Journal of Threatened Taxa |
www.threatenedtaxa.org | 26 August 2021 | 13(9): 19239–19245
ISSN 0974-7907 (Online) | ISSN 0974-7893
(Print)
https://doi.org/10.11609/jott.4618.13.9.19239-19245
#4618 | Received 05 July 2019 | Final
received 03 June 2021 | Finally accepted 14 July 2021
Diet of Leopards Panthera pardus fusca inhabiting protected areas and human-dominated
landscapes in Goa, India
Bipin S. Phal Desai 1,
Avelyno D’Costa 2, M.K. Praveen Kumar
3 & S.K. Shyama 4
1–4 Department of Zoology, Goa
University, Taleigao Plateau, Goa 403206, India.
1 Goa Forest Department, Government
of Goa, Goa 403001, India.
1 phaldesaibipin00@gmail.com, 2
avelynodc@gmail.com, 3 here.praveen@gmail.com, 4 skshyama@gmail.com
(corresponding author)
Abstract: The diet of leopards occupying
human-dominated and protected areas (PAs) in Goa, India was analyzed
through scat analysis. A total of 117 scats, 55 from wildlife sanctuaries/
national parks and 62 from human-dominated areas were collected and analyzed. Analysis of 55 scats from protected forests
revealed the presence of only wild prey in the leopard diet, whereas 61% of
scats collected from human-dominated areas consisted of only wild prey, 29% of
domesticated animals, and 10% a mixture of both wild prey & domesticated
animals. Of the prey biomass consumed in human-dominated areas, domestic
animals constituted only 33% of the leopard diet. Among all leopard scats, 71%
contained only one prey species, 28% contained two species, and 1% contained
three.
Keywords: Diet composition, hair medullary
pattern analysis, human-leopard interactions, scat analysis.
Editor: Shomita Mukherjee, Salim Ali Centre for
Ornithology and Natural History, Coimbatore, India. Date
of publication: 26 August 2021 (online & print)
Citation: Desai, B.S.P., A. D’Costa,
M.K.P. Kumar & S.K. Shyama (2021). Diet of
Leopards Panthera pardus
fusca inhabiting protected areas and
human-dominated landscapes in Goa, India. Journal of Threatened Taxa 13(9): 19239–19245. https://doi.org/10.11609/jott.4618.13.9.19239-19245
Copyright: © Desai et al. 2021. Creative Commons Attribution
4.0 International License. JoTT allows unrestricted use, reproduction, and
distribution of this article in any medium by providing adequate credit to the
author(s) and the source of publication.
Funding: None.
Competing interests: The authors
declare no competing interests.
Author details: Mr. Bipin S. Phal
Desai is a Range
Forest Officer of the Goa Forest Department, Government of Goa. His research
interests include wild cat biology and man-wildlife conflict studies. Dr. Avelyno H.
D’Costa is an Assistant Professor in the Department of Zoology, Goa University. He is an ecotoxicologist
and is interested in research related to genetics, ecology and conservation. Dr. M.K. Praveen Kumar is is a toxicologist
and is interested in research related to genetics and biotechnology. Dr.
S.K. Shyama is a Professor (Rtd.) and ex. head
of the Department of Zoology, Goa University. His area of expertise is genotoxicology and interests include ecotoxicology and
wildlife biology. He has guided several PhD students in toxicology and wildlife
and conservation.
Author contributions: BSPD collected the samples,
analyzed them and wrote the manuscript.
AD’C assisted with the analysis of the samples and revision of the
manuscript. MKPK assisted with the analysis of the specimens. SKS supervised
the study and helped in the revision of the manuscript.
Acknowledgements: The authors wish to acknowledge
the support and assistance offered by the Forest Department of Goa, Government
of Goa.
Introduction
Big cats
play an important role in maintaining the equilibrium of forest ecosystems, for
which they serve as indicators of health and integrity. Tigers Panthera tigris and
Leopards Panthera pardus
are integral parts of forest ecosystems (Karanth
& Sunquist 1995) and hence their conservation is
of prime importance. Leopards are widely distributed in India and they often
come into conflict with humans, indeed they are more frequently involved in
human conflict than other large cats (Holland et al. 2018). Many examples have
been reported from Sanjay Gandhi National Park (Mumbai), Baria
Forest Division (Gujarat), Junnar (Maharashtra), and Garhwal (Himalaya), and conflicts are becoming increasingly
prominent with an increasing human population and expanding developments
leading to competition for shrinking resources. This presents major obstacles
to the conservation of leopards, and a comprehensive region-specific study of their
ecology and biology is essential for long-term conservation.
