Livestock losses and hotspots of attack from tigers and leopards in Kanha Tiger Reserve, Central India

Abstract

Carnivore attacks on livestock are a primary driver of human–carnivore conflict and carnivore decline globally. Livestock depredation is particularly threatening to carnivore conservation in Central India, a priority landscape and stronghold for the endangered tiger. To strengthen the effectiveness of conflict mitigation strategies, we examined the spatial and temporal patterns and physical characteristics of livestock depredation in Kanha Tiger Reserve. We combined livestock compensation historical records (2001–2009) with ground surveys (2011–2012) and carnivore scat to identify when and where livestock species were most vulnerable. Between 400 and 600 livestock were reported for financial compensation each year, and most (91–95 %) were successfully reimbursed. Tigers and leopards were responsible for nearly all livestock losses and most often killed in the afternoon and early evening. Cattle and buffalo were most at risk in dense forests away from villages and roads, whereas goats were most often killed in open vegetation near villages. A spatial predation risk model for cattle revealed high-risk hotspots around the core zone boundary, confirming the significant risks to livestock grazing illegally in the core. Such ecological insights on carnivore–livestock interactions may help improve species-specific livestock husbandry for minimizing livestock losses and enabling coexistence between people and carnivores.

Full text

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Reg Environ Change (2016) 16 (Suppl 1):S17–S29
DOI 10.1007/s10113-015-0871-5
http://link.springer.com/article/10.1007/s10113-015-0871-5
Spatial and temporal patterns of livestock losses and hotspots of attack from tigers and
leopards in Kanha Tiger Reserve, central India
Jennifer R. B. Miller1,2*, Yadvendradev V. Jhala2 and Jyotirmay Jena3
1School of Forestry & Environmental Studies, Yale University, New Haven, Connecticut, 06511.
Current affiliation: Panthera, Lion and Leopard Programs, 8 West 40th Street, 18th Floor, New
York, NY 10018, USA
2Wildlife Institute of India, Dehradun, Uttarakhand, India, 248001
3Satpuda Maikal Landscape Programme, WWF-India, Mandla, Madhya Pradesh, India, 481661
*Corresponding author:
Jennifer R. B. Miller, Panthera, Lion and Leopard Programs, 8 West 40th Street, 18th Floor,
New York, NY 10018, USA. jmiller@panthera.org
Abstract
Carnivore attacks on livestock are a primary driver of human-carnivore conflict and carnivore
decline globally. Livestock depredation is particularly threatening to carnivore conservation in
central India, a priority landscape and stronghold for the endangered tiger. To strengthen the
effectiveness of conflict mitigation strategies, we examined the spatial and temporal patterns and
physical characteristics of livestock depredation in Kanha Tiger Reserve. We combined livestock
compensation historical records (2001-2009) with ground surveys (2011-2012) and carnivore
scat to identify when and where livestock species were most vulnerable. Between 400-600
livestock were reported for financial compensation each year and most (91-95%) were
successfully reimbursed. Tigers and leopards were responsible for nearly all livestock losses and
most often killed in the afternoon and early evening. Cattle and buffalo were most at risk in
dense forests away from villages and roads, whereas goats were most often killed in open
vegetation near villages. A spatial predation risk model for cattle revealed high-risk hotspots
around the core zone boundary, confirming the significant risks to livestock grazing illegally in
the core. Such ecological insights on carnivore-livestock interactions may help improve species-
specific livestock husbandry for minimizing livestock losses and enabling coexistence between
people and carnivores.
Keywords
Carnivore conservation; hotspot predation risk map; human-carnivore conflict; kill site; livestock
depredation; livestock compensation

