Biological Conservation 107 (2002) 155–164

www.elsevier.com/locate/biocon

Fragmentation and wildlife in montane evergreen forests, northern

Thailand
Anak Pattanavibool1, Philip Dearden*
Department of Geography, University of Victoria, Victoria, BC, Canada V8W 3P5

Received 29 November 2000; received in revised form 21 September 2001; accepted 5 November 2001

Abstract
Montane evergreen forests in northern Thailand are high in biodiversity and becoming increasingly fragmented. We studied
fragmentation and wildlife response in two contiguous wildlife sanctuaries. Om Koi still maintained relatively large patches ( >400
ha) with some connectivity while Mae Tuen was comprised mainly of small and isolated patches ( <100 ha). Mae Tuen lost 2640 ha
of montane evergreen forest between 1954 and 1996 compared to a loss of 888 ha in Om Koi. We compared the wildlife between
four forest patches in Mae Tuen, with four in Om Koi finding nine mammals and 89 birds in Mae Tuen and 19 mammals and 119
birds in Om Koi. Om Koi still supports populations of large mammals and frugivorous birds extirpated in Mae Tuen. The results
document the high rate of fragmentation in protected areas that often interact synergistically with other pressures to reduce biodi-
versity. Crown Copyright # 2002 Published by Elsevier Science Ltd. All rights reserved.
Keywords: Thailand; Birds; Mammals; Fragmentation; Hunting; Protected areas

1. Introduction 25% of the landscape in protected forest areas. Two

Tropical forest loss and degradation are among the
most important environmental issues now being faced
on Earth. Asia is of particular concern due to its higher
relative rates of deforestation and less remaining forest
(Laurance, 1999). Southeast Asia is a mega-biodiversity
region, yet as pointed out by Sodhi and Liow (2000)
there is a paucity of published research from this region
and yet deforestation rates are four times those found
elsewhere in the tropics (Laurance, 1999).
The situation in Thailand is symptomatic of the
region. In 1961, forests covered 53.3% of the land area.
By 1998, they covered only 25.28% of the land area, and
many of these forests were badly degraded (Rajani,
1999). The decline in forest area in northern Thailand
can be seen in Fig. 1. Thailand has now developed an
extensive protected area system, covering some 16%
of the land base. Government policy is to maintain
* Corresponding author. Tel.: +1-250-721-73327; fax: +1-250-
721-7326.
E-mail address: pdearden@office.geog.uvic.ca (P. Dearden).
1
Present address: Western Forest Complex Project, 84-Year Anni-
versary Building, Royal Forest Department, Bangkok, Thailand
10110.
0006-3207/02/$ - see front matter Crown Copyright # 2002 Published by Elsevier Science Ltd. All rights reserved.
PII: S0006-3207(02)00056-3
main categories of protected area forbid all extractive
use, national parks and wildlife sanctuaries. There are
currently 114 national parks, 75 of these are terrestrial
parks with a total land area of 4.31 million ha. Thirty-
nine proposed parks (2.03 million ha) are still in pro-
cess. There are also 47 wildlife sanctuaries with a total
area of 3.3 million ha. Another 12 wildlife sanctuaries
are proposed which would add 501,828 ha of protected
space (Rajani, 1999).
Despite these efforts many species in Thailand have
already been extirpated and others reduced to such low
abundance as to be considered ecologically extinct.
Schombergk’s deer (Cervus schomburgk) is now extinct, as
is probably the kouprey (Bos sauveli). Species such as the
Javan rhinoceros (Rhinoceros sondaicus), Sumatran
rhinoceros (Dicerorhinus sumatrensis) are no more to be
found in the wild. Small populations of brow-antlered
deer (Cervus eldi), and hog deer (Cervus porcinus anna-
miticus) were recently reintroduced after being long
extirpated. Of the 282 species of mammals about 40 are
classified as rare and endangered, and 190 of 916 bird
species, and 37 of 405 species of reptiles and amphibians
are threatened with extinction (OEPP, 1995). One of the
main problems is ongoing human impacts in the pro-
tected area system with fragmentation as a result of
156 A. Pattanavibool, P. Dearden / Biological Conservation 107 (2002) 155–164
Fig. 1. Deforestation in Thailand (1973–1998). Source: Royal Forest
Department statistics.
agricultural activities and hunting presenting major
challenges to management.
In northern Thailand, shifting cultivation together
with hunting and other forms of developments such as
roads and human settlements, commercial agriculture
and forestry have been cited as major causes of the
decline in biodiversity in montane forest areas (Dear-
den, 1996). The decline has been rapid. As late as the
1960s anthropologists, travelers, and missionaries were
still recording abundant wildlife in their travels (e.g.
Young, 1967; Shrock et al., 1970; Wongsprasert, 1975)
although there are few rigorous scientific studies from
this time (but see Bruver, 1973). Fragmentation has
been occurring in the area for centuries as a result of
traditional agricultural activities. However, over the last
30 years populations of ethnic hilltribe peoples have
more than quadrupled. Their traditional response to
declining agricultural and hunting yields, to move vil-
lages elsewhere to more fertile lands, is no longer possi-
ble. This has led to visually obvious increases in
fragmentation rates, an observation confirmed by quan-
titative measurement in one area by Fox et al. (1995).
Although previous studies in Thailand have examined
deforestation rates in national parks (e.g. Dearden et
al., 1996) and others, the responses of mammals to
fragmentation caused by flooding (Lynam and Billick,
1999), no studies have tried to understand the links
between a broad range of human impacts in fragmented
landscapes and wildlife response in a protected area.
This was the goal of this study.
Montane evergreen forest is the rarest evergreen for-
mation in Thailand and also one of the richest forma-
tions for biodiversity (Whitmore, 1984). Montane
species have also been found to be disproportionately
threatened both in Asia (e.g. Brooks et al., 1999) and
elsewhere (Renjifo et al., 1997). We wanted to find out
the rates of fragmentation of this forest type in a pro-
tected area and what impacts this might be having on
wildlife. Although aerial photographs and satellite ima-
ges allow comparison between past and present forest
extent, there is no data available on the original wildlife
diversity associated with these past conditions. One way
to surmount this problem is to select analogue sites
where conditions are similar but differ in extent of
human impact. This was the approach taken in this
study. We selected two comparable areas of evergreen
montane forest that were subject to different degrees of
human interference and compared fragmentation extent
and rates and wildlife response in these habitats. Two
contiguous wildlife sanctuaries were selected and these
will be discussed in more detail below.
2. Study area
Om Koi and Mae Tuen Wildlife Sanctuaries, estab-
lished in 1978, and together having a total area of 2397
sq. km were selected for study (Fig. 2). Reports (e.g.
Thailand Forest Research Center, 1991, 1992) and
reconnaissance field work indicated that Om Koi (OK)
possessed one of the most intact evergreen forest com-
munities in Northern Thailand and could be considered
a ‘‘hotspot’’ for biodiversity within the context of
Northern Thailand. Om Koi is home to the only herd
of wild elephants (Elephas maximus) remaining totally
within Northern Thailand. It is also home to the last
remaining herd of banteng (Bos javanicus), the largest
population of goral (Naemorhedus goral ), and other
endangered species such as tiger (Panthera tigris) and
leopard (Panthera pardus). Gaur (Bos gaurus) appear to
be the only large species that has been extirpated in
recent times. Their past presence is witnessed by trophies
on the walls of village houses around the sanctuary.
Contiguous with OK is another wildlife sanctuary,
Mae Tuen (MT), which possesses very similar ecological
characteristics, including areas of evergreen montane
forest with dominant tree species from the Fagaceae,
Lauraceae, and some upper Dipterocarpaceae families.
Reports, however, suggested that human impacts in MT
had increased dramatically, and the fauna in MT was
on the decline. Preliminary inspection of satellite images
and aerial photos coupled with field visits suggested that
the evergreen forests were very similar to those of OK in
all major aspects except human impacts. Salient char-
acteristics of both sanctuaries are shown in Table 1. It
should be noted that both precipitation stations are at
lower elevations and we estimate rainfall to be at least
double at the higher elevation of the montane forests.
3. Methodology
3.1. Landscape structure and change
We constructed digital maps of forest type and land
use of the sanctuaries through visual interpretation of
LANDSATTM imagery taken on 15 February 1996 at a
 A. Pattanavibool, P. Dearden / Biological Conservation 107 (2002) 155–164 157

