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Population densities, group size and biomass of ungulates in a lowland
tropical rainforest forest of the eastern Himalayas
K. Muthamizh Selvan, Salvador Lyngdoh, Govindan Veeraswami Gopi
⇑
, Bilal Habib, Syed Ainul Hussain
Wildlife Institute of India, Chandrabani, PO Box 18, Dehradun 248001, India
article info
Article history:
Received 24 July 2013
Revised 15 January 2014
Accepted 8 May 2014
Keywords:
Arunachal Pradesh
Biomass
Carnivores
Density
Distance
Eastern Himalayas
Ungulates
Group size
Sambar
Gaur
Muntjac
abstract
Large ungulate population monitoring is a crucial wildlife management tool as ungulates help in
structuring and maintaining the large carnivore populations. Reliable data on population status of major
ungulate prey species are still non-existent for most of the protected areas in the Indian part of the
eastern Himalayan biodiversity hotspot. Twenty transects were monitored over a period of three years
(2009–2011) totaling 600 km with an average length of 2 km. The estimated mean density of ungulates
was 17.5 km
2
with overall density of 48.7 km
2
. The wild pig Sus scrofa had the highest density
(6.7 ± 1.2 km
2
) among all the prey species followed by barking deer Muntiacus muntjak
(3.9 ± 0.6 km
2
), sambar Rusa unicolor (3.8 ± 0.5) and gaur Bos gaurus (3.5 ± 0.9 km
2
). The estimated total
ungulate biomass density was 2182.56 kg km
2
. This prey biomass can support up to 7.2 tigers per
100 km
2
. However, with two other sympatric carnivores sharing the same resources, the actual tiger
numbers that can be supported will be lower. The estimated minor prey species was 31 km
2
signifi-
cantly 30.6% crop damages were reported by wild pig (p= 0.01) and 35.4% was elephant (p= 0.004). This
data on ungulate densities and biomass will be crucial for carnivore conservation in this understudied
globally significant biodiversity hotspot.
Ó2014 Ecological Society of China. Published by Elsevier B.V. All rights reserved.
1. Introduction
Large ungulates are involved in fundamental ecological process
of seed dispersal, influence spatial patterns of vegetation and form
potential prey species for carnivores [48]. Abundance, distribution
and activity pattern of large carnivore is determined by different
size of the prey species [31] and the mean body size of large carni-
vores are mainly determined by the frequency distribution of
available prey [51]. Prey distribution, density and biomass within
a given area represent measurable amounts of energy potentially
available as food to carnivores. Predator–prey relationships
amongst large mammals have complex interactions in ecological
system [21]. Less preferred prey might have reduced mortality risk
when it co occurs with a favored prey if the predator concentrates
its attack on the preferred species [18]. The rate of carnivore pop-
ulation growth should be dependent on both the density of prey
and prey conspecifics [19]. Several studies have suggested main-
taining a healthy herbivore population both in terms of biomass
and community structure is essential for conserving a viable carni-
vore population [30,50]; Karanth and Stith, 1999; [29].
Overhunting in tropical forests compounded with little knowl-
edge on population ecology of large herbivores in South East Asia
is a major problem [20,48]. In Arunachal Pradesh, prey depletion
due to hunting and livestock depredation by carnivores that leads
to retaliatory killing of the latter is common [2,22]. This study pro-
vides baseline information on ungulate prey species available for
large carnivores in Pakke Tiger Reserve.
The recent study conducted by the Wildlife Institute of India in
Pakke–Nameri TR has documented a decline of tiger occupied areas
from 1100 km
2
to 371 km
2
[26]. Depletion of prey is the major
reason for carnivore decline especially for tiger throughout the
species range [32] and we suspect the same to be true for these
tiger reserves in Arunachal Pradesh.
Based on available studies in South East Asia ([24,7,44,45,34]),
in this paper we focus on biomass, density, population structure
and composition of the major prey species in PTR. Our study high-
lights certain important observations and we recommend likewise
key measures for long-term survival of prey species in the region.