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Introduction
Large carnivore populations worldwide are rapidly declining, in part due to retaliatory killing by
livestock owners following attacks on domestic animals (Woodroffe et al. 2005; Ripple et al.
2014). Much of this human-carnivore conflict occurs at the edges of protected areas where
carnivores, livestock, and people overlap (Woodroffe and Ginsberg 1998; Nyhus and Tilson
2004). Many non-lethal techniques exist to help reduce livestock and livelihood losses, including
livestock husbandry strategies, physical deterrents, and financial incentives for communities
(Treves and Karanth 2003; Shivik 2006). Yet effective implementation of these tools requires
detailed knowledge of when and where carnivores attack livestock and how risk differs between
livestock species. Understanding carnivore-livestock interactions is a crucial step towards
mitigating human-carnivore conflict and ultimately enabling coexistence between people and
carnivores (Treves and Karanth 2003; Goodrich 2010).
Ecological insights on the environmental factors and animal behaviors that lead to carnivore
depredation on particular livestock are particularly useful for strengthening livestock husbandry
techniques (Wikramanayake et al. 1998; Miller 2015). Many previous studies on human-
carnivore conflict have focused on depredation by a single carnivore species (usually a high-
priority species of conservation concern) on all livestock species generally, which can obscure
unique risk gradients for individual livestock species from specific carnivores (Treves et al.
2011; Lichtenfeld et al. 2014; Athreya et al. 2014; Miller et al. 2015). Differences between the
body sizes, anti-predator defenses, and grazing requirements of livestock species result in distinct
levels of vulnerability to wild carnivores (Seidensticker 1976; Sinclair et al. 2003). For instance,
in many areas smaller large carnivores like leopards, hyenas and wild dogs primarily kill
smaller-bodied livestock such as calves, sheep, and goats, whereas the largest carnivores like
tigers target larger-bodied livestock such as adult cattle, buffalo, and horses (Sangay and Vernes
2008). Likewise, large carnivore species employ unique hunting strategies and segregation
tactics to avoid interspecific competition that results in risks for livestock at different times and
locations (Laundré et al. 2009). For example, tigers and leopards often segregate temporally or
spatially to minimize interference competition (Odden et al. 2010; Harihar et al. 2011; Lovari et
al. 2013), with tigers mostly attacking livestock at night and in forest while leopards attacked in
open agricultural areas in mid-day (Katel et al. 2014; Malviya and Ramesh 2015). Understanding
the temporal and spatial patterns of interactions between different livestock and carnivores
species is necessary for developing ecologically informed strategies for conflict mitigation.
We focused our study in Kanha Tiger Reserve, a protected area in Madhya Pradesh, India
where 18% of households lose livestock to wild carnivores, primarily tigers (Panthera tigris) and
leopards (Panthera pardus; Karanth et al. 2013). Kanha also serves as a source site for tiger and
leopard populations throughout the central Indian landscape (Dutta et al. 2013; Sharma et al.
2013) and is thus a priority region for minimizing human-carnivore conflict. In Kanha, tigers and
leopards mainly kill cattle (Bos indicus), buffalo (Bubalus bubalis), and goats (Captra aegagrus
hircus; Kanha Tiger Reserve Forest Department 2012). The Forest Department permits people to

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graze livestock in the multiple-use buffer zone but bans grazing inside the interior core zone of
the park, except by livestock from several villages located in the core.
Local livestock owners implement distinct grazing regimes depending on the livestock
species and season, which reflect tradeoffs between livestock vulnerability to carnivores, herder
costs, and other environmental factors. Cattle and buffalo, which often graze side-by-side in
groups, are allowed to free-graze without a herder in the winter and summer months (November-
June). In the monsoon (July-October), herders accompany cattle and buffalo to prevent livestock
from consuming crops. In contrast, herders accompany goats year-round because goats can
browse on the low-quality forage around villages, are more vulnerable to wild carnivores, and
tend to wander off if unsupervised. These different temporal and spatial patterns of grazing
suggest that cattle and buffalo may experience different threats from carnivores than goats.
However, few studies have assessed how risk varies between domestic prey species, and this
information is not available for central India despite its importance as a Tiger Conservation
Landscape for the protection of the endangered tiger (Wikramanayake et al. 1998).
Our objective was to understand the temporal and spatial patterns of risk for different
livestock species and develop ecology-based insights for reducing livestock losses. Using cases
from the livestock compensation program, we examined historical records from 2001-2009 for
past trends in livestock losses. We obtained more detailed insight on the temporal and spatial
distribution and the physical characteristics of depredated livestock by conducting ground
surveys of livestock killed in 2011-2012. We also investigated the location and prey contents of
tiger and leopard scat to better understand the movement of the carnivores consuming livestock.
Through combining multiple data sources, we provide an ecological perspective on carnivore-
livestock interactions and develop ecologically informed recommendations for minimizing
livestock vulnerability to carnivores.
Materials and methods
The study was conducted in Kanha Tiger Reserve, Madhya Pradesh in central India. This 2,074
km2 protected area consists of a 940 km2 interior core zone, where human activities are
restricted, surrounded by a 1,134 km2 buffer zone, where human residences and activities such as
livestock grazing are permitted. The reserve supports stable populations of 70 tigers and 100
leopards and growing populations of 59,000 cattle, 22,000 buffalo, and 11,000 goats, which are
regularly attacked by wild carnivores (Kanha Tiger Reserve Forest Department 2012; Jhala et al.
2014b). In an effort to prevent livestock owners from retaliating against carnivores, the Indian
Forest Department financially compensates owners for domestic animals killed by wild
carnivores. To receive compensation, a livestock owner must locate and report the livestock
carcass to the Forest Department within 48 hours, after which an officer visits the site to record
evidence of the death. Although not all livestock owners choose to report lost livestock (Karanth
et al. 2012), many people living within the tiger reserve do, and between 400-600 livestock are
reported for compensation each year (Kanha Tiger Reserve Forest Department 2012).

