Notes on the
diet and habitat selection ofthe Sri Lankan Leopard Panthera pardus
kotiya (Mammalia: Felidae) in the central
highlands of Sri Lanka
Andrew M. Kittle1, Anjali C. Watson 2, P.H.S. ChanakaKumara 3, S.D. Kashmi C. Sandanayake 4, H.K. Nimalka Sanjeewani 5 & T. SamindaP. Fernando 6
1,2,3,4,5,6 The Leopard
Project, Wilderness & Wildlife Conservation Trust, 130 Reid Avenue, Colombo
4, Sri Lanka
1 Department of Integrative Biology, University of Guelph,
Guelph, Ontario, Canada
3,4,5 Department of
Natural Resources, Faculty of Applied Sciences, SabaragamuwaUniversity of Sri Lanka, Belihul Oya,
Sri Lanka
6 Department of Zoology,
University of Colombo, Colombo, Sri Lanka
1 akittle@uoguelph.ca
(corresponding author), 2 aalanka@sltnet.lk, 3 greatlife.lk@gmail.com,4 kcsandanayake@gmail.com, 5 nimalka.sanjeewani@gmail.com,6 samijaela@yahoo.com
Abstract: The endangered
Sri Lankan Leopard Panthera pardus kotiya occupies the
island’s highly fragmented central hills where data on its feeding ecology and
habitat use is largely absent. This
study’s objective was to investigate diet and resource selection of leopards
here with a focus on the extent of potential interactions with humans in this
heavily populated, largely unprotected landscape. Fecal sample analysis was undertaken to
investigate diet and sign index counts and selectivity index analysis to
determine habitat and landscape features important to fine scale leopard
utilization. Results indicated that
leopards in the central hills hunt a wide range of prey (at least 10 genera),
including larger species where available (e.g., Sambar Rusa unicolor) and smaller, more
specialized prey (e.g., Porcupine Hystrix indica) where necessary. No domestic species were recorded in
scat analysis (N=35) despite the availability of dogs Canis familiaris, suggesting such predation may be
atypical in Sri Lanka. Leopards use a range of landscapes within the region
including established and regenerating forests, plantation lands (e.g., pine,
eucalyptus, tea), and areas in close proximity to human settlement. At a fine scale, areas of dense
undergrowth including tall grasslands were preferred to more open forest, as
well as to Pine Pinus caribaeamonocultures. Avoidance of humans
may be influencing these patterns. This study has important implications as
researchers and managers necessarily expand beyond focusing on protected areas
toward integrated, landscape-level conservation strategies.
Keywords: Anthropogenic
disturbance, diet, habitat use, Panthera pardus kotiya, unprotected
areas.
doi: http://dx.doi.org/10.11609/JoTT.o3731.6214-21
Editor: Shomita Mukherjee, SACON,Coimbatore, India. Date
of publication: 26 August 2014 (online & print)
Manuscript details: Ms # o3731 |
Received 02 August 2013 | Final received 18 April 2014 | Finally accepted 11
August 2014
Citation: Kittle, A.M., A.C. Watson, P.H.S.C. Kumara, S.D.K.C. Sandanayake, H.K.N. Sanjeewani& T.S.P. Fernando (2014). Notes on the diet and habitat selection of the Sri
Lankan Leopard Panthera pardus kotiya (Mammalia: Felidae)
in the central highlands of Sri Lanka. Journal of Threatened
Taxa 6(9): 6214–6221; http://dx.doi.org/10.11609/JoTT.o3731.6214-21
Copyright: © Kittle et al. 2014. Creative
Commons Attribution 4.0 International License. JoTTallows unrestricted use of this article in any medium, reproduction and
distribution by providing adequate credit to the authors and the source of
publication.
Funding: PTES (People’s
Trust for Endangered Species) UK grant and contributions from CERZA
Conservation, France and Le parc des Félins, France.
Competing Interest: The authors
declare no competing interests.
