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Original Research Article
Characterizing and quantifying the wildlife trade network in
Sulawesi, Indonesia
Alice Latinne
a
,
*
, Suryo Saputro
b
, Jusuf Kalengkongan
a
, Citra Livi Kowel
c
,
Lidia Gaghiwu
c
, Tiltje Andretha Ransaleleh
d
, Meis Jacinta Nangoy
d
,
Indyah Wahyuni
d
, Tina Kusumaningrum
c
, Dodi Safari
c
, Yasha Feferholtz
a
,
Hongying Li
a
, Emily Hagan
a
, Maureen Miller
a
,
e
, Leilani Francisco
a
,
f
,
Peter Daszak
a
, Kevin J. Olival
a
, Joko Pamungkas
b
,
g
,
**
a
EcoHealth Alliance, New York, USA
b
Primate Research Center, IPB University, Bogor, Indonesia
c
Eijkman Institute of Molecular Biology, Jakarta, Indonesia
d
Faculty of Animal Science, Sam Ratulangi University, Manado, Indonesia
e
Mailman School of Public Health, Columbia University, New York, USA
f
Henry M. Jackson Foundation, Bethesda, USA
g
Faculty of Veterinary Medicine, IPB University, Bogor, Indonesia
article info
Article history:
Received 15 October 2019
Received in revised form 14 December 2019
Accepted 15 December 2019
Keywords:
Wildlife trade
Hunting
Wildlife markets
Flying foxes
Indonesia
Sulawesi
abstract
The island of Sulawesi in Indonesia is an important site for the wildlife trade that is
currently undergoing rapid exploitation of its local fauna to supply wild meat markets of
North Sulawesi. In this study, we used field surveys, ethnographic interviews, and daily
counts in markets to document species of terrestrial wildlife on sale in North Sulawesi
markets, and to identify the hunting sites, practices, and key actors within the wildlife
trade. We quantify the volume of wild meat traded and their prices, with a particular focus
on the flying fox trade.
Wildlife meat was routinely available for sale in 73% of the markets and supermarkets
surveyed in North Sulawesi. The wildlife taxa most commonly found in these markets were
flying foxes, wild pigs, rats and snakes. Wildlife hunting and trade networks extend to all
provinces of the island through a well-organized, dynamic and easy to access network
involving many actors. We identified 45 flying fox roosts in Sulawesi, 38 of which were
under active hunting pressure. A third of the active hunting sites are located in Southeast
Sulawesi, which acts as a hub for bat hunting and trade. We estimate that the number of
flying foxes annually traded in Sulawesi ranges from 662,551 to more than one million
individuals and conclude that current rates of flying fox harvest are unstainable. Stricter
law enforcement, implementation of hunting quotas, and further research efforts are
therefore urgently needed to improve the sustainability of the wildlife trade in Sulawesi.
©2019 The Authors. Published by Elsevier B.V. This is an open access article under the CC
BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
*Corresponding author.
** Corresponding author. Primate Research Center, IPB University, Bogor, Indonesia.
E-mail addresses: latinne@ecohealthalliance.org (A. Latinne), jpi-pssp@indo.net.id (J. Pamungkas).
Contents lists available at ScienceDirect
Global Ecology and Conservation
journal homepage: http://www.elsevier.com/locate/gecco
https://doi.org/10.1016/j.gecco.2019.e00887
2351-9894/©2019 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/
licenses/by-nc-nd/4.0/).
Global Ecology and Conservation 21 (2020) e00887
1. Introduction
Southeast Asia, one of the most biodiverse regions in the world, is currently facing a biodiversity crisis threatening
numerous species and ecosystems (Schipper et al., 2008;Wilcove et al., 2013). Overhunting and unstainable harvesting
constitutes one of the greatest threats to the biodiversity of the region (Di Minin et al., 2019;Gray et al., 2018;Harrison et al.,
2016). Wild animals are traded live as pets or dead as wild meat, ornaments or medicines to supply local, regional and global
markets (Gray et al., 2018;Nijman, 2010). Indonesia is an important wildlife trade hotspot in the region (Davies, 2005;Nijman,
2010). Widespread and poorly regulated trade in this country involves numerous species of wild birds (Bergin et al., 2018;
Shepherd et al., 2016), marine invertebrates (Nijman et al., 2016), reptiles (Natusch and Lyons, 2012), primates (Nijman et al.,
2017) and other mammals (Harrison et al., 2011;Lee et al., 2005;Luskin et al., 2014). Consequently, poaching and hunting
have already devastated populations of several wildlife species in Indonesia (Harrison et al., 2016).
During the past few decades, the fourth largest island of Indonesia, Sulawesi, has undergone rapid exploitation of its fauna
to supply wild meat markets in North Sulawesi province (Lee, 1997;Lee et al. 2005,2009;Sheherazade and Tsang, 2015).
Wildlife meat consumption in Sulawesi is closely associated with religious differences (Lee et al., 2005). The island is primarily
inhabited by Muslims but a large Christian community, called Minahasan people, inhabits the province of North Sulawesi.
Wild meat is not consumed by Muslim populations but is very popular among Minahasan people and has been part of their
cultural habits for decades. Minahasan people consume wildlife as well as other meat such as cats and dogs on a daily basis
but this consumption increases during the Christmas holidays and before Pengucapan, the “Minahasan Thanksgiving”(Lee,
2000;Sheherazade and Tsang, 2015). Wild meat has been sold in North Sulawesi markets since the 1970s and the wildlife
trade business, benefiting from road improvement, underwent rapid expansion in the 1980s and 1990s (Clayton and Milner-
Gulland, 2000). Studies in the 1990s and early 2000s showed that the wild animals for sale in North Sulawesi markets were
mainly rats, fruit bats, and wild pigs (Clayton and Milner-Gulland, 2000;Lee et al., 2005;Milner-Gulland and Clayton, 2002).
Legally protected species whose hunting and trade is prohibited such as macaques, babirusas, anoas, deers, cuscuses, and
tarsiers were also sold in smaller numbers (Lee et al., 2005;Milner-Gulland and Clayton, 2002).
Over-hunting has already extirpated several wild mammal species in North Sulawesi, including Pteropus and Acerodon spp.
fruit bats (flying foxes) and hunting activities have expanded to other provinces to supply the demand from the northern
province (Clayton and Milner-Gulland, 2000;Lee et al., 2005;Milner-Gulland and Clayton, 2002). Flying foxes are highly
vulnerable to over-hunting due to some of their ecological and life history traits (Epstein et al., 2009;Mickleburgh et al., 2009)
and this group of bats is facing higher risk of extinction than other bats (Jones et al., 2003). Island flying fox species are
suffering considerable and widespread decline worldwide (Vincenot et al., 2017) and their populations in Sulawesi might be
severely impacted by the wildlife trade (Sheherazade and Tsang, 2015). However, the extent and structure of the wildlife trade
network in Sulawesi remains unknown as previous studies focused on market vendors from North Sulawesi and not on other
actors within the trade. In this study, for the first time, we conducted field surveys and ethnographic interviews in all
provinces of Sulawesi and involving most wildlife trade actors to: (1) document the markets where wild meat is sold in North
Sulawesi and identify the taxa on sale, (2) identify hunting sites and key actors within the wildlife trade from capture to
consumption and (3) quantify the volume of wild meat traded and their prices. We focused particularly on the flying foxes,
which is one of the main taxa traded. These data are urgently needed to assess the current level of sustainability of the trade
and its impact on flying fox populations and to identify appropriate conservation measures to mitigate these impacts.
