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Interactions Between the , Livestock and Snow Leopards in the Sagarmatha National Park

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Series Title Chapter Title Interactions Between the Himalayan Tahr, Livestock and Snow Leopards in the Sagarmatha National Park Chapter SubTitle Copyright Year 2012 Copyright Holder Springer Science+Business Media B.V.

Corresponding Author Family Name Shrestha Particle Given Name Bikram Suffix Division Organization Forum of Natural Resource Managers Address Kathmandu, Nepal Email [email protected]

Author Family Name Kindlmann Particle Given Name Pavel Suffix Division Department of Biodiversity Research Organization Global Change Research Centre, AS CR Address Na sádkách 7, CZ-370 05, České Budějovice, Czech Republic Division Institute for Environmental Studies Organization Charles University Address Benátská 2, CZ-128 01, Prague 2, Czech Republic Email Author Family Name Jnawali Particle Given Name Shant Raj Suffix Division Program Terai Environment Organization National Trust for Nature Conservation Address Kathmandu, Nepal Email

Abstract Competition between wild and livestock for resources and interactions between these two and large predators are widely regarded as a major management issue in the Himalayas. Real data supporting these claims are scarce, but badly needed for developing good management strategies, which will effectively protect both wild ungulates and their predators in the Himalayas. Our study was done in August/September of 2006 in the Mongla and Phortse regions of the Sagarmatha National Park (SNP) with the aim of determining: (i) habitat overlap between tahr and domestic livestock, (ii) overlap in diets of tahr and domestic livestock, (iii) the effect of predators on tahr and (iv) explore the composition of vegetation in the region. Vantage points and regular monitoring from trails were used to observe the tahr and livestock. Direct observation and micro histological techniques were used to determine the overlap in diets of tahr and livestock. Diet of snow leopard was determined by scat analyses, which involved the microscopic identification of hair. There is overlap both in space and diet between tahr and livestock. Analysis of faecal samples revealed 24 species of plants in the faeces of tahr and 31 in those of livestock, of which 22 species were common to both. In total, 45 plant species were recorded at Mongla and 54 at Phortse. Two species of wild and four species of domestic were identified in the scats of snow leopard, with that of Himalayan tahr being the most frequent. In terms of domestic , the hair of was most frequently found in the faeces of snow leopard. The results of a questionnaire revealed that the snow leopard is the main predator of domestic livestock. We conclude that there is currently no serious competition between livestock and tahr for food, the main threat now comes from the decline in plant productivity in the region due to overgrazing. This trend could seriously change the situation, as tahr and livestock would then compete for food. The most effective way of reversing this trend is to introduce measures that limit the amount of grass that is harvested for feeding livestock during winter, which is rapidly increasing.

Keywords (separated by Himalayan tahr - Snow leopard - Livestock - Prey-predator dynamics '-') Chapter 5 1

Interactions Between the Himalayan Tahr, 2

Livestock and Snow Leopards 3

in the Sagarmatha National Park 4

Bikram Shrestha, Pavel Kindlmann, and Shant Raj Jnawali 5

Abstract Competition between wild ungulates and livestock for resources and 6 interactions between these two and large predators are widely regarded as a major 7 management issue in the Himalayas. Real data supporting these claims are scarce, 8 but badly needed for developing good management strategies, which will effectively 9 protect both wild ungulates and their predators in the Himalayas. Our study was done 10 in August/September of 2006 in the Mongla and Phortse regions of the Sagarmatha 11 National Park (SNP) with the aim of determining: (i) habitat overlap between tahr 12 and domestic livestock, (ii) overlap in diets of tahr and domestic livestock, (iii) the 13 effect of predators on tahr and (iv) explore the composition of vegetation in the 14 region. Vantage points and regular monitoring from trails were used to observe the 15 tahr and livestock. Direct observation and micro histological techniques were used to 16 determine the overlap in diets of tahr and livestock. Diet of snow leopard was deter- 17 mined by scat analyses, which involved the microscopic identification of hair. There 18 is overlap both in space and diet between tahr and livestock. Analysis of faecal sam- 19 ples revealed 24 species of plants in the faeces of tahr and 31 in those of livestock, of 20 which 22 species were common to both. In total, 45 plant species were recorded at 21 Mongla and 54 at Phortse. Two species of wild and four species of domestic mam- 22 mals were identified in the scats of snow leopard, with that of Himalayan tahr being 23 the most frequent. In terms of domestic animals, the hair of yak was most frequently 24

B. ShresthaUncorrected (*) Proof Forum of Natural Resource Managers, Kathmandu, Nepal e-mail: [email protected] P. Kindlmann Department of Biodiversity Research, Global Change Research Centre, AS CR, Na sádkách 7, CZ-370 05 České Budějovice, Czech Republic Institute for Environmental Studies, Charles University, Benátská 2, CZ-128 01 Prague 2, Czech Republic [AU1] S.R. Jnawali Program Terai Environment, National Trust for Nature Conservation, Kathmandu, Nepal

P. Kindlmann (ed.), Himalayan Biodiversity in the Changing World, DOI 10.1007/978-94-007-1802-9_5, © Springer Science+Business Media B.V. 2011 B. Shrestha et al.

25 found in the faeces of snow leopard. The results of a questionnaire revealed that the 26 snow leopard is the main predator of domestic livestock. We conclude that there is 27 currently no serious competition between livestock and tahr for food, the main threat 28 now comes from the decline in plant productivity in the region due to overgrazing. 29 This trend could seriously change the situation, as tahr and livestock would then 30 compete for food. The most effective way of reversing this trend is to introduce mea- 31 sures that limit the amount of grass that is harvested for feeding livestock during 32 winter, which is rapidly increasing.