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Data collection at livestock kill sites
We utilized the compensation program to investigate patterns of livestock depredation. We
analyzed the Kanha Tiger Reserve Forest Department historical records of compensation cases
from January 2001 through December 2009 to assess long-term trends in livestock losses. These
records provided information on the incident date, livestock species, carnivore, and
compensation amount. To obtain more detailed spatial, temporal, and demographic data on the
livestock killed, we conducted ground surveys of freshly killed livestock reported for
compensation from December 2011 through August 2012. Sampling methods are described in
detail in Miller et al. (2015) and overviewed here.
At each kill site, we recorded the incident date and time (if known by the owner), livestock
species and age, percent of carcass remaining, and GPS coordinates. We differentiated the kill
site (where the animal was killed) from the cache site (where the animal was dragged and
consumed) by trails of scuffmarks, blood, and hair. The death of each animal was attributed to a
specific carnivore based on fresh signs within 50 m of the kill and cache site. Researchers were
trained to identify differences in the size and shape of signs for each carnivore species following
the National Tiger Conservation Authority protocol (Jhala et al. 2009). We identified carnivore
signs conservatively and omitted from analysis any kill sites with ambiguous carnivore signs. A
total of 90% of all ‘confirmed’ kills were identified using direct sightings of the carnivore (25%
of kills), pugmarks (64% of kills), and/or scrapes (2% of kills), which can be clearly
distinguished between tigers and leopards (Karanth and Sunquist 1995). Because the methods
used to identify predators were unknown for historical records, we analyzed only ground survey
data when calculating carnivore-specific trends. Finally, we recorded the compensation amount
and the day payment was issued to livestock owners.
Carnivore scat
To study tiger and leopard diet and movement, we examined the prey contents and spatial
location of carnivore feces. We collected tiger and leopard scat opportunistically along roadsides
and foot trails, features which individuals use often for hunting and general movement (Smith et
al. 1989; Karanth and Sunquist 2000). Tiger and leopard scat are distinct in appearance from the
scat of other carnivores in our study area (Karanth and Sunquist 2000) but can be difficult to
distinguish between the two species. Scat was identified to carnivore species using genetic
analysis conducted by the Wildlife Institute of India (Yumnam et al. 2014). However, because
only a few scat samples (18%) contained viable genetic material, we did not associate carnivore
identity with scat for our final analysis. We identified the prey in feces by drying scat, sampling
hair contents, and microscopically comparing hair width, medullary structure, and other
characteristics to prey reference slides at the Wildlife Institute of India (Mukherjee et al. 1994;
Bahuguna et al. 2010). We mapped the GPS coordinates of each scat to examine where
carnivores moved after consuming livestock.
Landscape attributes

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We examined the landscape characteristics associated with kills by sampling environmental and
anthropogenic variables known to influence livestock depredation by large Felidae predators
(Seidensticker 1976; Shrader et al. 2008; Valeix et al. 2009; Kissling et al. 2009; Karanth et al.
2012; Zarco-González et al. 2013; Soh et al. 2014). For land-use, we utilized the Forest Survey
of India State of the Forests 2009 map of land cover, which included non-forest (i.e. agricultural
fields), water, scrubland, open forest, moderately dense forest, and very dense forest. Since the
land-use map did not distinguish villages, we used heads-up digitization with Google Earth
satellite imagery from 2007-2013 to outline village areas. We also quantified human presence
using roads digitized from Survey of India topographic maps and the boundary of the reserve
core zone provided by the Kanha Forest Department. Landscape variables were converted to
raster format at a 20-m spatial resolution using the Spatial Analyst toolset in ArcGIS (v.10.1,
ESRI, CA, USA). We then calculated the Euclidean distance between the center of each kill site
pixel to the center of the nearest pixel with the landscape attribute. We limited our study area to
within 4 km of village centers in the reserve since no livestock were killed beyond this distance
(Miller et al. 2015).
To contextualize kills against the available landscape, we also sampled the range of
landscape attribute values at randomly selected sites across the study area (Johnson et al., 2006;
Manly et al., 2002). The locations of these sites were determined by generating random points
stratified across a 200-m grid in ArcGIS, with one point per pixel separated by at least 200 m so
as not to repeatedly sample the same area. While ground surveying these random sites for
another study (Miller et al. 2015), no wild or domestic prey carcasses were observed.
We examined linkages between precipitation and livestock kills by comparing daily and
monthly rainfall measured by the Kanha Tiger Reserve Forest Department in 2011-2012 to kill
frequencies.
Statistical analysis and modeling
Since the 2011-2012 data on livestock characteristics and landscape characteristics were not
normally distributed, we used Mann-Whitney U tests to compare groups. For livestock species
with adequate sample sizes (cattle, goat, and buffalo but not pig since npig = 2), we investigated
differences by month in the historical frequencies of livestock kills using two-way ANOVAs.
We explored associations between cache distance and livestock age, and between the timing of
kills and daily or monthly rainfall, using linear regression models.
Using surveyed kill sites and landscape attribute data, we built a multivariate logistic
regression model to predict and map the probability of carnivore predation of cattle (Miller
2015). We focused on cattle exclusively because they are the most depredated livestock species
in Kanha, and because the numbers of buffalo and goats killed in 2011-2012 were not adequate
to build a validated model. We modeled combined risk from tigers and leopards for cattle
because we felt the output risk map would be most helpful to managers and owners for
understanding and mitigating risk to cattle generally. Following a use-availability design
(Johnson et al. 2006), the response variable in the model featured binary values, with 0 for