Fig. 2. Location of Om Koi, Mae Tuen and other protected areas in northern Thailand.

Table 1 We selected areas of approximately the same size that

Characteristics of Om Koi and Mae Tuen wildlife sanctuaries, north- encompassed the highest elevation sites in each sanc-
ern Thailand
tuary as our sample landscapes for more detailed ana-
Characteristics Om Koi Mae Tuen lysis. Aerial photographs taken in 1996 at the 1:50,000
scale were used to build more sensitive photomosaics
Size 1224 km2 1173 km2
Elevation 200–1929 m (MSL) 200–1664 m (MSL) for each of these landscapes. Photographs were scan-
Avg. Precipitation 1060 mm/year 1926 mm/year ned, interpreted and rectified with PCI remote sensing
Number of villages 11 (1993) 17 (1992) software and mosaics constructed and then digitized
Populations
2520 (1993)
3200 (1992)
using ARCVIEW. Due to the high relative relief
Ethnic group Dominantly Karen, Dominantly Hmong
Some Lahu & Karen between 20 and 40 ground control points were used for
each image along with a thin plate spline rectification
model (PCI, 1997). We used ‘‘FRAGSTAT version 2.0’’
1:50,000 scale. Ground checks were conducted along (McGarigal and Marks, 1995) to analyze the spatial
100 km of roads and trails during field surveys. Classi- configurations of these landscapes and the sample pat-
fication of vegetation types was based mainly on those ches selected and compare between the oldest air photo
suggested by the Thailand Forest Research Center records available (1954) and the 1996 record. Total core
(1992) and Juntakat 1999. Differences in forest types area for patches was defined by eliminating a 100 m
can be clearly identified on the ground by the difference wide buffer around the perimeter of each patch.
in dominant tree and other plant species. However, a
composite montane evergreen forest classification 3.2. Bird and mammal diversity and abundance
(Whitmore, 1984) was preferred over separate hill and
dry evergreen forests classifications (Thailand Forest Within both of the sample areas described above we
Research Center, 1991, 1992) as the two were hardly selected four patches of evergreen forest to represent the
distinguishable on the satellite image. An additional range of size and disturbance classes found within each
classification was also added to represent the clearly sanctuary. Within each patch two 500 m transects were
distinguishable disturbed montane evergreen sites. In established, one in the edge zone, defined as being
total, forest types and land use were classified into six within 100 m from the forest edge (McGarigal and
types: montane evergreen forest, open and disturbed McComb, 1995), and the other within the interior. Edge
montane forest, mixed deciduous forest, dry dipterocarp zone transects were further divided into five 100 m lines
forest, old clearing, and crop field. The maps produced and located perpendicular to the patch edge at 200–300
were used to identify the remaining tracts of montane m intervals. These intervals reduce the chance of count-
evergreen forest in each sanctuary. ing the same birds twice during sampling. Each transect
158 A. Pattanavibool, P. Dearden / Biological Conservation 107 (2002) 155–164

