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Review
Bats as bushmeat: a global review
Simon Mickleburgh,Kerry Waylen and Paul Racey
Abstract A questionnaire survey and literature review
revealed the extent of hunting of bats for bushmeat in
the Old World tropics. High levels of offtake were reported
throughout Asia, the Pacific islands and some Western
Indian Ocean islands, where fruit bats of the genus
Pteropus are eaten extensively. Most hunting in Africa was
reported in western states and the largest fruit bat Eidolon
helvum was preferred. Insectivorous bats are also eaten,
particularly Tadarida in Asia. Hunting is both for local
consumption and commercial, sometimes involving cross-
border transactions. The high levels of hunting reported
and the low reproductive rate of bats indicate there are
likely to be severe negative effects on bat populations, and
declines of several species are documented. Although there
has been only one reported attempt to manage offtake, this
indicates that it is possible and apparently successful. Fur-
thermore, voluntary controls on hunting have halted de-
clines in bat numbers. There have been several initiatives
to reduce hunting pressure and conserve threatened bat
species, mainly on islands that, when sustained, have been
successful. More education projects and community-based
conservation initiatives should be encouraged together
with further attempts at sustainable harvesting in situa-
tions where disease risk has been evaluated.
Keywords Bats, bushmeat, commercial hunting, Eidolon,
Pteropus, sustainability, Tadarida.
This paper contains supplementary material that can be
found online at http://journals.cambridge.org
Introduction
There is growing concern about the potential impact
of the bushmeat trade on a range of animal species
(Robinson & Bennett, 2000). Attention has often focused
on primates and other large mammals (Bowen-Jones &
Pendry, 1999) and to date there has been little information
on bats. There is some evidence that hunting and trade is
having a significant impact on bat populations in the Pacific
islands and South-East Asia (Mickleburgh et al., 2002) and
also in Madagascar (Jenkins & Racey, 2008) but there is no
overall view of its potential global impact on bats. Further-
more, recent reviews of emergent viral diseases in bats have
raised concerns that eating bats as bushmeat may transmit
such diseases (Messenger et al., 2003).
The low reproductive rate of bats makes them especially
vulnerable to harvesting for bushmeat. In several life-history
characteristics bats are similar to primates that are severely
impacted by the bushmeat trade (Bowen-Jones & Pendry,
1999). Bats are long-lived and often roost communally,
which increases their visibility and susceptibility to hunters.
Their vulnerability may be further increased by roost
location and fidelity. Roosts, such as caves and trees, are
vulnerable to disturbance and bats return seasonally to roost
sites, making them predictable targets. Many bat species
also face other threats, such as habitat loss, throughout their
range, and these threats may interact with hunting to
increase their vulnerability further. The global status of
bats is reviewed by Mickleburgh et al. (1992; for Old World
fruit bats) and Hutson et al. (2001; for all other bats).
Over 20% of all mammal species are bats (Simmons,
2005). On some islands bats may be the only native mam-
mals and may be keystone species in ecosystems (Cox et al.,
1992) as pollinators and seed dispersers of plants, many of
which are economically important (Fujita & Tuttle, 1991).
The hunting of bats for bushmeat could therefore be
a significant threat to their populations, and it is of crucial
conservation and economic importance to discover the
extent of hunting. We present here the results of a global
survey of the use of bats as bushmeat, discuss the potential
conservation problems this causes, and make recommen-
dations on how these problems could be tackled.
Methods
This study was initiated over April–September 2004 using
two principal methods. The first approach was a literature
review. This included literature accumulated by a Fauna &
Flora International review of the bushmeat trade (Bowen-
Jones et al., 2002) together with online publications and
media reports. Most material reviewed was in English; some
was in Spanish and French. The second approach involved
a questionnaire (Appendix 1) widely distributed by e-mail
and advertised in the journal Phelsuma and in African Bat
Conservation News. The questionnaire requested information
S
IMON
M
ICKLEBURGH
*and K
ERRY
W
AYLEN
y
Fauna & Flora International,
Cambridge, UK.
P
AUL
R
ACEY
(Corresponding author) School of Biological Sciences, Univer-
sity of Aberdeen, Aberdeen, AB24 2TZ, UK. E-mail p.racey@abdn.ac.uk
*Current address: The Rufford Maurice Laing Foundation, London, UK.
y
Current addresses: Centre for Environmental Policy and Division of Biology,
Imperial College London, Silwood Park Campus, Ascot, UK, and Macaulay
Institute, Aberdeen, UK.
Received 5June 2008. Revision requested 12 August 2008.
Accepted 12 December 2008.
ª2009 Fauna & Flora International,
Oryx
, 43(2), 217–234 doi:10.1017/S0030605308000938 Printed in the United Kingdom
about traditional and current use of bats as bushmeat, the
impact on bat populations, the techniques used to hunt bats
and the relative importance of bats in local bushmeat trade.
Where relevant, the basis of the information provided was
also queried: for example, the suggestion that hunting
adversely affected bat populations was only occasionally
supported by research. Where necessary, data from ques-
tionnaires were clarified and supplemented by further
enquiries. Anecdotes and other information from respond-
ents were also noted.
Results
Literature review
The literature review revealed 119 references with some
information indicating bat consumption, including 87
journal papers, seven media reports, three web articles,
four university theses or other manuscripts, nine govern-
ment reports and 20 NGO reports. Given the volume of
literature on bushmeat, bat consumption is not a prominent
topic. Thirty-one apparently comprehensive studies of
bushmeat hunting or trade did not mention bats. Whilst
additional reports may exist, especially in languages other
than English, the references reviewed are likely to give
a representative indication of the level of use of bats as
bushmeat. Information provided by respondents about the
use of bats in traditional medicine will appear elsewhere.
The terms fruit bat and flying fox are used interchangeably
for members of the Pteropodidae.
Questionnaire
A total of 109 individuals responded to the request for
information, with 90 completing questionnaires (Appendix 2).
The respondents provided 138 reports on bat consumption
(Table 1) and the geographical distribution of these is given
in Table 2. The methods used to hunt bats are listed in Table 3.
The information below only covers areas where significant
bat consumption has been reported (Table 2). Other areas,
such as Eurasia, North Africa, the Arabian Peninsula,
Australia, New Zealand, North and Central America and
the Caribbean are excluded because there was little or no
evidence of consumption of bats.
The use of bats as bushmeat
Generalizations concerning each area surveyed are pre-
sented in Table 1. Specific details are given below, by region.
South-East Asia
Cambodia Hunting has impacted bat populations, partic-
ularly Tadarida plicata.
Indonesia Bergmans & Rozendaal (1988) noted .100
Pteropus alecto for sale by one trader in a market in
Sulawesi in 1982 and several other species were also
regularly traded. Clayton & Milner-Gulland (2000) made
detailed observations of the meats, including bat meat, in
markets in north Sulawesi; in the early 1990s a single fruit
bat was worth USD 0.23–1.14, and 25–50 bats per week were
typically sold; this reached 300 in 1997. In each of two
markets in Ujung Pandang, Sulawesi, 100–200 flying foxes
were traded daily and c. 8% of bats from Ujang Pandang
were exported to supermarkets in Manado City (Heinrichs,
2004). In Jakarta Pteropus vampyrus were offered by at least
two market vendors for USD 10 each, who each sold c. five
per week (Fujita & Tuttle, 1991). Bats were also readily
available at Manadonese restaurants (Fujita, 1988). How-
ever, on Karakelang, most bats were eaten by the trappers
and hunting was not commercially driven (Riley, 1998).
