after the enactment of CITES restrictions. Heavy harvest pressure in many
parts of the region has led to significant population declines in a number of
pteropodids and at least one microchiropteran.
Regular human visitation of caves for guano mining, swiftlet nest collecting,
hunting, and other activities is believed to have harmed many populations of
cave-dwelling microchiropterans and megachiropterans, although regional
data are rare on the extent of this problem. Cave disturbance is perhaps most
widespread in the Philippines, Indonesia, and East Malaysia, but is probably
underdocumented elsewhere.
Exceptionally strong tropical cyclones occasionally strike a number of is-
lands in the region and can produce tremendous damage to forests, leaving
resident bat populations vulnerable to starvation, dehydration, and increased
hunting and predation. Several studies have documented declines of 57% to
possibly as high as 99% in flying fox numbers after major storms.
Relatively few examples of bat populations being harmed by invasive spe-
cies have been reported to date, probably because of a lack of study. This threat
will likely increase in the future, especially on smaller islands where impacts
of exotics are often most severe.
Acknowledgments
Matt Struebig deserves special thanks for useful comments and discussions on
the bats of Borneo and Indonesia. The following people kindly provided in-
formation on a number of topics: Mohamad Tajuddin Abdullah, James Ather-
ton, Don Drake, Jake Esselstyn, Janet Franklin, Gabriella Fredriksson, Melvin
Gumal, Larry Heaney, Kris Helgen, Ian Henson, Nina Ingle, Masako Izawa,
Dustin Janeke, Jill Key, Tigga Kingston, Kim McConkey, Tammy Mildenstein,
Keiko Osawa, Harold Ota, Jorge Palmeirim, and Ruth Utzurrum. We also thank
Colin O’Donnell, Ted Fleming, and two anonymous reviewers for their com-
ments on the manuscript, and Jan Sharkey for her support during manuscript
preparation.
APPENDIX 14.1
Table A14.1. Distribution of 354 bat species in 23 countries, territories, and island groups in the
tropical Pacific and insular Southeast Asia
Species IUCN statusaCountries, territories, or island
groups of occurrenceb, c
Megachiroptera
Pteropodidae
Acerodon celebensis LC Indonesia
Acerodon humilis EN Indonesia
Acerodon jubatus EN Philippines
Acerodon leucotis VU Philippines
Acerodon mackloti VU Indonesia, Timor-Leste
Aethalops aequalis LC Indonesia, E. Malaysia, Brunei
Aethalops alecto LC Indonesia*
Alionycteris paucidentata LC Philippines
Aproteles bulmerae CR PNG
Balionycteris maculata LC Indonesia, E. Malaysia, Brunei*
Chironax melanocephalus LC Indonesia, E. Malaysia, Brunei*
Cynopterus brachyotis LC Indonesia, E. Malaysia, Brunei*
Cynopterus horsfieldii LC Indonesia, E. Malaysia, Brunei*
Cynopterus luzoniensis LC Indonesia, Philippines
Cynopterus minutus LC Indonesia, E. Malaysia, Brunei
Cynopterus nusatenggara LC Indonesia
Cynopterus sphinx LC Indonesia, Taiwan*
Cynopterus titthaecheilus LC Indonesia, Timor-Leste
Desmalopex leucopterus LC Philippines
Desmalopex microleucopterus NE Philippines
Dobsonia anderseni LC PNG
Dobsonia beauforti LC Indonesia
Dobsonia chapmani CR Philippines
Dobsonia crenulata LC Indonesia
Dobsonia emersa VU Indonesia
Dobsonia exoleta LC Indonesia