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random sites and 1 for kill sites. Incidences where a carnivore killed multiple livestock during
the same predation event (n = 36) were treated as single kill sites to focus the spatial models on
units of kill sites rather than individual animals and avoid pseudoreplication. We generated
univariate linear regression models to examine which landscape attributes were strong predictors
of kill probability. Following standard methods (Garamszegi 2010; Treves et al. 2011), we ran
Spearman correlations between variables and built global models for livestock species that
included the variables that were significant in the univariate regression and not correlated with
more significant variables (rs < 0.6). These requirements excluded the variable distance to non-
forest, which was correlated with distance to village (rs = 0.7) and distance to very dense forest
(rs = 0.7). Based on field observations we suspected that roads, villages, scrubland, and very
dense forest would have a threshold relationship with kill risk such that effects might decrease in
a nonlinear direction at some distance. We found that including the quadratic structural form of
these variables lowered the global model AIC by ≥ 2 (Draper and Smith 1993; Burnham and
Anderson 2002). The global model were used to generate and rank models with all combinations
of the eligible variables based on the Corrected Akaike’s Information Criterion (AICc) to account
for small sample size (Burnham and Anderson 2002). Since no one top model emerged (AICc ≤
2), we averaged models to produce a final model.
We used the model to investigate the effect of each landscape attribute on the kill probability
by predicting risk while varying the attribute of interest and holding all other variables constant
at their means. We then mapped the model in ArcGIS to observe hotspots in carnivore kill risk
across the study area. Finally, we validated whether the model could predict future kills by
conducting a randomization test against an independent dataset of kills (detailed methods in
Appendix S1 and Figure S1). Statistical analyses were conducted using R (v.2.15.3, R Project
Development Team, www.r-project.org) with the MASS, MuMIN, and R DAAG packages.
Results
Historical records from 2001-2009 contained 4,561 livestock reported for compensation,
consisting of 72% cattle, 16% goats, 10% buffalo, 2% pigs, and <1% horses. All cases were
attributed to a specific carnivore: 64% were attributed to tiger attacks, 34% to leopard, 1% to
unknown carnivores, and <1% to wild dog and wolf. However, since the methods used to
identify carnivores is unknown and 30% of ground-surveyed kills in 2011-2012 did not contain
conclusive evidence about the carnivore, these data should be interpreted with care. Between
December 2011-August 2012, we ground surveyed 449 livestock carcasses, which totaled 92%
of all reported kills in Kanha during the study period. Livestock consisted of 76% cattle, 14%
goats, 9% buffalo, and <1% pigs. Based on carnivore signs, we were able to confidently identify
the predator at 71% of kills, of which we attributed 57% to tiger and 43% to leopard.
Ground surveys indicated that tigers and leopards selected different size classes for livestock
(U430 = 7357, P < 0.001; Fig. 1a-c). Leopards killed more young cattle (aged 1-4 years) and
buffalo (3 years) than tigers, whereas tigers killed more older cattle (5-11 years) and buffalo (6-

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10 years). Only leopards killed goats, which ranged from 1-6 years. The ages of depredated pigs
could not be identified.
Historically, 95% of livestock were accepted for compensation (the Forest Department does
not record cases that did not meet program requirements, such as injured livestock). The Forest
Department paid a total of INR 81,46,842 in compensation over the eight-year period, ranging
INR 8,78,471-16,28,150 per year. In 2011-2012 the Forest Department similarly paid INR
21,42,650 for 91% of reported kills, which it distributed to owners within an average of 17.4 ±
0.9 days (mean ± SE; based on 120 cases with data on payment date).
Fig. 1. Age (left, a-c) and attack time (right, d-f) of livestock killed by tigers (black) and leopards
(gray) for (a, d) cattle, (b, e) buffalo, and (c) goats (c, f).

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Tiger and leopard hunting behavior
Most livestock carcasses were cached away from the site where they were killed. Tigers cached
71% of cattle kills and leopards cached 63%, and both carnivores dragged carcasses similar
distances (mean ± SD for tigers was 50 ± 54 m and leopards was 51 ± 94 m; U155 = 2324, P =
0.154). Tigers and leopards cached 61% and 50% of buffalo kills, respectively, and varied more
in the length of their drags (average distance of 36 ± 26 m for tigers and 116 m ± 86 m for
leopards) but these differences were not statistically significant (U19 = 38, P = 0.203). Leopards
cached 52% of goat carcasses, dragging them an average of 192 ± 229 m, and cached both pig
kills (n = 2), moving one carcass 8 m and the other 470 m. For each carnivore, smaller-bodied
livestock species were cached farther from kill sites, and overall cache distance was significantly
but weakly correlated with livestock age (R2 = 0.066, F1,269 = 18.86, P < 0.001). In only 45% of
cases were carnivores able to consume more than half the carcass before the Forest Department
burned the body (Fig. S2).
We collected 133 tiger and leopard scat distributed across the reserve (Fig. 2), 69 (52%) in
the core zone and 64 (48%) in the buffer zone. The majority of scat (67%) contained only wild
prey animals, 29% contained only domestic animals (cattle and buffalo), and 4% contained both
wild and domestic prey (Fig. S3). Since hair from wild and domestic pig appear identical under
the microscope, we conservatively categorized all pig hairs as wild prey for our analysis. Out of
the 44 scat containing livestock, only eight were found in the park core zone, five of which were
located close (< 2 km) to the core-buffer boundary. Genetic analysis confirmed that two of the
five near the boundary were tiger and one was leopard. Three scats containing livestock were
found deep within the core interior (3.5-7.2 km from the boundary).
Fig. 2. Locations of tiger and leopard scat collected across Kanha Tiger Reserve, showing the
distribution of domestic and wild prey contents with respect to the reserve core and buffer zones.
Villages are primarily located in the buffer zone.