33 Keywords Himalayan tahr • Snow leopard • Livestock • Prey-predator dynamics

34 5.1 Introduction

35 5.1.1 General Background

36 The Himalayan tahr (Hemitragus jemlahicus) belongs to the family and is 37 a migratory herbivore common at high altitudes. The Red Data Book of the Fauna 38 of Nepal (BPP 1995) has categorized it as a species likely to become extinct, while 39 IUCN has listed it in its category (K): Insufficiently Known. The Himalayan tahr, 40 one of three species of tahr, is native to the Southern range of the Himalayan 41 Mountains, including the Sagarmatha National Park (SNP) and is one of the most 42 common species there. The other two species of tahr include the 43 (Hemitragus hylocrius) found in Southern India and (Hemitragus 44 jayakari) in Oman. 45 In our study area, the Khumbu region, which is a part of the Sagarmatha National 46 Park (SNP), agro-pastoralism is the main occupation of the majority of people. 47 Though tourism is emerging as an alternative source of income for people, those 48 living in remote areas are still dependent upon traditional agriculture. Yak and its 49 hybrids are the major livestock at high altitudes, while cows prevail at low altitudes. 50 Domestic livestock in grazing the mountain pastures are thought to compete with 51 wild herbivores by depleting resources and degrading the pastures (Schaller 1977; 52 Shah 1998; Richari et al. 1992). The presence of livestock intensifies the competi- 53 tion betweenUncorrected plant species and results either in the loss of speciesProof or in coexistence 54 by partitioning of resources between species, spatially or temporally (Gause 1934; 55 Begon et al. 1986). Buffa et al. (1998) and Shrestha (2006) note a spatial overlap in 56 the occurrence of wildlife (tahr) and domestic animals, which is likely to lead to 57 competition for food and habitat destruction due to overpopulation. Wildlife- 58 livestock competition for resources is therefore widely regarded as a major manage- 59 ment issue, particularly in the mountainous protected areas, such as the Shey 60 Phoksundo National Park, Rara National Park, Khaptad National Park, Makalu 61 Barun National Park, Dhorpatan Reserve, Kanchanjunga Conservation 5 Interactions Between the Himalayan Tahr, Livestock and Snow Leopards...

Area and the Annapurna Conservation Area (Shrestha et al. 1990; KMTNC 1997; 62 Richard et al. 1999; Basnet 2002). 63 The return of the snow leopard to the Mount Everest National Park (Ale and 64 Boesi 2005) is likely to lead to conflicts between local people and snow leopard, 65 if in the future this predator kills significant numbers of domestic animals. This 66 predator may already have had a significant ecological effect on the prey-predator 67 ­dynamics and community structure in the region. Shrestha (2004) and Ale and 68 Boesi (2005) report that the kid to female ratio of the Himalayan tahr is now 69 ­alarmingly low: 0.1, whereas it was about 0.6–0.8 in 1991–1992 (Lovari et al. 70 2006). This might be due to predation of juveniles by snow leopard, but this 71 remains to be ­confirmed. As snow leopard and common leopard are potential 72 predators of tahr, conservation of tahr results also in the conservation of pastures 73 and large predators, which in turn play a critical role in maintaining the ecologi- 74 cal integrity of the region. In order to preserve wildlife and improve the standard 75 of living of local people, it is crucial that managers understand the interactions 76 between humans and wildlife. Therefore, it is extremely important to develop a 77 proper conservation strategy for the tahr in the SNP. However, neither this prob- 78 lem, nor the tahr-livestock ­prey-predator relationships in this area have been 79 studied systematically and it is unknown, whether the grass cover in the area is 80 sufficient to support both tahr and livestock. 81

5.1.2 Study Objectives 82

The overall aim of this study was to investigate: (1) interactions between Himalayan 83 tahr and livestock, (2) effect of predators on both these ungulates and (3) the com- 84 position of the vegetation of alpine pastures in tahr habitats in the Sagarmatha 85 National Park. 86 The specific objectives were to: 87

1. investigate the habitat overlap between tahr and livestock by comparing their use 88 of this habitat in terms of altitude, aspect, slope, percentage vegetation and ter- 89 rain type; 90 2. determine the differences in the diet of tahr and livestock and relative proportions 91 of the Uncorrecteddifferent food plants in their diets; Proof 92 3. determine the effect of predators on tahr and livestock – for this the percentage 93 of scats containing hair of particular items of prey (Himalayan tahr and live- 94 stock) was determined and the loss of both tahr and livestock due to predators 95 estimated; 96 4. explore diversity, productivity and vegetation cover of alpine pastures/meadows 97 in tahr habitats; 98 5. determine conservation implications for tahr, pasturelands, main predators of 99 tahr and the associated high-altitude ecosystems. 100 B. Shrestha et al.

101 5.2 Study Area

102 Sagarmatha National Park (SNP, 27º45¢-28º07¢ N, 86º28¢-87º07¢ E) is in the Solo 103 Khumbu district (Fig. 5.1) in the north-eastern region of Nepal. This National Park 104 was founded in July 1976 and included on the World Heritage List in 1979. It has 105 an area of 1,148 km2. The park encompasses the upper catchments of the Dudh Kosi 106 river system, which is fan-shaped and forms a distinct geographical unit surrounded 107 by high mountain ranges. The northern boundary is defined by the main divide of 108 the Great Himalayan Range, which follows the international border with the Tibetan 109 Autonomous Region of China. In the south, the boundary extends almost as far as 110 Monjo on the Dudh Koshi. The natural environment of the SNP is strictly protected. 111 It hosts 28 species of mammals, 199 species of birds, 6 species of amphibians and 112 7 species of reptiles. The weather is usually sunny in autumn, October and November, 113 but in winter the weather is cold with frequent falls of snow. The local people, the 114 Sherpas, having originated from Salmo Gang in the eastern Tibetan province of 115 Kham, some 2,000 km apart from their present homeland, are of great cultural inter- 116 est. There were approximately 3,500 Sherpas in 63 settlements, mainly located in

Uncorrected Proof

Fig. 5.1 The Sagarmatha National Park 5 Interactions Between the Himalayan Tahr, Livestock and Snow Leopards...