Dobsonia inermis LC PNG, Solomons
Dobsonia magna NE Indonesia, PNG*
Dobsonia minor LC Indonesia, PNG
Table A14.1. (continued)
Species IUCN statusaCountries, territories, or island
groups of occurrenceb, c
Dobsonia moluccensis LC Indonesia, Timor-Leste
Dobsonia pannietensis NT PNG
Dobsonia peronii LC Indonesia, Timor-Leste
Dobsonia praedatrix LC PNG
Dobsonia viridis LC Indonesia
Dyacopterus brooksi VU Indonesia
Dyacopterus rickarti NE Philippines
Dyacopterus spadiceus NT Indonesia, E. Malaysia, Brunei*
Eonycteris major DD Indonesia, E. Malaysia, Brunei
Eonycteris robusta NT Philippines
Eonycteris spelaea LC Indonesia, E. Malaysia, Brunei,
Timor-Leste, Philippines*
Haplonycteris fischeri LC Philippines
Harpyionycteris celebensis VU Indonesia
Harpyionycteris whiteheadi LC Philippines
Macroglossus minimus LC Indonesia, E. Malaysia, Brunei,
Timor-Leste, Philippines, PNG,
Solomons*
Macroglossus sobrinus LC Indonesia*
Megaerops ecaudatus LC Indonesia, E. Malaysia, Brunei*
Megaerops kusnotoi VU Indonesia
Megaerops wetmorei VU Indonesia, Brunei, Philippines*
Melonycteris fardoulisi LC Solomons
Melonycteris melanops LC PNG
Melonycteris woodfordi LC PNG, Solomons
Mirimiri acrodonta CR Fiji
Neopteryx frosti EN Indonesia
Notopteris macdonaldi VU Vanuatu, Fiji
Notopteris neocaledonica VU New Caledonia
Nyctimene aello LC Indonesia, PNG
Nyctimene albiventer LC Indonesia, PNG
Nyctimene cephalotes LC Indonesia, Timor-Leste, PNG*
Nyctimene certans LC Indonesia, PNG
Table A14.1. (continued)
Species IUCN statusaCountries, territories, or island
groups of occurrenceb, c
Nyctimene cyclotis DD Indonesia
Nyctimene draconilla DD Indonesia, PNG
Nyctimene keasti VU Indonesia
Nyctimene major LC PNG, Solomons
Nyctimene malaitensis DD Solomons
Nyctimene masalai DD PNG
Nyctimene minutus VU Indonesia
Nyctimene rabori EN Indonesia, Philippines
Nyctimene sanctacrucis DD Solomon
Nyctimene vizcaccia LC PNG, Solomons
Otopteropus cartilagonodus LC Philippines
Paranyctimene raptor LC Indonesia, PNG
Paranyctimene tenax LC Indonesia, PNG
Penthetor lucasi LC Indonesia, E. Malaysia, Brunei*
Ptenochirus jagori LC Philippines
Ptenochirus minor LC Philippines
Pteralopex anceps EN PNG, Solomons
Pteralopex atrata EN Solomon
Pteralopex flanneryi CR PNG, Solomons
Pteralopex pulchra CR Solomons
Pteralopex taki EN Solomons
Pteropus admiralitatum LC PNG, Solomons
Pteropus alecto LC Indonesia, PNG*
Pteropus anetianus VU Vanuatu
Pteropus argentatus DD Indonesia
Pteropus aruensis CR Indonesia
Pteropus caniceps NT Indonesia
Pteropus capistratus EN PNG
Pteropus chrysoproctus NT Indonesia
Pteropus cognatus EN Solomons
Pteropus conspicillatus LC Indonesia, PNG*
Pteropus dasymallus NT Philippines, Taiwan, Ryukyus
Table A14.1. (continued)
Species IUCN statusaCountries, territories, or island
groups of occurrenceb, c
Pteropus fundatus EN Vanuatu
Pteropus gilliardorum DD PNG
Pteropus griseus DD Indonesia, Timor-Leste
Pteropus howensis DD Solomons
Pteropus hypomelanus LC Indonesia, E. Malaysia, Philippines,
PNG, Solomons*
Pteropus insularis DD FSM
Pteropus keyensis DD Indonesia
Pteropus lombocensis DD Indonesia, Timor-Leste
Pteropus loochoensis DD Ryukyus
Pteropus macrotis LC Indonesia, PNG*
Pteropus mahaganus VU PNG, Solomons
Pteropus mariannus EN Palau, FSM, Guam, N. Marianas