Author Contribution: AMK supervised data collection, analyzed data
and wrote the paper; ACW supervised data collection and wrote the paper; PHSCK
collected data and conducted laboratory analysis; SDKCS collected and analyzed
data; HKNS collected data and coordinated map design; TSPF conducted laboratory
analysis.
Author Details: Andrew M. Kittle is interested in
ecological research which can be used to inform
management and conservation policy. His main focus is carnivore behaviour and ecology, particularly spatial dynamics and
resource selection, as well as how these relate to carnivore co-existence. Anjali C. Watson is interested in the
ecology and conservation of the Sri Lankan Leopard as well as broader issues
including wildlife and ecosystem health. P.H.S.Chanaka Kumara is interested in conservation
biology. S.D. Kashmi C. Sandanayakeis undertaking her MPhil at the Post-Graduate Institute of Agriculture,
University of Peradeniya where her primary interests
are conservation biology and environmental science. H.K. Nimalka Sanjeewaniis interested and engaged in the conservation of Sri Lanka’s wildlife. T. Saminda P.
Fernando has an interest in wildlife ecology and parasitology
and is currently lecturing at the Open University of Sri Lanka.
Acknowledgements: We would like to thank the Kandy Municipal Council for permits andHemantha Ekanayake and
staff for support at Dunumadallawa forest reserve; Rohan Pethiyagoda for encouraging
and supporting our work in the Agrapatana Arboretum;
and Sri Lankan Department of Wildlife Conservation and Forestry Department for
continued support and permits. Laboratory space was generously provided by
Colombo University in liaison with Dr. Devaka Weerakoon.
For figures, images, tables -- click here
Introduction
The Sri Lankan
Leopard Panthera pardus kotiya is an Endangered (Kittle
& Watson 2008) endemic sub-species and the island’s apex predator (Miththapala et al. 1996; Uphyrkinaet al. 2001). It has evolved without intra-guild competition at least since Sri
Lanka split off from the Indian subcontinent ~5,000–10,000 ybp (Deraniyagala 1992; Yokoyama
et al. 2000), a unique scenario which may mean that this top carnivore is of
keystone importance to the system (Paine 1966).
Data on the
island-wide distribution and relative abundance of leopards is becoming
increasingly refined (Kittle & Watson 2008; Kittle et al. 2012) and it is
now acknowledged that these versatile predators occupy unprotected as well as
protected areas (Watson & Kittle 2004). Between these extremes are
semi-protected areas, small forest reserves and sanctuaries demarcated on paper
but not actively patrolled and routinely utilized by people. Wildlife populations in unprotected and
semi-protected areas are particularly susceptible to heightened levels of
human-wildlife conflict.
Sri Lanka is a
small (65,610km²) mostly rural (74–78 %) nation with substantial
spatial overlap between humans and wildlife (Dept of
Census and Statistics 2012). In the
central hills, high human population density (301–450 /km², Dept of Census and Statistics 2012), heterogeneous land
cover and a lack of protected areas makes understanding leopard diet and
resource selection vital. Minimal diet analysis has been conducted here (Ranawana et al. 1998) and leopard diet outside protected
areas remains unknown. Furthermore,
how leopards use this heavily fragmented habitat mosaic has not been investigated. This data deficiency has been recognized
as a barrier to designing conservation and management action plans to ensure
the leopard’s continued survival in the wild (Samarakoon1999). A minimum of 66 leopards
were killed island-wide from 2002–2012, mostly caught in snares set for
other species (Kittle & Watson, unpublished data). True mortality figures are unknown but
if leopards are preying on domestic species outside protected areas,
human-leopard conflict is likely to increase.
Our research
objective was to investigate leopard diet and habitat use at two semi- to
un-protected study sites in the central hills where leopards have been
previously documented (Kittle et al. 2012).
