BIOLOGICAL

CONSERVATION

Biological Conservation 123 (2005) 477–488

www.elsevier.com/locate/biocon

Wildlife trade and implications for law enforcement in Indonesia:

a case study from North Sulawesi
a,*
Robert J. Lee , Antonia J. Gorog a, Asri Dwiyahreni a, Stephan Siwu b, Jon Riley c

Harry Alexander a, Gary D. Paoli a, Widodo Ramono d

a
Wildlife Conservation Society, Indonesia Program, Jl. Pangrango No. 8, Bogor, Indonesia
b
Wildlife Conservation Society, Indonesia Program, Jl. Sea No. 105, Manado, Sulawesi, Indonesia
c
North Sunderland and Seahouses Development Trust, 62-64 Main Street, Seahouses, Northumberland NE68 2TP, UK
d
Department of Forestry, Manggala Wanabakti, Jl. Gatot Subroto, Jakarta 10065, Indonesia

Received 6 September 2004

Abstract

Excessive hunting pressure, due in large part to commercialization, has reduced the populations of many tropical large mammal
species. Wildlife over-exploitation is severe in Indonesia, especially on Sulawesi, where human resources and funding are inadequate
to monitor the wildlife trade and enforce existing protection laws. In response, the Wildlife Crimes Unit program was established in
December 2001 to: (i) monitor wildlife transportation into North Sulawesi and market sales; (ii) provide legal and technical support
to law enforcement agencies; and (iii) promote public awareness of wildlife and protection laws. Over a two-year period, 6963 wild
mammals en route to markets were encountered (8 individuals h 1) and 96,586 wild mammals were documented during market
surveys. The trade of some protected mammals declined significantly over this period, but overall trade in wild mammals increased
by 30%. High volume of trade in non-protected animals such as the Sulawesi pig Sus celebensis and large flying foxes (Pteropodidae),
raise concerns about the sustainability of current harvesting. To combat this problem, we recommend that: (1) efforts are continued
to reduce trade in protected species; (2) protected status is extended to heavily traded but non-protected taxa, such as flying foxes;
(3) the effects of hunting on rat and bat populations, as well as its impact on forest dynamics, are quantified; and (4) work is carried
out with local communities to strengthen awareness, set sustainable limits on wild mammal harvesting, and establish practical mech-
anisms for enforcing these limits.

2005 Elsevier Ltd. All rights reserved.

Keywords: Hunting; Indonesia; Law enforcement; Sulawesi; Wildlife trade

1. Introduction able levels, particularly in the humid tropics where an-

The world is facing a potentially massive loss of wild-
life due to over-hunting (Robinson and Bennett, 2000,
2002; Bennett et al., 2002). Hunting rates exceed sustain-
*
Corresponding author. Tel.: +62 251 321527; fax: +62 251 357347.
E-mail addresses: rlee@wcs.org (R.J. Lee), a.gorog@wcsip.org
(A.J. Gorog), a.dwiyahreni@wcsip.org (A. Dwiyahreni), s.siwu@wcsi-
p.org (S. Siwu), ninoxios@yahoo.co.uk (J. Riley), h.alexander@wcsi-
p.org (H. Alexander), gpaoli@umich.edu (G.D. Paoli),
cites@dephut.cbn.net.id (W. Ramono).
nual harvest rates are as high as 1–5 million tons in
Central Africa (Wilkie and Carpenter, 1999; Fa and
Peres, 2001; Wilkie et al., 2001), 67,000–164,000 tons in
the Brazilian Amazon (Robinson and Redford, 1991),
and 23,000 tons in Sarawak (Bennett et al., 2000). As a
consequence, wildlife populations are being extirpated
across their ranges (Bennett and Robinson, 2000; Wilkie
et al., 2001; Fa et al., 2004), and, in turn, people who de-
pend on wildlife meat for food and income suffer (Rob-
inson and Bennett, 2002; Wilkie and Lee, 2004). Recent
studies of hunting on the Indonesian island of Sulawesi