Whole roosts could be devastated by harvesting for markets
in north Sulawesi (Clayton & Milner-Gulland, 2000) and
five species, including one endemic, were threatened by
hunting on the islands of Sangihe and Talaud, with
Acerodon celebensis possibly warranting threatened status
(Riley, 2002). Fruit bats were heavily hunted on Karakelang
Island (Riley, 1998). Market surveys and colony observa-
tions indicated that c. 30,000 bats were killed per year in
southern Sulawesi, and overharvesting seriously threatens
flying fox populations (Heinrichs, 2004). Populations of
A.celebensis and Pteropus griseus mimus may have been
decimated and P. alecto populations reduced by 25%
(Heinrichs, 2004). In some regions smaller bat species
were hunted because flying foxes were no longer present
(Heinrichs, 2004). The impact of hunting in Indonesia may
be exacerbated because hunters leave obstructions in place
that are used to catch cave bats, which then preclude re-
establishment of some colonies. In Kalimantan c. 4,500
Pteropus vampyrus natunae were taken from one location
over a month, resulting in severe population decline
(Streubig et al., 2007). The bats were sold in markets for
USD 0.63–2.20 each. There has been some response to
problems in this region. Bat Conservation International
and Fauna & Flora International supported the work of
Scott Heinrichs to educate people in Sulawesi about the role
that bats, particularly flying foxes, play in the ecosystem,
and their conservation status, and he liaised with local
academics, students, restaurateurs and villagers.
Lao PDR The only confirmed large colony of the wrinkle-
lipped bat Tadarida plicata is heavily exploited, with several
thousand bats per day being sold in the market of Louang-
Namtha. Tadarida teniotis were also sold in markets at Ban
Lak (Francis et al., 1999) and bats may have been exported
to Thai markets (Robinson, 1994). In the Nam Ha protected
area trade was primarily local, with 97% of reported sales
being to people from the same province and 35%topeople
S. Mickleburgh et al.218
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, 43(2), 217–234
TABLE 1Summary of the use of bats as bushmeat by region and country, with information on levels of consumption, extent of trade, taxa targeted, hunting methods used, any legal
controls on hunting, references, and overall summary.
Region &
country Consumption
1
Extent
of
trade
2
Taxa
3
Hunting
methods
4
Legal
controls
5
References
6
Summary
South-East
Asia
All questionnaires & literature reports
mentioned bat consumption,
especially in karst regions where there
are likely to be many caves.
Cambodia 3 2 Cynopterus spp., Eonycteris spelaea,
Megaderma spp., Miniopterus spp.,
Pteropus lylei, Tadarida plicata
3 X Fruit bats that form large aggregations
are eaten. Bats locally traded & sold in
restaurants frequented by tourists.
Indonesia 4 2 Acerodon celebensis, Acerodon humilis,
Cynopterus spp., Dobsonia spp.,
Pteropus alecto, Pteropus griseus mimus,
Pteropus lombocensis, Pteropus
vampyrus, Pteropus vampyrus natunae,
Rousettus spp., Styloctenium wallacei,
Thoopterus nigrescens
2,4 X Owen et al., 1987; Bergmans &
Rozendaal, 1988; Fujita, 1988; Riley,
1988; Fujita & Tuttle, 1991; Hill, 1991;
Whitten, 1992; Clayton & Milner-
Gulland, 2000; Heinrichs, 2004;
Streubig et al., 2007
Bats eaten throughout the archipelago
& common in markets.
Lao PDR 4 3 Cynopterus spp., P. vampyrus,
T. plicata, Tadarida teniotis
2,3,4 X Robinson, 1994; Francis et al., 1999;
Johnson et al., 2003; Noreen &
Claridge, 2003
Consumption of bats common. Cave-
dwelling bats at risk of extirpation
chiefly because of exploitation for
food. Declines in amount of bat guano
collected support this.
Malaysia 4 2 Cheiromeles torquatus, Pteropus
hypomelanus, P. vampyrus
1,2,4 X Fujita, 1988; Fujita & Tuttle, 1991;
Mohd-Azlan et al., 2001
Consumption mainly by Chinese
population on special occasions.
Other indigenous groups occasionally
consume Cheiromeles as a delicacy.
Most bats hunted for family & friends
although some evidence of past trade
in local markets. Sabah & Sarawak
prohibit all hunting of fruit bats &
limit number of gun shells that can be
purchased, although enforcement of
regulations may be patchy.
Bats as bushmeat 219
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, 43(2), 217–234
Table 1 (Continued)
Region &
country Consumption
1
Extent
of
trade
2
Taxa
3
Hunting
methods
4
Legal
controls
5
References
6
Summary
Myanmar 3 2 Craseonycteris thonglongyai,
Rhinolophus marshalli, T. plicata
2 X Bates, 2003 All species hunted although smaller
bats preferred because bones can be
eaten with the meat. Bat populations
probably not threatened by hunting
although impacts vary with village
attitudes. Larger cave roosts protected
from overharvesting by local
communities to ensure supply of
guano.
New Guinea
(Papua
New
Guinea
& Irian
Jaya)
42Aproteles bulmerae, Dobsonia magna,
Dobsonia moluccensis, Miniopterus
schreibersii, Miniopterus magnater,
Miniopterus pusillus, Nyctimene aello,
Nyctimene cyclotis, P. hypomelanus,
Pteropus neohibernicus, Rousettus
amplexicaudatus, Syconycteris australis
1,3,4 X Lekagul & McNeely, 1977; Hladick
et al., 1993; Flannery, 1995a; Craven,
1998; Cuthbert, 2003a,b
Bats eaten throughout the highlands
& at least some populations adversely
affected by hunting. All major caves in
Irian Jaya showed signs of disturbance
from hunting.
Philippines 4 2 Acerodon jubatus, Acerodon leucotis,
Cynopterus brachyotis, Dobsonia
chapmani, E. spelaea, Harpyionycteris
whiteheadi, Macroglossus minimus,
Nyctimene rabori, Ptenochirus jagori,
P. hypomelanus, Pteropus pumilus,
P. vampyrus, R. amplexicaudatus,
insectivorous bats
1,2,3,4 X Kummer, 1992; Mickleburgh et al.,
1992; Heaney & Heideman, 1987;
Shively, 1997; Lacerna & Widmann,
1999; Mildenstein et al., 2002; Stier,
2003; Carin
˜o et al., 2006
Bats popular on special occasions but
not as subsistence food. However, bats
often the most popular bushmeat,
partly because other wild meats have
become scarce & bats are easily
hunted. When hunting is at its most
intense, in the fruiting seasons, bats
are targeted almost every night &
some hunters may take hundreds at
a time. Several bat populations,
especially tree-roosting or cave-
dwelling species, have been adversely
affected by hunting. Because loss of
habitat has been a major threat to bat
populations hunting now has
a proportionately greater impact on
those remaining. Cave-dwelling
species in Negros have declined
dramatically, almost certainly because
of the effects of hunting.