2. Material and methods
2.1. Market surveys
Local observers surveyed 31 markets and 14 supermarkets in nine districts of North Sulawesi province from November 2018
to February 2019 (Fig. 1 and Table A 1). Surveys consisted of a single visit to each marketor supermarket to assess the number of
vendors selling wild meatand identify the wildlife taxa being sold. We also conducted in-depth surveys in five markets, including
three of the largest wildlife markets in North Sulawesi, Langowan, Kawangkoan and Tomohon, in addition to Karombasan and
Tareran markets (Fig. 1). For these in-depth surveys, observers monitored each market two to three times per week during 12
consecutive weeks from November 2018 to January 2019. During each visit to Langowan, Kawangkoan, Tomohon, and Kar-
ombasan markets, they recorded the daily quantityand prices of large fruit bats, wild pigs, rats, and snakes sold by each vendor.
At Tareran market, only quantities of large fruit bats and rats were recorded. Prices were recorded in Indonesian rupiahs (IDR).
2.2. Mapping of bat roosts and hunting sites
As part of a multi-year wildlife viral surveillance project conducted in Sulawesi (USAID EPT-PREDICT project), we surveyed
large areas of Sulawesi for flying fox roost sites and bat hunting areas in 2016 and 2017. From April 2018 to March 2019, we
intensified this survey and systematically monitored suitable areas in five provinces of Sulawesi (Gorontalo, Southeast, South,
West, and Central Sulawesi).These surveys were carried out in collaborationwith our local network of collaborators including
researchers and officials from provincial offices of the Directorate General of Livestock and Animal Health Services (DGLAHS)
under the Indonesia Ministry of Agriculture. The GPS coordinates of each roost were recorded and hunting status assessed
through direct observation and discussion with local community members. We also collected detailed demographic data
A. Latinne et al. / Global Ecology and Conservation 21 (2020) e008872
(species, sex, weight, age class, forearm measurement) on bats caught by hunters in three hunting sites located in Gorontalo,
West Sulawesi, and Southeast Sulawesi provinces twice a year from 2016 to 2018.
2.3. Ethnographic interviews
We conducted 70 semi-structured interviews with market vendors (n ¼38), trade intermediaries (n ¼9) and wildlife
hunters (n ¼23) from March to August 2016 and from July 2018 to January 2019 in North, Southeast, and West Sulawesi.
Respondents were identified through discussions with local community members in markets and wildlife hunting areas of
Sulawesi. We targeted knowledgeable informants actively involved in the wildlife trade, mostly focusing on bats, to under-
stand the network configuration and identify key actors. Participation in interviews was anonymous and voluntary. Informed
consent was obtained before each interview. No questions about respondents’personal information or identity were asked.
Our research protocols were reviewed and approved by the Committee on Health Research Ethics of the National Institute of
Health Research and Development, Indonesia Ministry of Health (protocols LB.02.01/5.2/KE.040/2016 and LB.02.01/2/KE.306/
2018), by the Eijkman Institute Research Ethics Commission of the Eijkman Institute of Molecular Biology (protocol #117) and
by the Institutional Review Board at the University of California, Davis (protocol #754490). We obtained local research
permits from each province and district where the research was conducted.
Wildlife market vendors were asked which markets they sell their wildlife products in, the number of days they work per
week, the wildlife taxa they sell routinely or occasionally and their geographic origins, the quantity of bats sold routinely
every week and during the peak season and their corresponding prices. We asked trade intermediaries about the wildlife taxa
they trade, the geographic origin of these animals, the quantity of wild animals traded monthly and annually, the price they
pay to buy these animals and their selling price, the hunters and other intermediaries with whom they work, and information
about the transportation and logistics to North Sulawesi and its associated cost. Wildlife hunters were asked about when they
started their hunting activities, the hunting frequency, the quantity of animals they collect daily and weekly and its annual
variation, the hunting seasons, their hunting methods, the price they were paid by trade intermediaries, the cost of their
hunting activities, and the structure of the local group of hunters. All respondents were also asked about their perception of
bat population trends in Sulawesi and, lastly, their awareness of the Indonesian law for wildlife protected species.
2.4. Wild meat quantities traded
The average quantities of bats, rats, wild pigs, and snakes sold every week in Langowan, Kawangkoan, Tomohon, Kar-
ombasan and Tareran markets were extrapolated from the daily counts collected in these markets, using the sum of the
Fig. 1. Map of North Sulawesi province showing the 27 markets and 10 supermarkets where wild meat is sold routinely or occasionally (before Christmas, New
year and Thanksgiving only) according to the results of our survey in 31 markets and 14 supermarkets in the province from November 2018 to February 2019.
Dotted grey lines represent the provincial road network.
A. Latinne et al. / Global Ecology and Conservation 21 (2020) e00887 3
average quantities sold every market day. Regular weeklyaverage quantity (Q
WR
) was calculated from data collected between
November 12 and December 21, 2018 and between January 2 and 31, 2019. The weekly average quantity during Christmas/
New Year (Q
WC
) was estimated from the data collected between December 22 and 31, 2018. The annual quantity of wild
animals sold (Q
A
) was estimated by using the sum of the product of Q
WR
multiplied by 49 and Q
WC
multiplied by three
(corresponding to the two weeks before Christmas/New Year and one week before Thanksgiving). For the markets in which
we did not collect daily count data, we used weekly averages provided by interview respondents, multiplied by the number of
vendors in each market, to estimate Q
A
. For the markets in which we did not collect interview data, we used estimates of
weekly quantity of 50 kg for bats, 60 kg for wild pigs, and 15 kg of snakes per vendor. These quantities were the average
minimal weekly quantities sold by a single wildlife vendor in the 10 markets where daily counts or interview data were
available. We did not estimate the average weekly quantity of rats sold per vendor as it varied considerably among vendors.
We estimated the quantities of flying foxes caught and traded by hunters and intermediaries who responded to our in-
terviews using the data they provided. The hunters we interviewed were able to give information on the total number of bats
their hunting group took throughout the year. We used the product of the total daily or weekly quantities harvested and the
length of the hunting season to estimate the annual quantity harvested from each colony. For trade intermediaries, we
calculated the product of the number of trips to North Sulawesi per year, the average quantity of bats transported per trip
during the low and high season, and the length of each season.
3. Results
3.1. Wildlife diversity in markets of North Sulawesi
Wildlife meat was routinely available for sale in 23 of the 31 markets surveyed in North Sulawesi in 2018 and 2019 and was
also occasionally traded in four additional markets, mostly before Christmas and New Year (Table A1 and Fig.1). Most of these
markets are located in Minahasa and South Minahasa districts (Table A1). Some markets were active every day while some
were open only twice or three times per week. Many wildlife vendors shared their time among several markets from the same
region (Dumoga/Ibolian/Imandi, Motoling/Tompaso Baru/Modoinding/Poopo, Kawangkoan/Sonder/Tareran markets). We
also observed wildlife meat in 10 of the 14 supermarkets surveyed in Tomohon and Manado cities (Fig.1). Many restaurants in
North Sulawesi, called “Minahasan restaurants”, also served wildlife meat but were not systematically surveyed and mapped
for this study. Pictures are available in Appendix (Figs A1-A3).
The wildlife taxa most commonly found in these markets were bats, wild pigs, rats, and snakes (Table A1). Large fruit bats
(flying foxes), mostly Acerodon celebensis and Pteropus alecto, were commonly found in almost all wildlife markets (96%) we
identified in North Sulawesi. They were sold roasted. All market vendors we interviewed confirmed that they had not
received live bats for the past few years. Small fruit bats belonging to the genera Cynopterus,Rousettus,Nyctimene,Thoopterus,
and Styloctenium were also observed in the markets of the western part of the province (Ibolian, Dumoga, Imandi and
Modoinding markets). These small bats were sold freshly killed or roasted. Parts of wild pigs (Sus celebensis) and snakes
(Python sp.) were also found in most markets we surveyed in the province (96% and 91%, respectively). Two wild pigs that
were still alive and restrained were observed at the Tomohon market in January 2019. Roasted rats were also very common in
wildlife markets (91%). These are called “white tailed rats”by local people but they belong to a large diversity of mostly
endemic species including Taeromys celebensis,Rattus xanthurus,Maxomys hellwaldii,M. musschenbroekii,Paruromys domi-
nator,Echiothrix leucura, and Bunomys chrysocomus. These forest species were considered healthier and tastier than black
tailed rats hunted in paddy fields such as Rattus hoffmani,R. argentiventer,R. rattus, which are also available in some markets,
but at a lower price. Protected species were available in some of these markets (Table A1) including: Celebes crested macaque
(Macaca nigra) (39%), anoa (Bubalus depressicornis) (30%), cuscus (Strigocuscus celebensis and Ailurops ursinus) (17%), and
babirusa (Babyrousa celebensis) (13%). Monitor lizards, squirrels, and frogs were also noticed occasionally (Table A1). The other
taxa available in these markets were mostly domestic pig, dog, cat, and chicken as well as fish, all sold killed.