Pteropus melanopogon EN Indonesia
Pteropus melanotus VU Indonesia*
Pteropus molossinus VU FSM
Pteropus neohibernicus LC Indonesia, PNG
Pteropus nitendiensis EN Solomons
Pteropus ocularis VU Indonesia
Pteropus ornatus VU New Caledonia
Pteropus personatus LC Indonesia
Pteropus phaeocephalus DD FSM
Pteropus pilosus EX Palau
Pteropus pohlei EN Indonesia
Pteropus pselaphon CR Ogasawara and Iwo Islands
Pteropus pumilus NT Indonesia, Phillipines
Pteropus rayneri NT PNG, Solomons
Pteropus rennelli VU Solomons
Pteropus samoensis NT Fiji, Samoa, American Samoa
Pteropus scapulatus LC PNG*
Pteropus speciosus DD Indonesia, Philippines
Pteropus temminckii VU Indonesia
Table A14.1. (continued)
Species IUCN statusaCountries, territories, or island
groups of occurrenceb, c
Pteropus tokudae EX Guam
Pteropus tonganus LC PNG, Solomons, Vanuatu, New
Caledonia, Fiji, Wallis and Futuna,
Tonga, Samoa, American Samoa,
Niue, Cooks
Pteropus tuberculatus CR Solomons
Pteropus vampyrus NT Indonesia, E. Malaysia, Brunei,
Timor-Leste, Philippines*
Pteropus vetulus VU New Caledonia
Pteropus woodfordi VU Solomons
Rousettus amplexicaudatus LC Indonesia, E. Malaysia, Brunei,
Timor-Leste, Philippines, PNG,
Solomons*
Rousettus bidens VU Indonesia
Rousettus celebensis LC Indonesia
Rousettus leschenaultii LC Indonesia*
Rousettus linduensis DD Indonesia
Rousettus spinalatus VU Indonesia, E. Malaysia, Brunei
Styloctenium mindorensis DD Philippines
Styloctenium wallacei VU Indonesia
Syconycteris australis LC Indonesia, PNG*
Syconycteris carolinae VU Indonesia
Syconycteris hobbit VU Indonesia, PNG
Thoopterus nigrescens LC Indonesia
Microchiroptera
Rhinolophidae
Rhinolophus acuminatus LC Indonesia, E. Malaysia,
Philippines*
Rhinolophus affinis LC Indonesia, E. Malaysia*
Rhinolophus arcuatus LC Indonesia, E. Malaysia,
Philippines, PNG
Rhinolophus borneensis LC Indonesia, E. Malaysia, Brunei,
Philippines*
Rhinolophus canuti VU Indonesia, Timor-Leste
Rhinolophus celebensis LC Indonesia, Timor-Leste
Table A14.1. (continued)
Species IUCN statusaCountries, territories, or island
groups of occurrenceb, c
Rhinolophus creaghi LC Indonesia, E. Malaysia, Brunei,
Philippines
Rhinolophus euryotis LC Indonesia, PNG
Rhinolophus ferrumequinum LC Ryukyus*
Rhinolophus formosae NT Taiwan
Rhinolophus imaizumii NE Ryukyus
Rhinolophus inops LC Philippines
Rhinolophus keyensis DD Indonesia, Timor-Leste
Rhinolophus lepidus LC Indonesia*
Rhinolophus luctus LC Indonesia, E. Malaysia, Brunei*
Rhinolophus macrotis LC Indonesia, Philippines*
Rhinolophus madurensis EN Indonesia
Rhinolophus megaphyllus LC Indonesia, PNG*
Rhinolophus montanus DD Timor-Leste
Rhinolophus nereis DD Indonesia
Rhinolophus philippinensis LC Indonesia, E. Malaysia, Brunei,
Philippines, PNG*
Rhinolophus pusillus LC Indonesia, E. Malaysia, Taiwan,
Ryukyus*
Rhinolophus rufus NT Philippines
Rhinolophus sedulus NT Indonesia, E. Malaysia, Brunei*
Rhinolophus stheno LC Indonesia*
Rhinolophus subrufus DD Philippines
Rhinolophus trifoliatus LC Indonesia, E. Malaysia, Brunei*
Rhinolophus virgo LC Philippines
Hipposideridae
Anthops ornatus DD PNG, Solomons
Aselliscus tricuspidatus LC Indonesia, PNG, Solomons,
Vanuatu
Coelops frithii LC Indonesia, Taiwan*
Coelops robinsoni VU Indonesia, E. Malaysia,
Philippines*
