Does livestock benefit or harm snow leopards?

Biological Conservation 190 (2015) 8–13

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Short communication

Does livestock benefit or harm snow leopards? Rishi Kumar Sharma a,⇑, Yash Veer Bhatnagar a,b, Charudutt Mishra a,b a b

Nature Conservation Foundation, 3076/5, 4th Cross, Gokulam Park, Mysore 570002, India Snow Leopard Trust, 4649 Sunnyside Ave North, Suite 325, Seattle, WA 98103, USA

a r t i c l e

i n f o

Article history: Received 14 October 2014 Received in revised form 15 April 2015 Accepted 21 April 2015

Keywords: Panthera uncia Trans-Himalaya Pastoralism Large carnivores Livestock grazing Co-existence

a b s t r a c t Large carnivores commonly prey on livestock when their ranges overlap. Pastoralism is the dominant land use type across the distributional range of the endangered snow leopard Panthera uncia. Snow leopards are often killed in retaliation against livestock depredation. Whether livestock, by forming an alternative prey, could potentially benefit snow leopards, or, whether livestock use of an area is detrimental to snow leopards is poorly understood. We examined snow leopard habitat use in a multiple use landscape that was comprised of sites varying in livestock abundance, wild prey abundance and human population size. We photographically sampled ten sites (average size 70 sq. km) using ten camera traps in each site, deployed for a period of 60 days. Snow leopard habitat use was computed as a Relative Use Index based on the total independent photographic captures and the number of snow leopard individuals captured at each site. We quantified livestock abundance, wild prey abundance, human population size and terrain ruggedness in each of the sites. Key variables influencing snow leopard habitat use were identified using Information Theory based model selection approach. Snow leopard habitat use was best explained by wild prey density, and showed a positive linear relationship with the abundance of wild ungulates. We found a hump-shaped relationship between snow leopard habitat use and livestock stocking density, with an initial increase in habitat use followed by a decline beyond a threshold of livestock density. Our results suggest that in the absence of direct persecution of snow leopards, livestock grazing and snow leopard habitat use are potentially compatible up to a certain threshold of livestock density, beyond which habitat use declines, presumably due to depressed wild ungulate abundance and associated anthropogenic disturbance. Ó 2015 Elsevier Ltd. All rights reserved.

1. Introduction The impact of anthropogenic activities such as livestock grazing and resource extraction on large carnivores has been debated. Anthropogenic activities are reported to depress carnivore populations through direct persecution (Cardillo et al., 2004; Inskip and Zimmermann, 2009; Winterbach et al., 2012), hunting of wild prey (Karanth et al., 2004) and habitat loss (Harihar et al., 2009). On the other hand, some carnivores appear to adapt and even thrive in human modified environments (Athreya et al., 2013; Bouyer et al., 2014; Odden et al., 2014) and local communities and carnivores could even benefit from each other under specific conditions (Banerjee et al., 2013). Carnivores populations can persist despite high human populations when cultural tolerance is high (Karanth and Chellam, 2009), conflicts are managed properly (Treves and Karanth, 2003) and conservation policies are effectively implemented (Linnell et al., 2001). ⇑ Corresponding author. Tel.: +91 8212515601; fax: +91 8212513822. E-mail addresses: [email protected], [email protected] (R.K. Sharma). http://dx.doi.org/10.1016/j.biocon.2015.04.026 0006-3207/Ó 2015 Elsevier Ltd. All rights reserved.

Snow leopards (Panthera uncia) are endangered and face a diversity of threats across much of their range (IUCN Red List, 2014). Our understanding of snow leopard ecology remains limited due to their elusive nature, low densities and the remote mountainous terrain they inhabit. Snow leopard habitats throughout their distribution range are multiple use landscapes, supporting a variety of natural resource uses including livestock grazing, agriculture and extraction of medicinal plants and fuel-wood (Mishra et al., 2010). How snow leopards respond to these various anthropogenic factors within their landscapes continues to remain poorly understood. Livestock rearing is an important livelihood source for local people across the distributional range (Mishra, 1997), and livestock depredation by snow leopards brings their conservation into conflict with the goals of pastoral production (Bagchi and Mishra, 2006; Bagchi et al., 2004; Jackson et al., 2010). Many studies have reported rather high contribution of livestock (up to 70%) to the diet of the snow leopard (Anwar et al., 2011; Bagchi and Mishra, 2006), leading to the proposition that local livestock may be playing a role in sustaining populations of this endangered carnivore