S. Mickleburgh et al.220
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, 43(2), 217–234
Thailand 4 3 C. thonglongyai, Cynopterus spp.,
E. spelaea, P. hypomelanus, P. vampyrus,
R. amplexicaudatus, Rousettus
leschenaultii, T. plicata,Taphozous
melanopogon
2,3 X Stebbings, 1987; Lekagul & McNeely,
1988; Robinson, 1993, 1994, 1995, 1998;
Magnus, 2001
Bat consumption widely reported &
cave-dwelling species especially
threatened by hunting.
Vietnam 4 2 Cynopterus spp., E. spelaea, Pteropus
spp., Miniopterus spp., T. plicata
2,3 X Thong, 2004 Only immigrant minorities, such as
Thais, consume bats, & only on
special occasions. Some populations
probably threatened by hunting. In
the south fruit bat populations have
declined, probably because of hunting,
habitat loss & cave destruction.
East &
South-East
Asia
Heavy hunting pressure reported,
especially on Pteropus.
Bangladesh 2 1 Pteropus giganteus X X Eaten occasionally by tribal people.
China 4 2 Cynopterus sphinx, Hipposideros
pomona, P. giganteus, R. leschenaultii,
Rhinolophus spp.
1,2,4 3 Bat consumption reported, especially
in the southern provinces of
Guangdong, Guangxi, Sichuan &
Hainan & in the south-west. Hunting
for bushmeat likely to be detrimental
to some populations but entire
colonies of fruit bats may also be
destroyed because of their perceived
role in causing fruit crop damage.
India
(Andaman
& Nicobar
Islands)
41Pteropus faunulus, Pteropus melanotus 1 X Hunting seasonal, coinciding with the
reproductive aggregations of bats &
has adversely impacted populations,
especially in the Andamans, which has
a more sedentary non-tribal population
than the Nicobars.
India
(Mainland)
22P. giganteus 1,2,4 X Singaravelan et al., in press In south India P. giganteus eaten at
least as regularly as other types of
bushmeat & traded locally. Such trade
is not of great economic importance &
there is no evidence that the
consumption of bats poses any threat
to populations, particularly in
comparison with habitat loss & roost
disturbance.
Nepal 2 1 Not known X X Eaten by some tribes.
Bats as bushmeat 221
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, 43(2), 217–234
Table 1 (Continued)
Region &
country Consumption
1
Extent
of
trade
2
Taxa
3
Hunting
methods
4
Legal
controls
5
References
6
Summary
Pacific
Islands
Some species severely impacted by
hunting.
American
Samoa
41Pteropus samoensis, Pteropus tonganus X X Cox, 1983; Craig & Syron, 1992; Craig
et al., 1994a,b; American Samoa Code
Annotated, 1995; Brooke, 2001
Fruit bat populations declined by 80–
90% over 5 years, mainly near human
habitation, because of losses from
cyclones, land clearance & subsequent
overhunting by villagers.
Cook Islands
& Niue
41P. tonganus X X Krzanowski, 1977; Wodzicki & Felten,
1980; Brooke & Tschapka, 2002
Evidence of overhunting.
Federated
States of
Micronesia
21Pteropus mariannus ulithiensis 1,2 X Falanruw & Manmaw, 1992 Some harvesting for subsistence use.
Fiji 3 1 Notopteris macdonaldii, Pteralopex
acrodonta, P. samoensis, P. tonganus,
Pteropus vetulus, Tadarida bregullae
X X Flannery, 1995b Popular with locals & immigrant
Chinese.
Guam &
Common-
wealth of
Northern
Mariana
Islands
43Pteropus insularis, Pteropus mariannus,
Pteropus molossinus, Pteropus
phaeocephalus
1 X Wheeler, 1980; Lemke, 1986; Wiles &
Payne, 1986; Wiles, 1987; Wiles et al.,
1989; Wiles, 1990, 1992; Wiles et al.,
1997; Utzurrum et al., 2003; Banack
et al., in press
Combination of hunting & cyclones
resulted in population declines on
many of these islands in the last 20
years. The only large & stable bat
populations probably occur on the
remote & inhospitable islands north
of Farallon de Medinilla. Import of
bats to satisfy Guam’s needs for food
has caused population declines on
other Pacific islands.
New
Caledonia
42Pteropus ornatus, P. tonganus,
P. vetulus
1,3 X Province Sud Nouvelle-Caledonie, 2003 Local consumption of bat meat
considered a problem.
Solomon
Islands
21Pteropus rayneri, cave-dwelling bats X X Richardson, 1996; Bowen-Jones et al.,
1997
Cave-dwelling species hunted.
Vanuatu 2 1 12 native bat species 3 X Chambers & Esrom, 1991 Important element of the diet of
villagers.
Western
Indian
Ocean
On some islands extent of hunting of
bats for bushmeat remains a serious
concern.
S. Mickleburgh et al.222
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, 43(2), 217–234
Madagascar 4 2 Eidolon dupreanum, Hipposideros
commersoni, Pteropus rufus, Rousettus
madagascariensis, other large
insectivorous bats
1,2,4 3 Goodman, 2006; Durbin, 2007;
Goodman et al., 2008; Jenkins & Racey,
in press
In rural areas three endemic fruit bats
may be the most important bushmeat
species after tenrecs Tenrec ecaudatus
& high offtake poses a serious threat
to some bat bat populations.
Insectivorous bats also extensively
exploited, with some colonies reduced
in size or abandoned because of
hunting.
Comoros
Islands
21Pteropus livingstonii, Pteropus
seychellensis comorensis
3 X Trewhella et al., 1995; Sewall et al., 2004 Eaten only occasionally.
Mauritius 2 1 Pteropus niger X X Traditionally eaten but not currently
a source of food.
Rodrigues 2 1 Pteropus rodricensis 1 3 Traditionally eaten but not currently
a source of food.
Seychelles 3 2 Pteropus seychellensis 2 X Racey, 1979; Nicoll & Racey, 1981;
Hutson, 1997
Widely eaten, including in hotels &
restaurants.
Sub-Saharan
Africa
Although there are few published
studies most reports of bat
consumption are from West Africa,
where bats have been traditionally
eaten and hunted. Eidolon extensively
hunted & some populations
decreasing.
Benin 2 3 Eidolon helvum, Epomophorus
gambianus, Epomops franqueti
1,2,3 X Large fruit bats regularly eaten, but no
evidence of population decline.
Cameroon 2 2 E. helvum 1,2,3 X Hunting appears to have little impact
on bat populations.
Congo
Republic
22E. helvum, Lissonycteris angolensis,
Nycteris grandis
4 X Wilson & Wilson, 1991; Bennett
Hennessey, 1995
Fruit bats appear to be a less preferred
bushmeat.
Co
ˆte d’Ivoire 2 2 E. helvum X X Rarely eaten.
Democratic
Republic
of Congo
22E. helvum, E. franqueti, Hypsignathus
monstrosus
X X Colyn et al., 1987 Fruit bats eaten seasonally.