Hipposideros armiger LC Taiwan*
Hipposideros ater LC Indonesia, E. Malaysia, Brunei,
Philippines, PNG*
Table A14.1. (continued)
Species IUCN statusaCountries, territories, or island
groups of occurrenceb, c
Hipposideros bicolor LC Indonesia, E. Malaysia, Brunei,
Timor-Leste, Philippines*
Hipposideros boeadii DD Indonesia
Hipposideros breviceps DD Indonesia
Hipposideros calcaratus LC Indonesia, PNG, Solomons
Hipposideros cervinus LC Indonesia, E. Malaysia, Brunei,
Philippines, PNG, Solomons,
Vanuatu*
Hipposideros cineraceus LC Indonesia, E. Malaysia*
Hipposideros coronatus DD Philippines
Hipposideros corynophyllus DD Indonesia, PNG
Hipposideros coxi DD Indonesia, E. Malaysia
Hipposideros crumeniferus DD Indonesia, Timor-Leste
Hipposideros demissus VU Solomons
Hipposideros diadema LC Indonesia, E. Malaysia, Brunei,
Timor-Leste, Philippines, PNG,
Solomons*
Hipposideros dinops DD PNG, Solomons
Hipposideros doriae NT Indonesia, E. Malaysia, Brunei*
Hipposideros dyacorum LC Indonesia, E. Malaysia, Brunei*
Hipposideros edwardshilli DD PNG
Hipposideros galeritus LC Indonesia, E. Malaysia, Brunei*
Hipposideros inexpectatus DD Indonesia
Hipposideros larvatus LC Indonesia, E. Malaysia*
Hipposideros lekaguli NT Philippines*
Hipposideros macrobullatus DD Indonesia
Hipposideros madurae LC Indonesia
Hipposideros maggietaylorae LC Indonesia, PNG
Hipposideros muscinus DD Indonesia, PNG
Hipposideros obscurus LC Philippines
Hipposideros orbiculus EN Indonesia*
Hipposideros papua LC Indonesia
Hipposideros pelingensis NT Indonesia
Hipposideros pygmaeus LC Philippines
Table A14.1. (continued)
Species IUCN statusaCountries, territories, or island
groups of occurrenceb, c
Hipposideros ridleyi VU Indonesia, E. Malaysia, Brunei*
Hipposideros semoni DD PNG*
Hipposideros sorenseni VU Indonesia
Hipposideros sumbae LC Indonesia, Timor-Leste
Hipposideros turpis NT Ryukyus*
Hipposideros wollastoni LC Indonesia, PNG
Megadermatidae
Megaderma spasma LC Indonesia, E. Malaysia, Brunei,
Philippines*
Rhinopomatidae
Rhinopoma microphyllum LC Indonesia*
Emballonuridae
Saccolaimus flaviventris LC PNG*
Saccolaimus mixtus DD PNG*
Saccolaimus saccolaimus LC Indonesia, E. Malaysia, Brunei,
Timor-Leste, Philippines, PNG,
Solomons*
Taphozous achates DD Indonesia
Taphozous australis NT PNG*
Taphozous longimanus LC Indonesia, E. Malaysia*
Taphozous melanopogon LC Indonesia, E. Malaysia, Brunei,
Timor-Leste, Philippines*
Taphozous theobaldi LC Indonesia*
Emballonura alecto LC Indonesia, E. Malaysia, Brunei,
Philippines
Emballonura beccarii LC Indonesia, PNG
Emballonura dianae LC PNG, Solomons
Emballonura furax DD Indonesia, PNG
Emballonura monticola LC Indonesia, E. Malaysia, Brunei*
Emballonura raffrayana LC Indonesia, PNG, Solomons
Emballonura semicaudata EN Vanuatu, Fiji, Tonga, Samoa,
America Samoa, Palau, FSM,
Guam, N. Marianas
Emballonura serii LC Indonesia, PNG
Mosia nigrescens LC Indonesia, PNG, Solomons
Table A14.1. (continued)
Species IUCN statusaCountries, territories, or island
groups of occurrenceb, c
Nycteridae
Nycteris javanica VU Indonesia
Nycteris tragata NT Indonesia, E. Malaysia, Brunei*
Molossidae
Chaerephon bregullae EN Vanuatu, Fiji
Chaerephon jobensis LC Indonesia, PNG*
Chaerephon johorensis VU Indonesia*
Chaerephon plicatus LC Indonesia, E. Malaysia, Brunei,