0006-3207/$ - see front matter
2005 Elsevier Ltd. All rights reserved.
doi:10.1016/j.biocon.2005.01.009
478 R.J. Lee et al. / Biological Conservation 123 (2005) 477–488
have documented patterns of hunting, wildlife trade, and
the impact of these activities on target wildlife popula-
tions (Clayton, 1996; Lee, 1997, 2000a,b; Alvard, 2000;
Clayton and Milner-Gulland, 2000; OÕ Brien and Kin-
naird, 2000; Wildlife Conservation Society, 2000, report
on monitoring and control of trade in protected wildlife
in North Sulawesi, Indonesia in litt.). These studies have
shown that most hunting on Sulawesi is unsustainable,
and that heavy exploitation for human consumption is
the primary cause of wildlife declines on this island.
Terrestrial vertebrate species on Sulawesi are hunted
for subsistence and for the market. Consumption of
wildlife occurs throughout the island, but the practice
is most extensive in North Sulawesi Province, where
most of the population is Christian and, therefore, not
constrained by religious taboos held by much of Muslim
Indonesia against consuming wildlife species such as
monkeys, pigs, bats, and rodents. The people of North
Sulawesi have traditionally hunted wildlife locally, with-
in the province, but over-exploitation has extirpated
some mammal species including anoa (Bubalus spp.),
babirusa (Babyrousa babyrussa), and flying foxes from
once-productive hunting sites, such as the Tangkoko-
Duasudara Nature Reserve (Anon., 1981; OÕ Brien and
Kinnaird, 2000). As a result, market demand in the area
is met by transporting wildlife from other parts of
Sulawesi (Lee, 2000b; Wildlife Conservation Society,
2000, report on monitoring and control of trade in pro-
tected wildlife in North Sulawesi, Indonesia in litt.).
Completion of the Trans-Sulawesi Highway (Fig. 1) in
1980 made transportation more reliable and probably
increased importation of wildlife into North Sulawesi
from the rest of the island.
The high volume of wild meat consumed in North
Sulawesi appears to reflect a preference, rather than a
necessity. North Sulawesi is a well-developed province
of Indonesia, with relatively high levels of income, pro-
tein and caloric intake, education, and literacy, and its
people are neither nutritionally nor economically depen-
dent on wild meat (Lee, 2000a). Trade in North Sulawesi
markets has not previously been monitored throughout
the year, but one of the authors (Lee) has noted apparent
peaks in consumption of some wildlife species (e.g., ma-
caques) near holidays, further suggesting that wildlife
consumption reflects dietary choice rather than necessity.
Illegal hunting of wildlife is common, either because
hunters are taking protected species or are hunting in
protected areas. The Indonesian government and con-
servationists are, thus, working to reduce illegal hunting
pressure on wildlife populations. Two of the most
important approaches for protecting wildlife are creat-
ing a legal basis for protection and establishing a system
of law enforcement. Indonesia meets the first criterion,
with more than 150 existing national laws and regula-
tions to protect its wildlife species and areas. In addi-
tion, the country is a signatory to many international
conventions on biodiversity conservation, including the
Convention on International Trade for Endangered
Species of Wild Flora and Fauna (CITES) by Presiden-
tial Decree No. 43/1978 and the Convention on Biolog-
ical Diversity by Law No. 5/1994. Indonesia, however,
lacks an integrated system of law enforcement. Problems
include: lack of awareness among law enforcement offi-
cials about conservation laws and policies; low technical
capacity; limited financial and human resources; lack of
monitoring both where wild animals are captured and
sold and, thus, a lack of data on wildlife trade and its
impact on populations. These problems are com-
pounded by the vast, fragmented geography of the coun-
try. As a consequence of these problems, illegal hunting
and wildlife trade are rampant in Indonesia.
Recent changes in government policies, especially the
new decentralization laws (No. 22 and No. 25, both
passed in 2001), have had a major impact on the way
conservation and natural resource management are con-
ducted in Indonesia. Local governments, non-govern-
mental organizations, and local communities now
participate in managing and protecting natural re-
sources, requiring greater coordination between govern-
ment and non-government sectors, and between
national and local government agencies, to enforce laws.
To address the problems of illegal hunting and wild-
life trade, and to take advantage of recent political
changes, in 2001 the Indonesian Department of Forestry
and the Wildlife Conservation Society established a
Wildlife Crimes Unit in North Sulawesi, comprising
members of the Department of Forestry, local govern-
ment agencies and police, local and international non-
governmental organizations, and the media. This unit:
(i) trains law enforcement officers and local groups in
wildlife trade monitoring and law enforcement, (ii) mon-
itors wildlife sales and populations in source areas, (iii)
provides financial and technical support for legal action
against illegal hunters and wildlife traders, and (iv)
works with the media and local communities to heighten
public awareness of conservation issues. The long-term
goal of the Wildlife Crimes Unit is to develop a program
to curb illegal hunting and wildlife trade that can be ap-
plied widely throughout Indonesia.
In this paper, we report preliminary results following
two years of Wildlife Crimes Unit efforts to monitor and
reduce traffic in mammalian wildlife. We ask: (1) what is
the level of trade activity and taxonomic composition of
wildlife trade in markets of North Sulawesi; (2) has the
level of wildlife trade activity in markets, in particular
that concerning protected species, changed during the
two years since the establishment of Wildlife Crimes
Unit road blockades along the Trans-Sulawesi Highway;
(3) what is the origin, taxonomic composition, and min-
imum volume of wildlife currently transported along the
Trans-Sulawesi Highway in North Sulawesi; and (4)
does wildlife trade activity and taxonomic composition
 R.J. Lee et al. / Biological Conservation 123 (2005) 477–488
Fig. 1. Survey and road blockade locations (m, road blockades; n, markets in Manado; , markets in Minahasa).
vary with season or between regions in markets of North
Sulawesi? The market patrols and road blockades have
been conducted in tandem since the inception of the
Wildlife Crimes Unit program. Although prior data on
trade wildlife trade levels in North Sulawesi are not
available, the data presented here enable a quantifica-
tion of trends since establishment of the program and
provide a baseline for future comparisons and data
