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325
Chapter 12
Exploitation of Bats for Bushmeat and
Medicine
Tammy Mildenstein, Iroro Tanshi and Paul A. Racey
© The Author(s) 2016
C.C. Voigt and T. Kingston (eds.), Bats in the Anthropocene: Conservation
of Bats in a Changing World, DOI 10.1007/978-3-319-25220-9_12
Abstract Bat hunting for consumption as bushmeat and medicine is widespread
and affects at least 167 species of bats (or c. 13 % of the world’s bat species), in
Africa, Asia, across the islands of Oceania, and to a lesser extent in Central and
South America. Hunting is particularly prevalent among the large-bodied fruit bats
of the Old World tropics, where half (50 %, 92/183) the extant species in the family
Pteropodidae are hunted. Pteropodids that are hunted are six times more likely to
be Red Listed as threatened: 66 % of species in IUCN threatened categories (CR,
EN, VU, NT), compared to 11 % of species in the ‘Least Concern’ (LC) category.
However, there still appears to be an information gap at the international level. One
third of the hunted species on the Red List are not considered threatened by that
hunting, and nearly a quarter of the bat species included in this review are not listed
as hunted in IUCN Red List species accounts. This review has resulted in a com-
prehensive list of hunted bats that doubles the number of species known from either
the IUCN Red List species accounts or a questionnaire circulated in 2004. More
research is needed on the impacts of unregulated hunting, as well as on the sustain-
ability of regulated hunting programs. In the absence of population size and growth
data, legislators and managers should be precautionary in their attitude towards
T. Mildenstein (*)
Department of Biology, Cornell College, Mt. Vernon, IA, USA
e-mail: TMildenstein@cornellcollege.edu
I. Tanshi
Department of Biological Sciences, Texas Tech University, Lubbock, TX, USA
I. Tanshi
Department of Animal and Environmental Biology, University of Benin, Benin City, Nigeria
P.A. Racey
Centre for Ecology and Conservation, College of Life and Environmental Sciences,
University of Exeter, Penryn Campus, Penryn TR10 9FE, UK
326 T. Mildenstein et al.
hunting. Roost site protection should be a priority as it is both logistically simpler
than patrolling bat foraging grounds and reduces the comparatively larger scale
mortality and stress that hunting at the roost can cause. Education and awareness
campaigns within local communities should demonstrate how bats are a limited
resource and emphasize characteristics (nocturnal, slow reproducing and colonial)
that make them particularly vulnerable to hunting pressure.
12.1 Introduction
Most of the chapters in this book (Voigt and Kingston 2016) consider negative
consequences for bats from indirect effects of anthropogenic perturbations. In con-
trast, this chapter explores the direct exploitation of bats by humans for bushmeat
and medicine.
A global review of bats as bushmeat was published in 2009 providing an over-
view of bat hunting based on published literature and a questionnaire widely
distributed among bat biologists in 2004 (Mickleburgh et al. 2009). Here, we sum-
marize what is currently known about the exploitation of bats for consumption
and medicinal use, synthesizing the 2009 review with what has been published
since and unpublished information the authors have gathered from colleagues. The
result is a comprehensive list of hunted bats species that contains nearly twice as
many species as known from either the IUCN Red List species accounts or the
2009 review. It is unclear whether the increased concern about hunting is the result
of greater actual hunting pressure, or just represents our increased understanding
of hunting impacts. What is clear is that reviews now explicitly attribute species
declines and extinction risk to hunting pressure.
Most of the chapter discusses conservation needs in light of what is known
about bat hunting. We summarize regional patterns in bat hunting and protection
efforts and highlight areas of conservation concern. We provide details of current
research aimed at learning more about hunting impacts, and we give examples of
management activities to strengthen protection efforts against population-level
effects of hunting. We end with recommendations for research to better under-
stand the effects of hunting on bat populations as well as strategies for effectively
managing hunting to support bat conservation. All common and scientific species
names follow Simmons (2005).
12.2 Background
12.2.1 Overview of Bat Hunting
Bat hunting is widespread and affects at least 167 species of bats (or c. 13 % of
the world’s 1331+ bat species, Bat Conservation International 2015), occurring
in Africa, Asia, across the islands of Oceania, and in some parts of Central and
327
12 Exploitation of Bats for Bushmeat and Medicine
South America (compiled from IUCN 2014; Mickleburgh et al. 2009, and personal
communications by the authors; Appendix). Hunting is particularly prevalent on
the large-bodied fruit bats (family Pteropodidae) in the Old World tropics, where
half (50 %, 92/183) of all extant species experience hunting pressure (Mickleburgh
et al. 2009; IUCN 2014). A much smaller proportion of insectivorous (<8 %,
75/962 species) are hunted, particularly members of the Emballonuridae,
Hipposideridae and Molossidae in Asia and Southeast Asia, Vespertilionidae in
North Africa and West and Central Asia, and Phyllostomidae in Brazil, Peru, and
Bolivia as well as Rhinolophidae in sub-Saharan Africa, (Mickleburgh et al. 2009;
Lizarro et al. 2010; IUCN 2014) (Appendix).
Bats are hunted for a variety of reasons, from their perceived medicinal proper-
ties e.g. Nicobar flying fox, Pteropus faunulus, Kingston et al. (2008); ‘small bats’
in Nepal (Tuladhar-Douglas 2008); fat from pteropodid species in Pakistan (Roberts
1997) to their use in ornate decoration such as the teeth of the Makira flying fox
(Pteropus cognatus) used for necklaces (James et al. 2008). Bats are also hunted
for sport by urban residents seeking country pursuits (e.g. large fruit bat hunting
at Subic Bay, Philippines, S. Stier, pers. comm.) and tourists seeking exotic eat-
ing experiences (e.g. Pacific flying fox, Pteropus tonganus, hunting is offered as
a recreation option at hotels in Vanuatu; A. Brooke pers. comm. in Hamilton and
Helgen 2008). However, the most widespread reason for bat hunting, by far, is for
consumption; all 167 species that are hunted are, at least in part, wanted for their
meat as a source of protein. Bat meat ranges in value from a highly sought-after del-
icacy served at special ceremonies and traditional celebrations (e.g. Pteropus mari-
annus in the Mariana Islands) to “finger food” consumed in social drinking settings
(e.g. many bat species in Southeast Asia, Mildenstein 2012; and in West Africa, M.
Abedi-Lartey pers. comm.). Elsewhere, it provides an alternative source of protein
for local people for whom meat is an expensive commodity (Jenkins and Racey
2008) and in extreme cases, bats are consumed as starvation food (Goodman 2006).
The intensity and frequency of bat harvesting varies from year round to peri-
odical depending on the seasonality of the species, hunters’ lifestyles, and/or local
legislation. On the Islands of São Tomé and Príncipe, bats are hunted opportunisti-
cally for food all year round (Carvalho et al. 2014). In Southeast Asia, regular har-
vest of bats occurs in Indonesia and the Philippines (T. Mildenstein, unpublished
data). A migratory species, the African straw-colored fruit bat (Eidolon helvum)
is hunted whenever it is present in Accra (Fig. 12.1) and Kumasi, Ghana, between
November and March (Kamins et al. 2011). Reduced hunting intensity in March
is likely due to the northward migration of forest resident bats and/or the shift in
hunters’ occupation to farming. Similarly, in Madagascar, local legislation speci-
fies two hunting seasons—one for fruit bats, and another for Commerson’s leaf-
nosed bat (Hipposideros commersoni) (Jenkins and Racey 2008), though actual
hunting intensity is driven more by local availability (e.g. the lychee season for
fruit bats). In some localities in east and western Nigeria, year round harvest of
the Egyptian rousette (Rousettus aegyptiacus) is known (Fig. 12.1) (I. Tanshi pers.
obs.), and E. helvum was documented as hunted during peak population periods in
the southwest (Funmilayo 1978; Halstead 1977).
328 T. Mildenstein et al.
Bushmeat is preferred to domestic livestock in many places because of the taste
and perceived higher nutritional alue (Mbete et al. 2011, T. Mildenstein unpub-
lished data). In locations where domestic meats and fish are generally preferred,
such as Madagascar (Randrianandrianina et al. 2010), bushmeat becomes more
important in periods of food shortage (Jenkins and Racey 2008). Similarly, on the
island of Yap (Micronesia), hunting is socio-economically based, and bats are less
desirable than seafood. Only people of lower social ranks with no access to the
coast hunt fruit bats (Falanruw 1988). Consumption of bushmeat varies indirectly
with the availability of other protein sources (e.g. in west Africa: Brashares et al.
(2004)). In areas where bats are eaten, they are rarely the only available source
of protein. The exception to this is in times of food insecurity, when people turn
to bats as a food source, especially following natural disasters (e.g. typhoons:
Aldabra flying fox, Pteropus aldabrensis, Mickleburgh et al. 2008a; Vanuatu
flying fox, Pteropus anetianus, Helgen and Hamilton 2008a; Ontong Java fly-
ing fox, Pteropus howensis, Helgen and Allison 2008; Rodrigues flying fox,
Pteropus rodricensis, Mickleburgh et al. 2008b; Samoan Flying Fox, Pteropus
samoensis and P. tonganus, Brooke 2001, and P. mariannus, Esselstyn et al. 2006,
Fig. 12.1 Collection and sales of bats in Africa a R. aegyptiacus collected by a hunter with
sticks from a limestone cave in Etapkini near Calabar, Nigeria (credit I. Tanshi), b Fruit bat
kebab on sale in Kumasi, Ghana (credit M. Abedi-Lartey), c E. helvum and H. monstrosus on sale
in a small market by the River Congo in Kisangani, DRC (credit Guy-C. Gembu)
329
12 Exploitation of Bats for Bushmeat and Medicine
USFWS 2009) and during civil unrest (e.g., Bougainville monkey-faced fruit bat,
Pteralopex anceps antrata, S. Hamilton, pers. comm.). Similarly, species found
in low-lying areas (e.g. P. aldabrensis and P. howensis) may become increas-
ingly important food to local communities as rising sea-levels destroy other food
sources (Mickleburgh et al. 2008a; Helgen and Allison 2008).
Twenty years ago marked the end of a long period of international trade in
the Pacific with many pteropodids being imported into Guam and the Northern
Mariana Islands. Once local bat populations were depleted, bats were imported
from other island groups and mainland Southeast Asia (e.g. Wiles and Payne
1986; Wiles 1992; Stinson et al. 1992). Protracted international effort eventually
led in 1987–1989 to the addition of pteropodid species to the Appendices of the
Convention on International Trade of Endangered Species (CITES), which has
stopped legal trade of bats between nation states, although a black market still
occurs (e.g. into Europe, Samuel 2013).
Currently, hunting of bats for trade tends to be locally-based, and not inter-
national, but varies widely in intensity. An extensive commercial chain of bat
trade exists outside markets in Ghana (Kamins et al. 2011). Other high levels of
trade, include that of the large flying fox (Pteropus vampyrus) in Kalimantan,
Indonesia (Harrison et al. 2011) and of the Malagasy flying fox (Pteropus rufus) in
Madagascar (Jenkins et al. 2007; Oleksy et al. 2015b). More commonly, bats are
traded locally and on a lesser scale, with relatively few individuals sold in markets
(e.g. P. vampyrus in the Philippines, Sheffers et al. 2012; and in Southeast Asia,
Mickleburgh et al. 2009). Prices per bat range from <1 USD in Southeast Asia
(Indonesia: Heinrichs 2004; the Philippines: T. Mildenstein unpublished data) to
more than 130 USD when acquired through black market trading (e.g. P. marian-
nus on Guam and the Northern Mariana Islands, USFWS 2009).