Equatorial
Guinea
21E. helvum, E. franqueti, Hipposideros
spp., Micropteropus pusillus, Rousettus
aegyptiacus
X X Jones, 1972; Heymans, 1994; Fa et al.,
1995; Juste et al., 1995; Fa, 2000
Bats not the main bushmeat species.
Bats as bushmeat 223
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, 43(2), 217–234
Table 1 (Continued)
Region &
country Consumption
1
Extent
of
trade
2
Taxa
3
Hunting
methods
4
Legal
controls
5
References
6
Summary
Ethiopia 1 1 Not known X X Hunted because of negative
perception amongst local
communities. No evidence of
consumption.
Gabon 2 2 Not known X X Steel, 1994; Pearce, 1996 Eaten only occasionally & less popular
than other bushmeat.
Guinea 4 2 Rhinolophus maclaudi, Rhinolophus
ruwenzorii
X X Fahr et al., 2002; Fahr & Ebigbo, 2003 Cave-dwelling species particularly
threatened.
Liberia 3 2 E. helvum 3 X Anstey, 1991 Regularly consumed & traded.
Mali 2 1 Not known X X Single report of consumption.
Nigeria 3 2 E. helvum, E. franqueti, H. monstrosus X X Folorunso & Okpetu, 1975; Halstead,
1977; Adeola & Decker, 1987; Nigeria
Federal Department of Forestry, 1987
Consumption widespread. Some
controlled hunting.
South Africa 2 1 Rhinolophus spp. X X Occasional hunting.
Tanzania
(Pemba)
41E. helvum, Pteropus voeltzkowi 1,3,4 X Hutson, 1991; Seehausen, 1991;
Entwistle & Corp, 1996, 1997
Hunting widespread.
Uganda 2 2 E. helvum 4 X Ogilvie & Ogilvie, 1964; Monadjem
et al., 2007
Some local hunting in eastern
Uganda.
Zambia
(West)
21E. helvum X X Eaten by local people in the west.
South
America
Little evidence of use of bats as
bushmeat in the Americas.
Brazil 2 1 Three phyllostomid species X X Le
´vi-Strauss, 1979; Setz & Sazima, 1987;
Setz, 1991
Regular consumption by a few native
tribes.
1
1, No evidence of consumption; 2, Rarely eaten, no major threat; 3, Regularly eaten, no apparent threat; 4, Regularly eaten, probably serious threat
2
1, No internal or external trade; 2, Internal but no external trade; 3, Internal & external trade
3
Where known, individual species are identified. In some cases, the information given is more general, referring to genera or particular groups of bats (e.g. cave-dwelling species).
4
1, Firearms/airguns; 2, Nets; 3, Direct killing; 4, Other; X, No information (see also Table 3)
5
1, Well controlled; 2, Limited control; 3, No known control of hunting; X, No information
6
Refers to published information. Where no reference is shown the information has come from the returned questionnaires and anecdotal information.
S. Mickleburgh et al.224
ª2009 Fauna & Flora International,
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, 43(2), 217–234
from the same village (Johnson et al., 2003). However, some
bat species were not locally perceived as threatened; less than
1% of households surveyed in Nam Ha thought Cynopterus
sphinx was decreasing in abundance (Johnson et al., 2003).
Malaysia Bats are regarded as a luxury meat in cities. Most
vendors surveyed by Fujita & Tuttle (1991) reported selling
200–300 bats per season at USD 2.50–3.30 each. There have
been severe declines in the abundance and distribution of
P. vampyrus throughout Peninsular Malaysia, with most
extant colonies located deep in isolated and inaccessible
forests and in dense riparian vegetation. Populations of
P. vampyrus in eastern Malaysia are also declining (Mohd-
Azlan et al., 2001).
Malaysia (Peninsular Malaysia) The Department of Wild-
life and National Parks sells permits, each allowing 50
P. vampyrus to be shot, and about 40 such permits were
issued in 1year in Perak. Mohd-Azlan et al. (2001)calculated
that permits to shoot 56,000 P. vampyrus were issued between
1990 and 1996.
Malaysia (Sabah and Sarawak) In Sarawak hunting oc-
curred mostly during the bats’ reproductive season (Fujita,
1988). Cheiromeles torquatus was opportunistically cap-
tured at roosts by indigenous people. The Iban of Sarawak
could net up to 200 bats per night. Bats may also be shot for
sport or to eradicate them from fruit plantations (Fujita &
Tuttle, 1991). Although few traders specialized in bat meat,
one vendor in Sarawak sold 150–200 bats per week at c.
USD 3each, hunted by the Iban and sold to Chinese
customers. Non-specialist vendors were estimated to sell
a few thousand per year. Bat trade may thus be econom-
ically important to at least some Iban.
Myanmar Local boys at Nadkon village cave, and cement
factory workers at Saddan-Sin cave, both in Kayon state,
collect whatever species they can for consumption. In at
least one location (Nagamauk Cave) hunting stops during
the breeding season. Bats are only traded locally to sup-
plement income. For example, in Kyauk-Ta-Lone village,
1kg of bats may be sold for c. USD 1(Bates, 2003). Bat
populations are probably not threatened by hunting,
although impacts may vary according to attitudes in each
village. Many cave roosts in Myanmar, especially larger
ones, are protected from overharvesting by local commu-
nities, to ensure the supply of guano (Bates, 2003).
New Guinea (Papua New Guinea and Irian Jaya) The
Hatam, a local tribe of the northern Arfak Mountains of
Irian Jaya, eat both fruit and insectivorous bats. Bats are
also an additional source of protein for those practising
slash-and-burn agriculture. Fruit bats were probably un-
sustainably harvested by at least one group (Cuthbert,
2003a). In one village of c. 1,000 people 100–150 bats were
taken per year. When one large roost site was protected by
these villagers its bat population increased. Elsewhere,
harvesting levels by other groups, such as the Hagahai,
may be sustainable (Hladick et al., 1993). The decline and
possible extinction of Aproteles bulmerae at Luplupwintem
cave in the 1970s was probably because of the effects of
hunting with guns (Flannery, 1995a). In the early 1990s
hunting was thought to have caused the numbers of
Dobsonia moluccensis in a large cave near Herowana to
fall from many thousands to c. 200 (Cuthbert, 2003b).
Villagers voluntarily agreed to stop hunting bats in the
large cave and to harvest from small caves or take foraging
individuals. Counts of emergent individuals at the cave in
TABLE 2Geographical distribution of reports on bat consumption and any perceived problems. This is based on 138 records, including
questionnaires (Appendix 1) returned and anecdotal information provided. Only regions where there was evidence of significant
consumption of bats have been included in our analysis.
Geographical region Total no. of reports
Consumption reported Perceived problem?
