Biological Conservation 190 (2015) 8–13

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Biological Conservation

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Short communication

Does livestock benefit or harm snow leopards?

http://dx.doi.org/10.1016/j.biocon.2015.04.0260006-3207/� 2015 Elsevier Ltd. All rights reserved.

⇑ Corresponding author. Tel.: +91 8212515601; fax: +91 8212513822.E-mail addresses: rishi.eco@gmail.com, rishi@ncf-india.org (R.K. Sharma).

Rishi Kumar Sharma a,⇑, Yash Veer Bhatnagar a,b, Charudutt Mishra a,b

a Nature Conservation Foundation, 3076/5, 4th Cross, Gokulam Park, Mysore 570002, Indiab 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 2014Received in revised form 15 April 2015Accepted 21 April 2015

Keywords:Panthera unciaTrans-HimalayaPastoralismLarge carnivoresLivestock grazingCo-existence

a b s t r a c t

Large carnivores commonly prey on livestock when their ranges overlap. Pastoralism is the dominantland use type across the distributional range of the endangered snow leopard Panthera uncia. Snow leop-ards are often killed in retaliation against livestock depredation. Whether livestock, by forming an alter-native prey, could potentially benefit snow leopards, or, whether livestock use of an area is detrimental tosnow leopards is poorly understood. We examined snow leopard habitat use in a multiple use landscapethat was comprised of sites varying in livestock abundance, wild prey abundance and human populationsize. 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 basedon the total independent photographic captures and the number of snow leopard individuals captured ateach site. We quantified livestock abundance, wild prey abundance, human population size and terrainruggedness in each of the sites. Key variables influencing snow leopard habitat use were identified usingInformation Theory based model selection approach. Snow leopard habitat use was best explained bywild prey density, and showed a positive linear relationship with the abundance of wild ungulates. Wefound 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 snowleopard habitat use are potentially compatible up to a certain threshold of livestock density, beyondwhich habitat use declines, presumably due to depressed wild ungulate abundance and associatedanthropogenic disturbance.

� 2015 Elsevier Ltd. All rights reserved.

1. Introduction

The impact of anthropogenic activities such as livestock grazingand resource extraction on large carnivores has been debated.Anthropogenic activities are reported to depress carnivore popula-tions through direct persecution (Cardillo et al., 2004; Inskip andZimmermann, 2009; Winterbach et al., 2012), hunting of wild prey(Karanth et al., 2004) and habitat loss (Harihar et al., 2009). On theother hand, some carnivores appear to adapt and even thrive inhuman modified environments (Athreya et al., 2013; Bouyeret al., 2014; Odden et al., 2014) and local communities and carni-vores could even benefit from each other under specific conditions(Banerjee et al., 2013). Carnivores populations can persist despitehigh human populations when cultural tolerance is high (Karanthand Chellam, 2009), conflicts are managed properly (Treves andKaranth, 2003) and conservation policies are effectively imple-mented (Linnell et al., 2001).

Snow leopards (Panthera uncia) are endangered and face adiversity of threats across much of their range (IUCN Red List,2014). Our understanding of snow leopard ecology remains limiteddue to their elusive nature, low densities and the remote moun-tainous terrain they inhabit. Snow leopard habitats throughouttheir distribution range are multiple use landscapes, supporting avariety of natural resource uses including livestock grazing, agri-culture and extraction of medicinal plants and fuel-wood (Mishraet al., 2010). How snow leopards respond to these various anthro-pogenic factors within their landscapes continues to remain poorlyunderstood.

Livestock rearing is an important livelihood source for localpeople across the distributional range (Mishra, 1997), and livestockdepredation by snow leopards brings their conservation into con-flict with the goals of pastoral production (Bagchi and Mishra,2006; Bagchi et al., 2004; Jackson et al., 2010). Many studies havereported rather high contribution of livestock (up to 70%) to thediet of the snow leopard (Anwar et al., 2011; Bagchi and Mishra,2006), leading to the proposition that local livestock may be play-ing a role in sustaining populations of this endangered carnivore

R.K. Sharma et al. / Biological Conservation 190 (2015) 8–13 9

(Wegge et al., 2012). On the other hand, overstocking of rangelandswith livestock (Mishra et al., 2001) can lead to decline of wild preypopulations as a result of competition (Mishra et al., 2004; Namgailet al., 2006), which could further escalate the extent of livestockdepredation. For example, Bagchi and Mishra (2006) reported agreater presence of livestock in the diet of snow leopards in areaswith lower wild prey abundance. This creates a conservationdilemma. On the one hand, pastoralism has indirect negativeeffects on snow leopards as livestock depredation leads to retalia-tory persecution by herders (Jackson et al., 2010) and high live-stock abundance suppresses wild prey populations. On the otherhand, it is suspected that livestock might actually be an importantfood source for snow leopards. Whether livestock, by acting as anadditional food resource, is potentially beneficial to the snow leop-ard, or whether livestock use of an area negatively affects snowleopard habitat use, is not understood. We aimed to address thisquestion by examining snow leopard habitat use along gradientsof livestock and wild ungulate abundance.