was surveyed seven times during site visits spanning
from September 1997 to June 1998.
Bird surveys focused on diurnal species and were
undertaken between 07:00–11:00 and 14:00–17:00, when
birds are most active, on each transect by the same
observer when no appreciable rainfall was occurring.
Using the same observer standardized skill and effort
variability in bird identification (Bibby et al., 1992).
Bird species, number of individuals, and perpendicular
distance from the transect lines were recorded. The
software ‘‘DISTANCE version 2.0’’ (Laake et al., 1993)
was used for analysis of bird density (birds/ha). Species
abundance was indicated as ‘‘birds/site visit’’ as the
amount of data for individual species was not enough
for DISTANCE to produce reliable results. t-Tests were
used to test the null hypotheses that there were no sig-
nificant differences between species diversity and density
in: (1) forest patches with high and low rates of frag-
mentation/human disturbances, and (2) edges and
interior zones. The rate of fragmentation/human dis-
turbances was defined as the change in forest areas,
sizes, shapes, and other forms of developments (e.g.
roads) over time across the studied landscapes.
We recorded all large and medium sized mammals
encountered during the bird transect surveys. However,
the main data source for mammals was track counts.
Track counts have been used as indices of abundance
for species that are difficult to observe on the assump-
tion that track counts are related to population size in a
linear, or at least a monotonic, fashion (Wemmer et al.,
1996). In this study, ‘‘track recording stations’’, areas
51 m on the ground where litter was cleared and soil
tilled to create a soft zone for recording foot prints,
were established at 20–25 m intervals for every 100 m
along the transect. There were 40 stations in each patch.
Each was re-tilled after the tracks had been recorded. T-
tests were used to test the null hypothesis that there was
no significant difference of track abundance in each
mammal species between high and low fragmented/dis-
turbed landscapes.
In addition to the wildlife information recorded above
comprehensive field notes were also kept on any signs of
human activities, such as hunting (carcasses, poaching
platforms, poachers, gun shots etc), burning and cattle
grazing and the sites of these disturbances recorded by
GPS. This evidence helped construct a more qualitative
understanding and is used to supplement our quantita-
tive data in the discussion.
4. Results
4.1. Landscape structure and configuration
Table 2 shows the comparison between the two mon-
tane evergreen landscapes selected for more detailed
study in the two sanctuaries. The two landscapes are
approximately the same size (20,481ha OK, 18,530 MT)
and do not differ appreciably in the total amount of
evergreen forest remaining in 1996 with 3403 ha, in OK,
and 2475 in MT. However, they do differ in two
important respects. The first is the change over time.
Between 1954 and 1996, OK lost 5% of its montane
evergreen and MT 50%. Secondly, not only did MT
lose about 10 times as much forest, the remaining forest
is also much more highly fragmented than OK. One
critical indicator of the difference in fragmentation
between the sanctuaries is the largest patch index (LPI)
representing the percentage of the landscape occupied
by the largest patch. In OK this comprises almost 17%
of the landscape, but in MT only 4%. In 1954 the cor-
responding figure for MT was 23%. There are a larger
number of smaller patches of more uniform size in MT
than in OK. The total core area is also significantly
lower in MT, with only 35% of its extent in 1954,
whereas OK retained some 86% of total core area
since 1954.
4.2. Birds
Overall we made 2430 sightings of 149 species, with
1238 sightings of 89 species in Mae Tuen patches and
1192 sightings of 119 species in Om Koi (Table 3). The
mean number of species in OK per patch was 63.25

Table 2
Comparisons of montane evergreen forest change between 1954 and 1996 in Om Koi and Mae Tuen Wildlife Sanctuaries

Index Om Koi Mae Tuen

1954 1996 1954 1996

Total landscape area (ha) 20,481 20,481 18,530 18,530

Total area of remaining montane 3567 3403 4974 2475
evergreen forest (ha)
Largest patch index (%) 15.35 16.62 23.14 3.94
Number of patches 20 44 6 43
Mean patch size (ha) 178.35 77.36 829.15 57.57
Patch size SD (ha) 718.28 377.77 1,821.18 132.07
Total core area (ha) 1884.01 1623.26 2545.90 890.88
Mean core area (ha) 94.20 36.89 424.32 20.72
Core area SD (ha) 408.62 209.73 948.38 297.26
 A. Pattanavibool, P. Dearden / Biological Conservation 107 (2002) 155–164 159

(SE=3.20) with 54 species in Mae Tuen (SE=2.48).
This difference was not statistically significant and there
were no significant differences in bird species density
between patches in OK and MT or between richness
and patch size.
Abundance was calculated as birds/site visit and
based only on species found on transects to ensure the
same search efforts for both sites. Overall almost half of
the bird species at MT (40 out of 89) were found at an
abundance of < 1 bird/site visit while in OK this was
about one third of the species (42 from 119). Table 4
shows the five most abundant species at each site. The
five most abundant species in OK are all montane ever-
green forest obligates (Lekagul and Round, 1991). In
MT, however, two generalist species, the black bulbul
(Hypsipetes madagascariensis) and streaked spider-
hunter (Arachnothera magna) were found among the
five most abundant species. Large frugivorous birds
such as the brown hornbill (Ptilolaemus tickelli), and
great hornbill (Buceros bicornis) still existed in very low
abundance (1.43 and 0.43 birds/site visit respectively) in
OK but neither was found in MT where they appear to
have been extirpated, as they have throughout almost
all the rest of northern Thailand (Poonswad and Kemp,
1993; Vidhidharm et al., 1995).
In OK four species of ground insectivores, the slaty-
bellied tesia (Tesia olivea), pygmy wren-babbler (Pnoe-
pyga pusilla), streaked wren-babbler (Napothera brevi-
caudata), and dark-sided thrush (Zoothera marginata)
were found in interior zones, but only one, the dark-
sided thrush was found in the edge zone. These species
were absent from MT, presumably due to the lack of
interior zone in the smaller patches. Ground omnivores
in the Pheasant family such as the Kalij pheasant
(Lophura leucomelana) and rufous-throated partridge
(Arborophila rufogularis) were found in OK patches
with an abundance of 1 and 3 birds/site visit, respec-
tively. However, only the bar-backed partridge (Arbor-
ophila brunneopectus) with the same size and feeding
guild as the rufous-throated partridge was found, and
then with very low abundance (0.57 bird/site visit) in MT.
In terms of feeding guilds the mean abundance of
nectarivorous birds using MT patches was significantly
higher than in OK (Table 5). The streaked spiderhunter
(8 birds/site visit) and little spiderhunter (4.71 birds/site
visit) were the most abundant nectarivores in Mae Tuen
patches. Nectarivores in OK were more diverse but found
in lower abundance, with the chestnut-flanked white-eye
(3.57) and streaked spiderhunter (1.29) ranking first and
second. None of the other feeding guilds showed any
significant difference between the sites (Table 5).
4.3. Mammals
We found a total of 20 species, nine species in Mae
Tuen and 19 in OK (Table 6). OK fragments support large
mammals such as elephant (Elephas maximus), Asiatic
black bear (Selenartos thibetanus), leopard (Panthera
pardus), tiger (Panthera tigris), sambar (Cervus unicolor),
and primates such as assamese macaque (Macaca
assamensis), Phayre’s langur (Presbytis phayrei), and
white-handed gibbon (Hylobates lar). Except for the