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Temporal patterns
From 2001-2012, the frequency of livestock depredations varied substantially by year but did not
consistently increase or decrease over time (Fig. 3). The number of kills did not significantly
differ between month for cattle (F1,106 = 0.106, P = 0.746), buffalo (F1,106 = 0.039, P = 0.845),
goat (F1,106 = 1.664, P = 0.200), pig (F1,106 = 1.492, P = 0.225), or horse (F1,106 = 0.716, P =
0.399). A distinct peak in compensated livestock occurred each year sometime between July-
September during the monsoon (Fig. 3). However, the number of kills was not associated with
monthly (R2 = 0.137, F1,7 = 1.112, P = 0.327) or daily (R2 = 0.002, F1,210 = 0.392, P = 0.532)
rainfall.
Most livestock were killed in the afternoon and evening between 12:00-20:00 h (Fig. 1d-f).
Both tigers and leopards attacked cattle during this period and tigers also frequently killed cattle
throughout the morning (05:00-12:00 h). Leopards killed goats throughout the day and especially
in the early evening (16:00-20:00 h).
Fig. 3. Number of livestock killed each month in Kanha Tiger Reserve from 2001-2009. No
records were available for 2010.
Spatial patterns of livestock kills and attack risk
The majority of livestock were killed in the buffer zone of the reserve (82%), where most
villages and livestock grazing occur. The remaining 18% were killed in the core zone and were
concentrated around the villages in the core zone or close to the core-buffer boundary.
Comparisons between kill and random sites revealed that carnivores tended to kill cattle and
buffalo closer to forests and farther from non-forest (agricultural fields) and villages than random
(Table 1). In contrast, goats were killed closer to fields and villages. Both cattle and goats (but
not buffalo) were killed at farther distances from water and the park core boundary than random
sites. Carnivores killed cattle (but not goats or buffalo) farther from roads.

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Table 1 Mann-Whitney U test comparisons between mean values ± standard error of landscape
attribute variables at sites where tigers and leopards killed livestock (kill site) and random sites
in the study area in Kanha Tiger Reserve. P-values in bold are significant (P < 0.05).
Variable
Random
sites
(n = 435)
Cattle (n = 193)
Goat (n = 39)
Buffalo (n = 32)
Kill sites
P-
value
Kill sites
P-
value
Kill sites
P-
value
Distance to core
(km)
0.5 ± 0.1
2.0 ± 0.2
0.023
1.1 ± 0.2
0.023
2.1 ± 0.6
0.412
Distance to road
(km)
2.5 ± 0.3
0.7 ± 0.4
<0.001
0.5 ± 0.7
0.708
0.7 ± 0.9
0.058
Distance to village
(m)
827 ± 40
956 ± 47
<0.001
501 ± 79
0.042
975 ± 84
0.002
Distance to water
(km)
2.6 ± 0.8
2.9 ± 0.1
0.036
3.9 ± 0.2
<0.001
3.0 ± 0.3
0.195
Distance to non-
forest (m)
303 ± 23
362 ± 23
<0.001
83 ± 20
0.024
511 ± 80
<0.001
Distance to
scrubland (km)
6.7 ± 0.2
6.9 ± 0.2
0.128
5.8 ± 0.5
0.419
7.4 ± 0.7
0.251
Distance to open
forest (m)
362 ± 16
289 ± 19
0.048
285 ± 39
0.342
301 ± 58
0.189
Distance to
moderately dense
forest (m)
234 ± 17
68 ± 8
<0.001
197 ± 35
0.41
55 ± 11
0.003
Distance to very
dense forest (m)
495 ± 36
173 ± 39
<0.001
532 ± 101
0.182
104 ± 29
<0.001
We built a predation risk model using 435 random sites and 193 cattle kills with confirmed
predators. The model predicted the probability of a tiger or leopard killing a cattle given an
encounter, ranging from 0 (low risk) to 0.93 (high risk). The contribution of each variable to
predictions of predation risk was measured by its relative importance in the model. Most
variables played a strong role in predicting risk (importance > 0.70), including all human
presence and dense forest variables (Table 2). Randomization tests revealed that model
predictions performed better than random (Fig. S4). The model accurately identified 69% of
validation sites (88 out of 128 known kill sites) as kills, which is greater than would be expected
by random chance (P < 0.001).
The risk to cattle was greatest in moderate and very dense forests and at intermediate
distances from roads, villages, and scrubland (Fig. 4). Risk did not substantially change with
increasing distance from water or the core zone boundary. Kill probability showed a negative
quadratic relationship to the distance to road, village, and scrubland, with cattle vulnerability
increasing at farther distances up to a threshold point and thereafter decreasing. Cattle were most
vulnerable to carnivores 1.2 km from roads, 1.0 km from villages, 6.1 km from scrubland (Fig.
4a, 4c, 4e). The distance to very dense forest showed a negative quadratic relationship, with the