Fig. 5.2 The location of the two study areas at Mongla and Phortse

southern part of the park, in 1997. The study area is located in the Mongla and 117 Phortse rangelands in the Sagarmatha National Park (Fig. 5.2). 118

5.3 Materials and Methods 119 5.3.1 GeneralUncorrected Field Methods Proof 120 This study was done in August-September 2006. The areas selected were based on 121 the distribution of tahr, which was identified in a previous study carried out in 2004 122 (Shrestha 2006). 123 Vantage points and regular monitoring from trails (Jackson and Hunter 1996) 124 were the main methods used to record the numbers of tahr and livestock. Opportunistic 125 observations of tahr and livestock made during the vegetation survey provided addi- 126 tional data to those recorded in the fixed-point counts. 127 Once animals (tahr or livestock) were located, the area was searched and animals 128 identified using a 7 × 35 telescope. Each observation was treated as one group or 129 B. Shrestha et al.

130 sighting, irrespective of the number of individuals seen. For each sighting, the 131 ­following information was noted for an area with a radius of approximately 30 m 132 centered on the point of the highest aggregation of animals:

133 1. date and time of observation; 134 2. species of : tahr, yak hybrid (yak, nak, cow, dzom, zopkyo or ox); 135 3. group size and type (group types identified were: all males, females with young 136 or mixed groups); 137 4. sex and classification of tahr: female (adult female and yearling), male (class I, 138 class II and class III and kid, following Schaller (1977); [AU2] 139 5. slope; 140 6. aspect; 141 7. terrain type; 142 8. vegetation type; 143 9. distance to ridgeline; 144 10. distance to cliff; 145 11. GPS location of the point on the trail from where animals were sighted (wher- 146 ever possible) and any other remarks.

147 5.3.2 Habitat Overlap Between Tahr and Livestock

148 Habitat overlap between tahr and livestock was investigated by comparing the 149 ­habitats, in which these animals were observed, in terms of altitude, aspect, slope, 150 percentage vegetation cover and terrain. The altitude, aspect and slope were mea- 151 sured using GPS and a Brunton Compass meter. The vegetation categories were 152 grassland, rocks covered by vegetation and scrub. Similarly, the terrain was catego- 153 rized on the basis of ground surface morphology as smooth, rugged or very rugged. 154 A flat terrain with isolated undulating or small rocks was classified as smooth. 155 Distinctly undulating or rugged surface, including ravines and gullies with some 156 large rocks was classified as rugged. The presence of droppings was used to­estimate 157 the degree of spatial overlap. Uncorrected Proof

158 5.3.3 Dietary Overlap of Tahr and Livestock

159 5.3.3.1 Direct Observation

160 This was done in tahr areas using binoculars during the active feeding periods 161 between 7–12 a.m. and 3–5 p.m. Signs of plants having been recently eaten, such as 162 exudation of sap, crushed tissue, fresh clippings etc. were recorded. A herbarium 163 sheet for each plant species was prepared and taken to the Central Department of 5 Interactions Between the Himalayan Tahr, Livestock and Snow Leopards...

Environmental Science, Tribhuvan University, Kathmandu and The National 164 Herbarium, Godabari Centre, Kathmandu, for determination. 165

5.3.3.2 Micro Histological Analyses 166

A microscopic analysis of the indigestible plant fragments in dung, mainly epi- 167 dermal parts, which are characteristic of different plant groups (Metcalfe 1960), 168 was used to identify the plants eaten by the ungulates. This method is commonly 169 used for studying diets of ungulates, as it is simple, effective and accurate 170 (Baumgartner and Martin 1939; Anthony and Smith 1974; Dearden et al. 1975; 171 Holechek et al. 1982). It has one limitation: the amount forage consumed cannot 172 be quantified. 173

5.3.3.3 Faecal Analysis 174

Samples of faeces of tahr were collected from feeding sites in different habitats. 175 They were kept in paper bags and each day’s collection was labeled and air-dried 176 separately for a minimum of 72 h. After drying, individual faecal samples collected 177 on the same day were mixed thoroughly and packed in airtight polythene bags. 178 Later on, the samples were transported to the laboratory of the Department of 179 Environmental Sciences, Tribhuvan University, for further analyses. 180 Simultaneously, different plant species were collected and used to prepare refer- 181 ence slides for different tahr habitats. Slides were prepared following the method 182 used by Vavra and Holechek (1980) and Jnawali (1995), and used by Fjellstad and 183 Steinheim (1996) and Chetri (1999). The dried plant samples were separately 184 ground to a small size in an electric blender. The resultant powder was sieved 185 through two sieves (1 and 0.3 mm mesh size) placed one above the other. The mate- 186 rial that did not pass through the 0.3 mm sieve was used to prepare slides. A tea- 187 spoonful of each of the final samples was treated with warm 10% NaOH solution in 188 a test-tube and heated in a boiling water-bath for 4–6 min. The particles were 189 allowed to settle in a cold water-bath before the supernatant dark fluid was removed. 190 This procedure was repeated until a relatively clear supernatant solution was 191 obtained. Then, the material was washed 3–5 times with warm distilled water and 192 dehydratedUncorrected using 25%, 50%, 75%, 90% and then 100% alcohol. Proof The alcohol-treated 193 samples were finally treated using a series of solutions of xylene and alcohol in 194 which xylene gradually replaced the alcohol. A small amount of material was dried 195 between tissue paper and mounted in DPX under a 24 × 50 mm cover slip. The slide 196 was air dried for 5–6 days. 197 The slides of faecal samples were prepared in the same way as the reference 198 slides, except that 10% NaOH solution was replaced by a 5% NaOH solution, 199 before which the faecal samples were lightly washed with warm distilled water 200 to remove soil. A total of five slides were made of each composite faecal sample, 201 and labeled. 202 B. Shrestha et al.

