Biological Conservation 98 (2001) 11±18

www.elsevier.com/locate/biocon

Ecology of the red panda Ailurus fulgens in the

Singhalila National Park, Darjeeling, India
Sunita Pradhan a,*, Gautom K. Saha a, Jamal A. Khan b
a
P.G.Department of Zoology, Darjeeling Government College, Darjeeling-734102, India
b
Centre of Wildlife and Ornithology, Aligarh Muslim University, Aligarh-202002, India

Received 3 August 1999; received in revised form 16 February 2000; accepted 5 April 2000

Abstract
Red panda, Ailurus fulgens is a poorly known Himalayan member of Carnivora which has adapted to a herbivorous diet.
The present study conducted in the Singhalila National Park in the eastern Himalayas was initiated to gain information on the
ecology and conservation problems of the red panda. Indirect and direct evidence was used to assess its distribution, relative
abundance, habitat use and food habits in the National Park. Red panda was relatively more abundant within an altitudinal range
of 2800±3600 m. In study sites 1 and 2, it had a very narrow preferred range of 2800±3100 m. Higher bamboo cover, bamboo height
and canopy cover emerged as important habitat components in sites used by red panda (Animal centered plots) compared to ran-
dom plots. Red panda diet consisted chie¯y of bamboo leaves and both species of bamboo, Arundinaria maling and A. aristata
predominantly present as understorey in Singhalila National Park were eaten. The diet of bamboo was supplemented by seasonal
fruits and bamboo shoots. However, the composition of diet di€ered between the three study sites and the impact of this, if any, on
the overall ecology of red panda in the Singhalila National Park, needs further investigation. Some conservation problems are
discussed. # 2001 Elsevier Science Ltd. All rights reserved.
Keywords: Red panda (Ailurus fulgens); Singhalila National Park; Darjeeling (Eastern Himalaya); Ecology; Conservation

1. Introduction pattern of distribution, population status, preferred

There are more endangered taxa in the Himalayas
than anywhere else in India (Rodgers and Panwar,
1989). The red panda is one. It is an unusual member of
Carnivora, occupying a highly specialised niche as a
bamboo feeder like the giant panda (Alieuropoda mela-
noleuca). The distribution of the red panda extends from
Nepal in the west to a few provinces of China in the
east. In India, it is found in the eastern Himalaya that
includes Darjeeling, Sikkim and Arunachal Pradesh and
is protected as an endangered species under the Indian
Wildlife Protection Act (1972). Red panda is a protected
species in Nepal, China, Bhutan and Myanmar and is in
Appendix I of CITES (Brautigam, 1995). However, very
few studies (Johnson et al., 1988; Reid et al., 1991;
Yonzon and Hunter, 1991a,b) have been carried out on
the red panda and more needs to be known about the
* Corresponding author at present address: Forum for Himalayan
Studies and Research, Post Box 111, PO & Dist. Darjeeling - 734101,
West Bengal, India.
E-mail address: sunitaprad@yahoo.com (S. Pradham).
0006-3207/01/$ - see front matter # 2001 Elsevier Science Ltd. All rights reserved.
PII: S0006-3207(00)00079-3
habitats, availability and quality of such habitats in
relation to food, cover and safety. Population dynamics,
home range, movement pattern and their causal factors
and various problems of conservation in di€erent areas
of its distribution need further studies.
This paper presents the ®ndings of a 3 year study
(Pradhan, 1998) on the ecology of red panda in the
Singhalila National Park, Darjeeling (India). Use of
telemetry was not possible due to various logistic and
®nancial constraints, so that direct (sighting) and indir-
ect evidence (dung) was used to assess the distribution,
relative abundance, habitat use and food habits of red
panda in Singhalila National Park.
2. Study area
Singhalila National Park (87 590 ±88 530 E and 26 310 -
27 310 ±27 310 N) with an area of 108.8 km2 (core
area=78.6 km2, bu€er=30.17 km2) is located on the
northwestern border of Darjeeling, India (Fig.1). It has
12 S. Pradhan et al. / Biological Conservation 98 (2001) 11±18