11
greatest risk directly within (0 km) or far from (> 4 km) very dense forests (Fig. 4i). The
predation risk map revealed the highest risk levels in forest patches adjacent to the park core
boundary and the lowest levels in agricultural areas near villages and roads (Fig. 5).
Table 2 Statistics from the predation risk model for cattle, showing the relative importance,
coefficient (β), and standard error (SE) of variables in the final averaged model. Relative
importance values range from 0-1, with a value of 1 indicating a strong contribution to the
model.
Variables
Importance
β
SE
intercept
-1.16
0.63
distance to very dense forest2
1.00
8.24E-07
1.75E-07
distance to very dense forest
1.00
-3.02E-03
7.10E-04
distance to road
1.00
1.74E-03
5.00E-04
distance to road2
0.99
-6.98E-07
2.37E-07
distance to scrub2
0.96
-1.67E-08
8.15E-09
distance to village2
0.92
-3.31E-07
1.82E-07
distance to moderately dense
forest
0.88
-2.23E-03
9.43E-04
distance to scrub
0.84
2.36E-04
1.20E-04
distance to village
0.75
7.99E-04
4.70E-04
distance to open forest
0.43
-4.73E-04
4.02E-04
distance to core
0.35
-3.24E-05
3.67E-05
distance to water
0.28
2.26E-05
5.66E-05

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(a)
(b)
(c)
(d)
(f)
(g)
(h)
(e)
Fig. 4. The probability of carnivore depredation on cattle with increasing distances to landscape
attributes as predicted by the predation risk model. The 95% confidence intervals are shown in
grey.

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Predation risk
Fig. 5. Distribution of tiger and leopard predation risk for cattle in Kanha Tiger Reserve. Values
represent the kill probability given an encounter between a carnivore and cattle. Low-risk areas
primarily occur in agricultural fields and village areas whereas high-risk hotspots occur in dense
forest away from human activity. Notable villages are shown for perspective (not all villages are
marked). The study area was designated within 4 km of village centers (see methods for details).
Discussion
Kanha has one of the highest rates of livestock depredation from tigers and leopards in India
(Kala and Kothari 2013; Karanth et al. 2013; Singh et al. 2015) yet is also renowned as one of
the most successful and stable sites of tiger conservation (Post and Pandav 2013; Jhala et al.
2014a). The low frequency of retaliations against depredating carnivores in Kanha is largely due
to the Forest Department’s prompt livestock compensation program, which in 2011-2012
distributed payment on average about 2.5 weeks after livestock were attacked. This is
considerably faster than other reported compensation timeframes from India (Madhusudan 2003)
and on par with championed programs (Nyhus et al. 2005). The compensation program also
offers tractable long-term data that can be used for assessing the temporal and spatial patterns
and physical characteristics of livestock losses. Kill data offer exclusive fine-scale spatial
information about the sites where livestock are vulnerable to livestock, offering a unique
perspective not otherwise captured by household surveys, which have been the basis of many

14
previous studies on human-carnivore conflict (e.g. Wang and Macdonald 2006; Nugraha and
Sugardjito 2009; Karanth et al. 2012; Katel et al. 2014; Bhattarai and Fischer 2014).
Our study confirmed that tigers and leopards were the primary depredating carnivores, with
tigers responsible for killing slightly more livestock than leopards. This contrasts with reports
from Corbett Tiger Reserve in northern India and Bhutan, where leopards kill substantially more
livestock than tigers (Wang and Macdonald 2006; Sangay and Vernes 2008; Malviya and
Ramesh 2015). We suspect this difference may be related to the lower availability of free-
grazing adult cattle in Corbett (where ~30% of households stall-feed livestock; Malviya and
Ramesh 2015) and the lower density of tigers in Bhutan (Sanderson et al. 2006). We did not find
evidence of attacks from other carnivores, which the Forest Department had reported in previous
years, and we urge authorities to train field staff to make accurate predator identifications in
order to prevent false perceptions about threats from other carnivores (Dickman 2010;
Suryawanshi et al. 2013). Cattle were killed most frequently, followed by goats, buffalo, and
pigs, respectively. Compensation cases are likely biased against small-bodied livestock (goats
and pigs) because these animals are more often cached further and more completely eaten, and
thus more difficult for livestock owners to locate and report. These strong associations between
certain sized livestock and certain carnivore species suggests that segregating livestock by body
size and age, and grazing cohorts in habitats less conducive to attacks by their main predator
(e.g. open vegetation for tigers and dense forests for leopards), might discourage depredation
(Wang and Macdonald 2006; Goodrich 2010).
Temporal patterns
Tigers and leopards killed livestock at all hours of the day, particularly in the afternoon and
evening. Our analysis found a high number of attacks by tigers in the afternoon (12:00-16:00 h),
likely because this was when livestock were farthest from the village and in dense vegetation
where tigers often attack livestock (Soh et al. 2014; Malviya and Ramesh 2015; Miller et al.
2015). Furthermore, tigers (and leopards) are most active and hunting in the early morning, late
afternoon, evening and night and avoid much activity mid-day (Karanth and Sunquist 2000). Our
results did not reveal temporal separation between tiger and leopard or a tendency towards
nocturnal hunting in either predator as found in previous studies (Seidensticker 1976; Karanth
and Sunquist 2000; Malviya and Ramesh 2015), probably due to the nature of our dataset, which
depended on people witnessing attacks.
We observed a spike in livestock losses each year during the monsoon, which echoes similar
findings from other protected areas in South Asia (e.g. Bhadauria and Singh 1994; Sangay and
Vernes 2008; Singh et al. 2015). However, the number of livestock kills did not explicitly relate
to monthly rainfall as reported from Africa (Kolowski and Holekamp 2006). This may indicate
that the rise in depredations during monsoon is due to human-induced changes, such as herders
leading cattle and buffalo away from crops to graze in denser forests with greater predation risk.
If so, monsoon may be an ideal season in which to implement alternative grazing strategies since
humans have more control over livestock movement.