2. Study area

Spiti Valley, forming the catchment of the river Spiti, is a part ofthe Trans-Himalayas in the state of Himachal Pradesh, India. Lyingin 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 snowfallhas declined in the past decade (Singh et al., in press).Agro-pastoralist communities have inhabited this region for twoto three millennia. Parts of the landscape are visited by transhu-mant Gaddi herders accompanied by large flocks of sheep and goatand Tibetan Mastiff guard dogs. The local agro-pastoral

Fig. 1. Map of the study area showing sampling sites and camera trap locations. The m

community, largely Buddhist, do not use guard dogs and are con-centrated 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 extensivepastures 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 referredto as large stock and, (ii) medium and small-bodied herded, hence-forth referred to as herded stock (cow, donkey, cow–yak hybrid,goat and sheep; (Suryawanshi et al., 2013). Villages have clearguidelines on the grazing rights to pastures but some of the pas-tures are shared between neighboring villages. Herded stock istaken out to pastures every morning and brought back to villagesand kept in corrals by individual households. A designated shep-herd referred to as ‘Lugzi’ is responsible for livestock herding anddecides on which pastures to use on a daily basis (Singh et al., inpress). Every household helps the ‘Lugzi’ with livestock grazingon a rotational basis. Families mostly own small agricultural landholdings (1–2 ha).

The Snow Leopard Trust and Nature Conservation Foundationhave been working in the region for 15 years. A key characteristicof our study site is that there is almost no retaliatory persecutionof snow leopards in the region, partly due to these conservationprograms, prevalence of Buddhism, and awareness of laws. Thus,with human persecution being controlled for, this landscape wassuitable to answer our question.

Within this large ca. 4000 km2 landscape, we selected ten sitesfor our study. These ten sites represented a gradient of livestockdensity, wild prey abundance and human population size. Theirboundaries were demarcated based on natural boundaries of thetopographical features.

ap inset shows locations of the study area in the state of Himachal Pradesh, India.

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

The ratios of livestock to wild prey varied from zero livestock atone site to 1111:1 at the other extreme.

3. Methods

The ten sampled sites ranged from 42 to 112 km2 (Table 1). Wedeployed 10 Reconyx RM45 and RC55 camera traps at ten bestlocations within each site for a period of two months, guided bya 3 � 3 km grid cell based design. The camera traps were installedat 5 sites for 60 days from May to July and in the remaining 5 sitesfor another 60 days from July to September, thus the samplingbeing conducted from the beginning to the end of the summer sea-son. Trap locations were selected based on the frequency of signs ofsnow leopards such as pugmarks, scrapes and scent marks atprominent features such as ridgelines and cliffs. Camera traps wereprogrammed to take a picture every second and till the animalremained within the camera range.

We developed a relative use index RUI ¼ cC � n

N

� �: as a response

variable indicating snow leopard habitat use. The RUI assigns ahabitat use value to each sampled site based on number of cap-tures and number of adult snow leopards captured at each site.Where c = number of independent photographic captures of adultsnow leopards at a particular site, C = total number of independentphotographic captures of adult snow leopards across all sites,n = number of adult snow leopard individuals captured at a partic-ular site and N = total number of adult snow leopards capturedacross all sites’’. RUI thus assigns high value to sites having largerproportion of snow leopard captures and higher number of uniquesnow leopard individuals and can take values between 0 and 1.

We quantified four key influential variables that could poten-tially affect snow leopard habitat use. These included populationsize of large and small bodied livestock, determined using doorto door census at all the sites. This involved censuses in 51 villagesacross the ten sites. The abundance of wild ungulate prey, bluesheep Pseudois nayaur and ibex Capra sibirica, was estimated ineach site twice, using the capture–recapture based double observersurvey technique (Suryawanshi et al., 2012) and the average of thetwo estimates was used for analysis. Livestock stocking densitiesand wild prey densities for all ten sites were calculated by dividingtheir respective abundance estimates by the area of each site.Human population size was included as an indicator of anthro-pogenic activities (Harihar and Pandav, 2012; Woodroffe, 2000).Anthropogenic activities can influence activity patterns of carni-vores, with carnivores avoiding high human activity periods(Carter et al., 2015). Human population was derived from Censusof India data (http://censusindia.gov.in/).