2. Materials and methods
The study was carried out in Pakke Tiger Reserve (PTR) 26°54
0
–
27°16
0
N, 92°36
0
–93°09
0
E in the foot hills of east Himalayan region,
http://dx.doi.org/10.1016/j.chnaes.2014.05.003
1872-2032/Ó2014 Ecological Society of China. Published by Elsevier B.V. All rights reserved.
⇑
Corresponding author. Address: Department of Endangered Species Manage-
ment, Wildlife Institute of India, India.
E-mail address: gopigv@wii.gov.in (G.V. Gopi).
Acta Ecologica Sinica 34 (2014) 219–224
Contents lists available at ScienceDirect
Acta Ecologica Sinica
journal homepage: www.elsevier.com/locate/chnaes
East Kameng district of Arunachal Pradesh, India. Arunachal
Pradesh (26°28
0
–29°30
0
N and 91°30
0
–97°30
0
E; 83,743 sq. km) is
arguably the richest terrestrial biodiversity region of the country
[38]. Based on the biogeographic classification it falls in the east
Himalayan biogeographic region [41]. In the last few years Aruna-
chal witnessed many discoveries and unknown species [38]. The
Southern side of PTR is adjoining with Nameri National park and
Tiger Reserve which is in Assam and Northern side of the PTR is
contiguous with the forest. The Sanctuary is naturally protected
by the rivers in the east, west and south. The vegetation of the park
is classified as Assam valley tropical evergreen [13]. Rainfall is
received from both south-west (May–September) and north-east
monsoons (November–April) with average annual rainfall of
2500 mm [14]. Hilly terrain and dense cover has left many places
still inaccessible in many parts of Arunachal Pradesh. Around 20
Nyishi (indigenous community) villages are located surrounding
the park. The dominant form of cultivation that is practiced by
communities here is a form of slash and burn cultivation called
jhum. Though in valleys permanent agriculture fields exist, jhum
remains a common form of agriculture practiced. Hunting in
various forms such as recreational or ritualistic are traditionally
practiced largely along with fishing.
Apart from large carnivores (Tiger, leopard, Dhole and Clouded
leopard) smaller carnivores such as Asiatic golden cat Catopuma
temminckii [37], marbled cat Pardofelis marmorata [37] and leopard
cat Prionailurus bengalensis are also present in the PTR. Major
ungulate prey species are sambar Rusa unicolor, wild pig Sus scrofa,
gaur Bos gaurus, muntjac Muntiacus muntjak, and non-ungulate
prey species are capped langur Trachypithecus pileatus, rhesus
macaque Macaca mulatta, kalij pheasant Lophura leucomelanos,
red jungle fowl Gallus gallus and gray peacock pheasant Polyplec-
tron bicalcaratum.
We used distance sampling procedures [10,8] to estimate the
density of ungulate prey species in the study area as this method
is successfully used in South East Asia to estimate the animal den-
sities [28,6,23,3,54]. We gridded the low land forests of the Pakke
Tiger Reserve into 3 3 km grids and laid 1 transect in each grid
cell (Fig. 1). Twenty line transects were randomly chosen to enu-
merate the animals from September 2009 to January 2012 covering
a length of 600 km (3 years). Each transect was approximately
2 km in length and was walked 5 times in the morning hours
(5.30–9.00 am) by two observers. All transects were marked with
GPS coordinates and bearing using compass. For each detection
species, sighting distance, animal bearing, sighting angle, time,
group size, group composition, sex and age class of the individuals
were recorded. Distance and angle were recorded from the centre
of the cluster. To assess the crop damage by ungulates, question-
naire survey was conducted in the villages adjacent to the reserve,
to determine the extent of human-ungulate conflicts in terms of
crop depredation. The sampled households from each village were
selected randomly and more than 80% households interviewed
[33]. A total of 584 households were interviewed.