203 The reference slides were studied as recommended by Holechek and Gross 204 (1982). A diagnostic key for each plant species was prepared using free hand 205 sketches and any distinct character of the plant fragments in the faeces was noted 206 and photographed and compared with the histological features of the plant material 207 on the reference slides. The features include cell wall structure, shape and size of 208 cells, hairs and trichomes, shape and size of stomata and inner-stomatal cells, fiber 209 structure and arrangements of veins. 210 A compound microscope with 100× magnification lens and an ocular measuring 211 scale was used to observe and measure the plant fragments on the slides of faecal 212 material. On each slide, ten fragments were identified in at least one transect of the 213 slide using the identification key and photographs of the epidermis of the reference 214 material. Only fragments recognized as epidermal tissue and consisting of at least 215 four plant cells or with visible stomata were recorded. In total, 200 fragments from 216 faecal samples of both livestock and tahr were identified.

217 5.3.3.4 Statistical Analysis

218 To evaluate the niche breadth of plant species in the diet of each species, Levin’s 219 measure of niche breadth (B) was calculated using the following formula cited in 220 Krebs (1999):

1 B = n , P2 å i 221 i=1

222 in which Pi is the percentage of total samples belonging to species i (i = 1, 2, …, n) 223 and n is the total number of species in all samples. The value of B increases with 224 increasing number of species in the diet. A low value indicates that a species is 225 selective and eats only a few plant species. 226 In order to estimate dietary overlap between two species, the Simplified Morisita’s

227 index (CH) was calculated using:

n 2 PP Uncorrectedå ij ik Proof i=1 CH = nn, PP22+ ååij ik 228 ii==11

229 where Pij and Pik are the proportions of the resource used by the two species, j and 230 k. The degree of overlap varies from zero to one; zero indicates no overlap and one 231 complete overlap. 5 Interactions Between the Himalayan Tahr, Livestock and Snow Leopards...

The Relative Importance Value (RIV) for each plant species in a faecal sample 232 was calculated using the method described by Jnawali (1995): 233

RIVi= D ii f , 234

where RIVi is the relative importance value for species i, Di is the mean percentage 235 of species i in the sample an fi is the frequency of species in the sample. 236

5.3.4 Floristic Composition 237

5.3.4.1 Sampling 238

A detailed vegetation analysis of the floristic composition of the study area was 239 done based on a set of “modules”, which are cross sections across the valley, 240 ­centered on the valley floor and including the entire altitudinal range on either side. 241 There were three “modules” for each rangeland (Mongla and Phortse), evenly 242 spaced at intervals of 3 km throughout the expected range of the tahr in this region. 243 At each of these “modules”, six transects were established following the altitudinal 244 contour, each of which was 100 m long. These transects were located at a range of 245 altitudes, with 100–150 m separating them. Sites in the valley, which were covered 246 by bushes and shrubs, were excluded, since tahr spend most of the time feeding in 247 open areas (Schaler 1973). Along these transects, 1 × 1 m quadrats were placed 20 m 248 apart. The plant cover in each quadrat and the use by livestock of the area in the 249 vicinity of each quadrat were determined. The percentage cover of individual plant 250 species, percentage of bare soil and rock in each quadrat were estimated visually 251 following Smart et al. (1976). A sub-plot of 25 × 25 cm was randomly selected in 252 each quadrat and all the vegetation in this sub-plot was removed and weighed. The 253 fresh weight of the grass was then used to estimate wet biomass per unit area of 254 pasture. Species area curves were plotted to calculate the minimum number of 255 squares required for determining the floristic composition of the study area (Jnawali 256 1995), which was found to be 18. 257 Uncorrected Proof 5.3.4.2 Statistical Analysis 258

Simpson’s Index of Diversity (Krebs 1994) was used to measure floral diversity: 259

S DP=-1 2 å i i=1 260

where D is Simpson’s index of diversity, Pi the proportion of individuals of species 261 i in the community and S the number of species in the community. Simpson’s diversity 262 index ranges from 0 (low diversity) to a maximum of (1 – 1/S). 263 B. Shrestha et al.

264 Sørenson’s index of similarity (ISs, Krebs 1994) was used to compare ­similarity 265 of plant species in two habitats, A and B:

2C ISs 100, =´ 266 AB+

267 where C is the number of species common to both habitats, A the total number of 268 species in habitat A and B the total number in habitat B.

269 Vegetation analysis – the following values were calculated: Number of quadrats in which species A occurred t1.1 Frequency of f =´100 A Total number of quadrats t1.2 species A:

fA t1.3 Relative fre- RFA =´S 100 t1.4 quency of å fi t1.5 species A: i=1 Number of individuals of species A in all quadrats t1.6 Density of D = A Total number of quadrats ´ Size of quadrat t1.7 species A: Number of individuals of species A t1.8 Relative density RD = A Total number of individuals of all species t1.9 of species A: Total number of individuals of species A t1.10 Abundance of A = A Number of quadrats, where species A occurred t1.11 species A: t1.12 Relative abun- RAAA=´ A 100 t1.13 dance of t1.14 species A: t1.15 Importance IVIA=++ RF A RD AA RA

270t1.16 Value Index:

271 Frequency classes: based on the prominence value, PV, species were assigned to 272 one of five frequency classes (Sharma 2002) as follows: t2.1 UncorrectedPV Frequency class Proof t2.2 <1 very rare t2.3 1–5 rare t2.4 5–25 common t2.5 25–75 abundant

273t2.6 > 75 very abundant 5 Interactions Between the Himalayan Tahr, Livestock and Snow Leopards...