3.2. Wildlife trade network in Sulawesi and its key actors
Wildlife hunting and trade extends to all provinces of Sulawesi as most of the animals sold in North Sulawesi markets were
not locally sourced. Only rats, small fruit bats, and some wild pigs were provided to market vendors by a local network of
hunters while large fruit bats, snakes, most wild pigs, and some rats were hunted in other provinces and transported to North
Sulawesi (Fig. 2). Market vendors had poor knowledge of the origin of their wild meat and were only able to indicate the
province of origin of some taxa. However, our interviews with trade intermediaries and hunters, and our bat hunting site
survey allowed us to map the current wildlife trade network in further detail and identify its key actors (Fig. 2).
3.2.1. Hunters, hunting sites and hunting practices
Out of the 19 hunters interviewed in North Sulawesi, most (89%) said they hunted rats, followed by wild pigs (42%),
cuscuses (26%), small bats (16%), anoas (10%), snakes (5%), babirusas (5%) and monkeys (5%). Hunters interviewed in the other
provinces were exclusively flying fox hunters. Hunting practices and sites varied according to the taxa hunted.
A. Latinne et al. / Global Ecology and Conservation 21 (2020) e008874
3.2.1.1. Bats.
Small fruits bats were hunted only in North Sulawesi, mostly in the western part of the province, by local vil-
lagers for their own consumption or sale in markets (Fig. 2). Flying foxes were hunted in all other provinces of Sulawesi
(Fig. 2). Flying fox hunters were usually from Muslim communities living close to bat roosts and did not consume wild meat
for religious reasons. Therefore, bat hunting in these regions was commercially motivated. Hunters usually started their
activities after being encouraged and trained by trade intermediaries from North Sulawesi who pledged to buy the bats they
would catch. Local conflicts among villagers to exploit a bat colony was reported several times during our study as these bats
had an important economic value. Flying fox hunters interviewed in Gorontalo and West Sulawesi provinces were part of
hunter groups with a leader coordinating bat hunting and selling to trade intermediaries. In one of these groups, benefits from
bat sales were equally shared among all hunters while in the other group, each hunter was paid proportionally to the quantity
of bats caught. In Southeast Sulawesi, the hunters interviewed were working and dealing with trade intermediaries inde-
pendently. All hunters reported that bat hunting was the primary source of their annual income, although it was supple-
mented by fishing, farming (coconut, cashew nuts, corn), construction and other activities.
We identified 45 flying fox roosts in all provinces of Sulawesi except North Sulawesi (Fig. 3 and Table A2). Most roosts were
located in coastal mangrove swamps (Rhizophora sp., Sonneratia sp., Bruguiera sp., Ceriops sp., Avicennia sp., Lumnitzera sp.,
and Nypa fruticans) or coastal pine trees (Pinus sp.) (Table A2). A few roosts were located in swamps surrounding large lakes of
Central and Southeast Sulawesi. We also identified a few colonies roosting in emergent trees of inland forests (Table A2). Of
these 45 roosts, 38 (84%) were under active hunting pressure in 2018 and 2019, while the current hunting status of two of
them remained uncertain. Southeast Sulawesi seems to be the current hub of the bat hunting and trade in Sulawesi, with at
least 13 active hunting sites (34%) (Fig. 3). At least five hunting sites were located on the southernmost islandsof the province
(Wowoni, Muna and Buton islands). A bat roost in North Kolaka district, Southeast Sulawesi (#31 in Fig. 3), was reported as
one of the most productive bat hunting sites in Sulawesi by four respondents. Central Sulawesi, harbored the second highest
number (11, 29%) of active hunting sites. Only a few hunted roosts were identified in South Sulawesi (13%), Gorontalo (13%)
and West Sulawesi (11%). These numbers contrast with the information given by market vendors regarding the origin of the
flying foxes they sold: South Sulawesi was reported as the main source of flying foxes (32%), followed by Southeast Sulawesi
(24%), Gorontalo (24%), Central Sulawesi (13%), and West Sulawesi (2%). Other islands (Kalimantan, West Nusa Tenggara, East
Nusa Tenggara and North Maluku) were also reported as the source of flying foxes sold in North Sulawesi a few times during
our interviews.
Several respondents in hunting areas reported the disappearance of bat roosts in the past few years due to overhunting,
disturbance, or mangrove destruction for fisheries, and that local populations encouraged hunting because bats are
considered pests that damage fruit crops and mangroves. We identified only five flying fox colonies (11%) protected by local
villagers during our survey (Fig. 3), four of them protected for many years, and one colony in Toribulu, Central Sulawesi (#14 in
Fig. 3) exploited by hunters before being protected by villagers in late 2016. Some of these roosts have become eco-tourism
sites attracting local visitors, contributing to their protection.
Fig. 2. Overview of the wildlife trade network in Sulawesi and its key actors for the four taxa most commonly traded (bats, wild pigs, snakes and rats). Wild meat
selling places are highlighted in red. Sources of wildlife supply for middlemen based in North Sulawesi were located either in Gorontalo and Central Sulawesi
(green network) or in West, South, and Southeast Sulawesi (black network). Two to three middlemen based in Southeast Sulawesi were also identified (blue
network). (For interpretation of the references to colour in this figure legend, the reader is referred to the Web version of this article.)
A. Latinne et al. / Global Ecology and Conservation 21 (2020) e00887 5
The black flying fox (Pteropus alecto) and the Sulawesi flying fox (Acerodon celebensis) were the most commonly harvested
species in Sulawesi (Table A2). These often co-roost and were hunted without specific targeting of one or the other. According
to the hunting data we collected from 2016 to 2018, the proportion of Acerodon celebensis bats harvested varied from 11 to 26%
(Table A3). The sex ratio of hunted individuals is close to 1/1 and more than half of them were sexually mature adults (Table
A3). The average weight of the collected bats was 486 g (regardless of the species) (Table A3). Two colonies of Pteropus griseus,
called “white bats”by local hunters, were also harvested in Central Sulawesi (Fig. 3).
Most respondents reported that flying foxes in Sulawesi migrate seasonally, with a few colonies thought to never migrate.
Bat hunting is therefore a seasonal activity in most areas. In most provinces (West Sulawesi, South Sulawesi, Southeast
Sulawesi, and Gorontalo), flying fox colonies were reported to migrate during the rainy monsoon season, from January to
April. These colonies were back to their roost at the beginning of the dry monsoon season, from Mayto December, with a peak
of high population size from July to September. Several respondents in these provinces mentioned that hunting still happened
during the flying fox birthing season in December. However, opposite migration patternwas reported for two roosts of Central
Sulawesi (#12 and 16 in Fig. 3) with the period from November to June being reported as the high density season. In some
mixed colonies, several respondents mentioned that Pteropus bats migrated while Acerodon bats did not and occupied the
same roost throughout the year. However, many hunters paused their hunting activities during this low bat density season as
the hunting effort required is too high compared to the number of bats that could be harvested. Recent changes in bat
migration patterns, allegedly explained by a disrupted monsoon pattern due to climate change, were also reported by several
hunters and trade intermediaries.