against which to examine future estimates of wildlife
trade levels in North Sulawesi.
2. Methods
2.1. Study area
The study was conducted in North Sulawesi, a 20,670
km2 province occupying the eastern half of SulawesiÕs
northern peninsula (Fig. 1). Road blockades and market
479
surveys were carried out from December 2001 to
December 2003 to monitor an index of wildlife trans-
ported to and traded in the province. The Wildlife
Crimes Unit has identified at least 25 markets in the
province that sell wildlife. Animals are usually sold
dead. Large animals, such as pigs or other ungulates,
are sold in pieces. Small animals, like bats and rats,
are generally sold whole but have often been singed.
Many of these animals are captured in regions west
and south of the province, from where they are trans-
ported on the Trans-Sulawesi Highway to North Sulaw-
esi markets (Wildlife Conservation Society, 2000, report
on monitoring and control of trade in protected wildlife
in North Sulawesi, Indonesia in litt.; Fig. 1).
2.2. Road blockades
The transportation of wildlife into and within North
Sulawesi was monitored at road blockades established
480 R.J. Lee et al. / Biological Conservation 123 (2005) 477–488
along the Trans-Sulawesi Highway (Fig. 1). Road patrol
teams involved ten to twelve members of the Wildlife
Crimes Unit, including forestry staff, police officers,
media, and conservationists. During blockades, police
and forestry officers flagged down vehicles deemed likely
to be transporting wildlife. Vehicles were inspected, and
any wild animals or wild meat found were counted and
identified to at least rough taxonomic groups (e.g., large
or small bats, rats, cuscuses). The drivers were asked
where the animals originated. If protected species were
discovered (i.e., macaque Macaca spp., babirusa B. bab-
yrussa, anoa Bubalus spp., dwarf cuscus Strigocuscus
celebensis, bear cuscus Ailurops ursinus, tarsiers Tarsius
spp., and Timorese deer Cervus timorensis), the drivers
of the vehicles transporting them were interviewed on
the spot by police and forestry officials, the animals were
confiscated, and, in some cases, enforcement measures
taken. The drivers were asked the geographic origin of
animals being transported. While the driversÕ answers
may contain inaccuracies, they are currently our only
means of gathering information about the source loca-
tions of the animals recorded at road blocks.
Eighteen such blockades were conducted between 22
December 2001 and 22 December 2003. To undermine
the efforts of traders to avoid blockades, the timing of
blockades was intentionally irregular, taking place on
average every 40 days (median = 33 d; range = 11–
123). Duration of blockades varied, also to thwart at-
tempts at waiting out the blockades; blockades averaged
45.2 h and ranged from 24.5 to 87.6 h (median = 41.7 h).
2.3. Market surveys
During the same period, December 2001 to December
2003, the Wildlife Crimes Unit monitored wildlife trade
in six markets of North Sulawesi, including three in the
Manado area (Kanaka, Karombasan, and Paal Dua)
and three in the Minahasa area (Kawangkoan, Lango-
wan, and Tomohon) (Fig. 1).
During each market visit, one or two covert observers
working together walked the meat section of the market,
counting and identifying animals or animal parts dis-
played. Quantities too large or piled too deeply on tables
to be counted easily were estimated by counting a sub-
sample and extrapolating across the pile. At each mar-
ket stall selling multiple animal parts (rather than whole
animals), the number of individual animals was esti-
mated by calculating number of hind feet of that species
divided by two, number of fore feet divided by two, and
number of heads, and then taking the greatest number
of the three.
Due to a number of factors that prevented species-
level identifications for some individuals (e.g., under-
cover nature of the monitoring, charred or singed
conditions of some individuals), most animals were clas-
sified into general categories (Table 1). Each of the six
markets was visited at least once per month; on occasion
individual markets were surveyed two or three times in a
month. Survey duration ranged from 0.27 to 6.17 h
(mean = 1.55 h; median = 1.33 h).
2.4. Analysis of road blockade data
Linear correlation was used to test for positive or
negative trends over time in number of wildlife encoun-
tered (all taxa combined) and protected species only. We
also tested for differences in means between the first and
second years of the blockades for individual species and
various combinations of taxa. Independent sample t-
tests and non-parametric Kruskal–Wallis tests were used
as appropriate given the distribution of the data.
2.5. Analysis of market survey data
Due to the illegal nature of much of the wildlife trade
and the corresponding need to work undercover, com-
plete counts of animals being sold from markets could
not be conducted. As such, identification and counts
of animals were a sample of goods for sale during the
market survey. For this reason, we standardized the vol-
ume of wildlife counted in markets by hour and used
encounter frequency as our metric to compare patterns
of wildlife trade; this is the index of wildlife trade activ-
ity we use throughout this paper. The settings in differ-
ent markets were similar, and we assume that the
Wildlife Crimes UnitsÕ pace of walking through markets
did not differ systematically with market, thus enabling
direct comparisons of encounter rates among markets.
Wildlife encounter frequencies in individual markets vis-
ited multiple times in one month were averaged across
visits.
Encounter frequency from market surveys were ana-
lyzed in three ways. Differences in index of trade activity
between regions (Manado vs. Minahasa) and years were
examined using two-factor repeated measures ANOVA
with region and year as fixed factors and markets as
the replicate. This ANOVA was performed for all taxa
combined and for protected species only. Second, with-
in-year temporal variation in wildlife sales was examined
using two-way ANOVA with year and month as fixed
factors and market as the replicate. Standard ANOVA
rather than repeated measures was used, because the in-
dex of trade activity between adjacent months was not
significantly correlated (R = 0.144; p = 0.212), indicating
independence. The analysis was performed for all taxa
combined and protected species only, using the natural
log of encounter rate (3+ animals h 1) and z-scores, to
normalize large differences between markets. Finally,
we tested whether the index of wildlife trade activity
declined during the study by: (i) comparing trade index
between years using the ANOVAs describes above;
(ii) testing for significant declining trends over time
 R.J. Lee et al. / Biological Conservation 123 (2005) 477–488 481