a boundary with the State of Sikkim in the north and
Nepal in the west. A motorable road identi®es the
international border between Nepal and India (Fig.1).
Singhalila National Park ranges from an altitude of
2400 m±3636 m encompassing two broad vegetation
zones, viz. the temperate and the subalpine (Table 1).
The moist temperate climate of Singhalila varied with
altitude. Summer and winter temperatures ranged from
7 to 17 C and from 1 to 10 C, respectively in the tem-
perate zone. In the subalpine zone, average summer and
winter temperatures were 7 and 1 C, respectively, and
the area intermittently received snowfall from end of
November to end of March. Mean annual rainfall was
350 cm. and average humidity ranged from 83 to 96%.
The Park is surrounded by settlements (Fig. 1) the east-
ern side of which are referred to as the `bu€er settle-
ments' and the settlements on the western border as
`border settlements' in the text.

Fig. 1. Map of Singhalila National Park, Darjeeling.

 S. Pradhan et al. / Biological Conservation 98 (2001) 11±18 13

Table 1
The vegetation and their composition in the intensive study area of Singhalila National Park

Vegetation zones Altitude (m) Dominant upper canopy (IVI)a Dominant sub-canopy (IVI) Dominant shrubs

Temperate zone
Oak forest 2600±2800 Quercus pachyphylla (81.2) L. elongata (55.0) Arundinaria maling
Litesea sericera (20.9) Sche‚era impressa (36.4) V. erubescense
Magnolia campbellii (17.1) Rhododendron arboreum (32.0) Daphne cannabina
R. grithiana (17.2) Piptanthus nepalensis
Broad leaf >2800±3100 Sorbus cuspidata (49.6) Symplocus sp. (68.8) A. maling
deciduous forest Acer sp. (25.1) R. arboreum (33.3) A. aristata
L. sericera (17.0) Osmanthus sauvis (23.47) V. erubescense
Q. pachyphylla (10.8) R. falconeri (21.4) D. cannabina
Viburnum erubescens (10.3) Rosa sericera
Meliosma delleniaefolia (8.9) P. nepalensis
Eltzostia sp.
Subalpine zone
Broad-leaf coniferous >3100±3300 Abies densa (68.2) Betula utilisb (79.4) A. aristata
forest R. arboreum (71.0) V. erubescense
Tsuga brunonianab (7.1) R. sericera
S. cuspidatab (6.4) Berberis aristata
Buddleia sp. (6.4) B. angulosa
Andromeda formosa (5.9) R. lepidotum
Coniferous forest >3300±3600 A. densa (104.9) Betula utilisb (70.5) A. aristata
R. campanulatum (55.0) R. sericera
B. aristata
R. campanulatum
a
IVI, importance value index.
b
Species also form upper canopy.