Yes No Yes No
Regions included
South-East Asia 39 39 0 24 15
East & South Asia 13 8 5 2 11
Pacific Islands 6 4 2 4 2
Western Indian Ocean 14 11 3 5 9
Sub-Saharan Africa 25 16 9 9 16
South America 12 2 10 0 12
Regions excluded
Eurasia* 15 0 15 0 15
North Africa 2 0 2 0 2
Arabian Peninsula 1 0 1 0 1
Australia 1 1 0 0 1
New Zealand 2 0 2 0 2
North America 1 0 1 0 1
Central America 5 1 4 0 5
Caribbean 2 0 1 0 1
*Includes all of Europe, Russia and former Soviet Republics
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1992,1995,1996 and 2003 showed a clear rise in bat
numbers, demonstrating the possibility of sustainable bat
hunting (Cuthbert, 2003b).
Philippines P. vampyrus and Acerodon jubatus are pre-
ferred because of their larger size. Different species are
important in different areas: for example, Dobsonia chap-
mani may be preferred in Negros but insectivorous bats are
preferred in Montalban, Luzon. In south-western Negros
bat meat is second in importance to herpetofauna but in the
Tagbanua community in south Palawan it represents only
5% of game caught (Lacerna & Widmann, 1999). There is
some local bat meat trade that may involve people living far
away from roosts. Although 150,000 bats were counted at
roosts in the late 1920s, no group larger than a few hundred
was observed by Heaney & Heideman in 1987. After it was
thought to have been hunted out of caves in Negros .30
years ago (Heaney & Heideman, 1987)D. chapmani was
recently rediscovered (Simmons, 2005). Most hunters kill
only a few bats per trip but those operating at roost sites
take hundreds in a single hunt. In Palawan poorer house-
holds hunt more frequently, with bats being the target of
many hunting trips and harvest rates are high (Shively,
1997). Some communities in south-western Negros may
collect as many as 300–1,000 bats in a week during the rainy
season, amounting to 1,500–5,000 in a summer. A. jubatus
TABLE 3A summary of different methods used to hunt bats and the geographical areas where these are known to be deployed.
Method Where used
Nets or barbed burrs of Uncarina grandidieri Madagascar
Firearms (rifles, guns, etc.) Across much of Africa, Madagascar, Benin, Cameroon, Tanzania
(Pemba Island), China, India, India (Karnataka), Indonesia (Irian
Jaya), Malaysia, Philippines, Pacific States, New Caledonia
Airguns India (Andamans & Nicobars), Philippines, Federated States of
Micronesia (Ulithi Atoll)
Smoking out of roosts (E. dupreanum) Madagascar
Smoking out of roosts China, Indonesia, Philippines
Using birdlime (a sticky adhesive smeared on branches) India
Sticky forked sticks outside caves Malaysia (Sarawak)
Mist nets China, India (Karnataka), Lao PDR
Nets Indonesia (Sanguhe Island), Indonesia (north Sumatra), Indonesia
(Sulawesi), Malaysia (Sarawak), Thailand, Vietnam, Yap
Nets at roosts or fruit trees Madagascar, Benin, Cameroon (Mount Cameroon)
Nets set over cave entrances Myanmar, Philippines, Thailand
Fishing nets hoisted into feeding trees Philippines
Nets or nylon lines with fishing hooks suspended between trees Seychelles, Indonesia (north Sumatra)
Hooked ropes suspended through feeding trees Malaysia, Philippines
Kites with hooks Malaysia (Sarawak)
Nets tied to long poles to trap at boma trees (E. helvum) Cameroon (Bomboko)
Nets on end of bamboo poles India
Bamboo poles with thorns or fish hooks on the end Philippines (Palawan)
Catapaults or slingshots Comoros, Benin, Cameroon (Mount Cameroon), Cambodia, Lao
PDR, Tanzania (Pemba Island), Philippines
Killing whilst emerging from roost trees Congo (Kasai)
Killing whilst emerging from caves (D. moluccensis) Indonesia (Irian Jaya)
Bow & arrow (E. helvum) Liberia (Sinoe County)
Bow & arrow Vanuatu, New Caledonia
Crossbow New Caledonia
Thorny bushes Congo (Kasai), Tanzania (Pemba Island), Philippines
Hitting with sticks Tanzania (Pemba Island), Lao PDR (Nam Ha), Thailand (Phu
Kheio), Vietnam
Hitting with branches whilst roosting Indonesia (Irian Jaya)
Stoning Uganda (east)
Using snares Uganda (east), Indonesia (Sanguhe Island)
Using nylon rope to reach previously inaccessible roosts (A.
bulmerae)
New Guinea
Funnel trap to catch bats emerging from caves Lao PDR
Bait & hook Lao PDR
Sitting on posts & catching with hand nets Vietnam (Thanh Hoa)
Collecting by hand Lao PDR
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has also undergone a dramatic decline as it is both a forest
specialist and is heavily hunted (Mildenstein et al., 2002).
The government now recognizes the threat that hunting
poses to wildlife and it has been banned for all but in-
digenous minorities who use traditional methods, although
enforcement is weak. Some local people try to selectively
hunt the larger Pteropus species, and others may recognize
but ignore population declines. In areas of north-west
Panay and Boracay, Frankfurt Zoological Society’s Endemic
Species Conservation Project (PESCP) has encouraged local
recognition of the threats facing wildlife, including bats.
PESCP has also successfully reduced hunting through an
anti-gun campaign, where firearms can be exchanged for
rice or cash. In addition there is a flying fox conservation
and monitoring programme, with over 60 groups protect-
ing individual roosts. Dietary studies by Stier (2003) suggest
that although bats have catholic diets, A. jubatus prefer
Ficus subcordata, and educating hunters to reduce hunt-
ing at these fig trees could effectively reduce the take of
A. jubatus.
Thailand Bats were eaten by 7% of households in Phu
Kheio sanctuary (Magnus, 2001). Some, including the
world’s smallest species Craseonycteris thonglongyai, may
also be dried and sold as souvenirs (Robinson, 1993,1995).
The introduction of foreign aid and improvement in the
transport infrastructure in the 1960s meant Rousettus could
be hunted in caves in the Bangkok area and traded and sold
in restaurants. This resulted in dramatic declines and a drop
in guano production such that communities previously eco-
nomically reliant on it broke up (Stebbings, 1987). Several
species were sold (USD 0.20 each) in the daily market in
Chiang Khan, north-east Thailand, sometimes live, impaled
through their wings to prevent escape (Robinson, 1994);
some may have come from Lao PDR, indicating cross-
border trade. In remoter areas, such as villages in Phu
Kheio sanctuary, bats may only be sold if a hunter happens
to catch a surplus (Magnus, 2001). When limestone quarry-
ing and hunting came close to exterminating Eonycteris
spelaea, awareness of their role in the pollination of durian,
lobbying by conservationists, and the views of religious
leaders who regarded caves as sacred, resulted in legal
protection of the main caves and the bats (Stebbings, 1987).
All insectivorous bats and four species of Pteropus are
protected by the Wildlife Preservation and Protection Act
1992, although enforcement is weak and roost sites are
unprotected. Guano production may also provide an in-
centive for some communities to try to protect their bats at
a local level, although this may not always be successful
(Robinson, 1993).