5.3.5 Productivity and Forage Availability 274

The percentage cover of individual species in each quadrat was estimated visually 275 following Smart et al. (1976). Prominence values, PV, were calculated and used to 276 quantify the abundance of species at both of the rangelands following the method of 277 Dinerstein (1979): 278

PVA= M AA f , 279

where PVA is the prominence value for species A, MA is the mean percentage cover 280 of species A and fA is described above. Then the abundance of each species was 281 categorized as very rare (PVA £ 1), rare (1 < PVA £ 5), common (5 < PVA £ 40) or abun- 282 dant (PVA > 40). 283

5.3.6 Effect of Predators on Himalayan Tahr and Livestock 284

In order to estimate the effect of predators on Himalayan tahr and livestock, the 285 presence and identification of tahr or livestock hair in predator scat was determined 286 by examining the scat under a microscope. Economic loss of livestock to predators 287 was also estimated. 288

5.3.6.1 Microscopical Identification of Hair 289

The hairs in predator scats were used to determine the diet of the predators. The 290 analysis of scat samples required a reference collection of hair samples, an identifi- 291 cation key and slides of hair from scat. 292 Twenty scats of snow leopard were collected from the study area. The scats were 293 identified on the basis of their size and associated signs, such as scrapes and pug- 294 marks. The samples of scat were sun-dried, labelled and stored in polythene bags 295 for laboratory analysis. 296 Each sample was first examined macroscopically and the colour and texture 297 recorded.Uncorrected The sample was then put into 30% hydrogen peroxide Proof overnight prior to 298 microscopic examination. Each bleached sample was observed under a microscope at 299 400× magnification. The sample was then wet mounted in D.P.X. and the cortex and 300 medulla of the hairs and the details recorded. The micrometer measurement of both 301 the cortex and medulla were recorded at ten intervals along the shaft of each hair. 302 These measurements were then converted to millimetres and the mean calculated. 303 Similarly, the average diameter of the medulla was determined. In addition, medullar 304 indices and their averages were calculated. The above calculations were made only for 305 medullated hair. The medullary Index (MI) was calculated using the formula 306 B. Shrestha et al.

Width of medulla MI = .

307 Width of cortex

308 A tuft of hairs was inserted into a straw and then molten wax was sucked into the 309 straw. Once the wax solidified, the straw was cut open, the wax core and embedded 310 hair removed and cross sections obtained by transversally cutting it with a razor 311 blade. The sections were treated with xylene to remove the wax and viewed under a 312 microscope. 313 The characteristics of the cuticular scales along the shaft of each hair from the 314 root to the tip were noted. A thin layer of nitrocellulose lacquer (white nail polish) 315 was applied to the surface of microscope slides and the hair samples placed horizon- 316 tally on them. As the lacquer dried an impression of the surface of each hair formed, 317 then the hair was peeled off and the impression viewed under a microscope. 318 Microphotographs of representative cross sections, medulla and scale patterns 319 along the length of the hairs of each species were taken at a standard magnification. 320 Compared to direct comparison and using the dichotomous key, reference to photo- 321 graphs proved more convenient and easier for the routine identification of hair. 322 Therefore, a photographic reference key and microscopic examination of cuticular 323 scales and the medullary type, thickness of cortex and medulla, medullary index 324 etc., which are diagnostic tools for identifying species, were used in this study. 325 Descriptions of the hair in the key includes only the maximum diameter of the pri- 326 mary guard hair and the most diagnostic features of the hair of each species. The key 327 includes the eight wild and five domestic mammals found in the study area (not 328 presented here). 329 Each scat was soaked overnight in liquid dettol mixed with water and then 330 washed carefully over a sieve with a mesh width of 1 mm. Remains, like bones, 331 teeth, hooves, hair and feathers were removed, air dried and stored. Microscope 332 slide preparations of the hair in every scat were used to identify the species of prey 333 by using the key and the microphotographs of the hair of potential prey as was done 334 in preparing the key.

335 5.3.6.2 Statistical Analysis

336 The scat contents are presented here as “frequency of occurrence” (number of scats 337 containingUncorrected the hair of a particular species of prey) and “percentage Proof frequency of 338 occurrence” (% of scats containing the hair of a particular species of prey).

339 5.3.7 Depredation of Livestock

340 The questionnaires completed by the local herders at Phortse and the results of the 341 group discussions gave an indication of the economic loss due to the depredation of 342 livestock by snow and common leopards. 5 Interactions Between the Himalayan Tahr, Livestock and Snow Leopards...

5.4 Results 343

5.4.1 Abundance of Himalayan Tahr 344

During the observations at the Mongla and Phortse pasturelands there were a total 345 of 25 sightings of Himalayan tahr, which in total was 319 animals with an estimated 346 total population of about 125, whereas there were a total of 113 individuals in the 347 15 sightings of livestock (Tables 5.1 and 5.2). The male to female ratio was 0.72, kid 348 to female ratio 0.46 and yearling to kid ratio 0.65. Mean group size was 12.76 349 (range 1–38, standard deviation 10.65). 350

5.4.2 Composition and Numbers of Livestock 351

The 113 individuals of livestock included yak, jopkyo, cow, ox and calf. Yak made 352 up 56% of the livestock in the study area, followed by jopkyo, cow, ox and calf 353 (Table 5.2). 354

Table 5.1 Structure of the tahr population based on absolute numbers and all the animals seen at t3.1 Mongla and Phortse in the summer of 2006 (n = 25) t3.2 Mongla Phortse Known All animals Known All animals number tallied number tallied Himalayan tahr No. % No. % No. % No. % t3.3 Female 14 42 70 45 36 39 73 44 t3.4 Yearling 5 18 32 21 10 11 21 13 t3.5 Kid 5 15 22 14 18 19 32 19 t3.6 Total yearlings and kids 11 33 55 35 29 30 53 32 t3.7 Male I 2 6 6 4 10 11 13 8 t3.8 Male II 3 9 19 12 5 5 7 4 t3.9 Male III 3 9 6 4 13 15 18 11 t3.10 Total males 8 24 31 20 28 30 38 23 t3.11 Total Uncorrected33 100 155 100 92 Proof100 164 100 t3.12