The intensive study area encompassed three sites, viz.
site 1 (Gairibans), which ranges within 2558±3000 m,
site 2 (Kaiyakatta-Kalipokhari) within 2670±3100 m
and site 3 (Sandakphu) above 3150±3636 m.
3. Methods
A preliminary survey of the Singhalila National Park
was conducted from October 1993±1994 to establish the
presence/absence of the red panda. After the survey and
the con®rmation of the presence of the red panda in the
Singhalila National Park, the three study sites were
selected (Fig.1) for initiation of more intensive investi-
gations of red panda ecology.
As it was not possible to establish transects (Burnham
et al., 1980) due to the rugged terrain and often dense
bamboo undergrowth in the study area, the existing
paths and trails passing through di€erent habitat types
and altitudinal zones were carefully selected. A total of
12 such trails were monitored at regular intervals of 1
month for 2 years, for direct and indirect evidences of
the red panda. Apart from this, 91 random plots in dif-
ferent vegetation zones were marked and also mon-
itored on a monthly basis for red panda evidences.
Dung/pellet groups were found to be the most reliable
indirect evidence and were used to assess the distribu-
tion, abundance and habitat use.
The pellet groups encountered during the monitoring
were collected, dried and kept for subsequent analysis of
faeces to investigate the food habits of red panda. On
encountering pellet group/groups, information recorded
were on the number of pellets in a pellet group, number
of groups at one place and the type of substrates used.
For each sighting, notes on the number of animals, sex,
age and activity were taken. An encounter rate of pellet
groups/100 hours was used as an index to quantify the
relative abundance in the study area.
Each point of red panda sighting or faeces was
marked and habitat factors of the location was quanti-
®ed using ten-tree method (Mueller-Dombois and
Ellenberg, 1974). The sampled location where the ani-
mal evidence was found was named animal centered
plot to di€erentiate between the four additional random
plots established at four cardinal directions from the
animal centered plot. The four random plots were
established in order to compare the habitat factors of
the location found to be used by the red panda (animal
centered plot) and random locations. Random distances
(>30±250 paces) were generated for establishing these
random plots. Habitat variables such as altitude, habi-
tat type, topography, aspect, slope angle, tree species
14 S. Pradhan et al. / Biological Conservation 98 (2001) 11±18
Fig. 2. Classi®cation of the vegetation of the study area based on importance value index of tree species at di€erent altitudes.
and canopy cover were recorded. Shrub species, shrub
cover, bamboo species, number and average bamboo
height were quanti®ed in 3 m2 quadrats. The composi-
tion, structure and regeneration of vegetation and the
intensity and types of disturbances in the study area
comprising of sites 1±3 were assessed by placing 10 m2
quadrates at an interval of 100 m along seven altitudinal
gradients (2700, 2850, 3000, 3300, 3450 and 3600 m)
following Shrestha (1988). The variables recorded in the
quadrats were tree species present, their height, girth at
breast height (gbh) of each tree. Bamboo and other
shrub species, and their number were quanti®ed in a 3 m2
within the 10 m2 quadrats. Nomenclature for plant
species follow Cowan and Cowan (1929) and Polunin
and Stainton (1986). The importance value index (IVI)
for the tree species was determined by summing the
values of relative frequency, relative abundance and
relative dominance (Curtis, 1959). Cluster analysis was
performed on the data matrix of the IVI of the tree
species recorded at seven altitudes following Causton
(1988).
4. Results and discussion
The cluster analysis produced four distinct clusters of
vegetation communities corresponding to the vegetation
communities in the altitude zones (Fig.2). The vegeta-
tion zones are oak forest (2700±2800 m), broad-leaf
deciduous forest (>2800±3100 m), broad-leaf con-
iferous forest (>3100±3300 m) and the coniferous forest
(>3300±3600 m). The compositions of these zones are
presented in Table 1. Sites 1 and 2 of the intensive study
area represent the temperate zone with the broad-leaf
deciduous and oak forest. Site 3 represents the sub-
alpine zone with broad-leaf coniferous forest and the
coniferous forest.
4.1. Distribution and relative abundance
Sightings of red panda were extremely infrequent. Out
of the 32 sightings, 18 encounters occurred during the
actual monitoring of the trails and the rest during ®eld-
work other than the monitoring, details of which are
presented in Pradhan (1999). Encounter rate of pellet
groups and red panda sightings in the study sites 1, 2