Several
studies have documented the widespread distribution of leopards across India
(Daniel 1996; Vijayan & Pati 2002; Athreya et al. 2013), but few studies have focused on prey
availability and diet composition in human-dominated areas (Athreya
et al. 2013, 2014). Hence in this study, an effort has been made to compare the
diet of leopards in human habitations with those living in PAs in Goa, India
using scat analysis. Scat analysis is an indirect, non-destructive and
cost-effective method (Sunquist
1981; Johnsingh 1983) for recording the frequency of occurrence of prey
items in the diet of a carnivore. The hair of prey is relatively undamaged in
scat of leopards and tigers, hence it can be used as a tool to identify prey
species (Mukherjee et al. 1994a; Ramakrishnan et al. 1999). However, there is a
chance of error if molecular methods are not used to confirm species identity (Laguardia et al. 2015; Akrim et
al. 2018).
In Goa, the
Western Ghats run along the eastern border of the state which contains
protected forest areas. In addition to this, there are various small hill
ranges and plateaus stretching from Pernem in the
north to Canacona in the south that connect the Western
Ghats with the coastal landscape. Most of the old human settlements are
situated at the base of these hills and plateaus. In the last decade or so,
these areas have become prone to encroachment due to expansion of cities,
towns, villages and roads. These hills and plateaus primarily consist of
stunted cashew trees, thorn scrub jungle and coarse grass with dense
semi-evergreen forest patches in between (Jadhav & Pati
2012), which support a variety of wildlife, such as the Indian Leopard Panthera pardus fusca, Golden Jackal Canis
aurius, Dhole Cuon alpinus, Gaur Bos gaurus, Sambar
Rusa unicolor, Chital Axis axis, Northern Red Muntjac Muntiacus
vaginalis, Wild Boar Sus scrofa, Indian
Chevrotain Moschiola indica,
Bonnet Macaque Macaca radiata, Gray Langur Semnopithecus hypoleucos, and Indian Crested Porcupine Hystrix indica.
In this
work we have studied the diet composition of leopards in PAs as well as
human-dominated areas in Goa over a period of three years by collection and
analysis of scats, to identify potential human conflicts due to livestock
depredation, and to formulate management interventions and mitigating measures.
Study
area
Goa is
spread over the hilly region of Western Ghats towards the east, coastal plains
towards the west, a midland region with laterite plateaus and low-lying river
basins. The study area consisted of the entire state of Goa lying in between
latitudes 15.480–14.435N and 74.201–73.403E which included human-dominated
areas, with reported presence of leopards and wildlife sanctuaries and national
parks covering a total area of ~1,748.05 km2 (Figure 1). The average
altitude of Goa is approximately 511 m. The total geographical area is 3,702 km2
of which 2,219 km2 is covered with forests and 1,224 km2
represents state-owned forests, of which 649 km2 have been declared
protected areas in the form of a national parks and wildlife sanctuaries. The
overall human population density in Goa is 394 persons per km2. Goa
receives an average annual rainfall of 3,300 mm, and the major forest types are
tropical wet evergreen, tropical semi-evergreen, tropical moist deciduous, and
littoral & swamp forests.
Materials and Methods
Field
collection of leopard scats
Leopards
and tigers prefer use of forest road and footpaths/trails to move around and
also as a mechanism of inter and intra species social communication, hence they
are likely to defecate along such paths (Smith et al. 1989; Karanth
et al. 2004). Scat samples measuring larger than 20 mm in diameter (measured
using a custom-made 20 mm diameter metal ring) were collected to avoid
non-leopard predator scats (Norton et al. 1986; Rabinowitz 1989). The presence
of tigers was only reported from Mhadei Wildlife
Sanctuary (WS), where leopard scats were differentiated from tiger scats based
on size, shape, diameter, coiling and constriction patterns, along with
ancillary evidence such as pugmarks, scrapes, and claw marks (Lovari et al. 2014; Laguardia et
al. 2015; Rostro-Garcia et al. 2018). It is important to note that there is a
chance of identification error since molecular techniques were not used.