As snow leopards use steep and rugged terrain, we derived aTerrain Ruggedness Index (Riley et al., 1999) for each site using a30 � 30 m Digital Elevation Model Data from Aster Global Digital

Table 1Snow leopard habitat use estimated through photographic sampling of 10 sites in Spiti Vleopard photo-captures, the number of adult snow leopards captured in each site, their Rewild prey density, human population size, terrain ruggedness and altitudinal range at eac

Site name Area Humanpopulation

Ruggedness Livestockdensity (km2)

Wild preydensity (lm

Kibber-Chicham 112 841 26.35 20.04 3.83Dhankar-Lalung 42 853 37.7 47.61 2.11Sumra 47 402 42.94 35.79 3.86Demul 62 444 31.96 14.69 2.17Langza-Hikkim 92 1060 23.82 12.29 3.28Mane 62 354 39.43 13.68 1.25Lossar 91 739 39.13 57.23 0.1Poh-Sichling 30 0 35.46 0 1.63Tabo 71 1073 45.18 13 2.19Chandertal 90 50 27.7 55.84 0.05

Elevation Model data using the terrain analysis plugin inQuantum GIS 2.0 software (QGIS Development Team, 2013). Theterrain was categorized into six categories ranging from level toextremely rugged. Combined proportion of the highly rugged andextremely rugged category was used in the analysis.

We also examined snow leopard activity patterns in each site byextracting the date and time stamp information from the snowleopard camera trap images. Retributory killing of snow leopardsis a global cause of concern but in our landscape, long-termcommunity-based conservation programs (Mishra et al., 2003)and the Buddhist beliefs of local people have ensured that therehas been just one case of a snow leopard being killed over the last15 years. The current human population density in this landscapeis <1 person per km2 and agro-pastoralism is the dominant activityacross the landscape. Agricultural fields and orchards are situatedin the immediate vicinity of villages while villages themselvesare largely located in the valley along the Spiti River and not scat-tered across the landscape. At the current population density andthe kind and scale of anthropogenic activities, we did not considerhuman disturbance per se to have a major impact on snow leop-ards in our study area. However anthropogenic pressures wereconsiderably high in the two sites visited by nomadic pastoralistsas these sites were characterized by high livestock stocking densi-ties, use of guard dogs to protect livestock, and intensive use of theentire sites for livestock grazing. Since our study area is one con-tiguous landscape free from any natural or man-made barriersfor snow leopards, we considered it as a single population of snowleopards. We used the proportion of highly rugged and extremelyrugged terrain as an important variable in the analysis withthe assumption that good snow leopard habitats would becharacterized by a combination of rolling and rugged terrain whererolling terrain with pastures supports the prey species whilerugged terrain is essential for an ambush predator like snowleopard to hunt.

4. Data analysis

Snow leopard individuals in the photographs were identified byunique pelage patterns (Jackson et al., 2006). We used exploratoryanalysis using scatterplots and simple linear regression to under-stand the relationship between snow leopard habitat use and indi-vidual explanatory variables.

We used linear regressions to model the relationship betweensnow leopard habitat use and four of the key measured explana-tory variables. We scaled all continuous variables by normalizingto their means for the linear regression modeling. We used a quad-ratic term for livestock in the regression models since exploratorydata analysis showed that RUI initially increased with an increasein livestock density and then declined after a threshold (Fig. 2b).

alley, along with their topographic and biotic characteristics. The table details snowlative Use Index, and potential explanatory variables including the livestock density,h site.

2)Snow leopardphoto-captures

Snow LeopardIndividuals

RUI Altitudinalrange (m)

Presence ofguard dogs

21 8 0.0507 3970–5043 No20 6 0.0362 3724–5160 No20 5 0.0302 3137–4576 No18 5 0.0272 3684–4964 No21 4 0.0254 4049–5019 No15 3 0.0136 3551–5168 No

7 2 0.0042 3968–5139 Yes6 2 0.0036 3453–4663 No

10 1 0.003 3216–4760 No0 0 0 4034–5111 Yes

y = 0.0095xR² = 0.5982, P-value = 0.0086

0.00

0.01

0.02

0.03

0.04

0.05

0.06

0.00 0.50 1.00 1.50 2.00 2.50 3.00 3.50 4.00 4.50

RUI

Wild prey density

y = -4E-05x2 + 0.0024xR² = 0.534, P value = 0.064

0.00

0.01

0.02

0.03

0.04

0.05

0.06

0 10 20 30 40 50 60 70

RUI

Livestock stocking density

(a)