Table 3
Bird and mammal diversity, and bird density in different forest patches in Om Koi (OK) and Mae Tuen (MT)

Characteristic Patch data

MT OK

P1 P2 P3 P4 P5 P6 P7 P8

Area (ha) 38 29 89 85 96 796 27 462

Bird count 289 304 340 305 310 356 241 285
Bird species 50 54 51 61 68 64 54 67
Bird density (birds/ha) 9.51 8.67 8.92 8.10 5.92 7.89 9.06 4.75

Mammal species 7 2 6 5 9 18 7 10

Table 4
Five most abundant species (birds/site visit) in Om Koi and Mae Tuen wildlife sanctuaries

Om Koi Mae Tuen

Species Birds/site visit Species Birds/site visit

Gray-cheeked fulvetta (Alcippe morrisonia) 11.00 Gray-cheeded fulvetta (Alcippe morrisonia) 16.29
Mountain bulbul (Hypsipetes mcclellandii) 11.00 Black bulbul (Hypsipetes madagascariensis) 12.29
Golden-throated barbet (Megalaima franklinii) 10.86 Mountain bulbul (H. mcclellandii) 10.86
White-tailed leaf-warbler (Phylloscopus davisoni) 8.71 Streaked spiderhunter (Arachnothera magna) 8.00
Black-headed sibia (Heterophasia melanoleuca) 7.57 White-throated bulbul (Criniger flaveolus) 8.00
160 A. Pattanavibool, P. Dearden / Biological Conservation 107 (2002) 155–164

white-handed gibbon none of these animals were found tracks per site visit for all species combined was sig-
in MT. We found smaller size mammals such as the nificantly higher (t=2.74, 12 df, P=0.018) in OK
black giant squirrel (Ratufa bicolor), barking deer (x
=271.57 tracks/site visit, SE=46.48) than in MT
(Muntiacus muntjak), wild pig (Sus scrofa) and hog (x
=123.43 tracks/site visit, SE=27.78). Barking deer
badger (Arctonyx collaris) at both sites. Although fre- and wild pig tracks came first and second respectively in
quencies of observations were not high enough to abundance at both sites. In OK, elephants were the
undertake statistical tests, the relationship between third most abundant track. Barking deer were sig-
patch size and mammal diversity can be seen in Table 3. nificantly more abundant in OK than MT (t=2.21, 12
Smaller patches show little relationship between size df, P=0.047). No tigers or leopards were recorded at
and diversity, but as patch sizes get larger they provide the track stations.
suitable habitat for an increasing number of species.
Almost twice as many species (18) were found in the
largest patch (P6, 796 ha) when compared with the next 5. Discussion
largest patch (10 species in P8, 462 ha).
5.1. Landscape and patch configuration
4.3.1. Mammal track abundance
The total number of tracks for all species was 886 in These results clearly show that even within the pro-
MT and 2016 in OK (Table 7). The mean number of tected area system the amount of montane evergreen
forest has been severely degraded as a result of agri-
Table 5 cultural activities. Within the selected landscapes in the
Comparisons on bird abundance by feeding guild in montane two sanctuaries, OK lost 5% and MT an astonishing
evergreen forest patches between Om Koi and Mae Tuen wildlife 50% between 1954 and 1996. MT lost more than 10
sanctuaries
times as much montane evergreen as did OK over
Feeding guild Om Koi Mae Tuen t df P Powera this time period. Fragmentation statistics show MT to
x
SE x
SE have a larger number of smaller patches remaining with
a total core area of only 5% of the landscape, almost
Nectarivore 1.68 0.41 3.64 0.31 3.82 6 0.009* –
Frugivore 10.71 1.68 16.00 1.99 2.03 6 0.088 0.53 half of the 8% found in OK.
Insectivore 29.25 2.34 21.79 2.11 2.34 6 0.058 0.65 Five related causes probably underlie the more rapid
Omnivore 2.10 0.64 2.11 0.50 0.38 6 0.719 0.07 fragmentation in MT. First, there is a paved road run-
a
Statistical power at /=0.05. ning through the south part of the sanctuary that pro-
* Significant difference. vides ready access both for older established villages

Table 6
List of mammal species found in Om Koi and Mae Tuen wildlife sanctuariesa

Species Mammal species in Mae Tuen Mammal species in Om Koi

1. Assamese macaque (Macaca assamensis) N/F F

2. Phayre’s langur (Presbytis phayrei) N/F F
3. White-handed gibbon (Hylobates lar) F F
4. Black giant squirrel (Ratufa bicolor) F F
5. Malayan porcupine (Hystrix brachyura) F F
6. Asiatic black bear (Selenartos thibetanus) N/F F
7. Hog badger (Arctonyx collaris) F F
8. Siberian weasel (Mustela sibirica) F N/F
9. Yellow-throated marten (Martes flavigula) N/F F
10. Crab-eacting mongoose (Harpestes urva) N/F F
11. Civet (unknown) F F
12. Leopard (Panthera pardus) N/F F
13. Tiger (Panthera tigris) N/F F
14. Cat (Felis spp.) F F
15. Asiatic elephant (Elephas maximus) N/F F
16. Wild pig (Sus scrofa) F F
17. Common barking deer (Muntiacus muntjak) F F
18. Sambar (Cervus unicolor) N/F F
19. Serow (Capricornis sumatraensis) N/F F
20. Goral (Naemorhedus goral) N/F F