Table 5.2 Numbers of the t4.1 Livestock No. % t4.6 different kinds and percentage t4.2 Yak 63 56 t4.7 composition of the livestock t4.3 Jopkyo 33 30 t4.8 observed at Mongla and t4.4 Phortse in the summer of 2006 t4.5 Cow 8 7 t4.9 Ox 5 4 t4.10 Calf 4 3 t4.11 Total 113 100 t4.12 B. Shrestha et al. t5.1 Table 5.3 The habitat preferences (frequency) of Himalayan tahr and livestock at Mongla and t5.2 Phortse, in summer 2006 t5.3 Himalayan t5.4 Habitat variable tahr, % Livestock, Habitat variable Himalayan Livestock t5.5 and category (n = 25) % (n = 15) and category tahr, % (n = 25) % (n = 15) t5.6 Habitat type Slope t5.7 Barren 0 0 0°–30° 8 33 t5.8 Grassland 96 100 31°–60° 76 67 t5.9 Shrubland 4 0 61°–90° 16 0 t5.10 Forest 0 0 t5.11 Terrain type Aspect t5.12 Cliff 0 0 East (69°–113° ) 0 0 t5.13 Very rugged 36 0 Southeast 28 20 t5.14 (204°–248°) t5.15 Rugged 36 47 South (159°–203°) 52 73 t5.16 Rolling 28 53 Southwest 20 7 t5.17 (114°–158°) t5.18 Flat 0 0 West (249°–293°) 0 0 t5.19 Altitude (m) Distance to escape terrain (m) t5.20 3,601–3,800 24 0 0 8 0 t5.21 3,801–4,000 44 27 1–50 32 0 t5.22 4,001–4,200 20 66 51–100 48 13 t5.23 4,201–4,400 8 7 101–150 0 7 t5.24 4,401–4,600 4 0 > 150 12 80 t5.25 Position on slope Nearest water source (m) t5.26 Low 40 0 <200 68 7 t5.27 Middle 40 87 201–400 4 0 t5.28 Upper 20 13 > 400 28 93 t5.29 Preferred habitats are in bold

355 5.4.3 Preferred Habitat of Himalayan Tahr and Livestock

356 Grassland was the preferred habitat of both tahr (96% of cases) and livestock (100% 357 of cases).Uncorrected Tahr preferred very rugged (36%) and rugged (36%)Proof slopes, whereas 358 livestock preferred rolling (53%) and rugged slopes (47%). Tahr equally preferred 359 the lower and middle slopes (40%), while livestock preferred the middle slopes 360 (87%). Tahr preferred to graze at altitudes between 3,600 and 4,200 m and livestock 361 between 4,000 and 4,200 m. Tahr preferred mostly southern aspects (52%) as did 362 the livestock (73%). Both tahr and livestock preferred 31–60° slopes (76% and 67% 363 respectively). Tahr was also found on slopes greater than 61°, which were avoided 364 by livestock. Livestock was found on gentle slopes (0–30°) more frequently than 365 tahr. Livestock was found in areas more than 150 m from escape terrain, whereas 366 tahr occurred in areas less than 100 m from escape terrain (Table 5.3). 5 Interactions Between the Himalayan Tahr, Livestock and Snow Leopards...

Droppings of both these ungulates were found in 45% of the squares at Phortse 367 and 33% of those at Mongla, and there were no droppings of either in 5% 368 of the squares in Phortse and 17% at Mongla. There were droppings in the remain- 369 ing 50% (in both regions) of either tahr or livestock, but not of both. 370

5.4.4 Diet Composition 371

5.4.4.1 Diet of Himalayan Tahr 372

Of the 24 species of plant in the droppings of tahr, 53% were grasses and sedges 373 belonging to 6 taxa: Carex anomoea, Avena sp., Poa sp., Trisetum spicatum, 374 Cyperaceae sp. and Imperata sp. They were followed by Gueldenstaedtia himalaica 375 and Potentilla sp., whereas Pedicularis siphonantha, Persicaria capitata, Androsace 376 sarmentosa, Trachyspermum ammi, Habenaria aitchisonii and Ephedra gerardiana 377 made up only a small fraction of the diet (Table 5.4). Niche breadth of tahr was 378 B = 0.0137. 379 In the diet of tahr, the highest RIV was for Avena sp. followed by Carex 380 anomoea, Poa sp., Gueldenstaedtia himalaica, Trisetum spicatum, Cyperaceae sp. 381 and Potentilla sp. Remaining species had very low RIVs (Table 5.4). 382

5.4.4.2 Diet of Livestock 383

Of the 31 plant species found in the droppings of livestock, 38.5% were grasses 384 and sedges belonging to 6 taxa: Avena sp., Carex anomoea, Trisetum spicatum, 385 Cyperaceae sp., Poa sp., and Imperata sp. They were followed by Cotoneaster 386 microphyllus, Potentilla sp., Bistorta affinis, Gueldenstaedtia himalaica, Polygonatum 387 hookerii, Saxifraga brachypoda, Anaphalis contorta, Rhododendron lepidotum, 388 whereas Anaphalis triplinervis, Fragaria daltoniana and Gentiana sp. made up 389 only a small fraction of the diet (Table 5.5). Niche breadth of livestock was 390 B = 0.0175. 391 In the diet of livestock, the highest RIV was for Avena sp. followed by Carex 392 anomoea and Cotoneaster microphyllus. Among the species with low RIVs were 393 GentianaUncorrected sp., Habenaria aitchisonii and Androsace sarmentosa Proof (Table 5.5). 394

5.4.5 Dietary Overlap Between Tahr and Livestock 395

Remnants of 22 species of common plants were found in tahr and livestock droppings 396 (Tables 5.4 and 5.5). Morina nepalensis and Ephedra gerardiana were found only in 397 tahr droppings, while Gentiana sp., Gerbera gossypina, Notholirion macrophyllum, 398 Parnassia nubicola, Polygonatum hookerii, Polygonum sp., Saxifraga parnassifolia, 399 B. Shrestha et al.