Scat
collection from protected forest areas
A
preliminary scat survey was conducted to identify carnivore trails such as foot
paths and forest roads passing through the protected forest areas (wildlife
sanctuaries and a national park). A total of 25.5 km of forest roads were
sampled once every month with trained personnel.
Scat
collection from human-dominated areas
To identify
areas with potential human-leopard conflicts, complaint and rescue data was
collected from the Forest department of Goa for the years 2013–2016. Using this
data, areas prone to human-leopard interactions were identified. These areas
were later visited to study the geography, crop patterns and proximity with
dense forest areas of protected areas.
Preliminary
scat survey was conducted to identify carnivore trails such as foot paths and
unmetalled roads passing through areas having forested areas close to human
habitation (hamlets with small houses and fields) with maximum complaints on
leopards. These areas have medium to dense green cover and provide shelter to
leopards and wild prey species. Sampling at each site was carried out once a
month. A total of 34 km from such areas were sampled over a period of three
years from January 2016 to December 2018.
Scats were
measured and collected in polythene bags labeled with
the date of collection along with the GPS location of the site. Scat samples
were transported to the lab, sun dried and the dry weight recorded. A portion
of dried scats were soaked in water and passed through a metal sieve (1.5mm
mesh size), leaving only undigested prey remains which predominantly consisted
of hair and bone fragments. Hairs from the undigested remains were then separated
and 25 hair strands were randomly picked from each sample and analyzed. For hair medullary pattern analysis, hairs were
immersed in xylene for 24 hours and then mounted on permanent slides with a
cover slip using DPX mount (Mukherjee et al. 1994b). For observing cuticle
scale patterns of hair, an impression technique using gelatin
solution with eosin stain was used following Mukherjee et al. (1994b). Slides
were examined under 200x and 400x magnification based on the size of the hair
using a trinocular research microscope (Olympus BX53). A set of reference
slides were made from domesticated livestock and pet animals from the study
area and wild prey species in captivity, rescued animals and road kills.
Prey
species (identified from hair found in scat) were reported as the proportion of
scats that showed their presence. A species accumulation curve was also plotted
(Kshettry et al. 2018) to ascertain the number of
scats required to be analyzed for a reliable diet
estimate. To avoid bias due to variable prey body size, relative composition of
prey species varying in body size was calculated using Ackerman’s equation
(Ackerman et al. 1984), assuming leopards have similar digestive physiology to
Mountain Lions Puma concolor (Karanth & Sunquist 1995), as
follows:
Y=
1.980+0.035x
Where Y is
the kg of prey consumed per field collectible scat, and X is the average weight
of the particular prey species in kg (Ackerman et al. 1984). This method has
been used previously for leopards (Karanth & Sunquist 1995; Andheria et al.
2007; Khorozyan et al. 2008; Odden
& Wegge 2009; Mondal et al. 2012; Athreya et al. 2014). The body weights of probable predated
prey species were taken from literature (Mondal et al. 2012; Athreya et al. 2014).
The
relative biomass (D) and relative numbers (E) of each prey species consumed was
obtained using the equations:
D= (A x Y)/
∑(A x Y) x100
E= (D/X)/
∑(D/X) x 100
Where A is
the frequency of occurrence of the prey items in the scats, Y is the mass of
prey consumed per scat (kg) and X is the mean mass of the prey (kg) (Athreya et al. 2014).
Results
Protected
areas
Analysis of
scats collected from PAs revealed the presence of only wild prey (Indian
Crested Porcupine, Wild Boar, Northern Red Muntjac, Chital, Indian Hare Lepus
nigricollis, Bonnet Macaque, and Gray Langur) in the diet of leopards. No records of
domesticated animals (such as ox, dog, pig, goat, and cat) were found in the
scats from protected forest areas. Scat analysis of 55 scats collected from
these areas (Table 1) revealed that Wild Boar constituted a major proportion of
the prey biomass (29%), followed by Chital (25%), Indian Crested Porcupine
(15%), Barking Deer (13%), Gray Langur (5.6%), Bonnet
Macaque (5.4%), Sambar (4.1%), and Indian Hare (3.1%). Indian Hare was the most
preyed-upon species in relative numbers (21%) followed by the Indian Crested
Porcupine (18%), Bonnet Macaque (15%), Wild Boar (13%), Gray
Langur (12%), Northern Red Muntjac (11%), Chital (8.9%), and Sambar (1.1%). The
diet profile analysis also suggests that leopards preferred small-sized prey
(77%), over medium (33%), and large-sized prey (1.1%) (Table 1).