(b)

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

y = -0.0043x2 + 0.2492xR² = 0.6166, p-value = 0.017

0

0.5

1

1.5

2

2.5

3

3.5

4

4.5

0 10 20 30 40 50 60 70

Wild

pre

y de

nsity

Livestock stocking density

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

R.K. Sharma et al. / Biological Conservation 190 (2015) 8–13 11

We used an information-theoretic approach to develop aprioria set of models that best explained the habitat use of snowleopards. Our global model included wild prey density, livestockstocking density, human population size and terrain ruggednessas potential variables influencing snow leopard habitat use. Wethen developed 6 candidate sub-models using the variablesfrom this global model. Each sub-model represented a specifichypothesis explaining the relationship between explanatoryvariables and snow leopard habitat use. Akaike InformationCriterion adjusted for small samples (AICc) was used for modelselection (Anderson and Burnham, 2002). We implemented thisusing the package MuMIn (Barton, 2013) in program R (R CoreTeam, 2013).

5. Results

We obtained 138 independent captures of 24 adult snow leop-ards across ten sampling sites (Table 1). The estimates of livestock

Table 2Abundance estimates of wild prey (Blue sheep Pseudois nayaur and Ibex Capra sibirica) usinnumber of groups seen by first observer only, S2 is the number of groups seen by second obsP2 are detection probabilities for observer one and two respectively. The surveys were con

Survey One Survey Two

C S1 S2 bG N P1 P2 C S1

Chandertal 0 0 0 0.00 0 0.00 0.00 0 0Lossar 0 0 0 0.00 0 0.00 0.00 0 0Dhankar-Lalunga 0 0 0 0.00 0 0.00 0.00 7 0Mane 5 0 0 5.00 83 1.00 1.00 4 1Poh-Sichling 0 0 0 0.00 4 0.00 0.00 8 2Demul 6 3 0 9.00 125 1.00 0.67 6 0Tabo 8 5 0 13.00 131 1.00 0.62 6 3Langza-Hikkim 24 2 1 27.08 450 0.96 0.92 6 2Kibber-Chicham 22 1 3 26.13 509 0.88 0.96 16 0Sumra 10 1 1 12.09 216 0.91 0.91 6 1

a Estimation done only once.

abundance, wild prey abundance, livestock stocking densities, wildprey densities, human population size and ruggedness are pre-sented in Table 1.

The sites showed considerable variation in wild prey abundanceand density (abundance ranging from 5 to 433 and density rangingfrom 0.05 to 3.86/km2; Table 2) as well as livestock abundance andstocking densities (abundance ranging from zero at one site to5215 livestock heads at a site visited by nomadic herders; stockingdensity ranging from zero to 57.23 livestock heads per km2). Theobserver detection probability for the double observer surveysfor estimating wild prey abundance ranged from 0.61 to 1 acrossobservers and sites (Table 2).

Exploratory analysis using scatterplots indicated a monotonicrelationship of RUI with wild prey density and a quadratic relation-ship with livestock stocking density (Fig. 2a and b). There appearedto be a quadratic relationship between wild prey density/km2 andlivestock stocking density/km2 (Fig. 3).

5.1. Correlates of snow leopard habitat use

5.1.1. Model SelectionThe best fit model (xi = 0.92) suggested that wild prey density

influenced snow leopard habitat use (Table 3). The next best sup-ported model (D AICc = 0.92, xi = 0.23) included livestock stockingdensity. Based on distance from the lowest AICc (D AICc) andmodel weight (xi), there was little support for rest of the models.The parameter coefficients for the best models showed positiveeffect of wild prey (b = 0.013 ± 0.0038 SE) on snow leopard habitatuse.

g double observer surveys. C is the number of groups seen by both observers, S1 is theerver only, bG is the estimated number of groups, N is estimated population size and P1,ducted twice during the study period.