12.2.2 Hunting Overview by Region
12.2.2.1 Africa
In total, 55 species of bats are hunted in Africa, including mainly abundant large-
bodied fruit bats (Mickleburgh et al. 2009) such as E. helvum, Franquet’s epau-
letted fruit bat (Epomops franqueti), Gambian epauletted fruit bat (Epomophorus
gambianus), hammer-headed fruit bat (Hypsignathus monstrosus), R. aegyptiacus
and medium-sized species like Angolan soft-furred fruit bat (Myoncyteris ango-
lensis) (formerly Lissonycteris), Peter’s lesser epauletted fruit bat (Micropteropus
pusillus) and to a lesser degree insectivorous bats such as the large slit-faced bat
(Nycteris grandis), Maclaud’s horseshoe bat (Rhinolophus maclaudi), Ruwenzori
horseshoe bat (Rhinolophus ruwenzori) and Hipposideros species. Although insec-
tivorous bats are considered to be less palatable in many regions and may appear
to be under low hunting pressure, (Kamins et al. 2011; Dougnon et al. 2012) this
is not necessarily the case. Goodman (2006) showed that in addition to fruit bats,
330 T. Mildenstein et al.
mainly the Malagasy straw-colored fruit bat (Eidolon dupreanum), P. rufus, and
the Malagasy rousette (Rousettus madagascariensis), insectivorous H. commer-
soni is frequently hunted, especially during periods of food shortages. In addition,
while fruit bats are probably the most commonly hunted group, 64 % of the 55 bat
species hunted in Africa are animalivores (Appendix).
Hunting bats for food is common in West and Central African states where it
can be a major threat to their populations (Funmilayo 1978; Mickleburgh et al.
2009; Kamins et al. 2011). Frequent bat hunting is recorded from Benin Republic,
Ghana, Guinea, Liberia and Nigeria (Funmilayo 1978; Anstey 1991; Kamins et al.
2011; Dougnon et al. 2012) (Fig. 12.1), as well as in Cameroon, Congo Republic,
Democratic Republic of Congo (DRC), Equatorial Guinea and Gabon. High levels of
hunting have also been reported in the past from islands off Africa—the Comoros,
Madagascar, Mauritius and Rodrigues and São Tomé and Príncipe as well as Pemba
Island, Tanzania (Jenkins and Racey 2008; Carvalho et al. 2014), although conserva-
tion efforts have reduced this pressure in some of these islands (Trewhella et al. 2005).
While occasional bat hunting occurs in Mali and Zambia, there is almost no hunt-
ing in East Africa, except eastern Uganda, and bat hunting is rare in South Africa
(Mickleburgh et al. 2009). Bats are also persecuted because of negative perceptions
in Ethiopia (Mickleburgh et al. 2009) but that is not the focus of this chapter.
Although, Halstead (1977) reported the potential for sustainable harvesting of
bats at the Ile Ife campus, over-exploitation of E. helvum in southwestern Nigeria
was also evident (Funmilayo 1978). People in rural areas in southern Nigeria
admit to eating bats occasionally, whereas in parts of eastern Nigeria, R. aegyp-
tiacus is hunted intensively (Fig. 12.1). Over 3000 individuals of this species have
been collected in one night from a cave in Buanchor village by several hunters
who hunt more than once a month (I. Tanshi, unpublished).
North Africa and West and Central Asia. Bat hunting is less prevalent
in North Africa and West and Central Asia. Of the 98 bat species that occur in
this region, five are known to be hunted and these are for medicinal purposes, of
the 98 bat species that occur in this region, five (all Vespertilionidae) are known
to be hunted: long-fingered Myotis (Myotis capaccinii), Geoffroy’s myotis
(Myotis emarginatus), whiskered myotis (Myotis mystacinus), Natterer’s myotis
(Myotis nattereri), Maghrebian myotis (Myotis punicus) (Table 12.1, Appendix).
12.2.2.2 Asia
In Asia, hunting is known to affect 64 species, which represents the largest abso-
lute number of hunted bat species in a region.
Southeast Asia. The hunting pressure on bats is greatest in Southeast Asia,
where 56, or 17 % of the region’s bat species are hunted (Table 12.1, Appendix).
Bat hunting is widespread in 10 out of the 11 countries (Brunei, Cambodia,
Indonesia, East Timor, Laos, Malaysia, Myanmar, Philippines, Thailand,
and Vietnam). Only in Singapore are bats not thought to be hunted heavily
(Mildenstein 2012; IUCN 2014).
331
12 Exploitation of Bats for Bushmeat and Medicine
High levels of hunting occur in Indonesia, where there is a long history of bat
consumption (Fujita 1988) and large numbers of individuals are still sold in mar-
kets (e.g. P. vampyrus, Harrison et al. 2011; Sulawesi fruit bat, Acerodon celebensis,
gray flying fox, Pteropus griseus, black flying fox, Pteropus alecto, Heinrichs 2004).
Hunting pressure is also high in the Philippines, with a third (24/75) of its species
known to be hunted. Although Philippine bats are protected from hunting by the
Philippine Wildlife Act and the Philippine Cave Management Act, these laws are not
well enforced, and hunting for personal consumption and local trade is widespread.
In Malaysia, hunting of some species is regulated, which may curb some of the
hunting pressure but has not reduced hunting rates to sustainable levels (Epstein
et al. 2009). The laws and levels of enforcement are different for the different
regions of Malaysia. All bats are legally protected in Sarawak, but this is not the
case in Sabah and peninsular Malaysia. Illegal hunting still occurs in orchards and
by sport hunters in Sarawak at places where enforcement is lacking. Legal protec-
tion for Old World frugivorous bats is reviewed by Abdul-Aziz et al. (2015).
In Buddhist countries (Cambodia, Myanmar, Thailand and Vietnam), most roost
sites of large fruit bats are found in the gardens around temples and monasteries
because of the protection the monks provide (e.g. Ravon et al. 2014; T. Mildenstein
unpublished data). Whether this degree of protection is sufficient to maintain stable
populations of these species has yet to be investigated (Table 12.1, Appendix).
South Asia. In Bangladesh, large fruit bats are hunted for food by members of
tribal groups (Mickleburgh et al. 2009). In India and Pakistan, bats are classified
as vermin and are persecuted, although they are consumed infrequently, and more
often killed for medicinal purposes (Noureen 2014). The exception is the Indian
flying fox (Pteropus giganteus), which is eaten by indigenous forest-dwelling peo-
ple (Mickleburgh et al. 2009). On the Andaman and Nicobar islands, black-eared
Table 12.1 Proportion of bats hunted by region (Calculated by total number of extant bats spe-
cies hunted divided by the total number of bat species in the region)
Taxon Region Total# On Red
List
Not on
list
Total
hunted
%hunted
Chiroptera 1146 97 70 167 14.6
Caribbean islands 106 0 0 0 0.0
East Asia 130 3 4 7 5.4
Europe 42 0 0 0 0.0
Meso America 177 0 0 0 0.0
North Africa 41 3 1 4 9.8
North America 49 0 0 0 0.0
North Asia 43 0 0 0 0.0
Oceania 173 25 15 40 23.1
South America 249 0 8 8 3.2
South and Southeast Asia 365 43 20 63 17.3
SE 333 36 20 56 16.8
South 114 8 5 13 11.4
Sub-Saharan Africa 249 25 26 51 20.5
West and Central Asia 94 1 0 1 11
332 T. Mildenstein et al.
flying fox (Pteropus melanotus) and P. faunulus are hunted and eaten on special
occasions (Mickleburgh et al. 2009) (Table 12.1, Appendix).
North Asia. Bats are not specifically protected in China and many species are
eaten, especially in southern China, where bats are found regularly in markets
(Mickleburgh et al. 2009) (Table 12.1, Appendix). Requests from international
agencies following the SARS outbreak, (which resulted in several hundred human
deaths) that wildlife legislation be introduced in China prohibiting inter alia hunt-
ing and sale of bats have been ignored.
12.2.2.3 Pacific (Oceania)
Bats are often the only native mammals on remote Pacific Islands, and there is a
long history of bat species being hunted in many of these areas. Bats are eaten on
American Samoa, the Cook Islands and Niue, the Federated States of Micronesia,
Fiji, Guam and the Northern Mariana Islands, New Caledonia, Palau, the Solomon
Islands, and Vanuatu (Chambers and Esrom 1991; Mickleburgh et al. 2009). In
total, 40 bat species are affected, 23 % of Oceania’s bats, making this the region
with the highest proportion of hunted bat species on the planet. The value of bat
meat is highly variable in Oceania. It is a sought-after delicacy on Guam and
the Mariana Islands, where the bats are strictly protected by the United States’
Endangered Species Act (USFWS 2009). In contrast, in the nearby Federated
States of Micronesia, the same bat species are rarely eaten (Mickleburgh et al.
2009). In American Samoa, (another United States territory), bats were consumed
regularly in the past (Brooke 2001) but are now highly protected. Bat meat is also
a delicacy in the Cook Islands, Niue, and Raratonga (Brooke and Tshapka 2002)
and is a popular food on Fiji, New Caledonia, and Vanuatu (Mickleburgh et al.
2009).
12.2.2.4 South America
Bat hunting is much less common in South America, occurring in highly local-
ized areas and affecting eight species in the families Phyllostomidae (7 spp.) and
Vespertilionidae (1 sp.) (Table 12.1, Appendix).
12.3 Why Bat Hunting is a Conservation Problem
12.3.1 Negative Impacts on Bat Populations and Ecosystems
Bats are particularly vulnerable to the effects of hunting for a number of reasons.
They are long-lived for their body size (five species live >30 years, Racey 2015)
and reproduce slowly, with generally one young per year. They have a slow rate of
333
12 Exploitation of Bats for Bushmeat and Medicine
fetal growth and long gestation periods (Racey and Entwistle 2000). Females and
young bats are thus sensitive to hunting disturbance during a large portion of the
year.
Bats are nocturnal, making them susceptible to hunting at their roost sites
by day when humans can easily find them. This is especially a concern for the
highly sought-after fruit bats in the Old World, which tend to roost conspicuously,
aggregating in large numbers in the forest canopy (e.g. Mildenstein et al. 2008).
Whether roosting colonies are in caves, cliffs, or trees, hunting at the roost site is
likely to affect the entire colony. Hunting disturbance at the roost site causes injury
to many bats from the spread of shot gun pellets, large-scale infant mortality when
pups fall from fleeing mothers (Mildenstein and Stier unpublished data; R. Ulloa,
pers. comm.), and higher stress levels as resting bats are startled and forced to flee
from hunters (Van der Aa et al. 2006). In their survey of P. rufus in Madagascar,
Mackinnon et al. (2003) recorded a high incidence of abandonment of historical
roosts, which they attributed, at least in part, to high hunting pressure.
Finally, bat colonies are characterized by high roost site fidelity (e.g. Banack
1996; Brooke et al. 2000; Gumal 2004; Stier and Mildenstein 2005). Hence, bats
may be reluctant to leave when hunting starts and may find it difficult to find alter-
native roost sites after fleeing hunters. Because bats are likely to eventually return
to the preferred roost site, they are predictable prey for hunters. The overall effect
of hunting at roost sites is reduction of bat population densities to a fraction of
local carrying capacity (e.g. Mildenstein 2012).