Distance 6.0 [52] was used to estimate the line transect data for
prey density estimation. Data were checked for errors before using
program distance [25,54] and exploratory analysis was carried out
to check for the evidence of evasive movements before detections
[11,54]. To get the better model fit data was truncated and the best
model was selected on basis of lowest AIC (Akaike Information Cri-
teria) value [9,10]. To get the better density estimate a minimum
number of sightings are required in order to model the detection
function. In this case all the three years data were pooled together.
Density estimates obtained from transects were used to calcu-
late the biomass of prey species in the study area. A commonly
expressed version of density in terms of total biomass is the bio-
mass density that was calculated by multiplying the density of
prey species by their average individual weights. The average body
weight of each prey species required for biomass calculation was
taken from available literature [42,39,30]. The proportional repre-
sentation of individual age-sex classes of each prey was computed
(Table 1). Using these proportions, the average unit weight of each
prey species was calculated and was weighted by the proportions
of each age-sex class of that species. The overall densities of
animals for each species were multiplied by their average weight
following Berwick et al. [5] and Karanth and Sunquist [28] to calcu-
late the wild ungulate biomass. Kruskal–Wallis test were carried
out to assess the significant crop damages by ungulates and other
herbivores to calculate the wild ungulate biomass.
3. Results
A total of 417 sightings were made for four major prey species
(n= 292) and seven minor prey species (n= 125) along 600 km of
effort. Major prey species found were sambar (n= 93), barking deer
(n= 84), wild pig (n= 73), gaur (n= 42) and minor prey species
such as capped langur (n= 19), kalij pheasant (n= 19), red jungle
fowl (n= 39), gray peacock pheasant (n= 10), elephant (n= 20)
and Malayan giant squirrel (n= 13) were also encountered. Desir-
able precision level could not be achieved in some cases due to less
than 40 sightings as suggested by Burnham et al. [10]. In such cases
data were pooled for three years to get the density estimation of
that species. Overall prey density was 48.7 km
2
in which major
prey was 17.7 km
2
and minor prey was 31 km
2
. The density of
sambar was 3.8 ± 0.5 individuals/km
2
. Half Normal key Cosine
(AIC-115.1) was best fitted for sambar density estimation. Proba-
bility density function at zero (f0) was 0.7 and detection probabil-
ity (P) 0.2. Probability of a greater Chi-square value (P) 0.7 and
mean group size 1.7. Barking deer density was 3.9 ± 0.6 individu-
als/km
2
(Table 2).
Half Normal key was selected for density estimation (AIC
108.1). Probability density function at zero (f0) 0.8 and detection
probability (P) was 0.2. Probability of a greater Chi-square value
(P) 0.7 and mean group size 1.6. Wild pig had the highest density
(6.7 ± 1.2) among other major prey species. The selected model
was Half Normal Cosine with AIC value of 79.1. Probability density
function at zero (f0) was 0.6 and detection probability (P) was 0.2.
Probability of a greater Chi-square value (P) 0.5 and mean group
size was 2.1. The density of gaur was 3.5 ± 0.9 individuals/km
2
.
Half Normal Cosine was the best fitted key (AIC 68.03). Probability
density functions at zero (f0) was 0.3 and detection probability (P)
was 0.3. Probability of a greater Chi-square value (P) was 0.6 and
mean group size recorded was 2.9. Among minor prey species
red jungle fowl had the highest density (5.9 ± 1.1) followed by
capped langur (2.5 ± 0.1), kalij pheasant (2.1 ± 0.7) and Malayan
giant squirrel (2.1 ± 0.7). Half Normal Cosine was the best fitted
key for all the minor prey.
The population structure was derived for major prey species
from different sightings in 2009–2011. The adult sex ratio (male:-
female) for sambar 1:0.5, for barking deer adult sex ratio was 1:0.6,
wild pig 1:0.6 and gaur 1:2.1. The female-fawn ratio of sambar was
1:0.3, barking deer 1:0.2, wild pig 1:0.4 and for gaur 1:0.4. Capped
langur sex ratio was 1:0.5 and female infant ratio was 1:0.3 (Fig. 2).
Prey biomass density estimates shows gaur formed bulk of the
prey (61%) and barking deer formed lowest (63%) among major
prey of species of carnivores. Total prey biomass density was
2182.56 kg km
2
(Table 3).