Average N Survey area Density

S2 bG N P1 P2

0 0.00 9 0.00 0.00 4.5 89.54 0.050 0.00 20 0.00 0.00 10 91.11 0.110 7.00 88 1.00 1.00 88 41.65 2.110 5.00 71 1.00 0.80 77 61.52 1.250 10.00 93 1.00 0.80 48.5 30.01 1.622 8.00 145 0.75 1.00 135 62.14 2.170 9.00 180 1.00 0.67 155.5 70.99 2.191 9.29 158 0.86 0.75 304 92.45 3.292 18.00 357 0.89 1.00 433 112.98 3.832 9.29 150 0.75 0.86 183 47.33 3.87

Table 3Models 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) wild prey density 2 31.58 �53.20 0.00 0.91 0.55(2) human population density 3 28.51 �47.00 6.15 0.04 0.16(3) livestock density + I(livestock density^2) 3 30.94 �45.90 7.29 0.02 0.41(4) ruggedness 2 27.71 �45.4 7.75 0.02 0.019(5) wild prey density + livestock density + I(livestock density^2) 4 33.16 �41.31 11.84 0.00 0.56(6) human population density + livestock density + I(livestock density^2) 4 31.04 �37.10 16.09 0.00 0.32(7) wild prey density + human population density + ruggedness + livestock density + I(livestock density^2) 6 34.52 1.00 53.13 0.00 0.49

0%10%20%30%40%50%60%70%80%90%

100%

1073(10) 1060(21) 853(20) 841(21) 739(7) 444(18) 402(20) 354(15) 50(0) 0(6)

Human popula�on size

day night

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

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

5.1.2. Snow leopard activity patternsSnow leopard activity in the day (5 am to 8 pm; based on dawn

and dusk timings in the landscape) and night was similar across allten sites and did not show any discernible relationship withhuman population size (Fig. 4). The highest percent captures inthe day across sites was 67%, and day time captures were recordedat all sites where snow leopards were captured.

6. Discussion

The abundance of carnivores such as the tiger Panthera tigris isreported to primarily be driven by the abundance of their wild prey(Karanth et al., 2004). Our results show that wild prey density wasthe primary determinant of habitat use by snow leopards. We alsoinfer that in the absence of snow leopard persecution, productivehabitats with healthy wild prey populations seem to be suitablefor snow leopards even in the presence of livestock. Beyond athreshold of livestock density, however, reduced wild prey popula-tions due to competition for resources (Mishra et al., 2001, 2010;Namgail et al., 2006) and associated anthropogenic activitiesappear to negatively influence snow leopard habitat use. This isreflected in the quadratic relationships between snow leopardhabitat use and livestock density as well between wild prey den-sity and livestock density.

We recorded a notable difference in stocking density and herd-ing practices between the local resident livestock grazing andmigratory livestock grazing in our study area. Two of our studysites were used by migratory herders, and were marked by veryhigh livestock abundances. Another major difference was that themigratory herders traditionally use 4–5 Tibetan Mastiff dogs toprotect their livestock, while the local communities in Spiti donot use guard dogs. The Chandertal valley, one of our study sitesused by migratory livestock had 5000 sheep and goats. We did

not record a single snow leopard capture at this site, owing pre-sumably to high livestock densities, presence of dogs and lack ofwild prey (only 5 Ibex individuals recorded in our surveys).

Though terrain ruggedness is known to have a strong influenceon snow leopard habitat use, our model with ruggedness as theexplanatory 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 throughselection of locations that would maximize our chances of captur-ing snow leopards and such locations occur along rugged land-scape features such as cliffs and ridgelines, effectively reducingthe net variation in ruggedness in our dataset. Second the impor-tance of ruggedness may not have manifested itself at the scaleof our study.

Pastoralism is the dominant land use type in snow leopardhabitats across Central Asia. Our study, based on habitat use pat-terns of snow leopards, suggests that continued co-existencebetween people and snow leopards is possible till a thresholdwhere livestock and associated anthropogenic activities do notaffect wild prey populations negatively, and pastoral practicesare less disturbing. We have, however, not explored the potentialrole of disease prevalence and transfers between livestock, snowleopards, and wild ungulates, a subject that should be examinedin future studies. Assisting local communities with livelihooddiversification to reduce stocking density of livestock, and involv-ing them in conservation programmes appears important for snowleopard conservation and their coexistence with people and live-stock (Mishra et al., 2003).

Acknowledgments

Primary support for this project came through grants fromAssociation of Zoos and Aquariums Conservation EndowmentFund, Disney Conservation Fund, and Fondation Segré - WhitleyFund for Nature. We also thank Snow Leopard Network,Department of Science and Technology and Panthera. We arethankful to the Chief Wildlife Warden, Himachal, DivisionalForest Officer, Kaza and the Range Officer, Kaza, for permissionsand logistics. We thank Kulbhushansingh Suryawanshi for helpingwith data analysis and his comments on the manuscript. We arealso thankful to Suhel Qader for helping with data analysis.Chandrima Home helped immensely with snow leopardphoto-identification. Chunnit Kesang, Tenzin Thukten, RinchenTobgey, Sushil Dorje, Chudim, Takpa are thanked for tremendoussupport in fieldwork.

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