15
Spatial patterns
Results revealed distinct risk distributions by livestock species, which to our knowledge has not
been examined for tiger and leopard depredation. The predation risk model found that threats
from both carnivores combined were highest for cattle near dense forests and lowest near
agricultural fields, villages, and roads. These results are comparable to previous conflict studies
on tigers that likewise observed most attacks on livestock in forest and away from roads (Wang
and Macdonald 2006; Soh et al. 2014). The risk map for cattle closely resembles general tiger
risk for all livestock (Miller et al. 2015), likely because cattle are the most frequent species killed
by tiger and thus most strongly represented in the tiger risk model. Risk hotspots occurred 1-2
km from the core zone boundary in both the buffer zone and the interior of the core zone,
corroborating previous findings from central India that livestock depredation increases with close
proximity to protected areas (Karanth et al. 2013). The high kill probabilities inside the core zone
reiterates the need for strict enforcing to eliminate grazing livestock in the core, which has been
prioritized for reducing human-tiger in the past (Goodrich 2010). To reduce livestock losses,
livestock owners could minimize cattle presence in dense forests and favor grazing routes close
to open vegetation and human areas. If grazing routes are adapted to reduce risk, carnivores
should also be initially monitored for behavioral feedbacks to ensure that they are not drawn into
closer contact with people (Miller 2015).
Similarities in the landscape attributes associated with cattle and buffalo kill sites suggest
that buffalo may experience comparable distributions of predation risk as cattle. However, results
showed opposite trends for goats, which were more vulnerable in open vegetation and village
areas. These distinctions may be related to grazing patterns since herders may restrict goats to the
open vegetation and village areas that are most convenient for human access, whereas cattle and
buffalo graze unrestrained farther from villages for most of the year. Our data does not enable us
to discern whether these risk distributions are shaped more by carnivores or by livestock and
people but we encourage future studies to directly pursue the mechanisms behind depredation.
It is currently unknown whether the individual tigers and leopards predating on livestock are
resident or dispersing but our scat results offer some insight into their movement. Though scat
contents indicated that tiger and leopard diets primarily consisted of wild prey, 33% of all scat
(buffer and core zone) and 42% of scat found in the buffer contained domestic livestock, which
is a surprisingly large proportion considering the high abundance of wild prey available in the
core zone (Jhala et al. 2014b). Twelve percent of scat found in the core zone contained livestock
remains, and 10% was found 3-7 km from the core-buffer boundary in the interior of the core.
Although these scat may have been deposited by young tigers or leopards dispersing through the
reserve, it is also possible that resident carnivores may visit the buffer zone to supplement their
diet with livestock. Considering the extensive home ranges of tigers (~10-100 km2; Sharma et al.
2010) and leopards (~10-60 km2; Odden et al. 2014), livestock depredation may not be restricted
to transient individuals as commonly believed. Furthermore, if resident individuals are regularly
attracted out of the park core zone to kill livestock, they may be susceptible to human threats in