Philippines*
Chaerephon solomonis LC Solomons
Cheiromeles parvidens LC Indonesia, Philippines
Cheiromeles torquatus LC Indonesia, E. Malaysia, Brunei,
Philippines*
Mops mops NT Indonesia, E. Malaysia, Brunei*
Mops sarasinorum DD Indonesia, Philippines
Mormopterus beccarii LC Indonesia, PNG*
Mormopterus doriae DD Indonesia
Mormopterus loriae LC PNG*
Otomops formosus DD Indonesia
Otomops johnstonei DD Indonesia
Otomops papuensis DD PNG
Otomops secundus DD PNG
Tadarida insignis DD Taiwan*
Tadarida kuboriensis LC Indonesia, PNG
Tadarida latouchei DD Ryukyus*
Tadarida teniotis LC Indonesia*
Vespertilionidae
Arielulus circumdatus LC Indonesia*
Arielulus cuprosus DD E. Malaysia
Arielulus torquatus LC Taiwan
Eptesicus serotinus LC Taiwan*
Hesperoptenus blanfordi LC E Malaysia*
Hesperoptenus doriae DD E. Malaysia*
Table A14.1. (continued)
Species IUCN statusaCountries, territories, or island
groups of occurrenceb, c
Hesperoptenus gaskelli DD Indonesia
Hesperoptenus tomesi VU E. Malaysia*
Scotophilus celebensis DD Indonesia
Scotophilus collinus LC Indonesia, E. Malaysia, Timor-Leste
Scotophilus kuhlii LC Indonesia, E. Malaysia, Brunei,
Timor-Leste, Philippines, Taiwan*
Scotorepens sanborni LC Indonesia, PNG*
Nyctophilus bifax LC Indonesia, PNG*
Nyctophilus heran DD Indonesia, Timor-Leste
Nyctophilus microdon DD PNG
Nyctophilus microtis LC Indonesia, PNG
Nyctophilus nebulosus CR New Caledonia
Nyctophilus timoriensis DD Indonesia, Timor-Leste, PNG*
Pharotis imogene CR PNG
Glischropus javanus DD Indonesia
Glischropus tylopus LC Indonesia, E. Malaysia, Brunei,
Philippines*
Nyctalus aviator NT Ryukyus*
Nyctalus noctula LC Taiwan*
Nyctalus plancyi LC Taiwan*
Pipistrellus abramus LC Taiwan, Ryukyus*
Pipistrellus angulatus LC Indonesia, PNG, Solomons
Pipistrellus ceylonicus LC E. Malaysia*
Pipistrellus collinus LC Indonesia, PNG
Pipistrellus javanicus LC Indonesia, E. Malaysia,
Philippines*
Pipistrellus minahassae DD Indonesia
Pipistrellus papuanus LC Indonesia, PNG
Pipistrellus pipistrellus LC Taiwan*
Pipistrellus stenopterus LC Indonesia, E. Malaysia,
Philippines*
Pipistrellus studeei DD Ogasawara and Iwo Islands
Pipistrellus tenuis LC Indonesia, E. Malaysia,
Timor-Leste, Philippines*
Table A14.1. (continued)
Species IUCN statusaCountries, territories, or island
groups of occurrenceb, c
Pipistrellus wattsi LC PNG
Barbastella leucomelas LC Taiwan*
Plecotus taivanus NT Taiwan
Chalinolobus neocaledonicus EN New Caledonia
Chalinolobus nigrogriseus LC PNG*
Falsistrellus mordax DD Indonesia
Falsistrellus petersi DD Indonesia, E. Malaysia, Philippines
Hypsugo imbricatus LC Indonesia, E. Malaysia
Hypsugo kitcheneri DD Indonesia, E. Malaysia
Hypsugo macrotis DD Indonesia*
Hypsugo vordermanni DD Indonesia, E. Malaysia, Brunei
Philetor brachypterus LC Indonesia, E. Malaysia, Brunei,
Philippines, PNG*
Tylonycteris pachypus LC Indonesia, E. Malaysia, Brunei,
Philippines*
Tylonycteris robustula LC Indonesia, E. Malaysia, Brunei,
Timor-Leste, Philippines*
Vespertilio sinensis LC Taiwan*
Myotis adversus LC Indonesia, E. Malaysia,
Timor-Leste, Taiwan*
Myotis ater LC Indonesia, E. Malaysia, Philippines*
Myotis formosus LC Indonesia, Phillipines, Taiwan*
Myotis gomantongensis LC E. Malaysia
Myotis hasseltii LC Indonesia, E. Malaysia, Brunei*