and 3 di€ered signi®cantly (Table 2). Encounter rate of
both red panda and pellet groups was highest in broad-
leaf deciduous forest followed by subalpine and oak
forest (Table 3). The encounter rate of pellet groups
being a good index of abundance in the present study
was testi®ed by the fact that more pandas were seen in
areas with higher encounter rate of pellet groups. The
correlation between encounter rate of pellet groups/100
Table 2
Encounter rate of pellet groups/100 h and red panda sightings/100 h in
the study area
Site 1 Site 2 Site3
Pellet groups 24.5513.09 28.8332.16 37.8432.16 P<0.05
Red panda 0.440.03 3.042.0 2.982.1 P<0.05
 S. Pradhan et al. / Biological Conservation 98 (2001) 11±18
Table 3
Encounter rate of pellet groups/100 h and red panda sightings/100 h in the three vegetation zones
Oak forest Broad-leaf deciduous forest
(2600±2800) (>2800±3100)
Pellet group 20.3110.05 105.0629.59
Red panda 0 3.813.00
h and red panda sightings in di€erent transects in the
study area was also signi®cantly positive (rs=0.78, P
<0.05). Evidence of red panda was found in only
16.7% of the permanent random plots (n=91). The
density of pellet groups based on their presence in the
random plots showed lowest density in the oak forest
with 3.26/ha compared with 43.66/ha in the broad-leaf
deciduous forest and 39.19/ha in the subalpine forest.
The maximum number of panda individuals seen
together or in a group in the study area was three, on
three occasions. On the ®rst occasion, three adult red
pandas were seen together on a Rhododendron sp. tree in
late December (winter). As winter is the mating season
(Dittoe, 1944; Zuckerman, 1953; Mottershead, 1958,
1963; Erken and Jacobi, 1972; Roberts and Kessler,
1979; Keller, 1980; Roberts, 1981) these pandas could
be in a premating aggregation. In the wild, red pandas
are solitary outside their mating period (Yonzon and
Hunter, 1991a). On the next two occasions, two cubs
with a female red panda were seen in October in site 2.
All these sightings in site 2 are evidence of presence of a
breeding population in the area. A cub and a female
were sighted in site 3 during November along with
encounters of cub pellets in areas where direct sightings
were not made. No such evidence of breeding popula-
tion was found in site 1.
Results of the preliminary survey (Pradhan, 1998),
pellet-groups and red panda encounter rates supported
by the density of pellet groups in the random plots in
the vegetation zones and sighting records showed that
the red panda in the study area was distributed in the
entire study area but was relatively more abundant
within an altitudinal range of 2800±3600 m which
encompassed the broad-leaf deciduous forest and the
subalpine forest. The lowest altitude from where red
panda was reported in Singhalila National Park was
Upper Phedi (2400 m). Yonzon and Hunter (1991a)
reported the distribution of red panda in the Nepal
Himalaya within an altitudinal range of 3000±4000 m.
3063231m with a range of 2600±3600 m was the mean
altitude of red panda occurrence in Singhalila National
Park. A crude density of 1 red panda/1.67 km2 was
estimated in the Singhalila National Park which was
higher than the density of 1 adult panda/2.9 km2 esti-
mated in the Langtang National Park (Yonzon and
Hunter, 1991a). The studies of Johnson et al. (1988) and
Reid et al.(1991) reveal very little on the status and
15
Subalpine forest
(>3100±3600)
85.0042.00 P<0.05
2.782.2 P<0.05
abundance of the red panda in the Wolong Natural
Reserve, China.
4.2. Habitat use
There was no apparent relationship between high use
areas of the red panda and forest canopy other than the
presence of a patch of live bamboo in Wolong Nature
Reserve, China (Johnson et al., 1988). During the pre-
sent study, di€erences between the animal centered plot
(n=117) and the random plots (n=321) using the nine
habitat variables (altitude, bamboo cover, bamboo
density, bamboo height, tree density, canopy cover,
shrub cover, shrub density and shrub cover) showed
that the canopy cover, bamboo height and bamboo
cover di€ered signi®cantly between the animal centered
and random plots (Table 4). Logistic regression fol-
lowed by classi®cation table (Hamilton, 1998) showed
that the model classi®ed the plots by 76%. A classi®ca-
tion by only 76% could indicate the exclusion of habitat
variables that could have contributed to a higher per-
cent of classi®cation between the animal centered and
random plots. Presence of water bodies at a range of 0±
100 m from the point of animal centered plot was found
in 79% cases indicating the presence of water in the
preferred sites could be an important habitat requisite.
The di€erent substrates used by the red panda for
defecation were trees, ground or the forest ¯oor, fallen
logs, stumps, rock and tree holes (Table 5). Trees were