t6.1 Table 5.4 Species of plants found in the droppings of tahr, their t6.2 relative percentage and relative importance value (RIV) t6.3 Species name % RIV t6.4 Graminoids 28.0 146.1 t6.5 Avena sp. 19.0 117.1 t6.6 Imperata sp 2.5 5.6 t6.7 Poa sp. 6.5 23.4 t6.8 Sedges 25.0 109.2 t6.9 Carex anomoea 13.5 70.1 t6.10 Cyperaceae sp. 5.5 18.2 t6.11 Trisetum spicatum 6.0 20.8 t6.12 Herbaceous plants and shrubs 47.0 120.6 t6.13 Anaphalis contorta 2.5 5.6 t6.14 Anaphalis triplinervis 1.5 2.6 t6.15 Androsace sarmentosa 1.5 2.7 t6.16 Bistorta affinis 3.5 9.3 t6.17 Cotoneaster microphyllus 4.5 13.5 t6.18 Cyananthus hookerii 2.5 5.6 t6.19 Cypripedium himalaicum 1.0 2.6 t6.20 Ephedra gerardiana 1.0 1.4 t6.21 Fragaria daltoniana 2.5 5.6 t6.22 Gueldenstaedtia himalaica 6.0 20.8 t6.23 Habenaria aitchisonii 1.0 1.4 t6.24 Morina nepalensis 3.0 7.3 t6.25 Pedicularis siphonantha 1.5 2.6 t6.26 Persicaria capitata 1.5 2.6 t6.27 Potentilla sp. 5.5 18.2 t6.28 Rhododendron lepidotum 3.5 9.3 t6.29 Satyrium nepalense 2.0 4.0 t6.30 Saxifraga brachypoda 2.5 5.6

400 Salvia hians and Sedum sp. were found only the droppings of livestock. The Morisita

401 index of niche overlap between livestock and tahr was high: CH = 0.83.

402 5.4.6 FloristicUncorrected Composition and Vegetation Analysis Proof

403 5.4.6.1 Rangeland at Mongla

404 At Mongla, 71.48% of the ground is covered by vegetation, the remaining 18.29% 405 by bare soil and rock (10.23%). Vegetation consists of 45 species. The strongly 406 dominant taxon (taxon with the largest f-value) at Mongla was the sedge (grass) 407 Avena sp., followed by Cotoneaster microphyllus, Rhododendron lepidotum and 408 Carex anomoea – Annex 5.1. The Simpson’s diversity index for Mongla is 409 D = 0.941. 5 Interactions Between the Himalayan Tahr, Livestock and Snow Leopards...

Table 5.5 Species of plants found in the droppings of livestock, t7.1 their relative percentage and relative ­importance value (RIV) t7.2 Species name % RIV t7.3 Graminoids 18.5 84.7 t7.4 Avena sp. 14.0 74.1 t7.5 Imperata sp. 1.0 1.4 t7.6 Poa sp. 3.5 9.3 t7.7 Sedges 20.0 74.3 t7.8 Carex anomoea 8.5 35.0 t7.9 Cyperaceae sp. 5.0 15.8 t7.10 Trisetum spicatum 6.5 23.4 t7.11 Herbs and shrubs 61.5 161.3 t7.12 Anaphalis contorta 3.0 7.3 t7.13 Anaphalis triplinervis 2.5 5.6 t7.14 Androsace sarmentosa 0.5 0.5 t7.15 Bistorta affinis 4.0 11.3 t7.16 Cotoneaster microphyllus 8.0 32.0 t7.17 Cyananthus hookerii 1.5 2.6 t7.18 Cypripedium himalaicum 2.0 4.0 t7.19 Fragaria daltoniana 1.0 1.4 t7.20 Gentiana sp. 0.5 0.5 t7.21 Gerbera gossypina 1.0 1.4 t7.22 Gueldenstaedtia himalaica 3.5 9.3 t7.23 Habenaria aitchisonii 0.5 0.5 t7.24 Notholirion macrophyllum 1.5 2.6 t7.25 Parnassia nubicola 3.0 7.3 t7.26 Pedicularis siphonantha 1.5 2.6 t7.27 Persicaria capitata 1.0 1.4 t7.28 Polygonatum hookerii 3.5 9.3 t7.29 Polygonum sp. 2.5 5.6 t7.30 Potentilla sp. 6.5 23.4 t7.31 Rhododendron lepidotum 3.0 7.3 t7.32 Salvia hians 1.0 1.4 t7.33 Satyrium nepalense 2.0 4.0 t7.34 Saxifraga brachypoda 3.5 9.3 t7.35 Saxifraga parnassifolia 3.4 9.3 t7.36 UncorrectedSedum sp. 1.0 Proof1.4 t7.37

5.4.6.2 Rangeland at Phortse 410

At Phortse, 81.64% of the ground is covered by vegetation, 14.56% by bare soil and 411 3.8% by rock. Vegetation consists of 54 species. The dominant species (taxon with 412 the largest f-value) at Phortse is Carex anomoea, followed by Avena sp., Gerbera 413 gossypina and Pedicularis siphonantha – Annex 5.2. The Simpson’s diversity index 414 for Phortse is D = 0.937. 415 B. Shrestha et al.

416 5.4.6.3 Sørenson’s Index of Similarity (ISs)

417 The Sørenson’s index of similarity (ISs) for Mongla and Phorte is ISs = 0.83 (83%), 418 which indicates that the species at these two study sites are similar.

419 5.4.7 Productivity and Availability of Forage

420 The productivity of both rangelands was very similar: 2,643 kg/ha (wet weight) and 421 2,276 kg/ha (wet weight) at Mongla and Phortse, respectively. 422 Based on the prominence value, at Monga, 4 species were found in the drop- 423 pings of tahr and livestock that are very abundant or abundant (Avena sp., Carex 424 anomoea, Cotoneaster microphyllus and Rhododendron lepidotum), 12 species 425 that are common (Anaphalis contorta, Androsace sarmentosa, Cyananthus hook- 426 erii, Fragaria daltoniana, Gerbera gossypina, Habenaria aitchisonii, Notholirion 427 macrophyllum, Persicaria capitata, Potentilla sp., Satyrium nepalense, Saxifraga 428 brachypoda and Saxifraga parnassifolia), 9 that are rare (Bistorta affinis, 429 Gueldenstaedtia himalaica, Imperata sp., Cyperaceae sp., Poa sp., Polygonatum 430 hookerii, Salvia hians, Sedum sp. and Trachyspermum ammi) and 6 that are very 431 rare (Anaphalis triplinervis, Parnassia nubicola, Pedicularis siphonantha and 432 Trisetum spicatum) – Table 5.6. 433 At Phortse, 4 species found in droppings of tahr and livestock are abundant (Avena 434 sp. Carex anomoea, Polygonatum hookerii and Rhododendron lepidotum), 15 are 435 common (Bistorta affinis, Cotoneaster microphyllus, Cyananthus hookerii, Fragaria 436 daltoniana, Gentiana sp., Gerbera gossypina, Gueldenstaedtia himalaica, Imperata 437 sp., Notholirion macrophyllum, Parnassia nubicola, Pedicularis siphonantha, 438 Persicaria capitata, Polygonum sp. and Potentilla sp.), 7 are rare (Anaphalis con- 439 torta, Habenaria aitchisonii, Cyperaceae sp., Poa sp., Saxifraga brachypoda, 440 Saxifraga parnassifolia and Trachyspermum ammi), and 3 are very rare (Salvia 441 hians, Satyrium nepalense and Sedum sp.) – Table 5.6.