These population-level impacts may also have negative ecological conse-
quences. Some bat species play prominent roles in insect population control, polli-
nation, seed germination and dispersal, and in many areas, bat species are essential
to forest regeneration (e.g. large fruit bats are primary seed dispersers for hemi-
epiphytic figs, Shanahan 2001; Oleksy et al. 2015a). On isolated islands, where
there is little ecological redundancy, bats are often recognized as keystone species
due to their unique roles in seed dispersal (Shilton and Whittaker 2009). Mortality
due to hunting may, therefore, have cascading effects on ecological communities
(e.g. Mildenstein 2012) and ecosystem function (e.g. McConkey and Drake 2006).
12.3.2 Negative Impacts on Humans
The negative impacts of bat hunting extend beyond natural ecosystems to human
communities. Bats in their natural ecological roles perform valuable ecosystem
services beneficial to humans (e.g. insect suppression: Cleveland et al. 2006, pol-
lination: Bumrungsri et al. 2008b, 2009, seed dispersal maintaining local water-
sheds: Banack 1998; Stier and Mildenstein 2005), all of which are reduced when
bats are hunted. Bat colonies have also proved valuable as eco-tourism attrac-
tions supporting local economies (e.g., in Costa Rica, Lao People’s Democratic
Republic, Madagascar, the Philippines, and North America; examples in Pennisi
et al. 2004). However, hunted bats that are wary of human presence often do not
334 T. Mildenstein et al.
maintain colonies in locations easily viewed by people. Hence, reductions in bat
populations as a result of hunting could have expensive ramifications on local
communities’ water supplies, agriculture, and eco-tourism industries.
Finally, the hunting of bats may also expose human communities to poten-
tially zoonotic pathogens (Leroy et al. 2005). In the past decade, considerable
attention has been paid to bats as natural reservoirs of emerging infectious dis-
eases (Calisher et al. 2006). Studies that link infectious disease outbreaks to bats
demonstrate the spillover potential through contact with bats or exposure to faeces
and urine in bat habitats (reviewed by Plowright et al. 2015). Most notable are
the Ebola virus outbreaks, which have attracted international attention. Leroy et al.
(2009) suggest that the 2007 emergence of Ebola virus in the Occidental Kasai
province of DRC could be attributable to the consumption of freshly killed bats.
The authors trace the virus spread from a first patient with bat bushmeat contact
to an outbreak of the disease in 260 persons resulting in 186 deaths in 2007. The
re-emergence of the disease in 2014 may also have arisen from contact with bats
(Saéz et al. 2015) and has proven far more deadly.
12.4 Overhunting as a Growing Concern for Conservation
Human communities have long exploited bat populations for consumption.
Current hunting pressure, however, is likely to be much greater than historical
pressure with increases in human population density, greater accessibility to natu-
ral areas, technological advances in bat capture methods and transport options, and
relaxed adherence to cultural taboos (Brooke and Tschapka 2002; Millner-Gulland
and Bennett 2003). Hence, bat hunting is likely to be unsustainable (Bradshaw
et al. 2009), especially when coupled with other anthropogenic stressors (such as
those described throughout this book).
Overhunting (commonly also “unregulated” hunting, although not all unregu-
lated hunting is unsustainable, nor regulated hunting sustainable) is a globally-
recognized threat to many wild species of animals (Robinson and Bennett 2000;
Milner-Gulland and Bennett 2003). For bats, overhunting has been a conserva-
tion concern for over three decades (Lemke 1986; Mickleburgh et al. 1992, 2002,
2009; IUCN 2014). However, there has been a substantial lag time in our identifi-
cation of which species are affected and assessment of the impact of hunting.
Twenty years ago, the conservation status of nearly half (78/160) of the Old
World fruit bats was unknown due to lack of data (compiled from Mickleburgh
et al. 1992). Today only 11 % (21/183) of the extant Old World fruit bat species on
the Red List are considered data deficient (IUCN 2014). (These two reviews may
differ slightly in their definitions of data deficient species.) This general increase
in knowledge about bats includes a better understanding of the extent of hunting
pressure. In the first conservation review, 49 (31 % of the total 160 known) Old
World fruit bat species were recognized as hunted (Mickleburgh et al. 1992). Two
335
12 Exploitation of Bats for Bushmeat and Medicine
decades later, nearly twice as many species (N = 92) are known to be hunted, rep-
resenting over half of the 183 recognized species of Old World fruit bats (IUCN
2014) (Table 12.1).
Although there has been relatively little research explicitly focused on quanti-
fying hunting impacts, the general level of concern about hunting effects on bat
conservation has increased. Using Old World fruit bats as an example, in the first
review, hunting was not considered a threat for most (60 %) of the hunted spe-
cies (Mickleburgh et al. 1992). Now, all but five of these hunted species (25/30,
83 %) have been moved up to a higher threat status because of perceived pressures
that hunting causes (IUCN 2014). Overhunting is a recognized factor in the loss of
three (and probably also the little known fourth) of the now extinct fruit bat spe-
cies (IUCN 2014) and a cause behind local extirpations within species’ historic
distributions (e.g. Polynesian sheath-tailed bat, Emballonura semicaudata, from
Vanuatu, Helgen and Flannery 2002). Similarly, the declines of seven of the ten
fruit bat species listed as critically endangered are attributed directly to hunting;
the remaining three species are still virtually unknown (IUCN 2014) (Table 12.2).
The increased concern about bat hunting may be due to greater hunting pres-
sure, or may just represent our increased awareness of hunting impacts. What is
clear is that bat conservation biologists now explicitly attribute species declines
and increased extinction risk to hunting. Seven hunted bat species previously
assumed to be unaffected by hunting (Mickleburgh et al. 1992) now have hunting
listed as a major threat (IUCN 2014). Most (68 %) of the species that are hunted,
are listed as threatened by that hunting, while only 15 % of the hunted species
are expected not to be affected. However, it should be pointed out that for the
remaining 38 % of hunted species, reviews remain ambivalent about whether hunt-
ing is a problem. Similarly, in the review of bats as bushmeat carried out in 2004
(Mickleburgh et al. 2009), 59 % of questionnaire respondents said bat hunting
Table 12.2 Comparison of the conservation status of old world fruit bat species across two
decades from two sets of species accounts: 1992 IUCN Action Plan (Mickleburgh et al. 1992);
2014 IUCN Red List; and for comparison, the 2009 Bats as Bushmeat review (Mickleburgh et al.
2009)
IUCN Action Plan 1992 IUCN Red List 2014 Bushmeat Review 2009
# species considered 160 183 138 reports
# species hunted (% total) 49 (31 %) 92 (50 %) 82 (59 %)
# species perceived as
threatened by hunting (%
total hunted)
20 (40 %) 63 (68 %) 44 (54 %)
# data deficient species
(% total species)
78 (49 %) 21 (11 %)
# hunted species listed as
LC (% total hunted)
29 (59 %) 10 (5.3 %)
# hunted species not
listed as hunted on IUCN
list (% total hunted)
Unknown 18 (20 %)
336 T. Mildenstein et al.
occurred in their region, and over half (54 %) of those species hunted were per-
ceived to be negatively affected (Table 12.2).
The general consensus among biologists and managers is that hunting is a
major conservation threat. Despite lacking measures of hunting impacts, there
are many examples of population declines and extirpations of bats that are hunted
(e.g. loss of historical bat roosts and reduced population sizes in Madagascar,
Mackinnon et al. 2003; and in the Philippines, Heideman and Heaney 1992).
Biologists studying Old World fruit bats currently rank hunting as the top conser-
vation concern for this taxon (Mildenstein 2012). The IUCN Red List also reflects
this concern. Fruit bat species that are known by the IUCN to be hunted are almost
three times more likely to be listed as threatened (N = 58 spp. in IUCN catego-
ries: CR, EN, VU, NT compared to 21 species in the “LC” category) (Fig. 12.2).
However, there still appears to be an information gap at the international level.
Nearly 42 % (70/167) of the hunted species listed in this review are not listed by
the IUCN Red List as threatened by hunting. Half of these (35 spp.) may be for
lack of awareness, as they are not known to be hunted at all by the IUCN. The
other half are described as hunted by the IUCN but not considered to be threatened
by that hunting. In other words, these 35 species are described as hunted in their
Red List species accounts, but then hunting is not included in the list of threats
(compiled from IUCN 2014).
12.5 How Hunting Affects Bats
The least known area of bat biology is population dynamics, so it is difficult to
extrapolate from hunting mortality rates to a quantitative assessment of hunting
impacts on bat populations. Hence, one of the main conservation recommen-
dations for protecting hunted species is the direct study of the population–level
impacts of hunting (IUCN 2014).
Hunting does not necessarily lead to population declines in wild species. There
are some examples of hunted bats that appear to have stable population sizes or
Fig. 12.2 Proportion of
pteropodid species in IUCN
categories listed as hunted.
69 % of species in threatened
categories (CR, EN, VU,
NT) list hunting as a threat,
compared to 25 % of species
in the “least concern” (LC)
category
337
12 Exploitation of Bats for Bushmeat and Medicine
where the effects of hunting are minimal. In these cases, hunting pressure is small
relative to the bat population size due to effective law enforcement (e.g. P. mari-
annus on Rota, Mariana Islands, Mildenstein and Mills 2013), due to cultural/
religious taboos (e.g. related to Muslim beliefs: R. obliviosus and P. seychellen-
sis in the Comoros Islands, Sewall et al. 2003, 2007; P. vampyrus in the southern
Philippines, Mildenstein 2012), and/or for the reasons given for the 35 species on
the Red List that are hunted but not considered threatened by that hunting (see
Appendix).
To evaluate the impacts of hunting on a bat population, research must compare
the direct and indirect mortality rates of hunting with that population’s capacity
for growth. Falling short of these data-intensive lines of evidence, biologists have
found other ways to provide inferences of hunting impacts, e.g., expert opinion,
models of hunting and population growth, indices to measure population growth
and/or hunting mortality, and by comparing hunted to non-hunted populations.
Below, we describe the research that has contributed to knowledge of the impacts
of hunting on bat populations to date.
12.5.1 Expert Opinion
Expert opinion surveys can be an efficient means of gathering information on con-
servation priorities when research is lacking. Because of the paucity of data on
hunting impacts, much of the current concern about bat hunting is based largely
on expert opinion derived from anecdotal evidence and observations of bat hunt-
ing impacts on local scales. Red List risk assessments for lesser known bats are
often the result of consensus among biologists who have worked on the species.
Conservation recommendations for most bat species that are hunted are based on
perceived relationships between apparent bat population declines and levels of hunt-
ing that appear to be unsustainable (e.g. Pteropus flanneryi, Helgen et al. 2008a).
Surveys of bat biologists have been used to provide overviews of bats that are
hunted and where. Most recently, Mickleburgh et al. (2009) conducted a literature
review and global survey of bat biologists in 2004 to collate what is known about
bat hunting. From 109 questionnaire respondents, there were 138 reports of bat
consumption from which the authors provided a synthesis of bat hunting, identify-
ing West Africa and Asia as the principal regions of conservation concern.
Expert opinion surveys have also shed light on hunting as the main threat and
priority for conservation management to address. Mildenstein (2012) conducted
surveys through questionnaires at two Southeast Asia regional bat conferences to
learn about threats to fruit bat species. According to the 78 participants represent-
ing all Southeast Asian countries except East Timor, hunting is the main direct
threat to fruit bats across this region.
Caveats. While expert opinion is a readily available source of information to
identify conservation priorities in lieu of data, it does not replace systematically-
acquired knowledge. There are many examples of subsequent research leading to
338 T. Mildenstein et al.
recommendations that differ from expert opinion, especially when species-specific
ecological distinctions are concerned (e.g. how to conserve co-occurring special-
ist and generalist species, Mildenstein 2012). It is incumbent upon conservation
biologists to conduct research to verify priorities identified through expert opinion
to focus conservation resources and efforts on the most urgent issues.