Study Areas
The central
highlands occupy a ~2400km² section of Sri Lanka’s south-central interior
(Fig. 1), encompassing a vast expanse of often-rugged terrain, which varies
greatly in elevation, topography and climate.
The Dunumadalawa Forest Reserve is a secondary growth
mid-country wet zone forest situated at the northern end of Sri Lanka’s Central
massif (Figs. 1, 2a). This was an
active estate until the early 1900s when it was released from plantation use
(tea, coffee and cocoa) and allowed to naturally re-generate, with some active
reforestation of native plant species since 2000. It now protects the watershed of two
reservoirs providing water for Kandy, a town of ~100 000 within the limits of
which the reserve is located. Dunumadalawa is
characterized by high canopy mixed forest whichincludes Albizia sp. and jak Artocarpus heterophyllus remaining from the estate days. Large-leaf Mahogany Sweitinia macrophylla, Ceylon Almond Canarium zeylanicum and various Ficusspecies are also conspicuous. The
understory is dominated by unattended tea, coffee and cocoa plants grown wild
in sections and the ironwood tree Mesua ferrea in others. Patches of grassland, composed mostly of Imperata cylindrica are scattered throughout the reserve.
This grass can grow up to 3m tall and form dense monocultures. The abundant jakappears to provide an important continuous food source for potential leopard
prey species including Toque Macaques Macaca sinica aurifrons,
Barking Deer Muntiakus muntjak,
Wild Boar Sus scrofaand Porcupines Hystrix indica. The reserve includes a small (<40ha)
Caribbean Pine Pinus caribaea plantation at its highest point in the southeast. Anthropogenic disturbance is acute in
the surrounding area which is characterized by a mosaic of land-use types
including sub-montane forest, tea estates, pine and
eucalyptus plantations, open scrub, home gardens and villages. The reserve
falls under the purview of the Kandy Municipal Council and is extensively
utilized by a number of surrounding villages for firewood and fruit collection.
The Agrapatana Arboreatum is in the
heart of the tea industry’s vast plantation lands near the southern extent of
the central hills (Figs. 1, 2b; Image 1). A regenerating forest, this was part of a much larger tea estate until
the 1990s. It adjoins the Agra-Bopats Forest Reserve, a thickly vegetated 9,800ha sub-montane, montane and cloud forest
bordered by tea estates on three sides and connected to Horton Plains National
Park (3160ha) to the southeast. Sambar Deer Rusa unicolor, Wild Boar Sus Scrofa, Toque Macaque Macaca sinica, Purple-faced Langur Trachypithecus vetulusand Porcupine all reside within this forest reserve. The two study areas differ in a number
of important aspects (Table 1).
Methods
Diet: Leopard scat was collected opportunistically and during regular
sign index surveys (Rabinowitz 1997) conducted
between 2003 and 2009. These were
air dried and stored in re-sealable plastic bags. To differentiate leopard from Fishing
Cat Prionailurus viverrinusscat only samples with bolus width >2.5cm were retained (Henschel& Ray 2003). Samples were
washed and sieved to eliminate unwanted particles, and bones, quills, nails and
hooves were manually separated and identified. Remaining hairs were mixed and oven
dried at 600C for 24hr. Dried samples were placed in a shallow dish, a point grid overlaid and
50 hairs randomly sampled (Ciucci et al. 2004). Microscopic analysis followed Amerasinghe et al. (1990).
Habitat selection: We investigated fine scale leopard habitat selection
at Dunumadallawa by comparing habitat variables at
used and available locations (Table 2). Used locations were defined by the presence of leopard spoor (scat,
pugmarks, scrapes) on reserve trails detected during regular sign index surveys
between 2003 and 2009. Available
locations were determined from a systematic random sample conducted in November
2009, with points taken every 100m along the same reserve trails. Four trails (1300–2000 m) that
traverse the forest reserve were utilized for the use/availability analysis.