Table 1
Taxonomic categories used to group species encountered during road blockades and market surveys in North Sulawesi, Indonesia
Taxonomic category Representative species
Macaque Macaca hecki and M. nigrescens
Anoa Bubalus depressicornis and B. quarlesi
Civet Viverra tangalunga, Paradoxurus hermaphroditus and Macrogalidia musschenbroekii
Cuscus Strigocuscus celebensis, Ailurops ursinus
Rat Paruromys dominator
Squirrel Prosciurillus murinus and Rubrisciurus rubriventer
Small bat (mega- and microchiroptera) Cynopterus brachyotis and Megaderma spasma
Large bat (flying foxes) Pteropus hypomelanus, P. alecto, Acerodon celebensis and A. humilis
Tarsier Tarsius spectrum and T. pumilus
Sulawesi pig Sus celebensis
Babirusa Babyrousa babyrussa
Timorese deer Cervus timorensis

using linear correlation; and (iii) performing paired-

120
sample t-tests of months between years after averaging (a) All taxa combined
across all markets in a given month. 100

Individuals hour-1
80

3. Results 60
R=0.094NS

3.1. Road blockades 40

20
During a total of 813.7 h of road blockades over the
two-year period, 160 vehicles searched by the patrol 0

team contained 6963 wild mammals. Thirty-seven addi-

tional vehicles carried reptiles, wild birds, or domestic 1.2
animals (i.e., dogs, cats, or pigs), but no wild mammals.
Records were not kept of the total number of vehicles 1.0 (b) Protected species
searched and the total volume of traffic during the
0.8
patrol.
Individuals hr-1

Of mammalian wildlife recorded by road patrol 0.6

teams, large bats were the most frequently encountered
(66.4% of all animals), followed by the Sulawesi pig 0.4
(29.6%), and rats (2.6%). The remaining 1.4% of animals
recorded included small bats, babirusas, macaques, cus- 0.2 *
R = -0.492
cuses, and civets.
0.0
Pooling across all taxa, 64.3% of wildlife captures was
claimed by drivers to have originated from Gorontalo
Dec 2001
Feb 2002
Apr 2002
Jun 2002
Oct 2002
Dec 2002
Jan 2003
Feb 2003