Total found 9 19
a
F, Found; N/F, not found.
 A. Pattanavibool, P. Dearden / Biological Conservation 107 (2002) 155–164
Table 7
Relative abundance (average track counts per site visit) of mammal tracks in Mae Tuen and Om Koi, northern Thailand
Species
Overall spp
Malayan porcupine (Hystrix brachyuca)
Asiatic black bear (Selenarctos thibetanus)
Hog badger (Arctonyx collaris)
Civets in Family Viverridae
Felis spp.
Asiatic elephant (Elephas maximus)
Wild pig (Sus scrofa)
Common barking deer (Muntiacus muntjack)
Sambar (Cervus unicolor)
Goral (Naemorhedus goral)
a
Number of site visits.
b
No testing because of not enough data.
to get goods out to market and also for new settlers to
enter the sanctuary. There are no paved roads through
OK. Second, the accessibility of MT has made the area
prime growing land for temperate crops, particularly
cabbages, which are sent to Bangkok. Large profits can
be generated by such cash crops (e.g. see Tungittipla-
korn, 1998). In the 1997/1998 field season we saw many
new pick-up trucks as the market price for cabbage was
high. Fox et al. (1995) also note the huge impact that a
new paved road had on opening up villages to markets
in northern Thailand. Third, MT has become increas-
ingly settled by the Hmong ethnic group, who are well
known for their skill in cash crop cultivation. Large
numbers of illegal and poorly paid immigrants from
Burma can be witnessed during harvest time carrying
the cabbages to the waiting trucks of the Hmong. In
contrast, OK is occupied dominantly by the more
sedentary and longer established Karen ethnic group,
who have shown less interest in clearing the forests they
depend on to grow cash crops for distant markets. The
only cabbage growers in OK are the Lahu ethnic group.
Most of the ongoing fragmentation is from the expan-
sion of their fields. Fourth, human population numbers
are also higher in MT than OK, although little is known
of growth rates. Fifth, sanctuary authorities have been
more vigilant in OK in their efforts to control defor-
estation and other violations. Such vigilance in MT was
arrested when an officer was shot and killed at his guard
post. His surviving colleagues have been less vigilant
since that time.
As a result of the interaction among the factors men-
tioned above there are significant differences between
the two sanctuaries in the extent and rate of fragmenta-
tion. OK has retained a higher proportion of montane
evergreen forest, there are more large patches and the
average size of patches is larger with more core area
retained as a consequence.
However, difficulties still remain. All sites are unique
and human impacts are multi-dimensional. We have
documented the increased fragmentation in MT com-
161
Maetuen N=7a Omkoi N=7 T-test
123.43 271.57 0.018
0.29 2.43 n/ab
0 2.14 n/a
0.14 0.57 n/a
4.14 3.71 0.710
0.71 1.57 n/a
0 34.57 n/a
66.29 95.43 0.397
51.86 118.86 0.047
0 11.43 n/a
0 0.86 n/a
pared with OK over the last 40 years, but it is difficult to
assess other factors such as hunting impacts in any rig-
orous manner. Hence the following discussion is based
on both our quantitative results supplemented by
observations on human impacts and interviews with
local villagers and sanctuary guards.
5.2. Bird response
Bird species richness was higher in OK than MT.
Several factors could be responsible for this. Patch size
is greater in OK and several studies elsewhere (e.g.
Bierregaard et al., 1992; Kattan and Alvarez-Lopez,
1996, Cornelius et al., 2000) have shown relationships
between patch size and bird species richness. However,
no statistically significant relationships between patch
size and bird species richness, or density were detected.
Instead changes in the relative abundance between birds
with different habitat requirements were noted, with an
increase in nectarivorous species in the smaller patches
of MT and a decline in frugivorous species such as the
brown and great hornbills. Large frugivorous birds
require continuous habitat along altitudinal gradients
because fruit availability is variable in time and space,
and tracking these resources involves seasonal move-
ments that cover large areas. Forest fragmentation
separates the connection between foraging areas and
may severely restrict access to a year-round food supply
(Guindon, 1996). Research elsewhere has also suggested
that frugivorous birds might be more susceptible to
fragmentation and hence suitable species for indicating
minimum area requirements for conservation. Price et
al. (1999), for example found that pied imperial pigeons
were especially vulnerable to forest loss. Interestingly, in
our study, mountain imperial pigeons (Ducula badia)
were the eighth most abundant species in OK but were
not recorded in the heavily fragmented MT.
Forest destruction also reduces the number of large
trees which hornbills require for nesting. Reduction of
nesting sites can lead to change in nest competition and
162 A. Pattanavibool, P. Dearden / Biological Conservation 107 (2002) 155–164
reproductive success (Poonswad and Kemp, 1993).
Large frugivorous birds including three species of
hornbills were eliminated from Hong Kong and Singa-
pore by fragmentation and hunting (Corlett and Turner,
1997). Hornbills in this study have also suffered from
hunting. Casques of the Great hornbill (Buceros bicor-
nis) were commonly seen in hilltribe villages. Vidhid-
harm et al. (1995) reported that the species had been
extirpated from the central area of northern Thailand. It
remains to be seen whether this small relict population
can survive.
Another factor is the nature of the clearings in the
two sanctuaries. In OK, the fields tend to be abandoned
and in various stages of regrowth whereas in MT they
tend to be active fields dominated by cabbage growing.
The lower contrast between habitat types in OK could
help maintain the diversity of birds. Bierregaard and
Stouffer (1997) describe a number of primary rain forest