12.5.2 Determining Hunting Impacts on Bat Populations
To directly study the impacts of hunting, research must measure and compare
hunting mortality rates to a bat populations’ size and capacity for growth. To date,
there are only a few studies that have tried to evaluate the sustainability of bat
hunting. The first was Halstead’s (1977) on the Ile-Ife campus of the University
in Nigeria, which was unfortunately cut short by his return to the UK. Brooke and
Tschapka (2002) modeled what would be “sustainable take” on Niue, based on
the current bat population size and estimated reproductive rates. Comparing their
modeled sustainable take to the numbers of bats hunted on Niue, they determined
current hunting rates were unsustainable. Epstein et al. (2009) estimated potential
harvest rates of P. vampyrus in Malaysia as a function of the number of hunting
licenses issued. Incorporating these hunting mortality rates into their estimated bat
population growth matrix, they projected declines in the Malaysian bat population
using even the most conservative measures of hunting pressure. On Madagascar,
Goodman (2006) extrapolated total hunting pressure on hipposiderids from a sin-
gle hunter he interviewed. Comparing this estimated mortality rate to the local bat
population surveyed, he then inferred hunting levels were detrimental, because
take exceeded the breeding potential of the local bat populations.
Caveats. Rarely will information be available on harvest levels, popula-
tion sizes, and reproductive rates for the same bat species. The studies described
here work around missing information by using indices of bat harvest levels (e.g.
licenses), estimates of reproduction rates from better known congenerics, and
models of sustainable take based on rules of thumb from other harvested species
(e.g. “RR” production method, named after its authors, Robinson and Redford
1991).
When indices, estimates, and models based on other species are used, there are
caveats to consider. Researchers should be sure that the relationship between the
index and the measure of interest is known and does not vary. Estimates based
on similar species may differ from the species of interest. With population growth
rates in particular, ignoring density-dependent factors could lead to inaccurate
estimates of reproduction and population growth capability. Finally, models for
determining sustainable hunting rates, such as the RR method used by Brooke and
Tschapka (2002), predict a sustainable take rate of 40 % of the annual growth for
species with life spans the length of fruit bats. However, this rate is based on simi-
lar species with potentially different life histories and may not take into account
the other stressors that bats face today.
339
12 Exploitation of Bats for Bushmeat and Medicine
12.5.3 Measuring Hunting Mortality Rates
Surveys from local markets, hunters, and consumers can be used to gain insights
into hunting rates and trends. Studies use a variety of methods to estimate hunt-
ing pressure on bats. In the Mariana Islands, Esselstyn et al. (2006) interviewed
hunters of P. mariannus and found a 34 % increase in hunting pressure after a
recent typhoon. In Madagascar, Goodman (2006) used his interview of a single
bat hunter as a sample to extrapolate local hunting pressure on hipposiderids. In
Niue, Brooke and Tschapka (2002) used government permit records to identify
households with guns potentially used for hunting, and then interviewed a third of
these potential hunters about their bi-monthly harvest rates. The authors used their
results from this sample to extrapolate to hunting pressure over all hunters during
the two month hunting season on the island. Kamins et al. (2011) interviewed a
total of 551 Ghanians including hunters, vendors and consumers, demonstrating a
high off-take of >128,000 E. helvum per year. A recent study on the same popula-
tion used annulus markers in teeth to age bats and develop a static life table to
determine age structured survival rates (Hayman et al. 2012). Markets were sur-
veyed in Sulawesi to determine bat consumption rates of local people and num-
ber of bats exported to neighboring provinces for trade (Sheherazade and Tsang
2015). A questionnaire survey in 13 villages in Madagascar resulted in an estimate
of 6500 bats taken each year (Razakarivony 2003) and staff at a roadside restau-
rant in western Madagascar reported serving about 30 P. rufus each day, which
extrapolates to 10,000 a year (Racey et al. 2009). The largest roost counted during
surveys at that time consisted of 5000 individuals (Mackinnon et al. 2003) which
cast doubt on the reported rate of bat consumption. Nevertheless about 30 live P.
rufus were present in panniers in the food storage area of the restaurant during a
casual visit (Racey et al. 2009). Also in Madagascar, Oleksy et al. 2015b) inter-
viewed hunters to learn about bat numbers taken as well as the location, time of
night, and season in which the hunting occurred, to measure harvest rates.
Caveats. When using surveys of people’s knowledge and opinions to collect
information for conservation, it is important to remember the limitations of this
source of information. Hunters can provide insight into numbers of bats killed, but
not all of these are sold. So, when the study question involves bat trade, research-
ers must extend surveys to the end consumers of bats. To address this problem, the
surveys by both Kamins et al. (2011) and Harrison et al. (2011) employed ques-
tionnaire approaches where all actors at different stages of the commodity chain
were interviewed instead of restricting data collection only to market surveys.
There is a potential for market surveys to underrepresent the extent of fruit
bat hunting, especially when many bats are not sold in regular or bushmeat mar-
kets (Kamins et al. 2011). Mbete et al. (2011) interviewed householders in
Brazzaville and assembled a long list of bushmeat species consumed together with
details of the markets from which they were purchased. Bats were conspicuous
by their absence and enquiries (by PA Racey) revealed that they were ‘marketed
differently’.
340 T. Mildenstein et al.
In addition, hunter interviews have received conflicting reports on the accuracy
of information gathered. For example, some studies report that hunters overes-
timate what they take, as has been shown for geese in the United States where
goose hunting is legal (Andersen et al. 1996). Similarly, hunters who are being
interviewed for their unique traditional knowledge may want to please or impress
their interviewers, which could lead to overestimated harvest rates (e.g. indigenous
Aeta people in the Philippines, T. Mildenstein and S. Stier, unpublished reports).
However, in places where the species hunted is of perceived conservation concern,
hunters may underestimate their take levels to avoid scrutiny, especially if hunt-
ing is illegal (e.g. for Mariana fruit bats, P. mariannus, in the Mariana Islands, T.
Mildenstein unpublished reports).
To address the potential for inaccurate reports by hunters, some studies provide
methods for hunters to report take anonymously. The study of typhoon impacts on
hunting levels in the Mariana Islands, used a local hunter to collect data from other
hunters (Esselstyn et al. 2006). In Madagascar, Oleksy et al. (2015b) provided a
subpopulation of hunters with notebooks and monetary incentives to keep track of
hunting off-take over time and return the information anonymously at the end of
the study.
12.5.4 Estimating Hunting Impact from Population Declines
In cases where hunting mortality rates are unknown and/or difficult to measure,
study of population trends in hunted areas can provide an indication of hunt-
ing impacts. Hunting is often assumed to be the cause behind measured popula-
tion declines. For example, biologists interviewed local people at more than 30
bat roosting sites in Pakistan where bats were hunted, and the consensus of local
knowledge suggested there were large-scale declines in bat populations in areas
where hunting was common (Venkatesan 2007). The inference power of this type
of study is stronger, however, if a comparison can be made to areas free from
hunting. In comparisons of areas with and without hunting, bat population densi-
ties were 5–10 times larger when roost sites were protected (in the Philippines,
Mildenstein 2012) and up 100 times larger when their entire habitat was protected
(in the Mariana Islands, Mildenstein and Mills 2013).
Other studies have used indices of bat population size that are directly linked to
hunted bats for measuring trends. Harrison et al. (2011) used questionnaires to sur-
vey hunters and market vendors across 12 key population centers in Kalimantan,
Indonesia to gain insight into hunting intensity, seasonality, and market dynamics.
They used capture rates by hunters and sales rates in markets as indices of the pop-
ulation size of bats in the wild. From reductions in the numbers of bats captured
and the decreasing number of bats brought to market for sale, they inferred that the
bat population in the wild was declining.
Caveats. Again, caution is warranted when inferences are based on surveys
of people’s opinions and care should be taken in planning surveys and interviews
341
12 Exploitation of Bats for Bushmeat and Medicine
especially when researchers lack sociological training (St. John et al. 2013; see
also Nuno and St. John 2014 for a review of survey techniques). Similarly, it is
important to remember that population reductions in areas with hunting are the net
result of many stressors, natural and human-caused (e.g. chapters of this book),
and may not correlate directly with hunting pressure.
12.6 Conservation Management to Mitigate Hunting
Impacts
12.6.1 Enforcement of Hunting Prohibition
For threatened populations, the elimination of hunting as a threat can produce pop-
ulation-level results within decades (see also roost site protection). For areas that
have been not hunted, bat population sizes can be large, with densities (individu-
als/hectare of habitat) that are hundreds of times that of the same species in simi-
lar areas with hunting pressure (e.g. Mariana Islands, Mildenstein and Mills 2013;
Tacio 2015). Similarly, eliminating hunting allows bat populations to recover. In
American Samoa, typhoons and overhunting sent populations of P. samoensis and
P. tonganus into a sharp decline, eventually triggering a hunting ban in 1992 (Craig
et al. 1994a, b). Two decades later, the population sizes of these bats is much larger
and considered stable (Brooke and Wiles 2008, A. Miles pers. comm.).
12.6.2 Regulated Hunting
Theoretically, hunting can be sustainable if regulated tightly to ensure population
declines due to harvest do not exceed what bat populations can naturally replace
given the range of environmental variability to which they are subjected (meth-
ods in Mills 2012). Although several countries permit hunting, bat hunting has
generally proved difficult to regulate for sustainability for a number of reasons.
In some places, hunting laws are hard to understand and therefore not followed by
hunters. In Madagascar for example, bats can be hunted legally between May and
August or February to May, depending on the species (Jenkins and Racey 2008).
In addition, according to the regulations, hunting is allowed only during the day,
and game species cannot be hunted at their roost. Hence bat hunting is technically
impossible, but hunters harvest bats despite the rules (Racey et al. 2009).
In other countries, hunting regulations are poorly designed from a conserva-
tion perspective. In peninsular Malaysia, bat hunting is legal and numbers har-
vested are said to be regulated. However, while limits on the number killed are
issued by the provincial government where the hunter resides, the license owners
are allowed to hunt in multiple provinces which creates a potential for seasonal
take that far exceeds the bat populations’ capacities for regeneration (Epstein et al.
342 T. Mildenstein et al.
2009). In Niue, bat hunting is allowed for two months per year. Hunting levels are
not regulated because of the assumption that the bat supply is unlimited. However,
bats roost in sacred forest grounds that are taboo for humans to enter, making it
impossible to obtain a count of the population size. Based on models of simi-
lar species, harvest rates exceed what is expected to be sustainable (Brooke and
Tschapka 2002).
In other countries, harvest laws are not enforced or followed. For example,
the main threat to the ornate flying fox (Pteropus ornatus) in New Caledonia is
local hunting for food (Brescia 2007). There is a short legal fruit bat hunting sea-
son, which includes only the weekends of April with a quota of five bats/hunter.
However, based on reports, there is widespread and substantial illegal hunting,
including the commercial harvesting of these fruit bats (Brescia 2007).
Finally, there are a few countries where hunting is legal and unregulated. In
Pakistan and India, bats are considered “vermin” or “pests” and hunting is actu-
ally encouraged by the government without concern for long-term sustainability or
conservation (Noureen 2014).