Hunting techniques used were hooked ropes (42%), nets (33%), air rifles (17%), and kites with hooks (8%). Hooked ropes
suspended between bamboo poles or large trees around mangrove roost sites were used in Gorontalo, Central, and Southeast
Sulawesi provinces (Fig A4 in Appendix). These permanent structures remain in place throughout the year, are checked each
day during the high season or every two days during the low season. Up to 100 hooks may be placed on each 30 m rope.
Bamboo poles were replaced at least annually. Nets deployed in the early morning before the bats fly back to their roost were
also used in Gorontalo, Central, West, and South Sulawesi provinces (Fig A4 in Appendix). In some regions of Central Sulawesi
and Gorontalo, hunters also used nets to target fruit trees where bats feed nocturnally. Air rifles were used in Central and
West Sulawesi. The use of kites with hooks was only reported in Gorontalo province. Bats were usually killed directly after
capture, often by drowning, and then stored in cooler boxes. Accidents due to crocodile attacks during hunting activities were
reported several times in Central Sulawesi and several bat hunters were killed in this region.
3.2.1.2. Rodents.
Eighty-nine percent of hunters interviewed in North Sulawesi said they hunted rats in the surroundings
forests, both for their own consumption and for sale in markets or directly to consumers (Fig. 2). Central Sulawesi and
Fig. 3. Map of Sulawesi showing the flying fox roosts identified in this study and their hunting status. The two Pteropus griseus roosts in Central Sulawesi are
indicated by an asterisk. Other roosts are Pteropus alecto and/or Acerodon celebensis. Grey dotted lines correspond to the provincial road network in Sulawesi.
Numbers refer to Table A2 where more information about each roost is provided.
A. Latinne et al. / Global Ecology and Conservation 21 (2020) e008876
Gorontalo were also reported as the origin of many rodents sold in North Sulawesi markets, some of them coming from palm
oil plantations (Fig. 2). Rat hunters used traditional bamboo traps (69%) and air rifles (31%). In paddy fields, hunters also
caught rats overnight by digging holes and hitting them with a stick when they exited their burrows. A single hunter usually
caught around 20 to 40 rats per day but it could be as many as 100 during the most productive nights.
3.2.1.3. Wild pigs.
Wild pig hunting occurred in all provinces of Sulawesi (Fig. 2) but pigs were also imported from Kalimantan
province (Borneo island) according to a trade intermediary. This respondent reported that wild pigs from Southeast Sulawesi
were considered as the tastiest while those from Kalimantan (bearded pigs) were bigger but were not favored because of their
poor taste. According to the same respondent, wild pigs from Gorontalo province were smaller than those from other
provinces. Wild pigs were hunted using shotguns, wire traps, or mouth snares baited with a half coconut. Some pigs from
North and Central Sulawesi were transported alive with tied feet and mouth to the markets.
3.2.1.4. Snakes.
All snakes sold in North Sulawesi belonged to the genus Python. Pythons were mostly hunted in the swamps,
forests, and paddy fields of Central, South, and Southeast Sulawesi (Fig. 2), where they were abundant and easily caught. Most
were killed before being transported to North Sulawesi.
3.2.1.5. Babirusas and anoas.
Hunting of large animals such as babirusa and anoa occurred in Sulawesi from 2016 to 2018
despite being illegal since 1999 (Government Regulation No. 7,1999, Decree No. P.20/MENLHK/SETJEN/KUM.1/6/2018). Two of
our respondents reported trapping these large animals in the protected mountainous areas of North Sulawesi province.
Hunting these species required a significant effort because the deep forest sites required one day of walking to reach, and
hunting trips lasted three to four days. These large animals were slaughtered directly in the forest and meat pieces were
carried back to the village, often requiring several trips. Only a few of these animals were caught every month but their selling
price was very high (>5,000,000 IDR).
3.2.1.6. Monkeys.
Two respondents from North Sulawesi reported that macaque hunting was still active in the province. Both
mentioned that a preferred hunting practice was to cut down the sleeping tree on which a group of macaques spent the night.
This allowed them to capture 20 to 40 individuals in a single night.
3.2.2. Trade intermediaries
We identified trade intermediaries (which we call local dealers and middlemen) who played different roles in supplying
wild meat to North Sulawesi vendors. Most belonged to the Minahasan, Balinese, or Torajan ethnic groups. Local dealers
collected and stored bats and other wildlife (mostly wild pigs and snakes) directly from hunters. Many provided trans-
portation, hunting equipment and food to hunters in advance to ensure that they would not sell their animals to another
intermediary. When bat roosts are not exploited by local villagers, trade intermediaries sometimes sent their own hunters to
harvest a bat colony for several weeks or months before leaving the area. Local dealers were located in all regions of Sulawesi
where wildlife hunting occurred (Fig. 2). They did not organize the transportation of wild meat to North Sulawesi but sold it to
other trade intermediaries that we called middlemen.
Middlemen typically bought wildlife from local dealers or directly from hunters and coordinated the transportation lo-
gistics to North Sulawesi from other provinces (Fig. 2). We estimate around 10 to 12 middlemen have established in North
Sulawesi from our interviews. Their sources of wildlife were located either in Gorontalo and Central Sulawesi (green network
in Fig. 2) or in West, South, and Southeast Sulawesi (black network in Fig. 2). We also identified two to three middlemen based
in Southeast Sulawesi (blue network in Fig. 2). They all traded and transported bats, wild pigs, and snakes as well as dogs to
North Sulawesi. Middlemen usually owned one or several pickups and trucks and worked with their own team of drivers.
These drivers shuttled back and forth (two drivers per car) between North Sulawesi and other provinces to collect wild meat
from local dealers or hunters, transported on ice. Several middlemen also transported vegetables from North Sulawesi to sell
in South and Southeast Sulawesi to supplement income. A return trip from North Sulawesi took between 10 and 13 days. A
single car and two drivers were therefore able to perform two to three trips per month. Several respondents also mentioned
that bats were transported by plane from Southeast Sulawesi to Manado city in North Sulawesi but we were not able to
confirm this.
3.2.3. Wild meat sellers
In North Sulawesi, wildlife is available for sale to consumers in markets, supermarkets, and Minahasan restaurants (Figs. 1
and 2,Table A1,Figs A1-A3). We also identified a few markets and restaurants selling wildlife in Central Sulawesi, around Poso
and Parigi Moutong districts, where a large Christian community is established. These places were supplied by middlemen
drivers on their way back to North Sulawesi from Southeast and Central Sulawesi (Fig. 2).
3.3. Wild meat quantities traded and prices
We estimate that 9400 to 13,800 kg of flying foxes were harvested annually in 2018 and 2019 from each of the three roosts
we studied, representing around 20,680 to 26,634 individual bats removed from the colony each year (Table A3). Two
A. Latinne et al. / Global Ecology and Conservation 21 (2020) e00887 7
respondents also reported that up to four tons of bats, around 8000 bat individuals, could be harvested monthly from the bat
roost in North Kolaka, Southeast Sulawesi (#31 in Fig. 3). Four middlemen provided us with detailed numbers allowing us to
estimate the annual quantities of bats they traded in 2018 and 2019. Bats were the priority taxa for three of them; they
transported other taxa (wild pigs, snakes or dogs) only when there were not enough bats available to fill their vehicles
completely (a single pickup can carry up to 1700e1900 kg of wildlife meat). Each middleman traded 18,800 to 122,400 kg of
bats annually, all collected in West, South, and Southeast Sulawesi. The fourth middleman mostly focused on wild pigs and
transported around 9600 kg of bats from Southeast Sulawesi annually. We estimate that these four middlemen together
traded more than 176,500 kg of bats annually during the period 2018e2019. If we consider that their business represented a
third of the total business activities of the 10 to 12 middlemen active in Sulawesi, the quantity of bats traded on the island
annually could be as high as 529,500 kg, which would be more than one million bats hunted each year in Sulawesi when using
an average weight of 486 g per bat (Table A3).