birds in an Amazonian forest foraging in adjacent sec-
ondary forest, and using secondary forest to re-colonize
small primary forest fragments nearby. Some evergreen
forest species, such as the yellow-cheeked tit (Parus spi-
lonotus), and black-headed sibia (Heterophasia melano-
leuca), used the regrowth along the patch edges in OK.
Some primary forest specialists may avoid the matrix
of modified habitats, while others will adapt to the
changed landscape (Newmark, 1991; Laurance and
Bierregaard, 1997). However, no primary forest birds
were found in the cabbage fields, and clearing species,
such as the flavescent (Pycnonotus flavescens), and red-
whiskered bulbuls (Pycnonotus jocosus), were never
found in the patches far from the edge zones at either
site. Lovejoy et al. (1986) also found that very few sec-
ond growth bird species invaded tropical forest patches
in the Amazon.
The abundance of nectarivorous birds in MT patches
was striking and may be related to the abundance of
wild bananas surrounding most patches. These appear
to be one of the main sources of nectar for the birds,
and are not as abundant in OK. Nectarivores such as
hummingbirds generally prove to be less vulnerable to
fragmentation than insectivores and frugivores (Bierre-
gaard and Stouffer, 1997).
Hunting appears to be a greater threat in MT rather
than OK. We recorded over double the number of gun-
shots in MT as OK, encountered several poaching par-
ties in the former and also came across more carcasses
including the feathers of bird species such as barbets.
5.3. Mammal response
The conspicuously low numbers of mammal species
in MT patches compared to OK are probably a result
of the combined pressures of fragmentation and hunt-
ing. Large carnivores such as tiger (Panthera tigris),
Asiatic black bear (Selenartos thibetanus), and leopard
(Panthera pardus), and herbivores such as elephants
(Elephas maximus) and sambars (Cervus unicolor) were
not found in MT. Large mammals, especially top pred-
ators like tigers and leopards, normally have a large
home range. A single female tiger, for example, requires
at least 2000 ha and the male probably at least twice as
large (Sunquist, 1981). Home range sizes of Asian ele-
phants normally range between 3400 to 80,000 ha
(Stuwe et al., 1998). The fragmented forests of Mae
Tuen, with their intensive agricultural development may
be too small to support such animals. Laidlaw (2000),
working in protected areas in Malaysia, found that the
size of the remaining area of natural forest was the most
important factor affecting the nature of mammal com-
munities.
Hunting is another factor behind local extirpations.
All the ethnic hilltribes have a long history of hunting as
both an important source of food and also a part of
their culture and beliefs (e.g. see Garrett, 1926; Bruver,
1973; Dearden, 1995). As populations have increased,
villages have become sedentary and firearms more
effective, there has been simply too great a hunting
pressure for wild populations to be sustained. During
the course of fieldwork numerous poaching platforms
were found as were the carcasses of two banteng (Bos
javanicus), 25% of the herd. Development of the paved
road in MT has also contributed to species extirpations
through both disturbance and the access it provides for
poaching. In highly developed areas such as Singapore,
extirpations of large carnivores were caused mainly by
hunting (Corlett and Turner, 1997).
Tracks of a male tiger were found at high elevation in
OK, which is unusual habitat for tigers as mountainous
areas with severe seasonal fluctuations are less optimal
habitats (Rabinowitz, 1993). The presence of the tiger
could be an indicator of the availability of preferred
prey species such as sambar, which still exist in OK or it
could also mean that the tiger is avoiding more pre-
ferred habitat at lower elevations where human settle-
ments and encroachment are rampant. Rabinowitz
(1993) suggests that hunting, human settlements and
forest degradation from agricultural clearing are the
main threats to tiger populations in Thailand. All these
threats are more severe in MT patches where no tigers
were found.
Patch size and mammal diversity seem to be related in
OK but no such trend emerged in MT. The lack of
relationship in MT is probably because the intensity
of human activities overwhelms the effects of patch size
and diversity. As mentioned earlier, many authors (e.g.
Warburton, 1997; Kattan and Alvarez-Lopez, 1996;
Bierregaard and Lovejoy, 1989; Chiarello, 2000) have
reported positive relationship between patch size and
animal diversity. Greater diversity in larger patches can
be viewed as the result of having more habitat diversity
(Buckley, 1982) or larger area per se (Simberloff, 1976).
 A. Pattanavibool, P. Dearden / Biological Conservation 107 (2002) 155–164
6. Conclusions
From this study we can conclude that forest frag-
mentation is occurring at a high rate even within the
protected area system. There is need for an overall sur-
vey of fragmentation in the protected area system in
Thailand. Within the montane evergreen formation
in our sample landscapes, Om Koi has larger patches
and the greatest richness of mammal and bird species.
In Mae Tuen where human impacts are greater with
smaller remnant patches, more highly modified land-
scape matrix, greater hunting pressure and road dis-
turbance, then wildlife response is more extreme. Bird
populations show a change in species composition with
the elimination of more sensitive forest obligates and
large frugivores. Mammal diversity is also affected with
a marked decline in the numbers of species, particularly
large species, and also a reduced abundance of more
common species. It is difficult to determine the domin-
ant factor causing these changes for, as suggested by
other researchers (e.g. Turner and Corlett, 1996; Cullen
et al., 2000), disturbance factors tend to work together
and often synergistically.
Patch size is undoubtedly important for some species.