Human-dominated
areas
The results
of analysis of 62 scats collected from human-dominated areas revealed that
major proportion of leopard prey biomass comprised of wild prey (67%),
predominantly Wild Boar (26%), Indian Crested Porcupine (17%), Indian Hare
(14%), Bonnet Macaque (5.1%), Gray Langur (3.2%),
and Northern Red Muntjac (1.3%). Domestic animals (dog, pig, cat, and goat)
constituted only a minor portion (33%) of the leopard diet. The dog was the
most preyed-upon domestic animal (17%) followed by pig (11%), goat (2.7%), and
cat (2%) (Table 2). Of the nine wild prey species observed from scat analysis,
six were identified in scats collected from human-dominated areas.
Comparative
analysis of leopard habitats
A total of
117 leopard scats were collected during the period of the study, of which 55
were from PAs and 62 from
human-dominated areas; 62% of scats collected from human-dominated areas contained
only wild prey, 29% only domestic prey, and 9.7% had a mixture of both. A
majority of scats (71%) contained only one prey species, 28% contained two
species and 0.85% contained three (Figure 2). A total of 151 prey items were
identified, comprising of 12 prey species.
In both
habitats, Indian Hare remains were observed in the most scats (42%), followed
by Indian Crested Porcupine (13%) and Wild Boar (8.7%). Of the total prey biomass consumed,
domesticated animals constituted a minor fraction (17%) of the leopard diet,
the remainder consisted of Wild Boar (27%), Indian Crested Porcupine (16%),
Chital (12%), Indian Hare (8.9%), Northern Red Muntjac (7.1 %), Bonnet Macaque
(5.2%), Gray Langur (4.3%), and Sambar (2%).
The
cumulative curve (Figure 3) suggested that the proportion of scats with remains
of various prey species stabilized after 24 scats, with only one species being
added after 71 scats. From this analysis it can be also interpreted that 92% of
prey species were identified in the first 24 scats analyzed,
with an addition of just one species (cat) after analysis of the 71st
scat. Thus we consider the sample size adequate to interpret the overall diet
profile of leopards from the study area.
Discussion
The presence
of eight wild prey species in the study area may be attributed to the
availability of diverse vegetation from dry thorn forests to semi-evergreen
forests. The presence of leopards in human habitations is evident from the data
collected from the forest department regarding complaints received from a
majority of the human-dominated areas of North Goa and South Goa districts.
Hence it is likely that Leopards may be distributed throughout the state of
Goa.
Data
analysis from protected forest areas suggests that leopards consumed medium-
(Wild Boar, Northern Red Muntjac, Chital) and small-sized prey (Indian Hare,
Indian Crested Porcupine, Bonnet Macaque, Gray
Langur). The Indian Hare was found in the most scats, followed by Indian
Crested Porcupine, Wild Boar, Gray Langur, Bonnet
Macaque, Northern Red Muntjac, Chital, and Sambar. The preference of smaller to
medium-sized prey was also reported in the studies of Sunquist
& Sunquist (1989), Sankar
& Johnsingh (2002), Henschel et al. (2005), and
Ahmed & Khan (2008). Additionally, this preference may also be due to the
nocturnal feeding behaviour of Leopards as well as these small mammals, thus
making them more vulnerable to predation than the other species (Ahmed &
Khan 2008). Another point to be considered is that in the study area, wild prey
were also found to be present in human-dominated areas for the purpose of
grazing or foraging which could also be a reason for the leopards entering
these areas.
With regard
to the domestic animals, the predation of dogs and pigs was mostly due to an
increase in stray dog and pig population in human habitation probably due to
improper disposal of garbage in these areas. Very few households had a safe
night shelter for their domesticated pigs and dogs. During the study although
few complaints of leopard attacks on cattle calves were reported, no such
killings were found. Further no traces of cattle hair were found in any of the
scat samples.
From
informal observations and discussions with locals we realized that though
leopards came into conflict with humans almost throughout the year, this
conflict is significantly higher during the months of August, September, and
October and again intensifies in the months of January and February. This
pattern correlates with the breeding pattern of leopards (pre-breeding phase
during the monsoon months of August, September, and October) when wandering
males and sub-adult cubs (which have just left their mothers to fend for
themselves) come in conflict with humans. The conflict during the January and
February months could be mainly due to the movement of females in the
post-birth phase. These leopards, which continuously change their location for
the safety of the young cubs, come in contact with humans employed in cashew
plantations and other agricultural activities.