Oct 2003
Mar 2003
Apr 2003

Jun 2003
May-b 2003
May-a 2003

Aug 2003
Sept 2003

Nov 2003
Dec 2003

Province, just west of North Sulawesi, followed by

27.5% from Central Sulawesi, 2.3% from South Sulaw-
esi, 2% from North Sulawesi and 0.1% from Southeast
Sulawesi. The estimate for North Sulawesi probably
understates the relative intensity of hunting pressure
on mammals in this province, as our road blockades
were in the western half of North Sulawesi and therefore
under-sampled the province (see road blockade loca-
tions in Fig. 1). Species-specific patterns of origin were
similar, except in the case of the Sulawesi pig, for which
drivers claimed larger percentages coming from more
distant provinces: 36.8% from Central Sulawesi and
11.4% from Southeast Sulawesi.
For all taxa combined, encounter rate during block-
ades varied markedly over the two-year period, but
Month of patrol
Fig. 2. Variation over time in the frequency of encountering mamal-
lian wildlife during mandatory roadside vehicle inspections in North
Sulawesi. Data are shown for all taxa combined (a) and for protected
species only (b), as defined in Section 2. Significance values in this and
following figures: *, p < 0.05; **, p < 0.01; ***, p < 0.001.
was not significantly correlated with time (R = 0.094,
p = 0.794; Fig. 2(a)). In contrast, protected species com-
bined showed a weak but significant declining trend over
the two-year period (R = 0.492, p = 0.048; Fig. 2(b)).
In a comparison of blockade data from the years 2002
482 R.J. Lee et al. / Biological Conservation 123 (2005) 477–488
(N = 6) and 2003 (N = 12), neither the rate of encounter
of all animals nor all protected species combined differed
significantly. However, in species-specific comparisons,
the frequency of encountering babirusa, a protected spe-
cies, was lower in 2003 than 2002 (0.22 ± 0.015 vs.
0.0 ± 0.0; Kruskal–Wallis test, p = 0.009).
3.2. Market surveys
An estimated total of 96,586 individual wild mam-
mals was recorded during surveys of six markets in
North Sulawesi from January 2002 to December 2003.
The largest proportion of these was rats, which ac-
counted for 43.9% of all encounters, followed by large
bats (39.8%), small bats (7.5%), and Sulawesi pigs
(7.3%). Cuscuses, squirrels, babirusa, macaques, tar-
siers, civets, anoas, and deer together accounted for
the remaining 1.2% of the individual wild mammals re-
corded in markets. The index of trade in mammalian
wildlife sold in the six sampled markets of North Sulaw-
esi showed considerable variation across markets and
over time, from a consistent encounter frequency of
<10 individual animals h 1at Paal Dua to >1600 indi-
viduals h 1at Kawangkoan during October 2003 (Fig.
3(a)). In repeated measures ANOVA with region and
year as fixed factors, region (Manado vs. Minahasa)
had a significant effect on the trade activity index of
all wild mammals in markets (F1,8 = 1.354, p = 0.001;
Table 2), with an average across years of 63.6 individu-
als h 1 in Manado and 313.6 in Minahasa. Encounter
frequency of protected species did not differ significantly
between regions (Table 3), but was on average three
times higher in Minahasa than Manado (3.1 vs. 1.0 indi-
vidual h 1). Month-to-month variation was not signifi-
cant for both groups of animals (Tables 2 and 3).
Two-way ANOVA to test for monthly seasonality in
wild meat trade showed no seasonal pattern for all wild
mammals combined (Fig. 3(b); F11,143 = 0.240,
p = 0.994), but strong seasonality of trade in protected
species (Fig. 3(c); F11,143 = 2.000, p = 0.034; Table 4).
A clear peak occurred in both years in November. When
the analysis was repeated using z-scores (rather than
count data), the significance of seasonality was even
stronger (p = 0.01).
Repeated measures ANOVA to test for differences in
index of trade activity between years showed no signifi-
cant differences between years for either protected spe-
cies or all taxa combined (Tables 2 and 3). Linear
correlation with time, however, showed a weak but sig-
nificant increase in the total number of wild mammals in
markets (R = 0.594; p 6 0.001; Fig. 3(b)). No such cor-
relation with time was detected for protected species
only (R = 0.056; p = 0.874; Fig. 3(c)).
Paired-sample t-tests comparing months between
years (e.g., February 2002 with February 2003) showed
no significant changes between years in the index of
trade activity of all wildlife or protected taxa combined
(Fig. 4). However, among eight taxonomic groups ana-
lyzed separately, we found that trade declined signifi-
cantly in babirusa (p < 0.001) and showed a similar
trend in macaques (p = 0.081), both of which are pro-
tected species (Fig. 4). No significant change was de-
tected for the cuscuses, another protected group, (Fig.
4), and sample sizes for the protected taxa anoa, tarsier
and deer were too small to be analyzed. Trade in two
non-protected groups, large bats and squirrels, increased
between months in 2002 and 2003, but these trends were
not significant (p = 0.066 and 0.069, respectively; Fig. 4).
No significant changes were detected in the trade of
small bats, rats, and the Sulawesi pig from 2002 to
2003 (Fig. 4).
4. Discussion
4.1. Changes over time in wildlife trade
Transportation and market sales of protected species
declined over the two-year study period, and for some
species significantly, a pattern consistent with expecta-
tions that Wildlife Crimes Unit road blockades and
associated law enforcement measures should reduce
trade in these taxa. The near-complete drop in protected
species observed after four months of road blockades
(Fig. 2(b)) and the decrease in babirusa and macaques
recorded in market surveys (Fig. 4) may be an indication
that hunters, transporters, and traders are now selec-
tively avoiding dealing in protected species, so as not
to face potential legal action if they are caught by law
enforcers.
An alternative explanation for the decrease in trade
of protected species is that this pattern reflects declining
numbers of these taxa in the wild. To address this alter-
native, it would be necessary to have long-term data on
population status of protected species from all major
source areas supplying North SulawesiÕs markets. Such
data are beyond the scope of the current Wildlife Crimes
Unit program, but should become a high priority of ex-
panded efforts to study the sustainability of wildlife
trade.
Although the decrease in trade of protected species
since the institution of the Wildlife Crimes Unit pro-
gram is encouraging, patterns of trade in other taxa over
the last two years are cause for alarm. Between 2002 and
2003, overall trade increased by 30%, and trade in large
bats, rats and squirrels increased by 44.5%, 33% and
>300%, respectively, although absolute numbers of
squirrels are consistently very low (Fig. 4). It is possible
that the increasing number of large bats reflects a strat-
egy by hunters and traders to make up for revenues lost
by avoiding sales of protected species. Large bats are not
protected by law, are relatively inexpensive to transport,
 R.J. Lee et al. / Biological Conservation 123 (2005) 477–488 483

1800
(a)
1600 kanaka
karambasan
1400
pasar paal

Individuals hour-1
1200 kawangkoan
langowan
1000
tomohon
800
600
400
200
0
800
(b)
700

600 Month
Individuals hour-1

500

400
R=0.594***
300

200

100

0
14
(c)
12

10
Individuals hour-1

8
R=-0.056NS
6

0
Jan 02
Feb 02
Dec 01

Jan 03
Feb 03
Mar 02
April 02
May 02
June 02

Aug 02
Sept 02

Nov 02
Dec 02

Mar 03
Apr 03
May 03
June 03

Aug 03
Sept 03

Nov 03
Dec 03
July 02

Oct 02

July 03

Oct 03

Month

Fig. 3. The encouter rate of mamallian wildlife in bushmeat markets in North Sulawesi monitored over 25 months in six different markets. All taxa
combined are depicted by individual markets in (a) and averaged across markets in (b). Protected taxa only are shown in (c). Error bars are one
standard error. R indicates strength of linear correlation with time.