16
the buffer zone and non-protected areas (Balme et al. 2010). However, our limited sample size
and opportunistic (rather than systematic) sampling of scat limits the scope of our conclusions.
Greater efforts in the future must be dedicated to identifying which individual carnivores kill
livestock, especially since this answer may help more fully elucidate the drivers behind livestock
depredation.
Implications for human-carnivore conflict mitigation
The call to reduce human-carnivore conflict by avoiding predator hotspots has been sounded
before (Wang and Macdonald 2006; Goodrich 2010), and our study contributes insight to help
identify when and where different livestock species are most vulnerable to tigers and leopards.
Middle-aged cattle (4-8 years) were the most vulnerable to tigers and leopards and were attacked
primarily in the afternoon and early evening (12:00-20:00 h) near dense forests and at moderate
distances from road, village and scrub forests. Buffalo were mostly killed by tigers, which tended
to attack middle-aged individuals (6-8 years) during the afternoon and early evening (12:00-
20:00 h) in dense forests and away from open habitat and villages. To reduce losses we
recommend the use of herders year-round, instead of only during the monsoon, to enable greater
control over cattle and buffalo routes and timing to minimize high-risk grazing in forests.
Middle-aged goats (2-6 years) were most at risk from leopards during the early evening (16:00-
20:00 h) in open vegetation and village areas. Rather than shift goat grazing routes to denser
forest to reduce risk, which might increase threats from tigers, owners could consider protecting
goats earlier in the day (before 16:00 h) in reinforced, leopard-proof enclosures. Furthermore, to
decrease losses with all livestock species, owners living in high-risk areas could consider
implementing additional mitigation techniques such as trained guard dogs, predator-proof
enclosures and fencing, deterrents, and sensory stimulants, especially during the highest-risk
season (monsoon), to further reduce attacks from carnivores (Shivik 2006). Previous research in
central India (Karanth et al. 2013) and in east Africa (Kolowski and Holekamp 2006) found that
guard animals and fencing were especially useful in mitigating attacks. Finally, we encourage
managers worldwide to regularly update predation risk models and maps to monitor conflict, and
to develop results into relevant education and outreach materials to assist livestock owners in
understanding risks near their villages (Miller 2015).
This paper demonstrates that livestock compensation programs generate data that can be
useful for understanding and preventing conflict. Financial compensation systems play a
particularly important role in supporting livestock owners that live in the ‘diffuse edge’ buffer
zones of protected areas where the majority of human-carnivore conflict often occurs (this study;
Nyhus and Tilson 2004). This is the case in Kanha, one of the few tiger reserves in India with a
functional buffer zone, the livestock compensation program is key to minimizing retaliations
against carnivores. Compensation programs also present opportunities for villagers to develop
stronger relationships with the Forest Department, which can impact human-carnivore conflict
given that local trust in authority is directly linked to attitudes towards conservation (Treves et al.
2006; Dickman 2010; Carter et al. 2012). To build trust and local engagement, it is important

17
that authorities ensure that local people understand and can meet the regulations related to
livestock compensation (Nyhus et al. 2005). Most surveyed livestock losses (91%) in Kanha
from 2011-2012 were compensated within several weeks if basic requirements were met. These
levels greatly differ from the lower success rates (29%) reported just outside the buffer of Kanha
(Karanth et al. 2012), where compensation is mandated but often overlooked by governing
authorities in the absence of the high-profile tiger. This discrepancy has sparked confusion and
intolerance in some livestock owners (Karanth et al. 2013), who may be more prone to retaliate
against wild carnivores that depredate livestock. We encourage managers to maintain
consistency and generously award compensation whenever possible (while taking care to
avoiding false claims, corruption and perverse incentives; Nyhus et al. 2005). This is important
both within and outside of protected areas, especially since carnivore dispersal outside of parks is
essential for maintaining resilient populations (Yumnam et al. 2014).
The results of this paper face several potential limitations. First, compensation data may not
have evenly represented the spatial distribution of livestock depredations if village remoteness or
villager-authority relationships biased the reporting of kills. Because Forest Department beat
camps are evenly distributed across the Kanha buffer and core, and because we were not aware
of negative social tensions during nine months of extensive field visits, we do not expect that
results were significantly biased. Second, the presence of herders grazing cattle during the
monsoon months may change the distribution of predation risk from the rest of the year, yet our
cattle risk models did not address season-wise differences. While our goal in modeling cattle risk
was to understand year-round predation risk to offer managers simple guidance for decision-
making, we recognize that risk will shift with different grazing practices and resource
distribution and encourage future studies to more closely examine such short-term shifts. Finally,
the cattle risk model portrays combined risks for tigers and leopards, which offers managers and
owners a tool for strategizing protection for cattle but may limit inferences on the spatial
distribution of risk from each species.
Conclusions
The first increase in the tiger population recently reported from India (Jhala et al. 2014a) offers
hope that collective efforts worldwide can reverse carnivore declines. Yet even if carnivore
populations stabilize, expanding human development guarantees that natural resource managers
and livestock owners will continue to face challenges in mitigating human-carnivore conflict
(Treves and Karanth 2003; Ripple et al. 2014). Understanding the temporal and spatial factors
that underlie ecological interactions between specific carnivore and livestock species will be
essential for developing strong mitigating methods that ultimately make coexistence possible.
Acknowledgements
We thank Rajesh Gopal and the National Tiger Conservation Authority for permissions and
facilitation for carrying out this research. We acknowledge the Madhya Pradesh Principal Chief
Conservator of Forests, H. S. Pabla, for granting research permission and the Wildlife Institute of

18
India for institutional support. We are very grateful to the Kanha Forest Department for
providing historical records and field support, especially Field Director J. S. Chauhan, Research
Officer Rakesh Shukla, and the wireless controllers, forest guards, and chowkidars who helped
us survey sites. We thank Naseem Khan, Arvind Thakur, Ashish Bais, Amol Khumbar, and
Ashish Prasad for assisting with data collection. This manuscript benefited from discussions with
Oswald Schmitz, Anne Trainor, and Meghna Agarwala as well as feedback from several
anonymous reviewers. Maya Lim assisted with graphic design. We thank Ruth DeFries, Trishna
Dutta, and Sandeep Sharma for coordinating and editing this special issue. Funding was provided
by the American Institute for Indian Studies; American Philosophical Society Lewis and Clark;
Association of Zoos & Aquariums; John Ball Zoo Society; Yale Tropical Resources Institute;
and the National Science Foundation GRFP.
Electronic supplementary material
The online version of this article contains supplementary material, which is available to
authorized users.
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