the most preferred site for defecation (w2=180, df=3,
P<0.001). Use of forest ¯oor was high during monsoon
and winter. The higher use of forest ¯oor during mon-
soon was probably because the panda was seeking
bamboo shoots on the forest ¯oor. During winter, pel-
lets and pellet groups were mostly found on the forest
¯oor, paths on ridges and slopes and this could be a
mode of communication between the conspeci®cs as
winter happens to be a mating season for the red panda.
Table 4
Quantitative di€erence between variables in animal and random plots
Variable Animal Random Mann±Whitney P
plot plot (z)
Bamboo cover 32.051.4 23.730.87 4.20 0.000
Canopy cover 25.76 1.5 18.60.10 3.46 0.001
Bamboo height 2.5 .14 2.10.10 2.26 0.023
16 S. Pradhan et al. / Biological Conservation 98 (2001) 11±18
Table 5
Use of di€erent substrates for defecation by the red panda in the study area during di€erent seasons
Seasons
Pre-monsoon (March±May)
Monsoon (June±August)
Post-monsoon (September±November)
Winter (December±February)
a
Others include fallen logs, stumps and tree holes.
81.25% of sightings were made on trees and the rest on
the ground. Pellet groups or indirect evidences were
found on 13 species of trees namely Abies densa, Acer
sp., Betula utilis, Ilex sp., Lahare tenga plus any tree,
Magnolia campbellii, Osmanthis sauvis, Quercus pachy-
phylla, Rhododendron spp., Sche‚era impressa, Sorbus
cuspidata, Sorbus microphylla and Vitex heterophylla,
out of which only eight species (A. densa, B. utilis, Ilex
sp., M. campbellii, O. sauvis, Q. pachyphylla, Rhododen-
dron spp. and S. impressa) were recorded with direct
sightings of the red panda. Interestingly, the eight spe-
cies except for M. campbellii recorded with the sightings
were mainly evergreen species. The tree species used
frequently and which formed important components of
red panda habitat were A. densa, B. utilis, S. cuspidata,
Q. pachyphylla, M. campbellii and Rhododendron spp.
The ratio of direct evidence (sighting) to indirect evi-
dence (pellet groups) in A. densa was 2.8:1, 1:3 in B.
utilis, 2.7:1 in Q. pachyphylla, 4:1 in M. campbellii, 1:20
in S. cuspidata and 1.7:1 on Rhododendron spp. The
higher frequency of sightings in the evergreen trees
could be because of the need of the animal for better
cover and safety.
4.3. Food and feeding habits
Bamboo leaves of A. maling and A. aristata formed
the chief food items of the red panda diet especially
during the premonsoon and the winter seasons (Prad-
han et al., in press). A. maling dominated the under-
storey of the study area between an altitude of 2600±
3100 m. A. aristata was found to occur from 2850 m,
but dominated the understorey from above 3100±3600
m in the study area. Red panda in Singhalila National
Park consumed mainly A. maling (83%) in site 1 (2558±
3000 m) along with supplements of fruits of Actinidia
strigosa (5%) and bamboo shoots (12%). At site 2
(2670±3100 m), red panda was consuming A. maling
(40%) along with A. aristata (51%) supplemented by A.
strigosa (5%) and bamboo shoots (4%). At site 3 (3150±
3636 m), A. aristata, the dominant species of the area
was the only bamboo species (91.2%) that red panda
Substrates
Tree Forest ¯oor Rocks Othersa
(% used)
70 16.4 6.7 6
54 43.5 0 0
80.7 11.5 7.6 0
29 45.9 18.9 5.4
was found to be subsisting on. The diet of bamboo
leaves was supplemented by bamboo shoots (8%) and
seasonal fruits/berries of Sorbus microphylla (0.43%),
and Rosa sericera (trace). From these results, it could be
seen that the red panda in the Singhalila National Park
consumed leaves of A. maling, A. aristata, fruits of
A. strigosa and bamboo shoots in the temperate zone
whereas the food items in the subalpine zone consisted
of A. aristata, bamboo shoots and berries of S. micro-
phylla. Bamboo species such as Siandunaria fangiana,
and Fargesia spathecea were known to be taken by the
red panda, with a preference for the former in the
Wolong Nature Reserve, China (Schaller et al., 1985;
Johnson et al., 1988; Reid et al., 1991) and a species of
bamboo locally known as Jhapra in the Langtang
National Park, Nepal (Yonzon and Hunter, 1991a).
4.4. Past disturbances and other conservation problems
In Langtang National Park (Nepal Himalayas), the
presence of large numbers of cattle, herders and their
dogs caused immense disturbance to red panda and red
panda habitat (Yonzon and Hunter, 1991b).
In Singhalila National Park, major disturbances and
exploitation of the ¯ora and fauna of the area occurred
prior to 1993 and these disturbances can be referred to
as past disturbances. The Singhalila area was declared a
National Park in 1992 and the Park closed to cattle
grazing and cattle stations known as Goths between
1991 and 1993. The earliest records of cattle, goats and
sheep in these areas are by O'Malley (1907). It is known