and can be captured in very large numbers during a sin-
gle hunting trip. Although little is known about the
roosting patterns and colony size for most Sulawesi
bat species, colony size of other flying foxes in other
parts of Asia have reached up to 150,000 (see Pierson
and Rainey, 1992; Gumal, 2001).
4.2. Quantity of wild meat
Among the taxa representing the largest proportions
of animals traded in North Sulawesi markets in the last
two years, rats, bats, and Sulawesi pigs, we find special
cause for concern in large bats. This group, represented
in markets by only four species of flying foxes (Pteropus
hypomelanus, Pteropus alecto, Acerodon celebensis and
Acerodon humilis), is being exploited in astonishing
numbers: on average 445 individuals were counted per
hour across the six markets surveyed, representing
one-quarter of the wild meat markets operating in
North Sulawesi. At this rate, entire colonies can be
extinguished to supply demand for only a few days of
trade in these markets (Fujita and Tuttle, 1991).
SulawesiÕs flying foxes, animals with considerably
lower reproductive rates than similarly sized mammals
in other taxa, are unlikely to withstand such heavy hunt-
ing pressure without experiencing major population de-
clines (Pierson and Rainey, 1992; Racey and Entwistle,
2004). Evidence of such declines has already been noted
484 R.J. Lee et al. / Biological Conservation 123 (2005) 477–488

Table 2 Table 3
Two-way repeated measures ANOVA on the frequency of encounter- Two-way repeated measures ANOVA on the frequency of encounter-
ing mammalian wildlife (all species combined) in bushmeat markets in ing protected species of mammalian wildlife in bushmeat markets in
North Sulawesi North Sulawesi
Source df MS F p-Value Source df MS F p-Value
Within-subject effects Within-subject effects
Date Date
Greenhouse–Geisser 2.387 157931.12 1.327 0.292 Greenhouse–Geisser 1.890 175.10 2.419 0.125
Huynh–Feldt 4.747 79417.16 1.327 0.275 Huynh–Feldt 3.384 97.78 2.419 0.082
Date · year Date · year
Greenhouse–Geisser 2.387 153395.02 1.289 0.354 Greenhouse–Geisser 2.387 1.89 40.521 0.560
Huynh–Feldt 4.747 77136.12 1.289 0.290 Huynh–Feldt 4.747 3.38 22.628 0.560
Date · region Date · region
Greenhouse–Geisser 2.387 175030.50 1.471 0.255 Greenhouse–Geisser 1.890 20.80 0.287 0.742
Huynh–Feldt 4.747 88015.73 1.471 0.224 Huynh–Feldt 3.384 11.62 0.287 0.856
Date · year · region Date · year · region
Greenhouse–Geisser 2.387 94989.62 0.798 0.485 Greenhouse–Geisser 1.890 12.05 0.166 0.837
Huynh–Feldt 4.747 47766.42 0.798 0.552 Huynh–Feldt 3.384 6.73 0.166 0.934
Error (date) Error (date)
Greenhouse–Geisser 19.096 119001.86 Greenhouse–Geisser 15.118 72.40
Huynh–Feldt 37.975 59841.20 Huynh–Feldt 27.071 40.43
Between-subject effects Between-subject effects
Year 1 5123326.62 56.009 0.278 Year 1 7.034 0.120 0.738
Region 1 123888.75 1.354 0.001 Region 1 172.51 2.935 0.125
Year · region 1 2250250.01 24.600 0.221 Year · region 1 5.59 0.095 0.766
Error 8 161064.43 1.761 Error 8 58.77
Six markets in two regions were monitored over 24 months. Six markets in two regions were monitored over 24 months.
Greenhouse–Geisser and Huynh–Feldt adjusted probabilities are Greenhouse–Geisser and Huynh–Feldt adjusted probabilities are
reported for within-subject effects due to violations in the assumption reported for within-subject effects due to violations in the assumption
of sphericity. of sphericity.

in several Sulawesi locations. In the northern peninsula Table 4

of North Sulawesi, where flying foxes were once com- Two-way ANOVA to test for differences among months and across
years in the frequency of encountering of mammalian wildlife in
mon (Bergmans and Rozendaal, 1988), they are now ab- bushmeat markets in North Sulawesi
sent or occur in small numbers in Tangkoko-Duasudara
Source Total SS df MS F p-Value
Nature Reserve (Anon., 1981; Wildlife Conservation
Society, 1999, biological surveys and management (a) All taxa combined
Year 2.594 1 2.594 1.199 0.276
recommendations for Tangkoko Duasudara Nature Re- Month 5.707 11 0.519 0.240 0.994
serve, North Sulawesi, Indonesia in litt.), Manemobon- Year · month 6.271 11 0.570 0.263 0.991
embo Nature Reserve (Wildlife Conservation Society, Error 259.751 120 2.165
1999, biological surveys and management recommenda- (b) Protected
tions for Manembonembo nature reserve, North Sulaw- taxa combined
esi, Indonesia in litt.) and Gunung Ambang Nature Year 7.049 1 7.05 0.469 0.495
Reserve (Wildlife Conservation Society, 2000, biological Month 330.88 11 30.08 2.000 0.034
Year · month 76.54 11 6.96 0.463 0.923
surveys and management recommendations for Gunung
Error 1804.81 120 15.04
Ambang nature reserve, North Sulawesi, Indonesia in
Six markets in two regions were monitored monthly over 24 months.
litt.). Likewise, large colonies of flying foxes that used
Separate ANOVAs were performed for (a) all taxa combined and (b)
to occur in parts of Bogani Nani Wartabone National protected taxa combined.
Park in North Sulawesi and Gorontalo provinces are
no longer present (Walker pers. com.; Wildlife Conser-
vation Society, 2000, biological surveys and manage- 1994), Palau (Wiles et al., 1997), Niue Island in the South
ment recommendations for Bogani Nani Wartabone Pacific (Brooke and Tschapka, 2002), and Sarawak (Gu-
National Park, North Sulawesi, Indonesia in litt.) and mal, 2001), and suggest immediate action may be needed
populations of flying foxes in Talaud have plummeted to avoid similar losses in Sulawesi. The local disappear-
in recent years (Riley, 2002). ance of flying foxes will likely have major ecological im-
Local extinctions of flying foxes caused by hunting pacts, as these species or their close relatives have been
have been witnessed in American Samoa (Craig et al., shown to act as important seed dispersers and pollinators
 R.J. Lee et al. / Biological Conservation 123 (2005) 477–488 485