The quantities of wild meat sold daily varied in the five markets we studied (Fig A5 in Appendix), with Saturday usually the
busiest. Wild pig was the largest taxon by quantity sold in Langowan and Kawangkoan markets and in the combined data
from the five markets, followed by flying foxes (Fig A5 in Appendix). In the weeks before Christmas and New Year, wild meat
sales increased in all five markets and were two to five times higher than the regular weekly average (Fig A5 in Appendix).
Market vendors reported similar increases during the week before the “Minahasan Thanksgiving”celebrated in JuneeSep-
tember. Other than the sale increases observed during these three weeks, vendors reported that sale volumes remained
mostly similar to the regular weekly average throughout the year. We are therefore confident that our results are not
significantly biased by the study period.
We estimate that the total annual quantities of wildlife sold in all Sulawesi markets were 319 tons of bats, 419 tons of wild
pigs, and 72 tons of snakes (Table A4). According to the middlemen interviewed, the weight of wild pigs and snakes they
traded was around 10e50 kg and 7e80 kg, respectively. Therefore, these quantities could correspond to 662,551 bats,
8380e41,900 wild pigs, and 900e10,285 snakes sold annually in North Sulawesi markets. These quantities are likely un-
derestimates because wildlife meat sold in supermarkets and restaurants was not accounted for. Moreover, these numbers do
not include the wildlife sold in markets and restaurants from Central Sulawesi where we did not collect data.
Prices paid to hunters in 2018 for their flying foxes varied across regions (Fig. 4). In 2018, the price for flying foxes in
Southeast Sulawesi varied from 10 K to 30 K IDR/kg. Much lower prices, around 8e15 K IDR/kg, were paid in West Sulawesi. In
Central Sulawesi, flying fox prices varied from 10 to 22 K IDR/kg. The highest prices, around 35e40 K IDR/kg, were paid to
hunters in Gorontalo. Prices for wild pig and snake meat paid to hunters from Southeast Sulawesi were similar (Fig. 4). Several
middlemen reported that they always kept at least 10 K IDR difference between their buying and selling prices.
Flying fox prices in markets were two to five times higher than the prices paid to hunters. In markets and supermarkets,
bat meat was the most expensive wild meat (50e70 K/kg and 80e89 K/kg, respectively) and wild pigs the cheapest (35e40 K/
kg and 43e50 K/kg, respectively) (Fig. 4). The cost of macaque meat was around 40 K/kg or 200 K/individual in 2018. In
comparison, market prices for domestic meat were around 30e35 K/kg for chicken, 50e60 K/kg for domestic pigs, 100 K/
animal for cats and 35 K/kg for dogs in 2018. The weeks before Christmas and Thanksgiving, prices increased of 20e40 K IDR
per kg of wild meat (bats, wild pigs, and snakes).
Fig. 4. Price ranges for wild meat paid to the main actors of the wildlife trade network in Sulawesi in 2018e2019. Prices are given in Indonesia Rupiahs (IDR) per
kilogram of wild meat, 1 K IDR ¼1000 IDR (1 USD ¼14,000 IDR in July 2019).
A. Latinne et al. / Global Ecology and Conservation 21 (2020) e008878
3.4. Perception of bat population trends in Sulawesi
All wildlife hunters from North Sulawesi (n ¼19) reported that wildlife populations were seriously depleted and that only
rats, small bats, and in some places wild pigs remained sufficiently abundant for hunting. Respondents from the two groups of
flying fox hunters we interviewed in Gorontalo and West Sulawesi provinces did not notice a population size decrease in the
bat colony they harvested. However, flying fox hunters from Southeast Sulawesi observed a strong population decrease in
their colony over the past few years and reported that reduced harvests had forced some hunters in the village to stop
hunting. All flying fox hunters we interviewed said that they released young juveniles and babies caught accidentally by
placing them on tree branches in the mangrove close to the roost. In doing so, they believed that these babies would be taken
care of by other females from the colony. Several hunters and trade intermediaries from different regions also reported that
they never catch or trade bats from the genus Dobsonia. They believed these bats were legally protected and that catching or
trading them would attract police attention totheir business. These hunters said they always release the Dobsonia bats caught
accidentally. However, no Dobsonia species is currently legally protected in Indonesia. Most trade intermediaries reported
that bat populations in Central and South Sulawesi significantly decreased over the last 10 years attributed to the
displacement of bats to other regions of Sulawesi.
Several market vendors commented on difficulties obtaining wildlife from trade intermediaries. Some attributed this to
the road blockades organized by the Department of Forestry to monitor wildlife transportation into North Sulawesi. Others
mentioned that it was a seasonal effect.
3.5. Awareness of protected species law in Sulawesi
Eighty-nine percent of the vendors and 84% of the hunters interviewed were aware of the Indonesian law prohibiting the
hunting and trade of protected species including anoas, babirusas, and macaques. Most reported that these species were still
available in markets because the law was not enforced and the risk of getting caught was very low.
4. Discussion
4.1. Diversity and sustainability of the wildlife trade in Sulawesi
Species diversity of wildlife currently traded in North Sulawesi markets is quite similar to the diversity observed in these
markets in the early 2000s (Lee et al., 2005;Milner-Gulland and Clayton, 2002). Flying foxes, wild pigs, and rats are still
among the most common and abundant taxa traded in the region. However, the large quantities of snakes (pythons) currently
sold in North Sulawesi was not reported in previous work. It is possible that snakes are serving as a replacement for taxa that
are becoming rare. Tarsiers, civets, and deers were regularly traded in the early 2000s (Lee et al., 2005) but they were not
observed during our market surveys or reported in our hunteror vendor interviews. Protected species such as babirusa, anoa,
Celebes crested macaque, and cuscus species are still hunted and traded in low numbers in several markets.
Among the four wildlife taxa most commonly traded in North Sulawesi, flying foxes were sold in the highest number of
individuals and are the taxa most vulnerable to overhunting, due to their low reproductive rate and gregarious behavior
which exposes large numbers of bats to a single hunter (Mickleburgh et al., 2009). Our data provide a critical update to
previous studies (Lee et al., 2005;Sheherazade and Tsang, 2015). According to our estimates, the number of flying foxes
annually traded in Sulawesi ranges from 662,551 to more than one million individuals with up to 20,000 to 26,000 bats
harvested from a single roost annually. These quantities suggest that the Sulawesi trade in flying foxes is unsustainable.
Previous work on Pteropus populations suggest that a take of 3e10% of the population is sustainable (Brooke and Tschapka,
2002;Epstein et al., 2009). A wild population of 6,620,000 to 22,066,000 flying fox individuals would therefore be needed in
Sulawesi to sustain an annual harvest of 662,000 bats. However, there is a lack of data on baseline flying fox population size
and demography in Sulawesi.
Our systematic mapping of flying fox roosts in Sulawesi provides preliminary data for long-term monitoring of bat
populations on the island. We identified 45 flying fox roosts in Sulawesi, among which 38 were under active hunting pressure.
It is likely that there are additional unidentified roost sites. While there are no published data on flying fox colony size in
Sulawesi, colonies of P. alecto in Australia varied from 500 to 30,000 animals (Loughland, 1998;Vardon and Tidemann, 1999).