that grazing was restricted to areas above 2727 and 2879 m
in Tonglu and Singhalila Ranges for which grazing
permits were issued (Anon., 1967). A minimum of 80±90
Goths (cattle stations) were present inside Singhalila
National Park and wildlife poaching was widespread
prior to the establishment of the Park. Each goth had a
minimum average of 30±50 cattle. Some of the pressures
to the Singhalila forest were grazing, lopping, construc-
tion of roads by blasting of slopes, and ®re causing
major disturbances to the intactness of the ¯ora and
fauna of the region. Establishment of a cattle station
 S. Pradhan et al. / Biological Conservation 98 (2001) 11±18
required clearing of vegetation, thus causing gaps in the
canopy cover. The area was also bereft of bamboo, an
important component of red panda habitat. Interviews
with local people, trappers and agents indicated that
live-trapping of red panda from both Indian and Nepal
areas, to be supplied to the zoos was a lucrative business
which continued till the late 1970s to early 1980s. A
trader in Rimbick area had received as many as 16 red
pandas in a day. Among the animals received, there
would also be pregnant females and incidences of cubs
being born in the cages were also reported. Cases of red
panda cubs being still taken from their nests, to be kept
as pets in some of these remote areas were also found
during the tenure of this study. The Singhalila National
Park is surrounded by a number of settlements (Fig. 1).
A reserve forest bu€ered the eastern side of Singhalila
National Park and most of the bu€er settlements were
on the outer fringe of this reserve forest which kept
them at a considerable distance from the preferred
range of the red panda habitat in the core zone. How-
ever, the increasing human population, their relatively
unsound socio-economic condition and dependence on
the forest resources (Pradhan, 1998) were a threat which
should not be underestimated. It was also from these
areas that forest ®res started, damaging large tracks of
forest in Singhalila National Park. The border settle-
ments on the western border, had a comparatively lower
human population than the bu€er settlements, but
posed an immense threat to red panda and red panda
habitat, by being on the very range of the present dis-
tribution of red panda in Singhalila National Park. The
red pandas in sites 1 and 2 inhabited a very narrow
preferred altitudinal range of 2800±3100 m (broad-leaf
deciduous forest). Moreover, only one face of the slope
within Singhalila National Park encompassed this ele-
vation or vegetation zone in these sites. The other face
of the slope was in the Nepal and therefore the red
panda in Singhalila National Park is a€orded protec-
tion only within the western border. The case of sharing
the preferred vegetation zone of red panda with Nepal
in site 3 was also similar to that of the other two sites
although the preferred range of altitude was broader in
site 3 (<3150±3600 m).
The conservation measures in Singhalila National
Park would, therefore, require taking into consideration
the wellbeing of the red panda as well as the local com-
munity dependent on the habitat along and across the
borders of Singhalila National Park. Red panda uses
habitats in Sikkim, Nepal and Singhalila National Park
(Darjeeling). Concerted e€orts of all the three regions
(Nepal, Sikkim and Darjeeling) to protect the red panda
in this part of the Himalaya would be a better con-
servation strategy than conserving or protecting animals
within each region only. Talks of such co-operation and
understanding at a regional level have already started
between the Government of Nepal, State of Sikkim and
17
adjoining China which jointly proposed the formation
of a Kanchanjunga Mountain ecosystem (Rastogi et al.,
1997). This proposed complex could be extended further
also to include Singhalila National Park and the
adjoining areas of Nepal and Sikkim for a transbound-
ary conservation (Pradhan and Bhujel, 2000). This
would be helpful in managing and conserving the areas
along the borders more e€ectively with a common goal
of conserving the endangered red panda and its habitat
in this part of the Himalaya.
Acknowledgements
The research project on the ecology of the red panda
was sponsored by the Wildlife Division, Department of
Forest, Government of West Bengal. The manuscript
has bene®ted from the suggestions of Dr. Malcolm
Coulter, Dr. P.B.Yonzon, Dr. A.R. Rahmani and an
unknown referee. Dr. Angela R. Glatston is gratefully
acknowledged for her support and help throughout the
tenure of the Project. Dr. Rahul Kaul, Mr. Miles S.
Roberts, Dr. S. Kumar, Dr. S. Javed, Dr. S. Khaling,
research assistants and all the local people in the ®eld are
thanked for their support.
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