1.8 1.8
1.6 1.6 2002
Babyrussa Macaca 2003
1.4 1.4
p<0.001 p=0.081

Individuals hr-1
1.2 1.2
1.0 1.0
0.8 0.8
0.6 0.6
0.4 0.4
0.2 0.2
0.0 0.0

6 4

5 Cuscus Squirrels
3
p=0.410 p=0.069
Individuals hr-1

3 2

2
1
1

0 0

250 100

200 Large bats 80 Small bats

Individuals hr-1

p=0.066 p=0.897
60
150
40
100
20
50
0
0
30 250

25 Sulawesi pig Rats

200
p=0.418 p=0.262
Individuals hr-1

20
150
15
100
10

5 50

0 0
500 10

All species Protected species

400 8
p=0.232
Individuals hr-1

p=0.124
300 6

200 4

2
100

0
0
Jan Feb Mar Apr May June July Aug Sept Oct Nov Dec Jan Feb Mar Apr May June July Aug Sept Oct Nov Dec

Month Month

Fig. 4. Comparison between years of the encouter rate of mamallian wildlife in bushmeat markets in North Sulawesi. Data are averaged across six
markets for each month and compared between years using paired-sample t-tests. Solid line is 2002 and dashed line is 2003.

in lowland rain forest systems as well as agricultural
areas (Cox et al., 1991; Fujita and Tuttle, 1991; Howe,
1984; Cox and Elmqvist, 2000; Bollen and Van Elsacker,
2002; Meehan et al., 2002; Barclay and Harder, 2004).
Trade in the Sulawesi pig is alarmingly high for such a
large-bodied animal. On average, 8 individuals h 1
were observed during road blockades and 82 individuals
h 1 were recorded in the six markets surveyed (7% of all
market encounters). Furthermore, hunting pressure on
pigs can be highly localized: 250 pigs were counted in a
single truck originating from Buton Island, off the tip
of South Sulawesi, approximately 1270 km from Mana-
do by land and sea. Repeated localized harvests of this
scale would have severe impacts on local populations.
4.3. Origins of North SulawesiÕs wild meat
North Sulawesi wildlife markets are not supplied
wholly from local sources. Rather, wild meat is regularly
brought to markets in Minahasa and Manado from other
486 R.J. Lee et al. / Biological Conservation 123 (2005) 477–488
provinces several hundred kilometers away. This pattern
resembles others wherein local extirpations and high
market demands drive a very expansive trade network.
For example, hunters and traders across Indochina pro-
vide a steady volume of wildlife to supply VietnamÕs mar-
ket demands (Nash, 1997; Compton and Quang, 1998).
Differences in the taxonomic composition of animals
recorded in road blockades and those documented in
markets suggest that some animals, especially rats, orig-
inate locally. Rats comprised a relatively small percent-
age of animals detected during road blockades, but were
a large percentage of those seen in markets. If sampling
during road blockades and market surveys represents,
with reasonable accuracy, the taxonomic composition
of animals being transported and traded, this contrast
implies that rats are being harvested intensively within
North Sulawesi province. During surveys of all six mar-
kets combined, we encountered rats at an average rate of
430 individuals h 1 in 2002, and this increased to 580
individuals h 1 in 2003. If such enormous quantities of
rats are indeed being harvested locally, and if hunting
pressure remains at this level, then rats also are likely
to experience severe population declines in coming
years, despite relatively high reproductive rates. Declin-
ing populations of rats, which are important agents of
post-dispersal seed predation and dispersal in many rain
forest plant communities (Crawley, 1992; Terborgh et
al., 1993; Wright and Duber, 2001), would likely have
deleterious long-term impacts on plant community
dynamics, possibly leading to a gradual erosion of plant
species diversity in remaining fragments of North
Sulawesi rain forests.
In contrast to rats, it appears that most bats and pigs
are caught outside the province. This may reflect losses of
large mammals and flying foxes in many North Sulawesi
forests. During the 1980s and 1990s, hunting of large
mammals caused a precipitous decline, as much as 70%
in a six-year span and 95% in a 15-year span, in popula-
tions of these taxa throughout North Sulawesi (Lee,
2000a; OÕ Brien and Kinnaird, 2000). As a result of these
pressures, Lore Lindu National Park in Central Sulaw-
esi, a primary source of wildlife supplied to North Sulaw-
esi markets (Wildlife Conservation Society, 2000, report
on monitoring and control of the trade in protected wild-
life in North Sulawesi, Indonesia in litt.) may already be
experiencing wildlife declines (Waltert, pers. com.).
4.4. Geographic and seasonal variation in wildlife trade
An understanding of temporal and spatial variation
in wildlife sales in North Sulawesi markets provides use-
ful information to guide future Wildlife Crimes Unit ef-
forts. The index of wildlife trade is vastly higher in the
Minahasa region than in Manado, despite a larger hu-
man population in and more difficult access to the latter.
A probable explanation is that the Minahasa markets
supply meat to a large number of traditional restaurants
in the regionÕs uplands. These restaurants specialize in