Vietnam Bat consumption is probably less intense than in
neighbouring Thailand, especially in the north. Despite
extensive surveys into hunting and wildlife trade in Hanoi,
only one incident of bat consumption (in ConCuong district)
has been reported. In Kho Muong village in Thanh Hoa
Province, bats were collected for food from the local
bat cave for 10 days at the beginning of September, with
villagers sitting on posts to catch bats using hand-held
nets (Thong, 2004). Trade or sale of bats in restaurants is
generally not recorded in urban areas or in the north but it
has been noted in the Mekong region where there are
several restaurants selling fruit bats. At least one cave-roost
at Hang Doi Kho Muong in northern Vietnam has declined
because of hunting (Thong, 2004). Bat hunting is the
greatest threat to bat populations inhabiting Pu Luong
Nature Reserve in north-central Vietnam (Thong, 2004).
For example, in Kho Muong village local people estimate
that c. 1,000 kg of bats were collected from the local bat cave
during the annual harvest; the numbers collected declined
steadily from 1977 and a recent survey found only a small
colony in the cave (Thong, 2004). Perceptions of threats
posed by hunting vary; some consider it poses a threat to
bat populations, whereas others report that harvesting is
limited to avoid depleting large colonies and that Mini-
opterus spp. and Tadarida plicata have been regularly
harvested with no observed impact. The Fauna & Flora
International Vietnam programme has recorded cave-
roosting bat declines in northern Vietnam and is pro-
moting conservation action on this issue and awareness
amongst the local people. In Pu Luong Nature Reserve
measures to combat bat consumption were recommended
by Thong (2004). After the establishment of this reserve,
Kho Muong village was requested to end bat harvesting at
the local bat cave but recent surveys found few bats inside,
indicating either problems with compliance or ineffective
management.
East and South Asia
Bangladesh Some tribal people occasionally eat Pteropus
giganteus.
China In some areas bats are rarely consumed and always
less so than other bushmeat species. In southern China
however, bat meat is traded locally and regionally; it appears
on some restaurant menus in Guangdong and Guangxi
provinces, especially in Wuming County. Bats were seen in
markets during surveillance linked to the SARS epidemic in
2003. Bats are not specifically protected in mainland China
although proposed tougher wildlife laws, in response to
SARS, may ban the consumption of bushmeat.
India (mainland) Bats receive no statutory protection and
are classed as vermin (Singaravelan et al., in press). No bat
consumption has been observed in Arunachal Pradesh,
north-east India, despite extensive surveys that identified
the consumption of more than 40 bushmeat species. How-
ever, the aboriginal people of remote Indian forests may eat
P. giganteus, although in smaller quantities than other
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bushmeat. Anglo-Indians and aboriginal people regularly
eat bat meat in Visakhapatnam and Srikakulam Districts.
In India most bat meat is for private consumption.
India (Andaman & Nicobar Islands) Pteropus melanotus
and Pteropus faunulus are frequently hunted at night, at
their foraging and roosting trees, for consumption on special
occasions.
Nepal The Chepang, Newar, Tamang and Bahun Chetri
tribes use bats for food.
Pacific Islands
American Samoa There are several reports of consumption
of the two fruit bat species Pteropus tonganus and Pteropus
samoensis (Cox, 1983; Craig et al., 1994a,b; Brooke, 2001).
There are two relevant legislative measures: in 1986 expor-
tation and commercial hunting was prohibited and sub-
sistence hunting limited (Craig & Syron, 1992) and a 3-year
hunting ban was initiated in 1992, later extended to aid
population recovery after Cyclones Ofa and Val (American
Samoa Code Annotated, 1995). However, regulations may
be poorly known and enforced (Craig et al., 1994b) and
export of bats to Guam may continue illegally.
Cook Islands & Niue Bats are regarded as a great delicacy
in the Cook Islands, Niue and Mangaia (Krzanowski, 1977;
Brooke & Tschapka, 2002) and Rarotonga (Wodzicki &
Felten, 1980). On Niue fruit bats were hunted in an annual
2-month period that coincides with the bats’ reproductive
season (Wodzicki & Felten, 1980; Brooke & Tschapka, 2002).
On Niue Brooke & Tschapka (2002) suggested P. tonganus
was overhunted in 1998,when1,555 were shot by 60 hunters.
Federated States of Micronesia On Ulithi Atoll in the
Caroline Islands, although some people may harvest Pteropus
mariannus ulithiensis for subsistence use, only a few bats
were taken and they were not highly regarded (Falanruw &
Manmaw, 1992).
Fiji Bat consumption is popular with local people and
immigrant Chinese. The relationship between hunting and
population declines is challenged by reports from Fiji, which
suggest that although bats are intensively hunted Pteropus
vetulus remains common (Flannery, 1995b). Threats from
deforestation are considered more important.
Guam & the Commonwealth of the Northern Mariana
Islands Pteropus meat in Guam is in high demand (e.g.
Wiles, 1987,1992). Its popularity with wealthy residents, as
well as preparation methods, are described by Lemke
(1986). The Chamorros, indigenous to the Marianas,
consider it a great delicacy and serve it on special occasions
(Wiles, 1990), paying up to USD 30 for a single bat (Wiles &
Payne, 1986) despite the fact that eating it may result in
a neurogenerative disease (Banack et al., in press). On Guam
Pteropus tokudae has become extinct and populations of
Pteropus mariannus have declined severely (Wiles et al.,
1989). Many Pacific islands have supplied Guam with fruit
bats, including American Samoa, Palau, Commonwealth of
the Northern Mariana Islands, Federated States of Micro-
nesia, Fiji, Indonesia, Papua New Guinea and the Philip-
pines. The decline of fruit bats on Rota, Tinian and Saipan
was probably accelerated by harvests to supply Guam’s
market (Wheeler, 1980). Wiles et al. (1997) reported that
180,000 P. mariannus pelewensis were shipped from Palau
to Guam during 1975–1994.In1989 CITES regulations were
amended, with all Pteropus and Acerodon species included
in Appendix I or II. The seven most threatened species of
Pteropus, including P. mariannus, were placed in Appendix
I. These amendments were primarily a response to trade in
fruit bats across the Pacific to satisfy demand in Guam.
Across this area there are also several pieces of local
legislation that aim to regulate fruit bat harvesting or trade.
For example, in 1984 P. mariannus was declared Endan-
gered on Guam by the US Fish and Wildlife Service.
Initially, CITES regulations were not properly enforced
but with the arrival of a US Fish and Wildlife Service
Inspector, Guam’s imports in 1990 dropped from 3,989
during January–March to 292 in May–June (Wiles, 1990).
However, imports from other US territories then increased
as CITES could not regulate this internal trade. The use of
other legislation has been necessary to enforce the trade
limitation: for example, the situation in Palau was con-
trolled through application of the Lacey Act that prohibits
the inter-state transport of illegally killed wildlife (Wiles,
1990). Loopholes in the law have been exploited, to the
detriment of bat populations. In the Commonwealth of the
Northern Mariana Islands there are special authorizations
for bat hunts for two special festivals. However, at these
times the quota of 10–15 is exceeded, with hundreds being
killed, and there is evidence that bats harvested on Rota
may be traded to people on other islands. The small close-
knit communities of these islands also make it difficult for
local police to enforce this unpopular law (Lemke, 1986).