Our own estimate based on departure counts in a colony of Gorontalo province is around 15,000 individuals. Thus, assuming a
roost size of 15,000 and 30,000 individuals per site, the total flying fox population size in Sulawesi based on the 45 roosts we
observed, would range from 675,000 to 1,350,000 individuals. This is well below the sustainable population size required for a
3e10% offtake, and suggests that the current rates of harvest are not sustainable in the long term. Similar conclusions were
reached for other hunted Pteropodid species in Southeast Asia such as Pteropus vampyrus in Indonesia (Borneo) (Struebig
et al., 2007;Harrison et al., 2011), Malaysia (Epstein et al., 2009), and the Philippines (Scheffers et al., 2012); P. tonganus
on Niue Island (Brooke and Tschapka, 2002) and Solomon Islands (Lavery and Fasi, 2017) in the South Pacific Ocean; and in
Africa such as Eidolon helvum in Ghana (Kamins et al., 2011). Intense hunting during birthing season, as reported in most
provinces of Sulawesi during our study, may also increase the unstainability of the trade. Moreover, unintended collateral
mortality from bats may happen when using some hunting techniques such as air rifles which injure bats that escape and die
later without being caught by hunters. The total bat mortality due to the trade may therefore be higher than our estimated
A. Latinne et al. / Global Ecology and Conservation 21 (2020) e00887 9
number of bats sold in markets. Further, the lack of information on migration between Sulawesi and other major islands
suggests source-sink dynamics may artificially be inflating the sustainability of the bat harvest in Sulawesi (Breed et al., 2010).
While this is a key conservation concern, it may also have a significant economic impact, through loss of pollination and seed
dispersal ecosystem services (Aziz et al., 2017;Oleksy et al., 2015;Sheherazade et al., 2019).
The other taxa most commonly traded in North Sulawesi, wild pigs, snakes, and rats, present a lower risk of overhunting as
they are traded in lower numbers and have higher reproduction rates than flying foxes (Macdonald, 1993;Shine et al., 1999).
All are considered nuisance animals in most regions of Sulawesi. However, declines in their populations may have an impact
on the ecosystems that have not yet been assessed. Wild pigs and pythons are also commercially hunted and traded in other
regions of Southeast Asia where population declines were observed (Bennett et al., 2000;Kawanishi et al., 2013;Luskin et al.,
2014;Nijman et al., 2012;Scheffers et al., 2012;Shine et al., 1999).
4.2. Current extent of the wildlife trade network in Sulawesi
The trade network to supply wild meat to the markets of North Sulawesi is well organized and extends across the whole
island. Hunters and trade intermediaries are established in all provinces of the island. The network is dynamic, open, and
redundant, with numerous people playing similar roles (Phelps et al., 2016). There is no restriction on participation and any
hunter can access the market easily. Local dealers and middlemen actively seek out hunters and new sources of wild meat.
Hooked ropes, the most commonly used technique to hunt flying foxes, can be considered as passive hunting, easy to
implement in addition to other daily activities. It requires little effort, skill, and time and low economic investment. It is
therefore an opportunistic source of income for villagers living close to bat roosts. Similar hunting techniques using hooked
roped and kites have been observed in other regions of Southeast Asia, including Malaysia and the Philippines (Mickleburgh
et al., 2009;Mildenstein et al., 2016;Shively, 1997).
We clearly observed that middlemen reacted to wildlife population decline in North Sulawesi and Gorontalo provinces by
displacing effort to other provinces or by targeting other taxa. The results of our flying fox roost survey show that a third of the
flying fox hunting sites are now located in Southeast Sulawesi, mostly in the southern part of the province, while this province
was the least import source of flying foxes a few years ago (Sheherazade and Tsang, 2015). This region is the furthest and most
difficult to reach from North Sulawesi, which explains why it has been the last to be colonized by middlemen. Our study also
shows that, despite the seasonality of bat hunting in most provinces, the supply in North Sulawesi markets remains mostly
stable throughout the year as middlemen and trade intermediaries adapt to hunting seasonality and get bats from different
regions at different periods of the year.
4.3. Economics
Wildlife hunting in Sulawesi is mostly for commercial purposes and prices for flying foxes paid to hunters by middlemen
varied widely across regions. The highest prices were paid to hunters in Gorontalo, where the supply is low and the demand
from middlemen is high as this is the closest province from North Sulawesi and long trips are not required. Compared to the
prices in West and Central Sulawesi, prices paid in Southeast Sulawesi are quite high, despite the remoteness of this province.
This might be explained by strong competition between trade intermediaries in a region where a large number of local
dealers and middlemen are active. Hunters in Southeast Sulawesi can choose to sell their bats to trade intermediaries offering
the highest price after negotiations over the phone. This is likely notpossible in West Sulawesi as only one middlemanvisited
the region where we interviewed bat hunters. During the high season, a flying fox hunter in Southeast Sulawesi can earn
around 20 million IDR per month, which is 10 times the minimum monthly wage in the province based on government
standards.
Flying fox selling prices in markets also varied, Tomohon and Karombasan markets were the most expensive. Prices for
other wild meat were more similar across markets. Prices for wild meat in 2018e2019 were three to four times higher than
the prices in 1999 adjusted for inflation and bat meat was already more expensive than other meat at that time (Milner-
Gulland and Clayton, 2002). This price increase after adjustment for inflation over the last 20 years may reflect wildlife
population decrease in Sulawesi and the increased difficulty to supply markets of North Sulawesi. Future climate change may
increase this difficulty as the future distribution of Pteropus species is predicted to change in some regions of their distribution
range (Daszak et al., 2013).
4.4. Potential strategies for wildlife trade management in Sulawesi
Indonesian law states that anyone who trades, keeps, or kills a protected species may be jailed for up to five years and/or
fined up to 100 million Indonesia rupiahs (7000 USD) (Government Regulation No. 7, 1999). The trade of unprotected wildlife
species also requires a legal permit and is allowed only if a quota is allocated for the species. These quotas are determined
annually by the Natural Resources Conservation Agency (BKSDA) under the Directorate General of Forest Protection and
Nature Conservation (PHKA) at the Ministry of Environment and Forestry. However, a high number of wildlife species,
including flying foxes, do not have a hunting quota allocated so that their capture and trade is technically illegal (Bergin et al.,
2018;Shepherd et al., 2016). Three flying fox species (Acerodon humilis,Pteropus pumilus and Neopteryx frosti) are currently
legally protected (Environment and Forestry Ministry in 2018, Decree No. P.20/MENLHK/SETJEN/KUM.1/6/2018). However,
A. Latinne et al. / Global Ecology and Conservation 21 (2020) e0088710
the flying fox species most commonly traded in Sulawesi (Acerodon celebensis,Pteropus alecto and P. griseus) are not legally
protected and have no allocated hunting quotas. The inclusion of Acerodon celebensis and Pteropus griseus on the protected list
would likely be a very significant and simple conservation policy, as their distribution ranges are mostly limited to Sulawesi
where these species are massively hunted. Our study demonstrates that poaching and hunting of protected species occurs on
the island. A high proportion of respondents were aware of the protected species laws but suggested that the risk of getting
caught was low. This highlights a need for stricter law enforcement targeting market vendors and middlemen in coordination
with the BKSDA and the Ministry of Transportation. Hunting quotas for species that are not protected and their strict
implementation are also needed. Our study suggests that there would be high value in ecological research and field studies to
monitor wildlife populations in Sulawesi including the quantification of the impact of the trade on their demography and on
the ecosystem services they provide. This informationwould provide the basis for determining sustainable harvesting quotas
for each species.
Educational outreach to communities in North Sulawesi around wildlife conservation would likely help highlight the
importance of maintaining sustainable trading levels and it would be particularly useful targeted to those living close to
hunting sites for bats, wild pigs, and snakes. In particular, the pest status of these species could be addressed by highlighting
the ecosystem services they provide. This could include eco-tourism activities, which our study demonstrates has incite local
communities to conserve bat roosts. Educational campaigns are already ongoing in North Sulawesi, with mixed results
(Gursky and Fields, 2018). In particular, changing the wildlife consumption habits of Minahasan people is challenging as this
behavior is considered to be a source of cultural pride.
Reducing consumer demand through the use of economic incentives would also likely be a valuable approach to con-
servation. Our survey showed that the price for domestic pig meat is higher than the price for snake or wild pig meat and
similar to the price for flying fox meat. Strategies to provide a cheaper domestic alternative to wild meat in North Sulawesi
would therefore potentially reduce demand for wild meat.