wild meat, serving to weekend visitors from Manado
and other parts of Minahasa. Markets in this area also
provide meat to Minahasan people hosting social events
that typically feature wild meat. Future efforts to reduce
wild meat demand through awareness campaigns and
improved law enforcement should prioritize rural com-
munities in the Minahasa region.
In addition to regional differences, the data suggest a
clear pattern of seasonality in the trade activity of pro-
tected mammals, with peak sales in November. We are
unclear why the peak occurs around this time, as Novem-
ber falls between some of the larger cultural events in
Minahasa, including the harvest of agricultural crops
(panen) from September to October, and the Christmas
and New Year celebrations at yearÕs end. Whatever its
underlying causes, the peak in sales during November
implies that efforts to reduce demand through enforce-
ment and awareness campaigns should be intensified
during this period and include explicit reference to the
importance of avoiding trade in protected species and
non-protected taxa that are heavily harvested.
4.5. Future steps and recommendations to reduce wildlife
trade
This study took place during the pilot phase of the
Wildlife Crimes Unit program. Preliminary findings sug-
gest that road blockades are an effective enforcement
mechanism to reduce illegal wildlife trade. However, in
the long-term, this strategy may fail to preserve wildlife
populations, because it does not address the myriad
problems surrounding illegal hunting and trade, includ-
ing a paucity of data on wildlife populations, low aware-
ness of the problem, poor enforcement in protected areas,
and an unwieldy legal system. We recommend the follow-
ing more comprehensive set of actions be taken:
1. A nationwide Wildlife Crimes Unit Program should
be implemented to institutionalize and standardize
methods for the regulation and monitoring of wildlife
trade throughout Indonesia.
2. The national Wildlife Crimes Unit should establish a
system for regularly providing information on wild-
life hunting and trade to guide law enforcement
efforts.
3. Legal protection status should be assigned to several
species currently under threat. Based on the data pre-
sented above, these include the five flying foxes on
Sulawesi, P. hypomelanus, P. alecto, Pteropus cani-
ceps, A. celebensis and A. humilis.
4. A forensic center for wildlife crimes should be estab-
lished to enable the more accurate identification of
protected species in the wildlife trade, and perhaps
even the geographic origin of some animals.
 R.J. Lee et al. / Biological Conservation 123 (2005) 477–488
5. Laws that enable the prosecution of traders that
claim to sell protected wildlife should be formulated.
This strategy, by focusing on the intent of the vendor
to sell protected species rather than the actual selling
of a protected species, would lessen the need for time-
consuming and costly forensic work. Such an
approach is being used in Hong Kong, Sarawak State
in Malaysia and Africa (e.g., Gabon) (Bennett, pers.
com.).
6. Nationwide media and education campaigns should
be initiated to increase awareness among consumers
about the negative impacts of consuming wildlife spe-
cies and among hunters about the legal repercussions
of trading in protected species.
7. Ecological field studies should be conducted to: (1)
monitor fluctuations in wildlife populations inhabit-
ing source areas for wildlife trade; (2) describe the
reproductive biology of heavily harvested species
and identify limits of sustainable harvesting; and (3)
quantify the long-term effects of eliminating small
mammals, especially rats and bats, on the regenera-
tion dynamics of plant communities.
The people of North Sulawesi have high nutritional
standards and eat wild meat out of preference. Given
the current rapid wildlife declines, conservationists and
governments must find ways to work with people re-
align their consumption patterns so as to prevent extir-
pation of wildlife populations.
Acknowledgements
This study was supported by the East Asia Pacific
Environmental Initiative (a program supported through
the US State Department), the Wildlife Conservation
Society, and the Indonesian Department of Forestry.
We thank: Wildlife Conservation Society – Indonesia
Program, Sulawesi field staff (M.T. Soleman, B.S. Wi-
jaya, S. Ering, S. Mokodompit, M. Mahulette and M.
Mantiri); Department of Forestry field staff (Ir. Agus
Budiono, MSc, Ir. Dominggus); North Sulawesi Police
(Johnny H. Hutahuruk, Drs. A.P. Simanjuntak, S.H.
David Lembang, Laurens A. Wuisan); Konservasi Flora
dan Flora Sulawesi (Berty Pesik, Charles Bawode). Aslan
from the Wildlife Conservation Society – Indonesia Pro-
gram produced the map. We are grateful to Dr. Elizabeth
Bennett, Dr. David Wilkie, and anonymous reviewers
who provided insightful comments on the manuscript.
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