Whenever hunting regulation is being considered, managers should be aware
that sustainable off-take will have to be much lower than projected recruitment
for a number of reasons. Current bat population sizes, distribution, and number of
populations are a fraction of historical numbers. For example, mixed colonies of P.
vampyrus and golden-crowned flying fox (Acerodon jubatus) in the Philippines are
thought to once have been present on every major island in populations numbering
in the hundreds of thousands (Heideman and Heaney 1992). Now, these bats have
been extirpated from most islands. Fewer than 15 mixed colonies remain, often
with less than 2000 individuals and dangerously small numbers of the endangered
A. jubatus. (Mildenstein 2012). In addition to already being at historically low
population sizes, other human-caused stressors (persecution, habitat fragmenta-
tion, global climate change and other perturbations described in this book) con-
tinue to act on bat populations, exacerbating detrimental effects of harvest. Hence,
it is prudent for managers to be conservative when establishing regulated harvest
limits, and to carefully monitor populations and adapt regulatory management as
needed to meet sustainable goals.
12.6.3 Control of Guns, Ammunition, and Other Bat
Hunting Tools
Gun control is expected to have a positive effect on bats. In those countries where
private gun ownership is not allowed, Pteropus often benefits. After a coup d’état
in the Seychelles in 1977, all guns were confiscated and the numbers of Seychelles
flying fox (Pteropus seychellensis) rose having previously been of some conserva-
tion concern (Racey 1979; Nicoll and Racey 1981). A similar story occurred in
Palau following the death of the President by gunshot wounds (A. Brooke, pers.
comm.).
343
12 Exploitation of Bats for Bushmeat and Medicine
In Myanmar, private gun ownership is a crime with reportedly severe penalties.
Perhaps as a result, bat populations are large and easily approached. Bats are still
harvested. For example, Pteropus are catapulted to provide medicine for asthma,
Rousettus is often netted at cave entrances and sold in a market close to Mandalay,
and insectivorous bats are also caught at cave entrances, fried and sold as beer
snacks (U Khin Maung Gyi, pers. comm.). However, the harvest rates and overall
disturbance to bats in Myanmar are thought to be much lower without guns.
Bat hunting may also be regulated through control of capturing equipment.
For example, in Sarawak, as in many countries, it is illegal to sell or buy mist
nets without a permit. This method of protecting bats, however, is only effec-
tive if hunters use commercially-manufactured nets. Many bat hunters avoid the
high cost and regulation of mist nets and make their own nets or hook and line
traps from monofilament line and other inexpensive fishing materials (e.g. in the
Philippines, Mildenstein 2012).
12.6.4 Roost Site Protection
Bats are most vulnerable at their day roosting sites. So, it is not surprising that bat
populations settle in areas where they are most protected. In Buddhist countries
like Cambodia, Myanmar, Thailand, and Vietnam, large fruit bats are commonly
found in the gardens surrounding temples and monasteries (T. Mildenstein, unpub-
lished data). The presence of monks and religious activities turn these areas into
de facto sanctuaries for bats that would otherwise experience hunting pressure. In
nearby non-Buddhist countries, such as Indonesia and the Philippines, the same
fruit bat species colonize other “safe” spots such as privately protected lands and
parks, especially in the forest interior, using topographical features that afford pro-
tection from people (e.g. along rivers, within mangrove islands, and on cliff edges,
Mildenstein 2012). Active protection of roost sites alone (i.e. hunting still occurs
away from the roost) has been shown to result in as much as ten times the num-
ber of roosting fruit bats for the same amount of forest habitat, and is especially
important for sensitive species such as ecological specialists (Mildenstein 2012).
Because of this, and the fact that roost sites are geographically predictable, conser-
vation management by local government units and non-government organizations
often target roost site protection.
Case Studies. Conservation efforts for the Pemba flying fox
(Pteropus voeltzkowi) included roost protection through the setting up of com-
munity conservation clubs (Robinson et al. 2010). The recovery following these
conservation programs led to the downgrading of the species’ Red List threat
assessment from Critically Endangered to Vulnerable.
Until recently, permanent nets were a regular method of hunting in the roost
at Analalava, Madagascar by the people of Ambatondrazaka. The national NGO,
Madagasikara Voakajy, initiated community-based protection of the fruit bat roost
by incorporating payment for local rangers in a local peanut cooperative it funded.
344 T. Mildenstein et al.
Currently, hunting at the roost has ceased and the colony has increased from 200
to nearly 2000 individuals (Razafimanahaka 2013).
In Malaysia, the Wildlife Conservation Society has worked with local commu-
nities and the government in Sarawak to establish protected roosting areas of P.
vampyrus. (M. Gumal, pers. comm. 2015). Four out of the five maternity roosting
sites identified by Gumal (2004) are now protected for P. vampyrus, including:
Loagan Bunut National Park, Sedilu National Park, Limbang Mangroves National
Park, and Bruit National Park. A fifth maternity roost site at Bukit Sarang is in the
preliminary proclamation stage for a National Park (M. Gumal pers. comm.).
In the Philippines, the Filipinos for Flying Foxes project is building on the
successes of Bat Count-Philippines by developing bat roost site sanctuaries with
local governments. The collaborating organizations (Philippine Biodiversity
Conservation Foundation and Mabuwaya Foundation) are establishing commu-
nity-managed roost site sanctuaries across the distribution of the endemic and
endangered A. jubatus and studying bat population size increases and roost site
fidelity in these newly protected roost sites (SOS 2012).
12.6.5 Education and Awareness Raising
One of the first steps toward conservation management of hunted bats is educat-
ing local communities. Bats are important to human communities in a number of
ways, particularly for the valuable ecosystem services they provide, but local com-
munities are often unaware of these. Because hunting, like other human-caused
stressors, is tied directly to population declines in bats, it is important that human
communities are aware of the trade-offs between temporary gains from bat exploi-
tation and the risk of losing bats entirely from the region. Following a knowledge/
attitude/behavior approach to understanding responsible environmental behavior
(Hines et al. 1987), communities may come to appreciate bats and support bat
conservation only after understanding their role in the environment (see Kingston
2016).
12.6.5.1 Knowledge
People are generally aware of bats present near their local communities. Bats are
not cryptic animals, especially fruit bats that aggregate in large numbers by day
using conspicuous roosting sites, and they often forage at night in fruiting and
flowering trees on farms and in residential areas. Hence, local people’s knowl-
edge of bats often surpasses that of outside biologists, especially with respect to
bat roosting locations, foraging habits, seasonal behaviors, and even threats (e.g.
local community members’ awareness of subtle seasonal changes in fruit bat diet
of P. mariannus, Mildenstein and Mills 2013). It is, therefore, surprising how lit-
tle is known about bat conservation status in these same areas. Population size and
345
12 Exploitation of Bats for Bushmeat and Medicine
growth trends tend to be unknown by biologists and managers, much less by the
non-scientific members of the local community. So, even though local people are
aware of the disturbance they may be causing, they often have no idea of the sever-
ity of population-level consequences. Because bats appear to be numerous, popular
belief is that humans can have only minimal impact on their populations. For exam-
ple, the greater short-nosed fruit bat (Cynopterus sphinx) is believed by experts to
be threatened by hunting in parts of its range, but in other parts, <1 % of local peo-
ple surveyed believe the species could be threatened by their hunting (Johnson et al.
2003). Similarly, throughout the Philippines, bats are eaten regularly with little
understanding of the impacts that harvest is causing. Hunters who join biologists on
bat population counts commonly overestimate the population size by three orders
of magnitude prior to the count and then are shocked when the counted population
is in the hundreds or low thousands (Mildenstein et al. 2007; Mildenstein 2012).
Education and awareness programs. One of the most hunted bats in sub-
Saharan Africa, E. helvum is the focus of members of the Eidolon Monitoring
Network (EMN) who conduct education activities in areas near bat colonies (J.
Fahr, pers. comm.). In Kenya and Nigeria, scientists and volunteers of the EMN
carry out education programs in schools (Fig. 12.3) and among the general public
(Tanshi et al. 2013). Education on islands around Africa has proven effective in
drawing local attention to bat protection. Examples include the recovery program
Fig. 12.3 Conservation education and bat population monitoring by volunteers in Eidolon Mon-
itoring Network in Benin City, Nigeria, school students engage in conservation outreach event,
a volunteers prepare conservation outreach materials, b volunteers counting straw-coloured fruit
bats Eidolon helvum at King square, Ring Road, Benin City, c undergraduate student volunteers
Eidolon Population Monitoring team from University of Benin, Benin City, Nigeria
346 T. Mildenstein et al.
for the P. voeltzkowi in Pemba Tanzania, for P. rufus in Madagascar, P. rodricensis
in Rodrigues and the Comoro flying fox (Pteropus livingstonii) in the Comoros
(Wilson and Graham 1992; Trewhella et al. 2005; O’Connor et al. 2006; Robinson
et al. 2010; H. Doulton, pers. comm.).
12.6.5.2 Behavior—Local Commitment to Conservation of Bats and
Bat Habitat
Finally, once communities that value bats become aware of the threats bats
face, the may start to change their behaviors to support bat conservation (but
see Kingston 2016). A multi-faceted education and awareness program in the
Comoros Islands is a good example of how outreach can lead to changes in atti-
tude and behavior that support conservation. Local citizens became involved in
monitoring bat populations and directing conservation management (Trewhella
et al. 2005).
12.6.5.3 Capacity Building of Local Rangers/PA Managers
Many programs include training and capacity building in their bat conservation
efforts. Bat Count—Philippines held a national workshop in 2004 to train pro-
tected area managers in bat identification and monitoring techniques (Mildenstein
et al. 2007; Mildenstein 2011). The project, Filipinos for Flying Foxes, is now
working with local communities to establish sustainable management practices for
bats (SOS 2012). The project trains local rangers and management staff to monitor
their bat populations and encourages them to self-regulate their hunting pressure.
In Dalaguete, Cebu, rangers have continued forest protection despite the inconsist-
encies in availability of their modest stipends provided by the local government
(SOS 2012). In Divilacan, Northern Sierra Madre, Luzon, rangers receiving bat
conservation training have elevated bat roost protection to the top priority in their
regular monitoring activities (SOS 2012).
12.6.6 Stakeholder Engagement and Citizen Science
Collaborative conservation is more likely to be sustainable. In community-
based conservation management, stakeholders from a variety of factions within
the community are required to work together to implement effective conservation
practices. This often creates unlikely partnerships that bridge normal political,
socioeconomic and religious divides. For example, former rebels work with local
government officials to monitor bats in southern Mindanao, Philippines, a region
known for often violent stand-offs between the Philippine government and Islamic
separatists (LM Paguntalan pers. comm., SOS 2012). Uniting stakeholders toward
347
12 Exploitation of Bats for Bushmeat and Medicine
the common goal of bat conservation, however, creates collaborative programs that
prove to be robust to the changes that commonly lead to the demise of wildlife
conservation programs (e.g. change in political administrations).
Validity of data. A frequent concern when working with citizen scientists, is
that data gathered by untrained biologists may be less accurate and obscure the
signal that is being studied (reviewed in Johnson 2008). However, community-
based bat counts are perhaps a best case scenario for the use of citizen science.
The data gathered are the number of bats observed, requiring just the ability to
count and no other special training or equipment. Because bats, especially fruit
bats, tend to aggregate, the population being counted is all in one place, by-pass-
ing many sources of error arising from sampling approaches to abundance assess-
ment. Finally, the goal of community-based counts is detection of population
trends across time, so that local communities can track impacts that hunting may
be causing. Studies of count error in untrained observers show that while training
and experience has a positive effect on count accuracy, counts made by untrained
observers are as likely to detect population trend direction as those made by expe-
rienced biologists (Mildenstein 2012; Mildenstein and Mills 2013; Barlow et al.
2015).
Case studies. Population monitoring and roost protection for P. rufus in
Madagascar provides a good example of the effectiveness of citizen science and
participatory conservation efforts. Following the decline in populations of P. rufus
in Madagascar from overhunting and habitat loss, the NGO Madagasikara Voakajy
engaged local communities at four roost sites for the protection of the species.