Given that only the largest patch sampled (and, inci-
dentally, the largest remaining) had a full complement
of species, we suggest that management attempt to
recreate substantial connectivity between the smaller
patches through restoration activities, to approximate
the size of this largest patch, about 800 ha.
One important factor influencing both bird and
mammal responses is probably the nature of the land-
scape matrix. This is difficult to distinguish from the
effects of patch size as Om Koi not only has larger pat-
ches, but also a more inactive matrix with secondary
growth occurring, in contrast to the active agricultural
nature of the matrix in Mae Tuen. However, other
research on fragmentation effects (e.g. Gascon et al.,
1999; Umapathy and Kumar, 2000) has also suggested
that isolation-mediated habitat variation rather than
just patch size is an important determinant of species
survival. An important implication for management in
the wildlife sanctuaries following from this is the need to
place more value on protecting matrix habitat. At the
moment, disturbed areas are looked at more as man-
agement failures rather than important allies for con-
servation purposes. Fire, agricultural and grazing
activities in matrix habitat are considered minor man-
agement irritations and receive little attention.
A similar concern arises at a larger scale. The research
confirms the importance of Om Koi as a site of regional
significance for wildlife and a potential source for the
recolonization many of the faunistically impoverished
protected areas in northern Thailand. However, this will
require not only strong protection for these populations
but also a greater degree of habitat connectivity between
163
the protected areas in the region. The fragmentation
and other disturbance processes at work in northern
Thailand are ubiquitous throughout southeast Asia and
are resulting in the same faunistically impoverished
results in surrounding countries. Assessing remaining
‘‘hotspots’’ such as Om Koi Wildlife Sanctuary, pro-
viding management with adequate resources to ensure
the security of wildlife and planning for the reconnec-
tion of these key sites within the region is essential to the
future survival of the rich wildlife heritage of southeast
Asia.
Acknowledgements
The author would like to acknowledge funding
provided by SSHRC, Thai Biodiversity Fund and
Earthwatch. The Study could not have taken place
without the generous cooperation of the Royal Forest
Department and its Staff. Comments by two anony-
mous reviewers were very helpful in improving the
manuscript.
References
Bibby, C.J., Burgess, N.D., Hill, D.A., 1992. Bird Census Techniques.
Academy Press, London.
Bierregaard Jr., R.O., Lovejoy, T.E., 1989. Effects of forest fragmen-
tation on Amazonian understory bird communities. Acta Amazo-
nica 19, 215–241.
Bierregaard Jr., R.O., Lovejoy, T.E., Kapos, V., dos Santos, A.A.,
Hutchings, R.W., 1992. The biological dynamics of tropical rain-
forest fragments. BioScience 42, 859–866.
Bierregaard Jr., R.O., Stouffer, P.C., 1997. Understory birds and
dynamic habitat mosaics in amazonian rain forests. In: Laurance,
W.F., Bierregaard Jr., R.O. (Eds.), Tropical Forest Remnants:
Ecology, management, and conservation of fragmented commu-
nities. University of Chicago Press, Chicago, pp. 138–155.
Brooks, T.M., Pimm, S.L., Kapos, V., Ravilious, C.A., 1999. Threat
from deforestation to montane and lowland birds and mammals in
insular south-east Asia. Journal of Animal Ecology 68, 1061–1078.
Bruver, W., 1973. Fauna of north Thailand. Natural History Bulletin
of Siam Society 24, 463–466.
Buckley, R., 1982. The habitat model of island biogeography. Journal
of Biogeography 9, 334–339.
Chiarello, A.G., 2000. Density and population size of mammals in
remnants of Brazilian Atlantic forest. Conservation Biology 14,
1649–1657.
Corlett, R.T., Turner, I.M., 1997. Long-term survival in tropical forest
remnants in Singapore and Hong Kong. In: Laurance, W.F., Bier-
regaard Jr., R.O. (Eds.), Tropical Forest Remnants: Ecology, Man-
agement, and Conservation of Fragmented Communities.
University of Chicago Press, Chicago, pp. 333–345.
Cornelius, C., Cofre, H., Marquet, P.A., 2000. Effects of habitat frag-
mentation in a relict temperate forest in semiarid Chile. Conserva-
tion Biology 14, 534–543.
Cullen Jr., L., Bodmer, R.E., Padua, C.V., 2000. Effects of hunting in
habitat fragments of the Atlantic forests, Brazil. Biological Con-
servation 95, 49–56.
Dearden, P., 1995. Development and biocultural diversity in northern
Thailand. Applied Geography 15, 325–340.
164 A. Pattanavibool, P. Dearden / Biological Conservation 107 (2002) 155–164
Dearden, P., 1996. Development, the environment and social differ-
entiation in northern Thailand. In: Rigg, J. (Ed.), Counting the
Costs: Environmental Growth and Economic Change. ISEAS, Sin-
gapore, pp. 111–130.
Dearden, P., Chettamart, S., Emphandu, D., Tanakanjana, N., 1996.
National parks and hill tribes in northern Thailand: a case study of
Doi Inthanon. Society and Natural Resources 9, 125–141.
Fox, J., Krummel, J., Yarnasarn, S., Ekasingh, M., Podger, N., 1995.
Land use and landscape dynamics in northern Thailand: assessing
change in three upland watersheds. Ambio 24, 328–334.
Garrett, H.B., 1926. Notes on some traps made by the hill peoples of
Siam. Natural History Bulletin of Siam Society 9, 23–24.
Gascon, C., Lovejoy, T.E., Bierregaard Jr., R.O., Malcolm, J.R.,
Stouffer, P.C., Vasconcelos, H.L., Laurance, W.F., Zimmerman, B.,
Tocher, M., Borges, S., 1999. Matrix habitat and species richness in
tropical forest remnants. Biological Conservation 91, 223–229.
Guindon, C.F., 1996. The importance of forest fragments to the
maintenance of regional biodiversity in Costa Rica. In: Schelhas, J.,
Greenberg, R. (Eds.), Forest Patches in Tropical Landscapes. Island
Press, Washington, DC, pp. 168–186.
Juntakat, Y., 1999. The Study of Land Utilization Trend Analysis Using