A total of 584 households were interviewed in 11 villages.
People said the elephant was major crop raider in winter (35.4%)
followed by wild pig (30.6%), monkey (20.8%), rodents 10.2% and
other ungulates (4%). We found significant crop damages by ele-
phant (
v
2
= 5.6, p= 0.004) and wild pig (
v
2
= 4.9, p= 0.01) during
winter season (Fig. 3).
220 K. Muthamizh Selvan et al. / Acta Ecologica Sinica 34 (2014) 219–224
4. Discussions
The study is focused on a single protected tiger reserve i.e. the
Pakke Tiger Reserve (PTR) in the north east India which has seven
other designated tiger reserves viz., Namdapha in Arunachal Pra-
desh, Buxa in northern West Bengal, Manas, Kaziranga and Nameri
in Assam, and Dampa in Mizoram. Reliable data on population and
biomass estimation of major ungulate prey species are still non-
existent for most of these reserves. The approach and implications
of this research is applicable to other key reserves in the region.
Though the distance sampling had the limitation to be used in
the low density area [10], with sufficient effort it can be an effec-
tive tool to study the prey population [55]. Even though hunting
and jhum cultivation was widely practiced in Arunachal, the good
number of ungulate population was observed in the Intensive
Study area (ISA) due to better protection and management of hab-
itat in the south eastern boundary of the park. The estimate of
ungulate density (19.1 km
2
) was one of the lowest in other part
of South East Asia [43,31,55,40]. Sambar which is the potential
prey for major carnivores in Pakke Tiger Reserve [46] contributes
to 21% prey biomass next to the gaur (66.3%). Barking deer density
found to be high (3.6 km
2
) because small bodied barking deer are
more resilient to hunting pressure [15]. Wild pig was found in
almost all the habitats in Pakke Tiger Reserve. Respondents
claimed that wild pigs were responsible for most of the crop dam-
ages to adjacent villages. Better protection in PTR has attributed to
a relatively [36] good population of gaur. Capped langur population
was found to be low. Despite a ban on using animal parts for the
decoration of traditional tools, the rampant use parts such as its
fur for decorating local artifacts and tools may be one of the
Fig. 1. Map shows the distribution of transects along the study area.
Table 1
Proportional biomass and unit weights of major prey species in PTR (2009–2011).
Species Age/sex class Numbers Weight (kg) % in pop Prop wt (kg) Unit weight (kg)
Sambar Adult Male 61 225 21.63 48.67 121.97
Adult female 113 150 40.07 60.11
Sub adult 60 50 21.28 10.64
Fawn 48 15 17.02 2.55
Muntjac Adult Male 67 22 23.93 5.26 16.26
Adult female 117 20 41.79 8.36
Sub adult 52 10 18.57 1.86
Fawn 44 5 15.71 0.79
Wild pig Adult Male 67 60 20 12 31.19
Adult female 124 40 37.01 14.81
Sub adult 75 15 22.39 3.36
Piglets 69 5 20.6 1.03
Gaur Adult Male 37 745 13.03 97.06 413.34
Adult female 98 550 34.51 189.79
Sub adult 90 350 31.69 110.92
Calf 59 75 20.77 15.58
K. Muthamizh Selvan et al. / Acta Ecologica Sinica 34 (2014) 219–224 221
reasons. Various socio-cultural and religious practices in the region
have contributed to the decline the population of capped langur
[47,36].
Total biomass was estimated to be 2182.6 kg km
2
which was
enough to hold a good number of carnivore populations. Assuming
the annual prey requirement of tigers to be approximately 3000 kg
of prey/tiger [42,50] and tigers removes approximately 10% of the
prey every year [32]. Based on this, the estimated biomass of
2182.56 kg km
2
can support up to 7.2 tigers km
2
. However, with
two other sympatric carnivores sharing the same resources, the
actual tiger numbers that can be supported will be lower. Studies
in Bhutan have revealed a biomass of 379 kg km
2
which can
support up to 1.2 tigers per 100 km
2
[55]. Wild pig causes fore-
most damage to crop due to high density and other ungulate cause
negligible damage that too jhum cultivation.