We used Jacob’s
selectivity index to determine habitat selection where data was categorical
(Fig. 3):
D =(r – p)
/(r + p - 2rp)
where r is the
proportion of used locations of a given habitat/landscape class and p is the
proportional availability of that habitat/landscape class (Jacobs 1974). Values of D range from -1 indicating
maximum avoidance to 1 indicating maximum selection, with 0 value indicating
use in proportion to availability (Jacobs 1974). This selectivity index minimizes bias
from small sample sizes (e.g., rare food items or habitat proportions <10%;
Hayward & Kerley 2005). Continuous data was not normally
distributed but had equal variance (Levene’s test
P>0.05) so Mann-Whitney Test was used for analysis (Zar1999).
Results
Diet: Of the 42 scat samples collected, 35 produced results: 13 from Hantane and 22 from Agrapatana(Table 3). Porcupine was the most
frequently represented species found in scat at Dunumadalawa,
followed by Barking Deer, whereas Sambar was the most
common species in leopard scat in Agrapatana (Table
3). Domestic species were not
represented in samples from either location (Table 3).
Habitat selection: Thirty-nine used locations were compared to 68
available locations within Dunumadalawa. Leopards utilized the entire 5km²
study area but showed selection for grasslands (Jacob’s Index = 0.9963) and
avoidance of the Pinus plantation (Jacob’s
Index = -0.9867, Fig. 2a). Areas
with dense understory (Jacob’s Index = 0.8158) and no canopy cover (Jacob’s
Index = 0.7143) were selected, whereas areas with sparse canopy cover
(0–25 %) were avoided (Jacob’s Index = -0.6277). Areas far from permanent water (Jacob’s
Index = 0.5790) were preferred to areas close to permanent water (Jacob’s Index
= -0.5014, Fig. 2b).
Neither slope (Zw=0.537, p= 0.5907) or elevation (Zw=1.405, p= 0.1611) were significant factors
influencing fine scale habitat use, however leopards did prefer areas with
lower tree density (Zw=-2.443, p=.0147) and larger trees (Zw=-1.919, p= 0.0554).
Discussion
Diet: Sample size is
too small to definitively characterize leopard diet in these areas (Mukherjee
et al. 1994; Trites & Joy 2005), but given the
complete dearth of dietary data from un-protected areas in the central
highlands, even this preliminary information is valuable. The dietary breadth observed is wide,
including 10 genera ranging in size from 112g (Palm Squirrel; Phillips 1980) to
130kg (Sambar hind; Phillips 1984). This catholic
diet is consistent with leopard feeding ecology throughout its range reflecting
opportunistic predation and the leopard’s adaptive ability to broaden its diet
beyond the preferred range of medium-sized ungulates, 23–56 kg (Stander
et al. 1997; Hayward et al. 2006).
The Indian Porcupine was represented most frequently in scat
(61.5%) in the small, isolated forest reserve despite it being absent from
Hayward’s (2006) exhaustive analysis of leopard prey preferences (111 prey
species). Porcupines are
potentially dangerous prey, but the prey suite available to leopards in Dunumadalawa is relatively limited (Sanjeewani2010), so the likelihood of risking injury to secure adequate resources is
increased (Brown & Kotler 2007). Alternately, leopards here may have
adapted to hunting porcupines with minimum risk or a single leopard may be
responsible for the majority of the scat samples collected here, although three
individuals have been observed in the reserve (Kittle et al. 2012). Prey specialization by individuals
within a population is not uncommon across taxa (Araújoet al. 2011) including among apex terrestrial carnivores (Ross et al. 1997; Elbroch & Wittmer 2013). Establishing whether observations are
the reflection of a population-level adaptation to the suite of resources
available or a case of individual specialization requires additional
sampling. The implications of
either can be profound (Bolnick et al. 2003).