Lemke (1986) concluded that 75% of fruit bat colonies on
Rota had declined, and populations on Saipan had declined
to ,25 bats, and on Tinian and Aguijan there were ,10
bats on each island. All three islands supported hundreds, if
not thousands, of bats as recently as 10 years earlier.
New Caledonia In the Southern Province bat hunting is
allowed only on weekends in April, with a maximum
allowed catch of five per hunter per day (Province Sud
Nouvelle-Caledonie, 2003).
Solomon Islands About 1,000 bats per month have been
taken from limestone cave systems as an alternative protein
to fish (Richardson, 1996) and fruit bat is an ‘occasional
treat’ on the remote island of Choiseul (Bowen-Jones et al.,
1997).
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Vanuatu The 12 bat species are the only native mammals.
A questionnaire-based investigation (Chambers & Esrom,
1991) reported that fruit bats were eaten by villagers and
85% described them as important in the diet. Only one
group did not eat fruit bats, because they regarded them as
ancestors.
Western Indian Ocean
Madagascar Pteropus rufus was listed as vermin in 1961.In
1988, after the CITES listing of Pteropus, all Malagasy bats
were classed as game and can be legally hunted within
a defined season, which is May–October for fruit bats
(Durbin, 2007). Despite this, bats are hunted for food
throughout the year with both seasonal and geographical
patterns of exploitation. There is a small amount of regional
trade but most bats are sold locally or are eaten as
subsistence food. The extent of hunting of P. rufus has
resulted in a recently revised Red List categorization of
Vulnerable (Mackinnon et al., 2003; IUCN, 2008; Jenkins &
Racey, 2008). There is evidence that bat bushmeat is an
important source of food for people living with low food
security (Goodman, 2006) and that over a dozen species are
exploited for bushmeat throughout the island, although one
was cooked and fed to pigs (Goodman et al., 2008; Cardiff
et al., in press).
Islands & Archipelagos Pteropus subniger became extinct
in the Mascarene Islands in the 19th century due in part to
overhunting. Although P. rodricensis and P. niger have
traditionally been hunted for food on Rodrigues and
Mauritius, respectively, they are no longer eaten. As
a Critically Endangered species, P. rodricensis is protected,
although such protection may be removed from P. niger
because of its perceived role in damaging litchi crops. In
the Seychelles in the 1970s there was concern about the
declining numbers of P. seychellensis, which were hunted
for food. However, all guns were then confiscated and
bat numbers rose (Racey, 1979; Nicoll & Racey, 1981).
P. seychellensis remains a common item in supermarkets
(Hutson, 1997; S. Remie, pers. comm.) and was served by
a third of 65 hotels and restaurants surveyed (C. Uzice, pers.
comm.). Hutson (1997) also reported that although con-
sumption was the main threat to bats, populations were still
healthy. On the Comoros Islands P. seychellensis comorensis
is eaten only occasionally and not by all ethnic groups. The
threatened P. livingstonii is less likely to be eaten, because
of its rarity and because it is not perceived as so much of
a threat to crops (Trewhella et al., 1995; Sewall et al., 2004).
Sub-Saharan Africa Benin Larger fruit bats, such Eidolon
helvum and Epomophorus spp., are preferred and are seen
in markets, although they are probably not a regular dietary
component. Bats have also been bought in a Togolese
market near the Benin border by people from Benin.
Cameroon Consumption of fresh or smoked bat occurs,
usually by only a few people or as an irregular component
of the diet. E. helvum is a delicacy in the Bomboko area
where it can be a major source of income at peak harvesting
season, when it is sold for local consumption and to
restaurants. Bat meat is traded both locally and regionally.
Overall, bat consumption is considered negligible, espe-
cially in comparison to other bushmeat.
Congo Republic Most bat meat is hunted for family
consumption, although in south-west Congo three of five
market surveys conducted by Wilson & Wilson (1991)
found E. helvum being traded and eaten at a price lower
than any other bushmeat. Bennett Hennessey (1995) also
found that fruit bats retailed for a low price in Ouesso,
where they are still readily available in markets (R. Ruggiero,
pers. comm.).
Co
ˆte d’Ivoire E. helvum consumption was observed by one
correspondent in the 1980s.
Democratic Republic of Congo In a bushmeat survey in an
urban market in Kisangani, Colyn et al. (1987) made 2,475
observations of E. helvum and Epomops franqueti out
of a total of 73,948 observations of all types of bushmeat.
E. helvum is abundant seasonally in bushmeat markets in
Kisangani and Hypsignathus monstrosus is also sold (Plate 1).
Equatorial Guinea Jones (1972) noted that the only major
predator of E. helvum,E. franqueti and Micropteropus
pusillus was humans, and Heymans (1994) also described
hunting of E. helvum and Hipposideros spp., although bats
were not among the top 35 species preferred by consumers.
On Bioko E. helvum and Rousettus aegyptiacus were hunted
although they were not the main bushmeat species targeted
(Fa, 2000). Fa et al. (1995) and Juste et al. (1995) showed that
bat consumption could occur without trade (Jones, 1972),
perhaps reflecting the relative unpopularity of bat meat
compared with larger mammal species (Heymans, 1994).
PLATE 1 Straw-coloured fruit bats Eidolon helvum (with a single
hammer-headed bat Hypsignathus monstrosus in the centre) for
sale in the market in Kisangani, Democratic Republic of Congo
(photograph by Guy-Crispin Gembu Tungaluna).
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Ethiopia The only respondent mentioned bats being
heavily hunted because of peoples’ hatred of them, and
this may have contributed to local extinctions.
Gabon One respondent reported bat consumption. How-
ever, bats are eaten only occasionally and are less popular
than other bushmeat species.
Guinea One respondent noted that all species of bats are
hunted in caves for consumption on special occasions,
although no bat consumption was noted by Fahr & Ebigbo
(2003). Fahr et al. (2002) noted that cave roosts of Rhinolophus
maclaudi in Upper Guinea were ‘increasingly exploited’ and
R. ruwenzorii roosting in caves were vulnerable to exploita-
tion. Entire populations of cave-dwelling bats could be killed
in one visit. Although hunting is carried out only twice per
year a detrimental impact on bat populations is likely.
Liberia E. helvum is regularly consumed and traded but
they are the lowest priced item and constituted only 0.25%
of items recorded at market (Anstey, 1991).
Mali There is one report of bat consumption at Solo
village, near Manantali Dam, in Mali.
Nigeria Bat consumption seems widespread and Folorunso
& Okpetu (1975) published recipes for fruit bats. Halstead
(1977) described the harvest of E. helvum at the University
of Ife, Nigeria; bats were shot on a weekly basis over October–
March, with approximately 12,000 shot in a season. No
more than 600 bats were taken in any one shoot and there
was no apparent impact on the colony. Controlled hunting
appeared to allow the colony to grow, possibly because of
the lessened disturbance and harassment of the bats.
Adeola & Decker (1987) found E. helvum was harvested
by rural farmers during the rainy season. Bats were a cheap
meat, popular with women from Ife and surrounding areas.
South Africa There appears to be little bat consumption in
South Africa although occasional hunting of the small but
common Rhinolophus spp. may occur in the Limpopo and
Mpumalanga provinces.