Satellite Data: a Case Study of Om Koi Wildlife Sanctuary, Chiang
Mai and Tak Provinces. MS thesis, Kasetsart University, Thailand.
Kattan, G.H., Alvarez-Lopez, H., 1996. Preservation and manage-
ment of biodiversity in fragmented landscapes in the Columbian
Andes. In: Schelhas, J., Greenberg, R. (Eds.), Forest Patches in
Tropical Landscapes. Island Press, Washington, DC, pp. 3–18.
Laake, J.L., Buckland, S.T., Anderson, D.R., Burnham, K.P.,
1993. DISTANCE User’s Guide V2.0. Colorado Cooperative
Fish and Wildlife Research Unit. Colorado State University,
Fort Collins, CO.
Laidlaw, R., 2000. Effects of habitat disturbance and protected areas
on mammals of peninsular Malaysia. Conservation Biology 14,
1639–1648.
Laurance, W.F., 1999. Reflections on the tropical forest crisis. Biolo-
gical Conservation 91, 109–117.
Laurance, W.F., Bierregaard, R.O. (Eds.), 1997. Tropical Forest
Remnants: Ecology, Management, and Conservation of Frag-
mented Communities. University of Chicago Press, Chicago.
Lekagul, B., Round, P.D., 1991. A Guide to the Birds of Thailand.
Saha Karn Bhaet, Bangkok.
Lovejoy, T.E., Bierregaard Jr., R.O., Rylands, A.B., Malcolm, J.R.,
Quintela, C.E., Harper, L.H., Brown Jr., K.S., Powell, A.H.,
Powell, G.V.N., Schubart, H.O.R., Hays, M.B., 1986. Edges and
other effects of isolation on Amazon forest fragments. In: Soule,
M.E. (Ed.), Conservation Biology: The Science of Scarcity and
Diversity. Sinauer Associates, Sunderland, Massachusetts, pp.
257–285.
Lynam, A.J., Billick, I., 1999. Differential responses of small mammals
to fragmentation in a Thailand tropical forest. Biological Con-
servation 91, 191–200.
McGarigal, K. and Marks, B.J. 1995. FRAGSTATS: Spatial Pattern
Analysis Program for Quantifying Landscape Structure. US Forest
Service General Technical Report PNW351.
McGarigal, K., McComb, W., 1995. Relationships between landscape
structure and breeding birds in the Oregon Coast Range. Ecological
Monographs 65, 235–260.
Newmark, W.D., 1991. Tropical forest fragmentation and the local
extinction of understory birds in the eastern Usambara Mountains,
Tanzania. Conservation Biology 5, 67–78.
OEPP, 1995. Thailand’s Biodiversity. Natural Resources and Envir-
onmental Management Division, Bangkok.
PCI, 1997. GCP Works Reference Manual for PCI version 6.2. PCI,
Ontario.
Poonswad, P., Kemp, A.C., 1993. Manual to the Conservation of
Asian Hornbills. Hornbill Project, Thailand.
Price, O.F., Woinarski, J.C.Z., Robinson, D., 1999. Very large
requirements for frugivorous birds in monsoon rainforests of the
Northern Territory, Australia. Biological Conservation 91, 169–180.
Rabinowitz, A., 1993. Estimating the indochinese tiger Panthera
tigris corbetti population in Thailand. Biological Conservation
65, 213–217.
Rajani, M.R.B. 1999. System plan of protected area management in
Thailand. Paper delivered at ‘‘Roundtable on Environmental Pro-
tection and Rural Development in Thailand,’’ Marriott Hotel,
Bangkok, November. Faculty of Forestry, Kasetsart University,
Bangkok.
Renjifo, L.M., Servat, G.P., Goerck, G.M., Loiselle, B.A., Blake, J.G.,
1997. Patterns of species composition and endemism in the northern
Neotropics: a case for conservation of montane avifaunas. Orni-
thological Monographs 48, 577–594.
Shrock, J.L.I., Crowe, M., Fromme, D.E., Gosier, V.S., Mckenzie, R.,
Myers, W., Stegeman, P.L., 1970. Minority Groups in Thailand.
Department of the Army, Washington DC.
Simberloff, D.S., 1976. Experimental zoogeography of islands: effects
of island size. Ecology 57, 629–648.
Sodhi, N.S., Liow, L.H., 2000. Improving conservation biology
research in southeast Asia. Conservation Biology 14, 1211–1212.
Stuwe, M., Abdul, J.B., Mohd, B., Wemmer, C.M., 1998. Tracking the
movements of translocated elephants in Malaysia using satellite
telemetry. Oryx 32, 68–74.
Sunquist, M.E., 1981. The social organization of tigers (Panthera
tigris) in Royal Chitawan National Park, Napal. Smithsonian Con-
tributions to Zoology 336, 1–98.
Thailand Forest Research Center, 1991. The Master Plan for Mae-
Tuen Wildlife Sanctuary. The Forest Research Center, Faculty of
Forestry. Kasetsart University, Bangkok.
Thailand Forest Research Center, 1992. The Master Plan for Om Koi
Wildlife Sanctuary. The Forest Research Center, Faculty of For-
estry. Kasetsart University, Bangkok.
Tungittiplakorn, W. 1998. Highland Cash Crop Development and
Biodiversity Conservation: The Hmong in Northern Thailand. PhD
dissertation, University of Victoria, Victoria, BC.
Turner, I.M., Corlett, T.T., 1996. The conservation value of small,
isolated fragments of lowland tropical rain forest. Tree 11, 330–333.
Umapathy, G., Kumar, A., 2000. The occurrence of arboreal mam-
mals in the rain forest fragments in the Anamalai Hills, south India.
Biological Conservation 92, 311–319.
Vidhidharm, A., Nuttaro, S., Chimchome, P., Wangdilok, S., Chim-
chome, V., Suraparbmaitri, S., Eiadthong, W., Kanjanavanit, R.,
Kakkuntod, S., Jirawatkavi, N., Tunhikorn, S., Poonswad, P., 1995.
Current distribution and status of hornbills in Thailand. Journal of
Wildlife in Thailand 4, 1–11.
Warburton, N.H., 1997. Structure and conservation of forest avifauna
in isolated rainforest remnants in tropical Australia. In: Laurance,
W.F., Bierregaard Jr., R.O. (Eds.), Tropical Forest Remnants:
Ecology, management, and conservation of fragmented commu-
nities. University of Chicago Press, Chicago, pp. 190–206.
Wemmer, C.T., Kunz, T.H., Lundie-Jenkins, G., McShea, W.J., 1996.
Mammalian signs. In: Wilson, D.E., Cole, F.R., Nichols, J.D.,
Rudran, R., Foster, M.S. (Eds.), Measuring and Monitoring Biolo-
gical Diversity: Standard Methods for Mammals. Smithsonian
Institution Press, Washington DC, pp. 157–176.
Whitmore, T.C., 1984. Tropical Rain Forests of the Far East. Clar-
endon Press, Oxford.
Wongsprasert, S., 1975. Lahu trade and commerce. Journal Siam
Society 63, 199–219.
Young, G., 1967. Tracks of an Intruder. Souvenir Press, London.