With the help of the local government, roost sites were designated for protec-
tion and firebreaks with bare ground areas constructed around roost sites. Local
volunteers where trained to monitor the roosts of P. rufus using binoculars and
hand tally counters and have continued to do so. In addition, the engagement of
local people led to an interesting partnership where habitat restoration through tree
planting is ongoing, while local farmers receive support through a crop seed loan
system. Similarly, the local community is enforcing sustainable land use within
the protected roost areas. The project organizers ascribe the success of the project
to environmental education and outreach efforts, highlighting the benefits of local
community engagement through citizen science and partnerships that improve
local economies (Mahefatiana Ralisata pers. comm.).
In Asia, Filipinos for Flying Foxes also trains local bat stakeholders as citizen
scientists. By providing these community members with the skills and experi-
ence to monitor their bat populations, the project is encouraging local stakehold-
ers to conduct regular counts and to self-regulate their hunting pressure. So far,
the project has visited more than 35 communities near to fruit bat roosts, and
trained more than 200 local stakeholders in surveying and monitoring techniques.
It is encouraging that after training, monitoring has continued by the local com-
munities. Twelve communities have counted bats subsequent to training, and five
of these have regularly conducted annual counts for 10 years after their training
(Mildenstein 2011).
348 T. Mildenstein et al.
On Guam in the Mariana Islands, P. mariannus is a threatened species that must
be monitored regularly by the US government under the US Endangered Species
Act. Guam’s last colony of P. mariannus has declined precipitously since the
establishment of the invasive brown treesnake (Boigia irregularis, USFWS 2009).
In the past 10 years the bats have no longer been aggregating in the historical
colony location but rather are seemingly scattered in the forest, making popula-
tion abundance assessments using traditional roost counting methods impossible.
Given limited human resources and adherence to historical practices, biologists
contracting with the US government have conducted fruit bat surveys sequentially
using one or two observers from single observation sites in the forest on a survey
morning. These surveys yield occasional bat sightings and location information
but provide no basis on which to estimate the population size of the bats, which
is essential to generate funding and motivate protective management of this for-
mally-recognized USA national endangered species. In 2014, a different approach
to surveying was initiated. Using 85 trained citizen scientists placed at observa-
tion stations throughout the forest, simultaneous observation permitted a survey
of about 10 % of the forest habitat on Andersen Air Force Base. This collaborative
project between the University of Guam and the U.S. Navy resulted in the first
population size estimate for the threatened P. mariannus since the early 2000s.
The survey also brought together local stakeholders representing 25 government
and non-government organizations (including schools, environmental clubs, hunt-
ers, and local media) toward the common goal of supporting the conservation of a
local endangered species (Fig. 12.4; Mildenstein et al. 2014).
Fig. 12.4 Citizen science support enabled the first population count of Mariana fruit bats on
Andersen Air Force, Guam in nearly a decade. (Survey participants are showing the number of
bats they counted on their raised fingers) (credit SSgt. M. White)
349
12 Exploitation of Bats for Bushmeat and Medicine
12.7 Recommendations for Conservation of Hunted Bats
12.7.1 More Research is Needed to Understand Hunting
Impacts
One of the major challenges to bat conservation is lack of knowledge of how
hunting affects bats, their population size, and distribution. Collecting these data
requires trained biologists, sociologists, statisticians, and well-planned survey
techniques and questionnaires that can address sensitive questions. For many
regions and species, there is little or no population information on bats, so that
population trends are unknown. Managers are therefore encouraged to start moni-
toring programs by which hunting impacts on population size can be tracked over
time. Some bat conservation initiatives provide useful models for population size
assessment and monitoring (e.g. Southeast Asian Bat Conservation Research Unit,
Filipinos for Flying Foxes, WCS Malaysia, United States Department of Defense
in the Mariana Islands, FFI Cambodia). Using the population size estimation and
monitoring described, more studies are also needed that investigate the direct and
indirect impacts of hunting on bat populations. For example, long term monitoring
projects of both people and bats, could show trends in the correlations between
number of hunted bats and bat population responses.
Finally, more needs to be learned about the people hunting bats and the con-
ditions that lead to increased hunting (Cawthorn and Hoff 2015). If it is under-
stood why people hunt (e.g. for protein? for livelihoods? to vary their diet? for
Fig. 12.5 Members of the women’s peanut cooperative in Madagascar, which grows peanuts to
supplement local protein supplies and uses a portion of the proceeds to pay rangers to protect
fruit bat roosts (Razafimanahaka 2013)
350 T. Mildenstein et al.
tradition?) and what are the patterns in hunting intensity (e.g. seasonal, food inse-
curity), conservation managers can find creative solutions for mitigating hunting
impacts (e.g. Razafimanahaka 2013; Fig. 12.5).
12.7.2 Research to Understand How to Protect Bats
There is a need to evaluate methods employed in bat protection programs.
For example, roost site protection has been correlated with greater bat density
(Mildenstein 2012). It is important to follow this up with research that demon-
strates the effectiveness of this strategy and advises managers how to proceed.
Important questions are: what is the bats’ fidelity to protected roost sites? How
quickly do bat populations increase with roost site protection? How quickly do
bats become habituated to human presence?
Protection of habitat outside the roost area is also important, although much
more difficult to enforce. Studies of the relationship between foraging habitat and
bat population sizes would guide managers in their habitat-based conservation
strategies.
Equally important is to understand where protection efforts are failing. There
are many examples of regulated bat hunting leading to population declines. It is
important to understand where laws and implementation are falling short and why.
12.7.3 Education/Outreach
Education and outreach in local communities is essential to successful hunting
management campaigns. Many hunters do not perceive bats as a limited resource
and are unaware of the effect they may be having on bat species’ extinction risks.
An obvious first step to bat conservation in hunted areas is therefore the dissemi-
nation of information on the bat population size, basic biology, and monitoring
techniques, so that hunters can assess the impacts they are having.
Local communities should understand the benefits of bats and the valuable
ecological services they provide, including their contribution to forest regenera-
tion through seed dispersal. Communities that appreciate bats may be more likely
to pursue conservation management. If people understand the array of risks of
ignoring declining populations, they will be more inclined to exert effort to pro-
tect bats.
Local communities should also understand the human health risks of bat
hunting and consumption. Bat handling, trade, preparation, and consumption by
humans create a direct transmission route for disease spillover into human com-
munities and warrants consideration in bat protection programs. Education about
these risks is needed, as only a small percentage of participants in bat hunting and
351
12 Exploitation of Bats for Bushmeat and Medicine
trade are aware of their risk of exposure to disease in Asia (Harrison et al. 2011)
and Africa (Subramanian 2012). Finding a balance between the needs for bat con-
servation, sustainable harvesting and public health management is an important
approach in the regions where regulated bat hunting is a goal (Halstead 1977).
Change in perceptions and attitudes towards conservation and wildlife in gen-
eral require an effective outreach approach. Thus, education and public awareness
projects should be designed to engage the audience as has been demonstrated to
be effective in Madagascar (Racey 2013) and Latin America (Navarro 2013). The
involvement of all stakeholders and policy makers in conservation outreach pro-
jects is crucial to the effectiveness of education programs by facilitating the enact-
ment and enforcement of protective legislation (Robinson et al. 2010).
12.7.4 Protect Colony Locations at the Roost
Bat conservation through roost protection by local communities has been dem-
onstrated to be effective for the recovery of previously declining populations
(Mildenstein 2012; Fig. 12.6). The adoption of such roost protection programs in
other countries could hold the key to sustaining populations. This is especially true
for areas where fruit bat hunting is intense. If successful roost site protection pro-
grams could be demonstrated and published, these could be used as models for
other areas (e.g. P. rufus populations in Madagascar—M. Ralisata pers. comm.; P.
vampyrus in Malaysia, M. Gumal, pers. comm.; P. vampyrus and A. jubatus popu-
lations in the Philippines, SOS 2012).
Fig. 12.6 Tourists viewing formerly hunted fruit bats at their protected roost site in Mambukal
Resort, Negros Occidental, Philippines (credit LM Paguntalan)
352 T. Mildenstein et al.
12.7.5 Regulated Hunting
In many areas where hunted bats are threatened, hunters do not want to extir-
pate bat populations, but they also do not want to lose the ability to hunt bats
(Mildenstein 2012; Cawthorn and Hoffman 2015). In fact, some roost site protec-
tion campaigns are successful, because hunting outside the roost site is not dis-
cussed or prohibited, making it easier for hunters to respect roost site sanctuaries
(T. Mildenstein pers. obs.; SOS 2012). Once communities understand that human
disturbance has population-level impacts and that conservation management must
balance negative impacts with the bats’ innate ability to add to their population,
community-level planning of a sustainable hunting program can ensue. Targets
must be established for minimum population sizes and numbers of viable popu-
lations before hunting can be allowed. After thresholds are reached, sustainable
harvest levels must be determined using adaptations of the well-developed harvest
management practices for other species.
Finally, an effective enforcement and harvest regulation program must be
designed that starts out conservatively, carefully tracking impacts of hunting
on bat populations and making adjustments to hunting allowances as needed.
Halstead (1977) described how regulated hunting of E. helvum at the University
of Ile Ife in south western Nigeria can be mutually beneficial to the bat population,
local community livelihoods, and managers of property where roosts are present.
In places where hunting laws are in place but not respected or enforced, educa-
tion and outreach are instrumental in garnering public support (as Madagasikara
Voakajy has done for roosts of P. rufus in Madagascar).
12.7.6 Encourage Local Researchers and NGO’s
A key to effective and sustainable conservation is to develop the capacity of local
people, including local researchers and the establishment of local NGOs (Racey
2013). Few detailed studies report reliable estimates of bat hunting impacts on bat
populations. Some studies may indeed have been conducted but remain as Masters
or PhD theses or published as grey literature or in local journals, thereby limiting
the distribution of such information. Because valuable results are not often pub-
lished or accessible, current efforts to revise species account entries in the IUCN
Red List have had to rely heavily on experts gathering unpublished information to
determine conservation priorities for hunted species (T. Mildenstein, unpublished
data). It is important that biologists are encouraged to publish their findings, even
in lesser developed countries where there are few if any personal incentives for
doing so (Milner-Gulland et al. 2010).
Finally, the establishment of local non-profit organizations creates a network
for stakeholders and a bridge between local interests and conservation manage-
ment. Such organizations play a critical role in ensuring the sustainability of con-
servation projects across political administration changes by engaging the local
353
12 Exploitation of Bats for Bushmeat and Medicine
stakeholders and coordinating conservation activities in harmony with local needs
(e.g. Figs. 12.5 and 12.6).
12.8 Conclusion
Conservation biologists’ understanding of the role hunting plays in bat popula-
tion declines has changed over the last three decades. Conservation concerns were
originally focused on large scale hunting operations and especially international
commercial trade (e.g. Pteropus spp. in the Pacific and Southeast Asia). After
international trade was largely shut down in the late 1980s, conservation managers
turned their attention to hunting within countries, still focusing on commercially
hunted species as a highest priority (e.g. Mickleburgh et al. 1992).
Although commercially harvested species are still a high priority today, con-
servationists’ concerns are no longer limited to species found in markets. With
more research and experience, conservation managers have become increasingly
aware of the negative impacts caused by hunting even on small scales, i.e. for per-
sonal use and/or local trade. Especially detrimental is hunting at roost sites, which
can lead to a tenfold increase in population declines (Mildenstein 2012). This is
probably because hunting disturbance at roosts also affects non-target individu-
als, including mothers and babies which are especially sensitive to disturbance.