R.K. Sharma et al. / Biological Conservation 190 (2015) 8–13

(Wegge et al., 2012). On the other hand, overstocking of rangelands with livestock (Mishra et al., 2001) can lead to decline of wild prey populations as a result of competition (Mishra et al., 2004; Namgail et al., 2006), which could further escalate the extent of livestock depredation. For example, Bagchi and Mishra (2006) reported a greater presence of livestock in the diet of snow leopards in areas with lower wild prey abundance. This creates a conservation dilemma. On the one hand, pastoralism has indirect negative effects on snow leopards as livestock depredation leads to retaliatory persecution by herders (Jackson et al., 2010) and high livestock abundance suppresses wild prey populations. On the other hand, it is suspected that livestock might actually be an important food source for snow leopards. Whether livestock, by acting as an additional food resource, is potentially beneficial to the snow leopard, or whether livestock use of an area negatively affects snow leopard habitat use, is not understood. We aimed to address this question by examining snow leopard habitat use along gradients of livestock and wild ungulate abundance.

2. Study area Spiti Valley, forming the catchment of the river Spiti, is a part of the Trans-Himalayas in the state of Himachal Pradesh, India. Lying in the rain shadow of the Himalayas, the region is cold and arid (3000–6700 m) with most of the precipitation in the form of snow, though rainfall events have increased in frequency while snowfall has declined in the past decade (Singh et al., in press). Agro-pastoralist communities have inhabited this region for two to three millennia. Parts of the landscape are visited by transhumant Gaddi herders accompanied by large flocks of sheep and goat and Tibetan Mastiff guard dogs. The local agro-pastoral

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community, largely Buddhist, do not use guard dogs and are concentrated in village clusters largely along the main Spiti River (see Fig. 1). The livestock assemblage includes sheep, goat, donkey, cow, cow–yak hybrid, horse and yak. Livestock graze in the extensive pastures except during extreme winter when they are stall-fed. Based on herding practices, local livestock can be classified as (i) large-bodied free ranging (yaks and horses), henceforth referred to as large stock and, (ii) medium and small-bodied herded, henceforth referred to as herded stock (cow, donkey, cow–yak hybrid, goat and sheep; (Suryawanshi et al., 2013). Villages have clear guidelines on the grazing rights to pastures but some of the pastures are shared between neighboring villages. Herded stock is taken out to pastures every morning and brought back to villages and kept in corrals by individual households. A designated shepherd referred to as ‘Lugzi’ is responsible for livestock herding and decides on which pastures to use on a daily basis (Singh et al., in press). Every household helps the ‘Lugzi’ with livestock grazing on a rotational basis. Families mostly own small agricultural land holdings (1–2 ha). The Snow Leopard Trust and Nature Conservation Foundation have been working in the region for 15 years. A key characteristic of our study site is that there is almost no retaliatory persecution of snow leopards in the region, partly due to these conservation programs, prevalence of Buddhism, and awareness of laws. Thus, with human persecution being controlled for, this landscape was suitable to answer our question. Within this large ca. 4000 km2 landscape, we selected ten sites for our study. These ten sites represented a gradient of livestock density, wild prey abundance and human population size. Their boundaries were demarcated based on natural boundaries of the topographical features.

Fig. 1. Map of the study area showing sampling sites and camera trap locations. The map inset shows locations of the study area in the state of Himachal Pradesh, India.