Prey biomass estimates have ranged from 266 to 426 kg km
2
in the tropical rainforests of peninsular Malaysia [34]. Estimates
of prey biomass from Indonesia ranged from 200 to 400 kg km
2
[24,7,44,45]. Seidensticker et al. [44] has concluded that the bio-
mass of essential ungulate prey species for tigers in Asian rainfor-
ests does not exceed 500 kg km
2
. Eisenberg [16] have suggested
that tropical humid forests are generally considered to be poorer
habitats than savanna in terms of supporting higher ungulate
biomass. Such generalizations have been earlier questioned and
have proposed that several factors like strict enforcement, absence
of climate induced mortalities, presence of diverse flora due to bio-
climatic variables and maintenance of successional vegetation
determine the prey densities [28]. Our estimated biomass was
comparatively higher as the Intensive Study Area (ISA) is a pro-
tected area and is ensured good protection and management. The
results of the present study is comparable with results of other
studies in the Indian subcontinent, 4937 kg km
2
in Ranthambore
[35], 7638 kg km
2
in Nagerhole NP [28] and 6013 kg km
2
in
Pench TR [6]. The lowest ungulate biomass (379 kg km
2
) have
been reported in Jigme Singye Wang chuck National Park [55] in
Bhutan (Table 4), though Jigme Singye Wang chuck National Park
closer to PTR the estimated biomass was too lower than PTR which
Table 2
Density estimation for major prey species in Pakke Tiger Reserve, (2009–2011).
Years Species Number of observations Model ESW GS ± SE Dg ± SE %CV D ± SE %CV Confidence interval
Lower Upper
2009 Sambar 21 HF/normal 12.1 ± 1.9 1.9 ± 0.2 2.2 ± 0.5 27 4.3 ± 1.2 30.1 2.3 7.7
2010 44 HF/normal 26.1 ± 3.01 1.8 ± 0.1 2.6 ± 0.4 17 4.9 ± 0.9 19.1 3.4 7.2
2011 28 HF/normal 16.8 ± 2.3 1.8 ± 0.2 3.2 ± 0.8 27 5.9 ± 1.7 20.1 3.3 10.5
CD 93 HF/normal 16.1 ± 1.3 1.7 ± 9 2.2 ± 0.2 13 3.8 ± 0.5 13.9 2.9 5.1
2009 Muntjac 25 HF/normal 11.6 ± 1.7 1.2 ± 0.1 2.7 ± 0.7 27 3.5 ± 1.02 29.1 1.9 6.1
2010 40 HF/normal 16.0 ± 2.03 0.9 ± 0.1 3.8 ± 0.8 21 3.8 ± 0.8 20.01 2.5 5.8
2011 19 HF/normal 14.2 ± 2.4 1.1 ± 0.5 2.5 ± 0.7 30 2.7 ± 0.8 30.4 1.5 5
CD 84 HF/normal 11.8 ± 1.2 1.0 ± 0.1 3 ± 0.5 15 3.9 ± 0.6 15.5 2.9 5.3
2009 Wild pig 17 HF/normal 7.8 ± 1.5 1.4 ± 0.1 2.7 ± 0.8 30 4.1 ± 1.3 32.8 2.1 7.6
2010 41 HF/normal 18.5 ± 2.2 1.9 ± 0.1 3.4 ± 0.7 20 6.6 ± 1.4 22.1 4.3 10.3
2011 15 HF/normal 12.3 ± 0.3 2.8 ± 0.3 3.3 ± 1.3 32 9.3 ± 3.5 40 4.21 21.1
CD 73 HF/normal 14.4 ± 1.4 2.1 ± 0.1 3.1 ± 0.4 16 6.7 ± 1.2 18.1 4.7 7.9
2009 Gaur 15 Uniform/cosine 35.0 ± 2.5 3.6 ± 0.6 0.32 ± 0.1 36 1.2 ± 0.5 39.02 0.6 2.7
2010 18 HF/normal 34.2 ± 6.1 4.2 ± 0.4 0.8 ± 0.2 31 3.4 ± 1.1 32.8 1.8 6.6
2011 9 Uniform/cosine 49.0 ± 0.00 4.8 ± 1.1 0.5 ± 0.1 22 2.4 ± 0.9 40 1.1 5.4
CD 42 HF/normal 26.2 ± 5.3 2.9 ± 0.3 1.2 ± 0.2 24 3.5 ± 0.9 21.1 2.1 6.1
ESW – Estimated Strip Width, GS – Group Size, SE – Standard Error, SE – Standard Error, Dg – Group density/km
2
, D – Individual density/km
2
, %CV 3 – Percentage of Co-
efficient Variation.