Tanzania (Pemba) Although bats are not consumed in
mainland Tanzania bat meat is popular on the island of
Pemba (Entwistle & Corp, 1997). There are four fruit bat
species and Pteropus voeltzkowi is endemic. It is considered
a delicacy and is particularly prominent in the diet in
June–July (Entwistle & Corp, 1996). Entwistle & Corp (1997)
found no evidence that bats were sold but were rather
distributed among members of a hunting party for private
consumption. Seehausen (1991) described the dramatic
decline of P. voeltzkowi, comparing past reports of large
colonies with the small groups seen in 1989, and attributed
this to increasing intensity of hunting as well as habitat
destruction. Entwistle & Corp (1997) found hunting was
widespread throughout Pemba, occurring at 13 of 19
occupied roost sites, with signs of recent hunting at five
deserted roosts. An education programme led one village to
protect their flying fox roost and the number of bats at this
roost then apparently increased to become one of the
largest on the island (Entwistle & Corp, 1996). The
expansion of this education programme through Environ-
ment Clubs has resulted in a decline in hunting and an
increase in bat numbers to an estimated 21,000 (S.J. Ali, S.K.
Haji & F.M. Saleh, pers. comm.; Robinson, 2008).
Uganda There was local demand for E. helvum when they
were sold at Makerere University in Kampala after a storm
blew them to the ground (Ogilvie & Ogilvie, 1964). How-
ever, numbers of E. helvum there declined from 250,000 to
40,000 in 40 years (Monadjem et al., 2007). This species,
along with others, is eaten by the Bagisu people in eastern
Uganda, where bats are hunted for private consumption.
Zambia In western Zambia local people eat E. helvum
(C. Kandunga, pers. comm.).
South America
Despite the diversity and abundance of bats across tropical
America the majority of reports suggest bat consumption is
rare. However, a perception of bats as being dangerous can
lead to their being hunted with the aim of exterminating
populations. There is one report of regular bat consump-
tion by some native tribes, most notably the Nambiquara of
western Brazil, who consume three species of phyllosto-
mids (Le
´vi-Strauss, 1979; Setz & Sazima, 1987; Setz, 1991). In
Brazil all native species are legally protected but bats have
traditionally been regarded as dangerous and have been
exterminated in many areas.
Discussion
This review presents clear evidence that bat populations are
seriously threatened by hunting for bushmeat in several
countries, particularly Indonesia, Malaysia and the Philip-
pines, and several islands in the Indian and Pacific Oceans.
Unsurprisingly, the main target species are fruit bats of the
genus Pteropus because they are the largest, roost in trees
and forage on flowers, leaves or ripe fruit, so that their
whereabouts are predictable. Reported levels of offtake in
species with such a low reproductive rate is a matter of
serious concern. In those mainland African countries in
which bats are hunted, it is also the largest fruit bat species,
E. helvum, that is targeted, and population declines have
also been reported. Although bat faunas are richer in
Central and South America the constituent species are
small-bodied and there is little to suggest that hunting is
widespread or is having a significant effect on populations.
Legislation to protect wildlife in the developing world is
seldom effective by itself because of the difficulties of en-
forcement, and other approaches are necessary. However,
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national legislation to protect bats is preferable because its
absence in some countries, such as India, where bats are
classed as vermin, inhibits conservation action at some levels
(Singaravelan et al., in press). Occasionally, the control of
guns and ammunition has proved effective in preventing
population declines of bats hunted for food, as in the
Seychelles. Education programmes that emphasize the role
of bats in providing ecosystem services are now widely
implemented (Trewhella et al., 2005; O’Connor et al., 2006,
Jenkins et al., 2007). Local communities often resent com-
mercial hunters from outside the community taking bats
from their neighbourhood, and community conservation
programmes have proved particularly effective in Pemba and
Madagascar. The most pressing requirement is for research
into sustainable harvesting, particularly because the only
managed harvest reported to date, that organized by Halstead
(1977) in the campus of Ife University, Nigeria, appeared to
have been effective. Voluntary controls of hunting in New
Guinea (Cuthbert, 2003b) and Madagascar (Jenkins et al.,
2007) have also been successful in halting declines in bat
numbers.
The provision of reliable data on changes in population
size, to measure the impact of hunting, requires well-
designed monitoring programmes, which are particularly
challenging to implement for bats forming large aggrega-
tions and moving between roosts. Where such long-term
programmes are established, as in Uganda (Monadjem
et al., 2007), they have documented declines that are seldom
dramatic from year to year but over longer periods become
a matter of serious concern. Similar monitoring pro-
grammes have been in operation over shorter periods in
Madagascar (Jenkins et al., 2007). Long-term monitoring
programmes are difficult to sustain in the tropics but are
clearly needed.
Considering the widespread nature of hunting, there
have been few studies on the relative importance of bats in
the diet, i.e. to determine the extent to which it is an
expensive luxury reserved for special occasions, as with the
Chamorro on Guam, or adds variety to a diet that is not
protein deficient. Elsewhere it may be a starvation food, as
in south-west Madagascar (Goodman, 2006). Bat con-
sumption driven by preference, rather than need, may
require different interventions if conservation is to be
successful.
The threat of pathogen transfer from bats to people is of
growing concern. The discovery of asymptomatic Ebola
virus infections in three species of pteropodids in West
Africa (Leroy et al., 2005) raises concerns about the risk to
humans of preparing bats for consumption. Appropriate
surveillance for the presence of disease should be carried
out before sustainable harvesting programmes are
encouraged.
In conclusion we recommend: (1) Continuing surveys of
the extent to which bats are taken as bushmeat and the
incorporation of bats in surveys of general bushmeat
consumption, recognizing that the supply chain for bat
meat may differ from that for other bushmeat; these
surveys should also evaluate the relative importance of
bat bushmeat in the diet and the possible health risks of bat
consumption. (2) Where none exists, national legislation
should be introduced to protect bats or establish closed
seasons for hunting, depending on the conservation status
of the species concerned. (3) More education projects
aimed at publicizing the importance of bats as pollinators
and seed dispersers, and their role in forest ecology. (4)
More community-based projects aimed at conserving local
bat populations or, where appropriate, harvesting them
sustainably.
Acknowledgements
The authors would like to thank all of those who provided
information for this review. Their names and geographical
areas of expertise are listed in Appendix 2. We would also
like to thank Richard Jenkins, Steven Goodman and an
anonymous referee for comments on earlier drafts and
Nora Morrison at the University of Aberdeen for help in
the production of the final manuscript.
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Appendices 1–2
The appendices for this article are available online at http://
journals.cambridge.org
Biographical sketches
SIMON MICKLEBURGH has been involved in bat conservation at
a national and international level since 1984, and co-authored the two
IUCN action plans on bats. KERRY WAYLEN is concerned with
improving conservation outcomes in developing countries, using both
socio-economic and biological knowledge. She is currently carrying
out research on the evaluation of community-based conservation
projects. PAUL RACEY has worked on the ecology, and reproductive
and conservation biology of bats for over 40 years. He is Vice
Chairman of Fauna & Flora International and Co-Chair of IUCN’s
Chiroptera Specialist Group.
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