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The ratios of livestock to wild prey varied from zero livestock at one site to 1111:1 at the other extreme. 3. Methods The ten sampled sites ranged from 42 to 112 km2 (Table 1). We deployed 10 Reconyx RM45 and RC55 camera traps at ten best locations within each site for a period of two months, guided by a 3  3 km grid cell based design. The camera traps were installed at 5 sites for 60 days from May to July and in the remaining 5 sites for another 60 days from July to September, thus the sampling being conducted from the beginning to the end of the summer season. Trap locations were selected based on the frequency of signs of snow leopards such as pugmarks, scrapes and scent marks at prominent features such as ridgelines and cliffs. Camera traps were programmed to take a picture every second and till the animal remained within the camera range.   We developed a relative use index RUI ¼ Cc  Nn : as a response variable indicating snow leopard habitat use. The RUI assigns a habitat use value to each sampled site based on number of captures and number of adult snow leopards captured at each site. Where c = number of independent photographic captures of adult snow leopards at a particular site, C = total number of independent photographic captures of adult snow leopards across all sites, n = number of adult snow leopard individuals captured at a particular site and N = total number of adult snow leopards captured across all sites’’. RUI thus assigns high value to sites having larger proportion of snow leopard captures and higher number of unique snow leopard individuals and can take values between 0 and 1. We quantified four key influential variables that could potentially affect snow leopard habitat use. These included population size of large and small bodied livestock, determined using door to door census at all the sites. This involved censuses in 51 villages across the ten sites. The abundance of wild ungulate prey, blue sheep Pseudois nayaur and ibex Capra sibirica, was estimated in each site twice, using the capture–recapture based double observer survey technique (Suryawanshi et al., 2012) and the average of the two estimates was used for analysis. Livestock stocking densities and wild prey densities for all ten sites were calculated by dividing their respective abundance estimates by the area of each site. Human population size was included as an indicator of anthropogenic activities (Harihar and Pandav, 2012; Woodroffe, 2000). Anthropogenic activities can influence activity patterns of carnivores, with carnivores avoiding high human activity periods (Carter et al., 2015). Human population was derived from Census of India data (http://censusindia.gov.in/). As snow leopards use steep and rugged terrain, we derived a Terrain Ruggedness Index (Riley et al., 1999) for each site using a 30  30 m Digital Elevation Model Data from Aster Global Digital

Elevation Model data using the terrain analysis plugin in Quantum GIS 2.0 software (QGIS Development Team, 2013). The terrain was categorized into six categories ranging from level to extremely rugged. Combined proportion of the highly rugged and extremely rugged category was used in the analysis. We also examined snow leopard activity patterns in each site by extracting the date and time stamp information from the snow leopard camera trap images. Retributory killing of snow leopards is a global cause of concern but in our landscape, long-term community-based conservation programs (Mishra et al., 2003) and the Buddhist beliefs of local people have ensured that there has been just one case of a snow leopard being killed over the last 15 years. The current human population density in this landscape is <1 person per km2 and agro-pastoralism is the dominant activity across the landscape. Agricultural fields and orchards are situated in the immediate vicinity of villages while villages themselves are largely located in the valley along the Spiti River and not scattered across the landscape. At the current population density and the kind and scale of anthropogenic activities, we did not consider human disturbance per se to have a major impact on snow leopards in our study area. However anthropogenic pressures were considerably high in the two sites visited by nomadic pastoralists as these sites were characterized by high livestock stocking densities, use of guard dogs to protect livestock, and intensive use of the entire sites for livestock grazing. Since our study area is one contiguous landscape free from any natural or man-made barriers for snow leopards, we considered it as a single population of snow leopards. We used the proportion of highly rugged and extremely rugged terrain as an important variable in the analysis with the assumption that good snow leopard habitats would be characterized by a combination of rolling and rugged terrain where rolling terrain with pastures supports the prey species while rugged terrain is essential for an ambush predator like snow leopard to hunt. 4. Data analysis Snow leopard individuals in the photographs were identified by unique pelage patterns (Jackson et al., 2006). We used exploratory analysis using scatterplots and simple linear regression to understand the relationship between snow leopard habitat use and individual explanatory variables. We used linear regressions to model the relationship between snow leopard habitat use and four of the key measured explanatory variables. We scaled all continuous variables by normalizing to their means for the linear regression modeling. We used a quadratic term for livestock in the regression models since exploratory data analysis showed that RUI initially increased with an increase in livestock density and then declined after a threshold (Fig. 2b).