Finally, providing opportunities for alternative livelihoods to hunters may also be a solution to overhunting. Bat hunting is
considered as “easy money”and is an important source of income for hunters, butmost of them also have other activities such
as farming and fishing that could be further developed. We observed that hunters in regions of Southeast Sulawesi where bat
populations are declining and where hunting is less profitable than a few years ago were willing to find an alternative to
hunting with the support of the local government. Middlemen could be a key target for intervention as a small number of
them trade high volume of animals.
5. Conclusion
Our study showed that hundreds of thousands of bats, wild pigs, snakes, and rats are still hunted and traded annually in
Sulawesi through a well-organized and dynamic network involving many actors. Flying fox hunting is an important source of
income in Sulawesi, as it is easy to implement in conjunction with other daily activities, and hunters can readily access to the
market. However, our analysis suggests current rates of flying fox harvests are significantly unstainable, and that population
declines have already affected the availability of wild meat. Trade intermediaries have reacted to the declines in North
Sulawesi and Gorontalo provinces by displacing effort to the furthest provinces, where populations are now declining. We
also demonstrate that trade of protected species is still ongoing too, but a lower scale, in North Sulawesi where there is an
urgent need for stricter law enforcement. Implementation of hunting quotas as well as research and conservation efforts are
also urgently needed to improve the sustainability of the wildlife trade in Sulawesi as the impact of this trade on Sulawesi
local ecosystems remains mostly unknown.
Declaration of competing interest
The authors declare that they have no known competing financial interests or personal relationships that could have
appeared to influence the work reported in this paper.
Acknowledgements
We thank all respondents who participated in this study and kindly agreed to share their knowledge with us. We also
thank our local collaborators, Dr. Safriyanto Dako, Dr. Agustinus Lomboan and drh Rakhwana, and drivers who provided us
with valuable information and assisted us in data collection in the field. This study was funded by the United States Agency for
International Development (USAID) Emerging Pandemic Threats PREDICT project (cooperative agreement number GHN-A-
OO-09-00010-00).
Appendices
A. Latinne et al. / Global Ecology and Conservation 21 (2020) e00887 11
Fig. A1. (A) Wild meat for sale at Tareran market; (B) Wild meat (bats, wild pigs and birds) for sale at Dumoga market; (C) Wild meat (macaque, snakes and wild
pigs) for sale at Dumoga market (pictures: Suryo Saputro/Alice Latinne).
Fig. A2. (A) Rat roasting in Kawangkoan market; (B) Roasted rats for sale at Tomohon market; (C) Roasted bats for sale at Tomohon market; (D) Wild meat (wild
pigs, snakes, and bats) and dogs for sale at Tomohon market (pictures: Suryo Saputro/Alice Latinne).
A. Latinne et al. / Global Ecology and Conservation 21 (2020) e0088712
Fig. A4. Flying fox hunting techniques in Sulawesi: (A) Drone photo showing poles holding nets around a roost of Sulawesi flying foxes in South Sulawesi; (B)
Hooked ropes and bamboo poles in Southeast Sulawesi; (C) Hunter removing a black flying fox from hooked ropes in Southeast Sulawesi (pictures: Suryo
Saputro/Alice Latinne).
Fig. A3. (A) Wild meat (wild pigs, rats, and bats) for sale in a supermarket in Manado; (B) Wild meat cooked in a Minahasan restaurant in North Sulawesi; (C)
Pythons in a freezer in a middleman’s house in Tomohon (pictures: Suryo Saputro/Alice Latinne).
A. Latinne et al. / Global Ecology and Conservation 21 (2020) e00887 13
Fig. A5. Quantities of large fruit bats, wild pigs, rats and snakes sold every day in Langowan (A), Kawangkoan (B), Tomohon (C), Karombasan (D) and Tareran (E)
markets and total quantities sold in these five markets every week (F) from November 12, 2018 to January 31, 2019. Quantities are in kg, except for rats where
quantities are given in numbers of individuals sold. The three days before Christmas (December 22e24, 2018) and New Year (December 29e31, 2018) are
highlighted in grey.
A. Latinne et al. / Global Ecology and Conservation 21 (2020) e0088714
Table A1
Markets surveyed in North Sulawesi province in 2018e2019 and wildlife taxa sold in these markets
District Subdistrict City/Market Wildlife
for sale
Taxa Market days Number of
wildlife
vendors
Bolaang Mongondow West Dumoga Ibolian Yes Bats, Snakes, Wild pigs, Rats,
Monkeys, Anoa, Squirrels,
Monitor Lizards
Tue-Thu-Sun 7
Bolaang Mongondow East Dumoga Dumoga Yes Bats, Snakes, Wild pigs,
Rats, Monkeys, Anoa,
Squirrels, Monitor Lizards
Tue-Fri 7
Bolaang Mongondow East Dumoga Imandi Yes Bats, Snakes, Wild pigs,
Rats, Monkeys, Anoa,
Squirrels, Monitor Lizards
Mon-Wed-Sat 7
South Minahasa Modoinding Modoinding Yes Bats, Snakes, Wild pigs,
Rats, Monkeys, Cuscus
Mon-Wed-Fri 10
South Minahasa Tompaso Baru Tompaso Baru Yes Bats, Rats, Wild pigs,
Anoa, Snakes,
Monkeys, Cuscus
Tue-Sat 10
South Minahasa Ranoyapo Poopo Yes Bats, Rats Mon-Tue-Wed-Fri 5
South Minahasa Motoling Motoling Yes Bats, Snakes, Wild pigs,
Rats, Monkeys, Anoa,
Cuscus, Babirusa
Thu 11
South Minahasa Sinon Sayang Ongkaw Yes Bats, Snakes,
Wild pigs, Rats
Mon-Sat 3
South Minahasa Tenga Tenga Yes Wild pigs, Rats, (Snakes) Mon-Wed-Fri 2
South Minahasa Amurang Amurang Yes Bats, Snakes, Wild pigs,
Rats, (Monkeys)
Mon-Tue-Wed-Thu-Fri-Sat-Sun 6
South Minahasa Tumpaan Tumpaan Yes Bats, Snakes, Wild pigs Mon-Tue-Wed-Thu-Fri-Sat-Sun 6
South Minahasa Tareran Tareran Yes Bats, Snakes, Wild pigs, Rats Mon-Wed-Fri 2
Southeast Minahasa Tombatu Tombatu Yes Bats, Snakes, Wild pigs, Rats Mon-Wed-Sat 1e2
Southeast Minahasa Ratahan Ratahan Yes Bats, Snakes, Wild pigs, Rats Mon-Tue-Wed-Thu-Fri-Sat-Sun 5
Minahasa North Langowan Langowan Yes Bats, Snakes, Wild pigs, Rats Tue-Thu-Sat 7e11
Minahasa North Kawangkoan Kawangkoan Yes Bats, Snakes, Wild pigs, Rats,
Anoa, Babirusa, Frogs, (Monkeys)
Mon-Thu-Sat 7e11
Minahasa Sonder Sonder Yes Bats, Snakes, Wild pigs, Rats Mon-Wed-Fri 4
Minahasa West Tompaso Tompaso Yes Bats, Snakes, Wild pigs, Rats Wed-Fri 2
Tomohon City East Tomohon Tomohon Yes Bats, Snakes, Wild pigs, Rats,
Monitor Lizards, Frogs,
Cuscus, Monkeys, Anoa, Babirusa
Mon-Tue-Wed-Thu-Fri-Sat-Sun 6e10
North Minahasa Airmadidi Airmadidi Yes Bats, Snakes, Wild pigs, Rats Tue-Thu-Sat 2
Manado City Wanea Karombasan Yes Bats, Snakes, Wild pigs, Rats Mon-Tue-Wed-Thu-Fri-Sat-Sun 2
Manado City Wenang Bersehati Yes Wild pigs, other taxa if ordered Mon-Tue-Wed-Thu-Fri-Sat-Sun 1
Manado City Mapanget Paniki Yes Bats, Snakes, Wild pigs, Rats Mon-Tue-Wed-Thu-Fri-Sat-Sun 1
Minahasa East Langowan Old Langowan Occasion. Bats Mon-Wed-Fri Unknown
Bitung City Girian Girian Occasion. Wild pigs, other taxa if ordered Mon-Tue-Wed-Thu-Fri-Sat-Sun Unknown
Minahasa Remboken Remboken Occasion. Wild pigs, other taxa if ordered Mon-Tue-Wed-Thu-Fri-Sat-Sun Unknown
North Minahasa Dimembe Tatelu Occasion. Wild pigs, other taxa if ordered Mon-Tue-Wed-Thu-Fri-Sat-Sun Unknown
Minahasa West Tondano Tondano No NA Mon-Tue-Wed-Thu-Fri-Sat NA
North Minahasa Airmadidi Sukur No NA Tue-Thu-Sat NA
North Minahasa Kauditan Kauditan No NA Mon-Wed-Fri-Sun NA
North Minahasa Dimembe Matungkas No NA Mon-Wed-Fri-Sun NA
A. Latinne et al. / Global Ecology and Conservation 21 (2020) e00887 15
Table A2
Flying fox roosts identified in Sulawesi and their hunting status in 2018e2019. Map numbers refer to Fig. 3. Pa, Pteropus alecto; Ac, Acerodon celebensis; Pg,
Pteropus griseus.