Myotis hermani DD Indonesia*
Myotis horsfieldii LC Indonesia, E. Malaysia, Brunei,
Philippines*
Myotis insularum DD Samoa
Myotis macrodactylus LC Ryukyus*
Myotis macrotarsus NT E. Malaysia, Philippines
Myotis moluccarum LC Indonesia, PNG, Solomons,
Vanuatu*
Myotis montivagus LC Indonesia, E. Malaysia*
Myotis muricola LC Indonesia, E. Malaysia, Brunei,
Philippines, Taiwan*
Table A14.1. (continued)
Species IUCN statusaCountries, territories, or island
groups of occurrenceb, c
Myotis ridleyi NT Indonesia, E. Malaysia, Brunei*
Myotis siligorensis LC E. Malaysia*
Myotis stalkeri DD Indonesia
Myotis yanbarensis CR Ryukyus
Miniopterus australis LC Indonesia, E. Malaysia, Brunei,
Timor-Leste, Philippines, PNG,
Solomons, Vanuatu, New
Caledonia*
Miniopterus fuscus EN Ryukyus
Miniopterus macrocneme DD Indonesia, PNG, Solomons,
Vanuatu, New Caledonia
Miniopterus magnater LC Indonesia, E. Malaysia, Timor-
Leste, PNG*
Miniopterus medius LC Indonesia, E. Malaysia, PNG*
Miniopterus paululus DD Indonesia, E. Malaysia, Philippines
Miniopterus pusillus LC Indonesia, Timor-Leste*
Miniopterus robustior EN New Caledonia
Miniopterus schreibersii NT Indonesia, E. Malaysia, Brunei,
Philippines, PNG, Solomons,
Taiwan*
Miniopterus shortridgei DD Indonesia
Miniopterus tristis LC Indonesia, Philippines, PNG,
Solomons, Vanuatu
Harpiocephalus harpia LC Indonesia, E. Malaysia, Philippines,
Taiwan*
Harpiocephalus mordax DD E. Malaysia*
Harpiola isodon DD Taiwan
Murina aenea VU Indonesia, E. Malaysia*
Murina cyclotis LC Indonesia, E. Malaysia, Brunei,
Philippines*
Murina florium LC Indonesia, PNG*
Murina puta NT Taiwan
Murina rozendaali VU Indonesia, E. Malaysia*
Murina ryukyuana EN Ryukyus
Murina suilla LC Indonesia, E. Malaysia, Brunei*
Murina tubinaris LC Philippines*
448 G. J. Wiles and A. P. Brooke
Table A14.1. (continued)
Species IUCN statusaCountries, territories, or island
groups of occurrenceb, c
Kerivoula agnella DD PNG
Kerivoula flora VU Indonesia, E. Malaysia
Kerivoula hardwickii LC Indonesia, E. Malaysia, Brunei,
Philippines*
Kerivoula intermedia NT Indonesia, E. Malaysia, Brunei*
Kerivoula lenis LC Indonesia, E. Malaysia*
Kerivoula minuta NT Indonesia, E. Malaysia, Brunei*
Kerivoula muscina LC PNG
Kerivoula myrella DD PNG
Kerivoula papillosa LC Indonesia, E. Malaysia, Brunei*
Kerivoula pellucida NT Indonesia, E. Malaysia, Brunei,
Philippines*
Kerivoula picta LC Indonesia*
Kerivoula whiteheadi LC E. Malaysia, Philippines*
Phoniscus atrox NT Indonesia, E. Malaysia*
Phoniscus jagorii LC Indonesia, E. Malaysia,
Philippines*
Phoniscus papuensis LC Indonesia, PNG*
Note: Taxonomy largely follows Simmons 2005, with information on distribution taken from the sources
listed in table 14.1. Threatened categories are from IUCN 2008.
a Status abbreviations: CR = critically endangered; DD = data deficient; EN = endangered; EX = extinct;
LC = least concern; NE = not evaluated; NT = near threatened; VU = vulnerable.
bAsterisks denote species with geographic ranges extending outside the region. Geographic abbreviations:
E. Malaysia = Sarawak and Sabah (East Malaysia); FSM = Federated States of Micronesia; N. Marianas = Com-
monwealth of the Northern Mariana Islands; PNG = Papua New Guinea.
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