Table 1 Snow leopard habitat use estimated through photographic sampling of 10 sites in Spiti Valley, along with their topographic and biotic characteristics. The table details snow leopard photo-captures, the number of adult snow leopards captured in each site, their Relative Use Index, and potential explanatory variables including the livestock density, wild prey density, human population size, terrain ruggedness and altitudinal range at each site. Site name

Area

Human population

Ruggedness

Livestock density (km2)

Wild prey density (lm2)

Snow leopard photo-captures

Snow Leopard Individuals

RUI

Altitudinal range (m)

Presence of guard dogs

Kibber-Chicham Dhankar-Lalung Sumra Demul Langza-Hikkim Mane Lossar Poh-Sichling Tabo Chandertal

112 42 47 62 92 62 91 30 71 90

841 853 402 444 1060 354 739 0 1073 50

26.35 37.7 42.94 31.96 23.82 39.43 39.13 35.46 45.18 27.7

20.04 47.61 35.79 14.69 12.29 13.68 57.23 0 13 55.84

3.83 2.11 3.86 2.17 3.28 1.25 0.1 1.63 2.19 0.05

21 20 20 18 21 15 7 6 10 0

8 6 5 5 4 3 2 2 1 0

0.0507 0.0362 0.0302 0.0272 0.0254 0.0136 0.0042 0.0036 0.003 0

3970–5043 3724–5160 3137–4576 3684–4964 4049–5019 3551–5168 3968–5139 3453–4663 3216–4760 4034–5111

No No No No No No Yes No No Yes

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R.K. Sharma et al. / Biological Conservation 190 (2015) 8–13

(a)

0.06

4.5

y = 0.0095x R² = 0.5982, P-value = 0.0086

0.04

RUI

y = -0.0043x2 + 0.2492x R² = 0.6166, p-value = 0.017

4

Wild prey density

0.05

0.03 0.02 0.01

3.5 3 2.5 2 1.5 1

0.00 0.00

0.50

1.00

1.50

2.00

2.50

3.00

3.50

4.00

0.5

4.50

0

Wild prey density

(b)

0

y = -4E-05x2 + 0.0024x R² = 0.534, P value = 0.064

0.05

20

30

40

50

60

70

Fig. 3. Relationship between livestock and wild prey density. It appeared that sites with relatively productive habitats could support a greater abundance of both livestock and wild prey, but that wild prey abundance declined as livestock abundance increased beyond a threshold.

0.04

RUI

10

Livestock stocking density

0.06

0.03 0.02

abundance, wild prey abundance, livestock stocking densities, wild prey densities, human population size and ruggedness are presented in Table 1. The sites showed considerable variation in wild prey abundance and density (abundance ranging from 5 to 433 and density ranging from 0.05 to 3.86/km2; Table 2) as well as livestock abundance and stocking densities (abundance ranging from zero at one site to 5215 livestock heads at a site visited by nomadic herders; stocking density ranging from zero to 57.23 livestock heads per km2). The observer detection probability for the double observer surveys for estimating wild prey abundance ranged from 0.61 to 1 across observers and sites (Table 2). Exploratory analysis using scatterplots indicated a monotonic relationship of RUI with wild prey density and a quadratic relationship with livestock stocking density (Fig. 2a and b). There appeared to be a quadratic relationship between wild prey density/km2 and livestock stocking density/km2 (Fig. 3).

0.01 0.00 0

10

20

30

40

50

60

70

Livestock stocking density Fig. 2. (a) Relationship between Relative Use Index (RUI) representing snow leopard habitat use and the density of wild prey in Spiti Valley depicted with a linear regression passing though the origin. (b) Relationship between RUI and livestock stocking density fitted with a quadratic model. Snow leopard habitat use showed an initial positive relationship with livestock, but appeared to decline beyond a threshold of livestock abundance.

We used an information-theoretic approach to develop apriori a set of models that best explained the habitat use of snow leopards. Our global model included wild prey density, livestock stocking density, human population size and terrain ruggedness as potential variables influencing snow leopard habitat use. We then developed 6 candidate sub-models using the variables from this global model. Each sub-model represented a specific hypothesis explaining the relationship between explanatory variables and snow leopard habitat use. Akaike Information Criterion adjusted for small samples (AICc) was used for model selection (Anderson and Burnham, 2002). We implemented this using the package MuMIn (Barton, 2013) in program R (R Core Team, 2013).