For hunted bat taxa (e.g. Old World fruit bats), hunting now ranks as a top threat
among bat conservation biologists.
Research that quantifies the relationship between hunting rates and bat popula-
tion declines is still lacking, but general awareness about the breadth of bat hunt-
ing effects has increased. The number of bat species known to be hunted is larger
than in earlier reviews. Similarly, biologists now recognize that hunting is usually
a threat to bats; the number of species documented as threatened by hunting is
much larger now and includes many species that are not commercially hunted.
However, for 28 % of the species known to be hunted according to IUCN Red List
species accounts, hunting was either not considered to be a threat or not evaluated
at all. Finally, information about a quarter (38/167) of the species listed as hunted
by this review has come from sources other than the IUCN Red List, where there
is no mention of hunting for those species. More research on population sizes and
trends, hunting impacts, and effective management tools will provide very impor-
tant information for bat conservation.
Research should also address the role of bat meat in local people’s diets. Studies
that focus on seasonal patterns in bat consumption and the dependency on bats as a
source of protein would provide managers with information that would guide pol-
icy and conservation actions complementary to the dietary needs of local communi-
ties. Similarly, research on the use of bats for medicinal purposes should investigate
patterns of bat use and ailments that bat consumption is said to cure. Study of the
effects of bat use on ailing consumers in situ as well as in randomized trials could
play a significant role in conservation by helping tease out whether the medicinal
354 T. Mildenstein et al.
properties of bats have a real effect or are a myth. These investigations should
begin with asthma and other chest complaints since anecdotes about the curative
effects of bats for such conditions are widespread across the Old World tropics.
Along with conservation-focused research, public education and capacity build-
ing of local managers must also be encouraged to counter what is clearly a major
threat to bat populations in the Old World tropics. Greater awareness about bats’
reproductive characteristics of one young a year together with the ecological ser-
vices bats provide will strengthen local communities’ commitment to supporting
conservation management. Hunters, in particular, begin to cooperate, even track-
ing their harvest rates as a community, when they realize that bats are a limited
resource and that populations may be extirpated altogether if hunting pressure is
not halted or highly regulated (Mildenstein 2011). Local managers can be empow-
ered to track bat conservation and hunting with training in the simple and inexpen-
sive field techniques needed to monitor bat population abundance, and these local
stakeholders are key to creating sustainable monitoring programs.
Hunting has already led to the loss of four bat species in the last few decades.
Without research, public education and awareness, and bolstering local managers’
capacity to protect bats, unregulated hunting may well claim more many species.
Acknowledgment We thank Anne Brooke and three anonymous reviewers for helpful
comments that improved the manuscript.
Open Access This chapter is distributed under the terms of the Creative Commons Attribution
Noncommercial License, which permits any noncommercial use, distribution, and reproduction
in any medium, provided the original author(s) and source are credited.
Appendix. List of Hunted Bat Species Showing Primary Use
(Food or Medicine), Summarized by Region and Country.
We Followed IUCN Regional Classification
IUCN
region
Country in
which hunted
Species Status Food Medicine Source
East Asia China Cynopterus
sphinx
LC x Bates et al. (2008d),
Mickleburgh et al. (2009)
China Eonycteris
spelaea
LC x Francis et al. (2008c),
Stebbings (1987)
China Hipposideros
pomona
LC x x Bates et al. (2008a)
China Pteropus
giganteus
LC x x Molur et al. (2008a),
Mickleburgh et al. (2009)
China Rousettus
leschenaultii
LC x x Molur et al. (2002),
Bates and Helgen (2008),
Mickleburgh et al. (2009)
Japan Pteropus
pselaphon
CR x Ishii and Maeda (2008)
Taiwan Pteropus
dasymallus
NT x Heaney et al. (2008d)
355
12 Exploitation of Bats for Bushmeat and Medicine
IUCN
region
Country in
which hunted
Species Status Food Medicine Source
North
Africa
Algeria,
Morocco,
Tunisia, Libya
Myotis punicus NT x Aulagnier et al. (2008)
Algeria, North
Africa
Myotis
emarginatus
LC x Hutson et al. (2008h)
North Africa Myotis
capaccinii
VU x Hutson et al. (2008g)
North Africa Myotis nattereri LC x Hutson et al. (2008a)
Oceania American
Samoa, Cook
Islands and
Niue, Fiji, New
Caledonia,
Vanuatu
Pteropus
tonganus
VU x Hamilton and Helgen
(2008),
American
Samoa, Fiji,
Samoa
Pteropus
samoensis
NT x Brooke and Wiles (2008)
Fiji Mirimiri acro-
donta/
Pteralopex
acrodonta
CR Flannery (1995b)
Fiji, Vanuatu Chaerephon
bregullae/
Tadarida
bregullae
EN x Flannery (1995b),
Palmeirim (2014)
Fiji, Vanuatu Notopteris
macdonaldi
VU x Flannery (1995b),
Palmeirim et al. (2007),
Palmeirim (2008)
Pteropus
ualanus
VU x Wiles et al. (2008)
Indonesia, Papua
New Guinea
Dobsonia
moluccensis
LC x Hutson et al. (2008j)
Micronesia,
Federated
States of
Pteropus
yapensis
VU x Wiles et al. (2008b)
Micronesia,
Federated
States of;
Caroline Islands
Pteropus insu-
laris/
Pteropus
phaeocephalus
CR x Helgen and Wiles (2010)
Micronesia Pteropus
molossinus
VU x Buden et al. (2008)
Micronesia,
Guam and
Commonweatlh
of Northern
Mariana Islands
Pteropus
mariannus
EN x Falanruw and Manmaw
(1992), Allison et al.
(2008), Lemke (1992)
New Caledonia Notopteris
neocaledonica
VU x Brescia (2008a), Boissenin
and Brescia (2007)
New Caledonia Pteropus
ornatus
VU x Brescia (2008b)
New Caledonia Pteropus vetulus VU x x? Brescia (2008c), Flannery
(1995b)
Palau Pteropus
pelewensis
NT x Wiles (2008), Wiles et al.
(1997)
356 T. Mildenstein et al.
IUCN
region
Country in
which hunted
Species Status Food Medicine Source
Papau New
Guinea
Pteropus
hypomelanus
LC x Francis et al. (2008a),
Fujita and Tuttle (1991),
Fujita (1988)
Papua New
Guinea
Aproteles
bulmerae
CR x Flannery (1995b), Hutson
et al. (2008m)
Papua New
Guinea
Miniopterus
magnater
LC x Bonaccorso and Reardon
(2008b), Cuthbert (2003a,
b)
Papua New
Guinea
Nyctimene aello LC x Bonaccorso and Helgen
(2008), Cuthbert (2003a, b)
Papua New
Guinea
Nyctimene
cyclotis
DD x Cuthbert (2003a)
Papua New
Guinea
Pteralopex
flanneryi
CR x Helgen et al. (2008a)
Papua New
Guinea
Pteropus
neohibernicus
LC x Salas et al. (2008)
Papua New
Guinea
Rousettus
amplexicaudatus
LC x Csorba et al. (2008g),
Utzurrum (1992)
Papua New
Guinea
Syconycteris
australis
LC x Cuthbert (2003a)
Papua New
Guinea, pos-
sibly Cambodia
and Vietnam
Miniopterus
pusillus
LC x Cuthbert (2003a, b)
Papua New
Guinea,
Solomon
Islands
Pteralopex
anceps
EN x Helgen et al. (2008c)
Papua New
Guinea,
Vanuatu
Miniopterus
macrocneme
DD x Bonaccorso and Reardon
(2008a)
Solomon
Islands
Pteralopex
atrata
EN x Helgen and Hamilton
(2008b)
Solomon
Islands
Pteralopex taki EN x Hamilton et al. (2008a)
Solomon
Islands
Pteropus
cognatus
EN x James et al. (2008)
Solomon
Islands
Pteropus
nitendiensis
EN x Leary et al. (2008a)
Solomon
Islands
Pteropus
rennelli
VU x Hamilton et al. (2008b)
Solomon
Islands
Pteropus
tuberculatus
CR x Leary et al. (2008b)
Solomon
Islands, Papau
New Guinea
Pteropus rayneri NT xHamilton and Leary (2008),
Bowen et al. (1997)
Vanuatu Emballonura
semicaudata
EN x Bonaccorso et al. (2008),
Chambers and Esrom
(1991)
Vanuatu Miniopterus
tristis
LC x Chambers and Esrom
(1991)
357
12 Exploitation of Bats for Bushmeat and Medicine
IUCN
region
Country in
which hunted
Species Status Food Medicine Source
Vanuatu Pteropus
anetianus
VU x Mickleburgh et al. (1992),
Helgen and Hamilton
(2008a), Chambers and
Esrom (1991)
Vanuatu Pteropus
fundatus
EN x Helgen and Hamilton
(2008c), Chambers and
Esrom (1991)
Vanuatu,
others?
Aselliscus
tricuspidatus
LC x Bonaccorso et al. (2008),
Chambers and Esrom
(1991)
Vanuatu, Papua
New Guinea
Miniopterus
australis
LC x Chambers and Esrom
(1991)
South
America
Bolivia Artibeus sp. LC Lizarro et al. (2010)
Bolivia Carollia
perspicillata
LC x Lizarro et al. (2010)
Bolivia Desmodus
rotundus
LC Lizarro et al. (2010)
Bolivia Myotis sp. ? Lizarro et al. (2010)
Brazil Glossophaga sp. LC or
DD
x Mickleburgh et al. (2009)
Brazil sp. 1 x Lévi-Strauss (1979), Setz
and Sazima (1987), Setz
(1991)
Brazil sp. 2 x Lévi-Strauss (1979), Setz
and Sazima (1987), Setz
(1991)
Brazil sp. 3 x Lévi-Strauss (1979), Setz
and Sazima (1987), Setz
(1991)
South
Asia
Bangladesh Pteropus
giganteus
LC x x Mickleburgh et al. (2009),
Molur et al. (2008a)
India Hipposideros
speoris
LC x Molur et al. (2008b)
India Latidens
salimalii
EN xMolur and Vanitharani
(2008)
India Megaderma lyra LC x x Csorba et al. (2008a)
India Megaderma
spasma
LC x x Csorba (2008e)
India Nyctalus
montanus
LC x Molur and Srinivasulu
(2008)
India Pteropus
faunulus
VU x Kingston et al. (2008);
Singaravelan et al. (2009)
India Pteropus
melanotus
VU x Hutson et al. (2008d),
Mickleburgh et al. (2009)
India Taphozous
melanopogon
LC x Csorba et al. (2008f),
Molur et al. (2002)
India Taphozous
theobaldi
LC x x Bates et al. (2008e), Molur
et al. (2002)
India, Sri
Lanka
Hipposideros
lankadiva
LC x x Molur et al. (2008c)
358 T. Mildenstein et al.
IUCN
region
Country in
which hunted
Species Status Food Medicine Source
South Asia
wide
Rousettus
leschenaultii
LC x Molur et al. (2002),
Bates and Helgen (2008),
Mickleburgh et al. (2009)
South Asia
wide
Cynopterus
sphinx
LC x Bates et al. (2008d), Molur
et al. (2002)
South
East
Asia
Brunei
(Borneo),
Cambodia, Lao
PDR, Malaysia,
Thailand,
Vietnam
Chaerephon
plicatus/
Tadarida plicata
LC x Csorba et al. (2014)
Brunei,
Indonesia,
Malaysia,
Philippines,
Thailand
Pteropus
vampyrus
NT x Bates et al. (2008f),
Clayton and Milner-
Gulland (2000)
Cambodia,
Indonesia,
Lao PDR,
Philippines,
Thailand,
Vietnam
Cynopterus
brachyotis
LC xLacerna and Widmann
(1999)
Cambodia,
Indonesia, Lao
PDR, Thailand,
Vietnam
Cynopterus
sphinx
LC x x Bates et al. (2008d),
Johnson et al. (2003)
Cambodia,
Philippines,
Thailand,
Vietnam
Eonycteris
spelaea
LC x Mickleburgh et al. (2009)
Cambodia,
Thailand
Pteropus lylei VU x Bumrungsri et al. (2008b);
Mickleburgh et al. (2009)
Indonesia Acerodon
celebensis
LC x Hutson et al. (2008c),
Clayton and Milner-
Gulland (2000)
Indonesia Acerodon
humilis
EN x Hutson et al. (2008b),
Clayton and Milner-
Gulland (2000)
Indonesia Acerodon
mackloti
VU x Hutson et al. (2008i)
Indonesia Cheiromeles
parvidens
LC x Csorba et al. (2008b)
Indonesia Harpyionycteris
celebensis
VU x Hutson et al. (2008l)
Indonesia Neopteryx frosti EN x Hutson et al. (2008k)
Indonesia Pteropus alecto LC x Bergmans and Rozendaal
(1988), Hutson et al.