Conclusion
It can be
interpreted from our data that although leopards were reported close to human
habitations throughout the year, their dependence on domestic animals was low.
This study also indicates that the wild species that the leopards preyed upon
in PAs were also present in forested areas close to human habitations. This
could be the reason for the presence of leopards in human-dominated areas with
a low dependence on domestic animals. Hence it is of utmost importance to
create awareness about the role of these large cats in ecosystems and their
feeding and behavioral patterns, and to adopt
mitigating and precautionary methods in case of human-leopard conflicts.
Table 1. Diet composition of Leopards inhabiting
protected forest areas in Goa through analysis of scat samples (55) during
January 2016 to December 2018.
|
Prey species |
N (Percent occurrence) |
Average body weight (X) |
A (%) (Percent frequency) |
Y (Kg/scat) |
D (%) (Relative biomass) |
E (%) (Relative number of individuals consumed ) |
1 |
Indian Crested Porcupine |
12 |
14 |
17.39 |
2.47 |
14.57 |
17.69 |
2 |
Wild Boar |
18 |
37 |
26.08 |
3.27 |
28.98 |
13.31 |
3 |
Northern Red Muntjac |
10 |
20 |
14.49 |
2.68 |
13.18 |
11.19 |
4 |
Gray Langur |
5 |
8 |
7.25 |
2.26 |
5.56 |
11.80 |
5 |
Indian Hare |
3 |
2.5 |
4.35 |
2.07 |
3.05 |
20.73 |
6 |
Bonnet Macaque |
5 |
6 |
7.25 |
2.19 |
5.38 |
15.25 |
7 |
Dog |
0 |
18 |
0 |
2.61 |
0 |
0 |
8 |
Chital |
14 |
48 |
20.29 |
3.66 |
25.19 |
8.92 |
9 |
Pig |
0 |
30 |
0 |
3.03 |
0 |
0 |
10 |
Sambar |
2 |
62 |
2.90 |
4.15 |
4.08 |
1.12 |
11 |
Cat |
0 |
3.5 |
0 |
2.10 |
0 |
0 |
12 |
Goat |
0 |
25 |
0 |
2.85 |
0 |
0 |
Total |
69 |
|
|
|
|
|
Y—estimated
weight of the prey consumed per collectable scat produced.
Table 2. Diet composition of Leopards inhabiting
human-dominated areas in Goa through analysis of scat samples (62) during
January 2016 to December 2018
|
Prey species |
Percent occurrence (N) |
Average body weight (X) |
A (%) (Percent frequency) |
Y (Kg/scat) |
D (%) (Relative biomass) |
E (%) (Relative number of individuals consumed ) |
1 |
Indian Crested Porcupine |
15 |
14 |
18.29 |
2.47 |
17.24 |
11.20 |
2 |
Wild Boar |
17 |
37 |
20.73 |
3.27 |
25.91 |
6.37 |
3 |
Northern Red Muntjac |
1 |
20 |
1.22 |
2.68 |
1.25 |
0.57 |
4 |
Gray Langur |
3 |
8 |
3.66 |
2.26 |
3.15 |
3.59 |
5 |
Indian Hare |
15 |
2.5 |
18.29 |
2.07 |
14.43 |
52.49 |
6 |
Bonnet Macaque |
5 |
6 |
6.10 |
2.19 |
5.10 |
7.72 |
7 |
Dog |
14 |
18 |
17.07 |
2.61 |
17.01 |
8.59 |
8 |
Chital |
0 |
48 |
0 |
3.66 |
0 |
0 |
9 |
Pig |
8 |
30 |
9.76 |
3.03 |
11.28 |
3.42 |
10 |
Sambar |
0 |
62 |
0 |
4.15 |
0 |
0 |
11 |
Cat |
2 |
3.5 |
2.44 |
2.10 |
1.96 |
5.08 |
11 |
Goat |
2 |
25 |
2.44 |
2.85 |
2.66 |
0.97 |
Total |
82 |
|
|
|
|
|
Y—estimated
weight of the prey consumed per collectable scat produced.
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