Map # Province District Subdistrict Species Habitat Hunting
1 Gorontalo North Gorontalo Kwandang Pa, Ac Coastal mangrove swamp Yes
2 Gorontalo North Gorontalo Sumalata Coastal mangrove swamp Unknown
3 Gorontalo Boalemo Paguyaman Pantai Pa, Ac Coastal mangrove swamp Yes
4 Gorontalo Boalemo Tilamuta Ac Coastal mangrove swamp Yes
5 Gorontalo Boalemo Tilamuta Pa Coastal mangrove swamp Yes
6 Gorontalo Pohuwato Lemito Pa Coastal mangrove swamp Yes
7 Central Buol Lakea Coastal mangrove swamp Yes
8 Central Toli Toli Basidondo Pa Mainland forest Yes
9 Central Toli Toli Dondo Coastal mangrove swamp (Nipa palm) Yes
10 Central Donggala Sojol Coastal mangrove swamp (Nipa palm) Yes
11 Central Donggala Sojol Pa, Ac Coastal mangrove swamp (Nipa palm) Yes
12 Central Donggala Damsol Lake mangrove swamp (Nipa palm) Yes
13 Central Donggala Balaesang Coastal mangrove swamp (Nipa palm) Yes
14 Central Parigi Moutong Toribulu Pa Coastal mangrove swamp No
15 Central Banggai Bualemo Pg Coastal mangrove swamp Yes
16 Central Banggai West Toili Pa, Ac Lake mangrove swamp Yes
17 Central Morowali North Bungku Pa, Ac Coastal mangrove swamp Yes
18 Central Morowali Bungku Pg Coastal mangrove swamp Yes
19 West Mamuju Karossa Coastal mangrove swamp Yes
20 West Mamuju Mamuju Pa, Ac Island forest Yes
21 West Majene Tubo Pa, Ac Coastal mangrove swamp Yes
22 West Majene Sendana Pa, Ac Coastal mangrove swamp Yes
23 South Pinrang Suppa Coastal mangrove swamp Unknown
24 South Barru Balusu Pa, Ac Coastal mangrove swamp Yes
25 South Pangkajene Kepulauan Liukang Tupabbiring Coastal mangrove swamp Yes
26 South Maros Simbang Pa, Ac Residential area No
27 South East Sinjai Tongke Tongke Coastal mangrove swamp Yes
28 South Soppeng Lalabata Pa Residential area No
29 South Wajo Sajoanging Coastal mangrove swamp Yes
30 South North Luwu Bonebone Coastal mangrove swamp Yes
31 Southeast North Kolaka Ngapa Pa Coastal mangrove swamp Yes
32 Southeast Konawe Puriala Pa Lake mangrove swamp Yes
33 Southeast Kolaka Watubangga Pa, Ac Coastal pine trees No
34 Southeast Bombana Poleang Coastal mangrove swamp Yes
35 Southeast Konawe Selatan Tinanggea Pa, Ac Coastal mangrove swamp Yes
36 Southeast Konawe Selatan Laeya Pa Coastal mangrove swamp Yes
37 Southeast Muna North Tiworo Coastal mangrove swamp Yes
38 Southeast Bau Bau Coastal mangrove swamp Yes
39 Southeast Wakatobi South Wangi-Wangi Ac Coastal pine trees No
40 Southeast Buton Kapuntori Coastal mangrove swamp Yes
41 Southeast Muna South Wakorumba Pa Coastal mangrove swamp Yes
42 Southeast Konawe Selatan Kolono Pa, Ac Coastal mangrove swamp Yes
43 Southeast Konawe Kepulauan Wowoni Coastal mangrove swamp Yes
44 Southeast Konawe Selatan Moramo Coastal mangrove swamp Yes
45 Southeast Konawe Bondoala Pa Coastal mangrove swamp Yes
Table A3
Demographic data from bats hunted in three flying fox roosts in Gorontalo, West Sulawesi and Southeast Sulawesi and estimates of the quantity of bats
harvested annually in each roost. Pa, Pteropus alecto; Ac, Acerodon celebensis. A, Adults; SA; Subadults; J, Juveniles; B, babies.
Province n Numbers of Ac/Pa
hunted (proportion in %)
Numbers of males/females
hunted (proportion in %)
Average
weight (g)
Numbers of A/SA/J/B
hunted (proportion in %)
Estimates of annual
quantity harvested
(in kg/in individuals)
Gorontalo 479 116/363 (24.2/75.8) 225/254 (47/53) 493.62 298/176/2/0 (62.2/36.7/1.1/0) 11,200kg/22,736 bats
West Sulawesi 382 41/341 (10.7/89.3) 173/209 (45.3/54.7) 454.34 140/186/54/3 (36.6/48.7/14.1/0.6) 9400 kg/20,680 bats
Southeast Sulawesi 363 96/267 (26.4/73.6) 208/155 (57.3/42.7) 517.68 252/78/28/4 (69.4/21.5/7.7/1.4) 13,800 kg/26,634 bats
Global 1224 253/971 (20.7/79.3) 636/618 (52/48) 486.54 690/440/84/7 (56.4/35.9/6.9/0.8)
A. Latinne et al. / Global Ecology and Conservation 21 (2020) e0088716
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Table A4
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Markets Large fruit bats (kg) Wild pigs (kg) Rats (individuals) Snakes (kg) Source
Tomohon 46,692 26,710 5594 6199 Daily obs.
Karombasan 16,743 16,413 149 8935 Daily obs.
Kawangkoan 25,410 41,956 2675 3636 Daily obs.
Langowan 30,505 134,312 3373 6080 Daily obs.
Tareran 3671 6240 1413 1890 Daily obs.
Ibolian/Dumogo/Imandi 8372 21,840 7800 5460 Interviews
Modoinding 48,360 31,200 NA 7800 Interviews
Motoling/Poopo/Tompaso Baru 57,200 34,320 NA 8580 Interviews
Sonder 24,960 15,600 NA 3120 Interviews
Amurang 31,200 18,720 NA 4680 Interviews
Ongkaw 2600 9360 NA 2340 Estimate
Tumpaan 5200 18,720 NA 4680 Estimate
Tombatu 5200 3120 NA 780 Estimate
Ratahan 2600 15,600 NA 3900 Estimate
Tompaso 2600 6240 NA 1560 Estimate
Airmadidi 5200 6240 NA 1560 Estimate
Bersehati Not sold 3120 NA Not sold Estimate
Paniki 2600 3120 NA 780 Estimate
Tenga Not sold 6240 NA Not sold Estimate
Total 319,113 419,071 21,004 71,980
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