5.1. Correlates of snow leopard habitat use 5.1.1. Model Selection The best fit model (xi = 0.92) suggested that wild prey density influenced snow leopard habitat use (Table 3). The next best supported model (D AICc = 0.92, xi = 0.23) included livestock stocking density. Based on distance from the lowest AICc (D AICc) and model weight (xi), there was little support for rest of the models. The parameter coefficients for the best models showed positive effect of wild prey (b = 0.013 ± 0.0038 SE) on snow leopard habitat use.

5. Results We obtained 138 independent captures of 24 adult snow leopards across ten sampling sites (Table 1). The estimates of livestock

Table 2 Abundance estimates of wild prey (Blue sheep Pseudois nayaur and Ibex Capra sibirica) using double observer surveys. C is the number of groups seen by both observers, S1 is the b is the estimated number of groups, N is estimated population size and P1, number of groups seen by first observer only, S2 is the number of groups seen by second observer only, G P2 are detection probabilities for observer one and two respectively. The surveys were conducted twice during the study period. Survey One

Chandertal Lossar Dhankar-Lalunga Mane Poh-Sichling Demul Tabo Langza-Hikkim Kibber-Chicham Sumra a

Survey Two

C

S1

S2

b G

N

P1

P2

C

S1

S2

b G

N

P1

P2

0 0 0 5 0 6 8 24 22 10

0 0 0 0 0 3 5 2 1 1

0 0 0 0 0 0 0 1 3 1

0.00 0.00 0.00 5.00 0.00 9.00 13.00 27.08 26.13 12.09

0 0 0 83 4 125 131 450 509 216

0.00 0.00 0.00 1.00 0.00 1.00 1.00 0.96 0.88 0.91

0.00 0.00 0.00 1.00 0.00 0.67 0.62 0.92 0.96 0.91

0 0 7 4 8 6 6 6 16 6

0 0 0 1 2 0 3 2 0 1

0 0 0 0 0 2 0 1 2 2

0.00 0.00 7.00 5.00 10.00 8.00 9.00 9.29 18.00 9.29

9 20 88 71 93 145 180 158 357 150

0.00 0.00 1.00 1.00 1.00 0.75 1.00 0.86 0.89 0.75

0.00 0.00 1.00 0.80 0.80 1.00 0.67 0.75 1.00 0.86

Estimation done only once.

Average N

Survey area

Density

4.5 10 88 77 48.5 135 155.5 304 433 183

89.54 91.11 41.65 61.52 30.01 62.14 70.99 92.45 112.98 47.33

0.05 0.11 2.11 1.25 1.62 2.17 2.19 3.29 3.83 3.87

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Table 3 Models that best explained snow leopard habitat use in a multiple use landscape, ranked using the Akaike’s Information Criterion values corrected for small sample sizes. Columns include number of parameters (K), Log Likelihood, AICc values, distance from lowest AICc (D AICc), model weight xi and adjusted R-squared values (r-sq.). Model

K

logLik

AICc

D AICc

xi

r-sq.

(1) (2) (3) (4) (5) (6) (7)

2 3 3 2 4 4 6

31.58 28.51 30.94 27.71 33.16 31.04 34.52

53.20 47.00 45.90 45.4 41.31 37.10 1.00

0.00 6.15 7.29 7.75 11.84 16.09 53.13

0.91 0.04 0.02 0.02 0.00 0.00 0.00

0.55 0.16 0.41 0.019 0.56 0.32 0.49

wild prey density human population density livestock density + I(livestock density^2) ruggedness wild prey density + livestock density + I(livestock density^2) human population density + livestock density + I(livestock density^2) wild prey density + human population density + ruggedness + livestock density + I(livestock density^2)

day

night

100% 90% 80% 70% 60% 50% 40% 30% 20% 10% 0% 1073(10) 1060(21) 853(20) 841(21)

739(7)

444(18) 402(20) 354(15)

50(0)

0(6)

Human populaon size Fig. 4. Snow leopard activity patterns in ten sampled sites across a gradient of human population size. One of the two sites used by migratory pastoralists recorded the minimum percent day captures (Site 5 from left in the figure) while the other one (Site 9 from left in the figure) did not record any snow leopard captures. Numbers on x-axis show human population size, numbers in brackets are total number of snow leopard captures at each site.