(2008n)
Indonesia Pteropus
caniceps
NT x Hutson and Helgen (2008a)
Indonesia Pteropus
chrysoproctus
NT x Hutson and Helgen (2008b)
Indonesia Pteropus griseus DD x Heinrichs and Zahnke
(1997)
359
12 Exploitation of Bats for Bushmeat and Medicine
IUCN
region
Country in
which hunted
Species Status Food Medicine Source
Indonesia Pteropus
lombocensis
DD x Clayton and Milner-
Gulland (2000), Helgen and
Salas (2008a)
Indonesia Pteropus
melanopogon
EN x Helgen and Salas (2008b)
Indonesia Pteropus
ocularis
VU x Helgen and Salas (2008c)
Indonesia Pteropus pohlei EN x Helgen and Bonaccorso
(2008a)
Indonesia Pteropus
temminckii
VU x Helgen and Bonaccorso
(2008b)
Indonesia Rousettus bidens VU x Helgen et al. (2008d)
Indonesia Rousettus
celebensis
LC x Ruedas et al. (2008b)
Indonesia Styloctenium
wallacei
NT x Ruedas et al. (2010),
Mickleburgh et al. (2009)
Indonesia Thoopterus
nigrescens
LC x Ruedas et al. (2008a),
Mickleburgh et al. (2009)
Indonesia,
Thailand
Rousettus
leschenaultii
LC x x Mickleburgh et al. (2009)
Lao PDR Hipposideros
scutinares
VU x Francis and Bates (2008)
Lao PDR Tadarida
latouchei
DD x Francis and Maeda (2008)
Lao PDR,
Philippines,
Thailand
Taphozous
melanopogon
LC x Csorba (2008f), Magnus
(2001)
Lao PDR,
Thailand,
Vietnam
Hipposideros
armiger
LC x Bates et al. (2008b)
Malaysia Cheiromeles
torquatus
LC x Csorba et al. (2008c)
Malaysia,
Philippines,
Thailand
Pteropus
hypomelanus
LC x Francis et al. (2008a),
Fujita and Tuttle (1991),
Fujita (1988)
Myanmar Hipposideros
pomona
LC x x Bates et al. (2008a)
Myanmar Rhinolophus
marshalli
LC x Bates (2003)
Myanmar,
Thailand
Craseonycteris
thonglongyai
VU x Bates (2003)
Philippines Acerodon
jubatus
EN x Mildenstein et al. (2008),
Heaney and Heideman
(1987)
Philippines Acerodon
leucotis
VU x Ong et al. (2008b)
Philippines Dobsonia
chapmani
CR x Heaney et al. (2008b),
Heaney and Heideman
(1987)
Philippines Eonycteris
robusta
NT x Ong et al. (2008e)
Philippines Harpyionycteris
whiteheadi
LC x Ong et al. (2008d),
Mickleburgh et al. (2009)
360 T. Mildenstein et al.
IUCN
region
Country in
which hunted
Species Status Food Medicine Source
Philippines Hipposideros
coronatus
DD x Gomez et al. (2008)
Philippines Hipposideros
pygmaeus
LC x Heaney et al. (2008a)
Philippines Macroglossus
minimus
LC x Mickleburgh et al. 2009,
Francis et al. (2008b)
Philippines Nyctimene
rabori
EN x Mickleburgh et al. (2009)
Philippines Ptenochirus
jagori
LC x Heaney and Heideman
(1987)
Philippines Pteropus
dasymallus
NT x Heaney et al. (2008d)
Philippines Pteropus leu-
copterus/
Desmalopex
leucopterus
LC x Ong et al. (2008a)
Philippines Pteropus
pumilus
NT x Heaney et al. (2008c),
Mickleburgh et al. (2009)
Philippines Pteropus
speciosus
DD x Rosell-Ambal et al. (2008)
Philippines Rhinolophus
rufus
NT x Ong et al. (2008c)
Philippines Styloctenium
mindorensis
DD x Esselstyn (2008)
Philippines,
Thailand
Hipposideros
lekaguli
NT x Csorba (2008d)
Philippines,
Thailand
Rousettus
amplexicaudatus
LC x Csorba (2008g), Utzurrum
(1992)
SE Asia Megaderma
spasma
LC x x Csorba (2008e)
Thailand Hipposideros
halophyllus
EN x Bates et al. (2008b)
Vietnam,
Cambodia
Megaderma lyra LC x x Csorba et al. (2008a)
Sub
Saharan
Africa
Unspecified Epomophorus
labiatus
LC x Mickleburgh et al. (2008b)
Unspecified Hipposideros
gigas
LC x Mickleburgh et al. (2008p)
Unspecified Hipposideros
jonesi
NT x Mickleburgh et al. (2008g)
Unspecified Hipposideros
marisae
VU x Mickleburgh et al. (2008h)
Unspecified Hipposideros
ruber
LC x Mickleburgh et al. (2008i)
Unspecified Hipposideros
vittatus
NT x Mickleburgh et al. (2008f)
Unspecified Mops
midas/Tadarida
midas
LC x Jenkins et al. (2014)
Unspecified Myotis morrisi DD x Jacobs et al. (2008a)
Unspecified Rhinolophus
alcyone
LC x Jacobs et al. (2008b)
361
12 Exploitation of Bats for Bushmeat and Medicine
IUCN
region
Country in
which hunted
Species Status Food Medicine Source
Unspecified Rhinolophus
guineensis
VU x Fahr (2008a)
Unspecified Rhinolophus
hillorum
NT x Jacobs et al. (2010)
Unspecified Rhinolophus
silvestris
DD x Cotterill (2008)
Unspecified Rhinolophus
ziama
EN x Fahr (2008d)
Unspecified Rousettus
lanosus
LC x Mickleburgh et al. (2008n)
Unspecified Taphozous
mauritianus
LC Hutson et al. (2008e)
Benin Epomophorus
gambianus
LC x Mickleburgh et al. (2008r)
Benin,
Cameroon,
Congo Rep.,
Cote d’Ivoire,
DRC, Equatorial
Guinea, Ghana,
Liberia, Nigeria,
Tanzania,
Uganda, Zambia
Eidolon helvum NT x x Halstead (1977), Kamins
et al. (2011), Mickleburgh
et al. (2008l)
Benin, The
Democratic
Republic of
the Congo,
Equatorial
Guinea, Nigeria
Epomops
franqueti
LC xColyn et al. (1987), Fa et al.
(1995), Juste et al. (1995),
Bennett Hennessey (1995)
Comoros
Islands
Pteropus
livingstonii
EN x Trewhella et al. (1995)
Congo
Republic
Lissonycteris
angolensis
LC x Wilson and Wilson (1991),
Mickleburgh et al. (2008c)
Congo
Republic
Nycteris grandis LC x Mickleburgh et al. (2008k),
Bennett Hennessey (1995)
Côte d’Ivoire Chaerephon
ansorgei/
Tadarida
ansorgei
LC x Mickleburgh et al. (2008e)
Equatorial
Guinea
Micropteropus
pusillus
LC x Juste et al. (1995), Fa
(2000)
Equatorial
Guinea, Nigeria
Rousettus
aegyptiacus
LC x Fa et al. (1995), Fa (2000),
Benda et al. (2008)
Guinea Hipposideros
lamottei
CR x Mickleburgh et al. (2008q)
Guinea Rhinolophus
maclaudi
EN x Fahr (2008b), Fahr et al.
(2002), Fahr and Ebigbo
(2003)
Guinea Rhinolophus
ruwenzorii
VU x Fahr et al. (2002), Fahr
and Ebigbo (2003), Fahr
(2008c)
Madagascar Chaerephon
jobimena/
Tadarida jobimena
LC x Andriafidison et al. (2014a)
362 T. Mildenstein et al.
IUCN
region
Country in
which hunted
Species Status Food Medicine Source
Madagascar Eidolon
dupreanum
VU x Andriafidison et al.
(2008a), Jenkins and Racey
(2008)
Madagascar Emballonura
atrata
LC x Jenkins et al. (2008b)
Madagascar Hipposideros
commersoni
NT x Goodman (2006), Jenkins
and Racey (2008)
Madagascar Miniopterus
gleni
LC x Andriafidison et al.
(2008b), Goodman (2006),
Goodman et al. (2008)
Madagascar Miniopterus
majori
LC x Jenkins and Rakotoarivelo
(2008)
Madagascar Miniopterus
manavi
LC x Andriafidison et al.
(2008c), Golden (2005)
Madagascar Mops
leucostigma
LC x Andriafidison et al. (2014b)
Madagascar Mormopterus
jugularis
LC x Andriafidison et al. (2008d)
Madagascar Myzopoda
aurita
LC x Jenkins et al. (2008a)
Madagascar Pteropus rufus VU x Andriafidison et al.
(2008e), Jenkins and Racey
(2008)
Madagascar Rousettus mada-
gascariensis
NT x Andriafidison et al. (2008f),
Jenkins and Racey (2008)
Madagascar Scotophilus
robustus
LC x Andriafidison et al. (2008g)
Madagascar Triaenops
furculus
LC x J. Razafimanahaka pers.
comm
Madagascar Triaenops rufus LC x Goodman (2006),
Andriafidison et al. (2008h)
Mauritius Pteropus niger VU x Hutson and Racey (2013),
Mickleburgh et al. (2009)
Mauritius Pteropus
rodricensis
CR x Mickleburgh et al. (2008d),
Mickleburgh et al. (2009)
Rwanda Rhinolophus
hilli
CR x Fahr (2010)
São Tomé and
Príncipe
Chaerephon
tomensis
EN x Carvalho et al. (2014)
São Tomé and
Príncipe
Miniopterus
newtoni
DD x Carvalho et al. (2014)
São Tomé and
Príncipe
Myonycteris
brachycephala
EN x Carvalho et al. (2014)
Seychelles Pteropus
seychellensis
LC x Carvalho et al. (2014)
Tanzania
(Pemba)
Pteropus
voeltzkowi
VU x Mickleburgh et al. (2008m)
The
Democratic
Republic of the
Congo, Nigeria
Hypsignathus
monstrosus
LC x Mickleburgh et al. (2008j),
Mickleburgh et al. (2009)
Unspecified Myotis
mystacinus
LC x Hutson et al. (2008f)
363
12 Exploitation of Bats for Bushmeat and Medicine
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