In Agrapatana Sambarwas most frequently represented in scat samples (59.1%), consistent with
probable prey availability, given that the study area is contiguous with Horton
Plains National Park (HPNP) where sambar density is
estimated to be 66.5/km² (Rajapakse 2003). Previous scat analysis from HPNP found sambar in 75.8% of samples (N=22) (Ranawanaet al. 1998). That fewer samples
contained Sambar here than in HPNP, and more
contained Black-naped Hare and Purple-faced Langurs (Table 3; 13.6% compared to 6.8% and 3.4% in HPNP, Ranawana et al. 1998) hints at variation in prey
availability between the core of a protected area such as HPNP and an adjoining
yet peripheral area such as Agrapatana. More samples are needed to validate this
hypothesis.
Scat samples included no domestic species at either site despite
dogs being common at both. This
suggests low human-leopard conflict in these regions despite close proximity to
human populations, but given the low sample size, interpretations must be made
with caution. In Dunumadalawa dogs have been predated in the past, however,these incidents may be uncommon. Questionnaire surveys here show fishing cats responsible for some predation events
blamed on leopards (Kittle & Watson, unpublished data). That predation on domestic animals
appears atypical is relevant from a management perspective since predators
viewed as a menace tend to be increasingly persecuted (Cavalcantiet al. 2010).
Habitat selection: Leopards in the Dunumadalawa Forest Reserve may be selecting fine scale
habitats offering improved hunting efficiency. Leopards and lions select areas of high
prey vulnerability not high prey density at the fine scale (Hopcraftet al. 2005; Balme et al. 2007; Davidson et al.
2012). As ambush predators,
leopards require at least 20cm of cover for concealment (Bothma& LeRiche 1984), so tall grasses (>30cm) and
thick vegetation are sufficient. However, in South Africa hunting leopards select intermediate cover,
presumably because dense cover obscures their perception of prey (Balme et al. 2007). Alternatively then, where anthropogenic disturbance is high such as in
isolated forests within densely populated urban areas, dense undergrowth and
long grass might provide leopards concealment from people, a potential risk in
this environment. Avoidance of
permanent water may also indicate avoidance of people as the
reserve’s two large permanent water sources are regularly monitored by Water
Department personnel. However, leopards did not avoid other high human-use areas (e.g., border
villages). The reserve is replete
with seasonal streams and pools which allow leopards
to effectively exploit areas far from permanent sources.
In summary, this research provides valuable insight into the diet
and habitat selection of an endangered apex predator, in an area of its range with
little existing information and where it is afforded limited protection.
Avoidance of humans in these heavily disturbed landscapes may be an important
consideration for leopards when making space-use decisions, as it is in parts
of their African range where bush meat is an important commodity and humans and
leopards engage in a form of exploitative competition (Henschelet al. 2011). Camera trap images
revealed leopards and humans engaged in temporal niche partitioning in both
sites, with humans detected in the day and leopards at night at the same
location (Kittle et al. 2012). Temporal niche partitioning is a mechanism used to explain carnivore
co-existence in Africa (Hayward & Slotow 2009)
and might similarly be important in South Asia where top carnivores necessarily
share space with humans (Athreya et al. 2013).
As human populations continue to increase and protected areas
become more and more isolated (DeFries et al. 2005),
the effective conservation of biodiversity becomes increasingly reliant on
processes outside protected areas (Hansen & DeFries2007). These areas tend towards
habitat mosaics with forested areas heavily fragmented and surrounded by a
patchwork of alternate land use types. Large, wide-ranging species such as top carnivores nevertheless often
persist (Lyra-Jorge et al. 2008) and can be adept at
exploiting semi and unprotected areas even where human density is high (Athreya at al. 2013). We are increasingly recognizing the value of these sub-optimal, shared
habitats in terms of connectivity and as supplements to remaining forest
fragments (Caryl et al. 2012), as well as
appreciating the important role played by predators in maintaining biodiversity
within such fragments (Terborgh et al. 2001). As such, the need to increase our understanding
of how apex predators utilize these landscape mosaics is of paramount
importance.
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