5.1.2. Snow leopard activity patterns Snow leopard activity in the day (5 am to 8 pm; based on dawn and dusk timings in the landscape) and night was similar across all ten sites and did not show any discernible relationship with human population size (Fig. 4). The highest percent captures in the day across sites was 67%, and day time captures were recorded at all sites where snow leopards were captured.

not record a single snow leopard capture at this site, owing presumably to high livestock densities, presence of dogs and lack of wild prey (only 5 Ibex individuals recorded in our surveys). Though terrain ruggedness is known to have a strong influence on snow leopard habitat use, our model with ruggedness as the explanatory variable did not receive support in model selection (D AICc = 8.08, xi = 0.02). There are likely two reasons for this. First because our camera trap placement was optimized through selection of locations that would maximize our chances of capturing snow leopards and such locations occur along rugged landscape features such as cliffs and ridgelines, effectively reducing the net variation in ruggedness in our dataset. Second the importance of ruggedness may not have manifested itself at the scale of our study. Pastoralism is the dominant land use type in snow leopard habitats across Central Asia. Our study, based on habitat use patterns of snow leopards, suggests that continued co-existence between people and snow leopards is possible till a threshold where livestock and associated anthropogenic activities do not affect wild prey populations negatively, and pastoral practices are less disturbing. We have, however, not explored the potential role of disease prevalence and transfers between livestock, snow leopards, and wild ungulates, a subject that should be examined in future studies. Assisting local communities with livelihood diversification to reduce stocking density of livestock, and involving them in conservation programmes appears important for snow leopard conservation and their coexistence with people and livestock (Mishra et al., 2003).

6. Discussion The abundance of carnivores such as the tiger Panthera tigris is reported to primarily be driven by the abundance of their wild prey (Karanth et al., 2004). Our results show that wild prey density was the primary determinant of habitat use by snow leopards. We also infer that in the absence of snow leopard persecution, productive habitats with healthy wild prey populations seem to be suitable for snow leopards even in the presence of livestock. Beyond a threshold of livestock density, however, reduced wild prey populations due to competition for resources (Mishra et al., 2001, 2010; Namgail et al., 2006) and associated anthropogenic activities appear to negatively influence snow leopard habitat use. This is reflected in the quadratic relationships between snow leopard habitat use and livestock density as well between wild prey density and livestock density. We recorded a notable difference in stocking density and herding practices between the local resident livestock grazing and migratory livestock grazing in our study area. Two of our study sites were used by migratory herders, and were marked by very high livestock abundances. Another major difference was that the migratory herders traditionally use 4–5 Tibetan Mastiff dogs to protect their livestock, while the local communities in Spiti do not use guard dogs. The Chandertal valley, one of our study sites used by migratory livestock had 5000 sheep and goats. We did

Acknowledgments Primary support for this project came through grants from Association of Zoos and Aquariums Conservation Endowment Fund, Disney Conservation Fund, and Fondation Segré - Whitley Fund for Nature. We also thank Snow Leopard Network, Department of Science and Technology and Panthera. We are thankful to the Chief Wildlife Warden, Himachal, Divisional Forest Officer, Kaza and the Range Officer, Kaza, for permissions and logistics. We thank Kulbhushansingh Suryawanshi for helping with data analysis and his comments on the manuscript. We are also thankful to Suhel Qader for helping with data analysis. Chandrima Home helped immensely with snow leopard photo-identification. Chunnit Kesang, Tenzin Thukten, Rinchen Tobgey, Sushil Dorje, Chudim, Takpa are thanked for tremendous support in fieldwork. References Anderson, D., Burnham, K., 2002. Avoiding pitfalls when using informationtheoretic methods. J. Wildl. Manage. Athreya, V., Odden, M., Linnell, J.D.C., Krishnaswamy, J., Karanth, K.U., 2013. Big Cats in our backyards: persistence of large carnivores in a human dominated landscape in India. PLoS ONE 8, e57872. http://dx.doi.org/10.1371/ journal.pone.0057872.

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