21.63
40.07
21.28
17.02
23.9
41.8
18.6
15.7
0.00
5.00
10.00
15.00
20.00
25.00
30.00
35.00
40.00
45.00
Adult Male Adult
female
Sub adult Fawn
No.of Individuals (%)
Age/Sex
SAMBAR
BARKING DEER
Fig. 2. Group structure and composition of Sambar (n= 282) and Barking deer
(n= 280) in PTR from 2009–2011.
Table 3
Biomass density and total biomass estimates of major prey species in PTR (2009–
2011).
Species D Unit weight kg/km
2
Biomass (%)
Sambar 3.8 121.97 463.486 21.3
Muntjac 3.9 16.26 63.414 2.9
Wild pig 6.7 31.19 208.973 9.6
Gaur 3.5 413.34 1446.69 66.3
Total 17.9 2182.56
D – Individual prey density.
35%
31%
10%
20%
4%
Elephant
W.pig
Rodent
Primates
Other ungulate
Fig. 3. Crop damages by wild animals in PTR.
222 K. Muthamizh Selvan et al. / Acta Ecologica Sinica 34 (2014) 219–224
may attributed to the absence of large bodied ungulates such as
gaur, and also JSWNP comes under the temperate forest where as
PTR is influenced by tropical climate.
Large carnivore conservation is critically linked with prey
density which is the key determinant of large carnivore abundance
[12,32]. To conserve a viable carnivore population and reduce the
livestock depredation, wild prey species density needs to be man-
aged effectively [54]. The ungulate density and biomass in the
tropical forests have always been considered to be poor; however
such generalization may always hold true as several factors like
strict enforcement, absence of climate induced mortalities, pres-
ence of diverse flora due to bioclimatic variables and maintenance
of successional vegetation determine the prey densities and
biomass in any particular area. The estimated ungulate density
and biomass in Pakke Tiger Reserve is higher in comparison to
other reserves in such similar habitats. This is primarily due to
strict enforcement inside the park. The park ensures effective
enforcement and protection by having 1 antipoaching staff per
8.56 km
2
with a total of 24 antipoaching camps operating inside
the park [53]. Better compensation scheme for crop damages
would be helpful for minimizing the retaliatory killing of depredat-
ing ungulates by the local people.
Acknowledgements
We thank Dr. V.B. Mathur Director of WII and Dr. P.K. Mathur,
Dean FWS of WII for providing the Institutional support. We would
like to sincerely thank the Department of Science and Technology,
Govt. of India and the Rufford Small Grant Foundation for provid-
ing the necessary fund support. We thank Dr. A.J.T. Johnsingh
and Dr. Claudio Sillero, Chair IUCN canid SG for their generous
support. We sincerely thank the Department of Environment and
Forest, Government of Arunachal Pradesh, particularly to Mr. J.L.
Singh, Principal Chief Conservator of Forests and Chief Wildlife
Warden (PCCF and CWW); C. Loma, CCF; P. Ringu; and Tana Tapi,
DFO Pakke. We are grateful to our field assistants, Joli Weli and
Gangaram Chiri for their immense support in data collection.
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Gaur 3.5 9.4 9.6 0.5 0.7 A A NA 0.4 1.48 – A –
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