442 5.4.8 EffectUncorrected of Predators on Tahr and Livestock Proof

443 In total, 20 scats were analyzed. The prey consumed included two species of wild 444 and four species of domestic mammals. The most frequent prey of snow leopard is 445 Himalayan tahr, which was detected in 55% of the scats, followed by yak (25%), 446 cow (20%), musk (20%), dog (10%) and horse (10%) (Table 5.7). Wild species 447 were present in 75% and domestic species in 65% of scats. 448 The results of the questionnaire indicate that snow leopard is the main predator 449 of domestic livestock. Between January 2005 and September 2006, snow leopard 450 killed 16 animals belonging to 8 farming families (Table 5.8). 5 Interactions Between the Himalayan Tahr, Livestock and Snow Leopards...

Table 5.6 Prominence Value (PV) of the species most frequently eaten by tahr and t8.1 livestock at Mongla and Phortse t8.2 Species prominence t8.3 Value (PV) t8.4 Sp. No. Species name Mongla Phortse t8.5 1. Anaphalis contorta 6.9 1.3 t8.6 2. Anaphalis triplinervis 0.7 – t8.7 3. Androsace sarmentosa 5.8 – t8.8 4. Avena sp. 84.9 58.2 t8.9 5. Bistorta affinis 3.2 7.1 t8.10 6. Carex anomoea 44.1 67.7 t8.11 7. Cotoneaster microphyllus 102.6 21.9 t8.12 8. Cyananthus hookerii 11.1 17.1 t8.13 9. Fragaria daltoniana 7.8 8.2 t8.14 10. Gentiana sp. – 10.3 t8.15 11. Gerbera gossypina 10.1 11.9 t8.16 12. Gueldenstaedtia himalaica 4.3 6.3 t8.17 13. Habenaria aitchisonii 5.7 1.2 t8.18 14. Imperata sp. 1.9 6.0 t8.19 15. Cyperaceae sp. 1.2 5.0 t8.20 16. Notholirion macrophyllum 9.3 10.3 t8.21 17. Parnassia nubicola 0.8 18.7 t8.22 18. Pedicularis siphonantha 0.7 8.2 t8.23 19. Persicaria capitata 17.1 16.4 t8.24 20. Poa sp. 1.3 0.8 t8.25 21. Polygonatum hookerii 4.3 40.6 t8.26 22. Polygonum sp. – 8.6 t8.27 23. Potentilla sp. 10.8 11.6 t8.28 24. Rhododendron lepidotum 55.7 63.6 t8.29 25. Salvia hians 2.1 0.3 t8.30 26. Satyrium nepalense 8.8 0.1 t8.31 27. Saxifraga brachypoda 9.7 2.0 t8.32 28. Saxifraga parnassifolia 5.6 2.5 t8.33 29. Sedum sp. 2.1 0.4 t8.34 30. Trachyspermum ammi 1.6 1.5 t8.35 31. Trisetum spicatum 0.2 – t8.36 Uncorrected Proof Table 5.7 Absolute and relative frequencies of occurrence of prey items in the diet of t9.1 snow leopard (n = 20) t9.2 Prey species Frequency of occurrence % Frequency of occurrence t9.3 Himalayan tahr 11 55 t9.4 Yak 5 25 t9.5 4 20 t9.6 Cow 4 20 t9.7 Dog 2 10 t9.8 Horse 2 10 t9.9 B. Shrestha et al. t10.1 Table 5.8 Percentage t10.6 % killed by snow t10.2 composition of the six types t10.7 Livestock type leopard (N = 16) t10.3 of livestock killed by snow t10.8 Yak 43.7 t10.4 leopard at Phortse during t10.9 Nak 12.5 t10.5 2005–2006 t10.10 Jom 0.0 t10.11 Jopkyo 0.0 t10.12 Cow 31.2 t10.13 Ox 12.5

451 5.5 Discussion

452 5.5.1 Status and Population of Himalayan Tahr and Livestock

453 Based on these results it is estimated that 125 tahr inhabit the rangelands at Mongla 454 and Phortse. Lovari (1992) estimated that there were 300 tahr in 1989 present in the 455 SNP at Namche, which includes the Mongla and Phortse valleys. He also estimated 456 there were 27 individuals/km2 in the area between Phortse and Pangboche. Shrestha 457 (2006) surveyed tahr in 2004 and records 205 Himalayan tahr in the Namche 458 (including Mongla), Phorche and Thame valleys of the SNP. In October-November 459 2004 there were 104 tahr in Gokyo, Phortse and Namche (including Mongla), and 460 3.2 and 5.1 individuals/km2 in Phortse and Namche valleys, respectively, and in 461 August-November 2005, 277 tahr in Gokyo, Phortse, Namche and Thame valleys 462 and the number of individuals/km2 of tahr ranged from cca. one in the Gokyo to as 463 many as seven in the Namche valley (Ale 2006). Based on these data, it is likely that 464 the population of tahr is smaller than was recorded in 1989 by Lovari (1992). The 465 ratio of kid to female recorded in this study was 0.46, which is similar to that 466 recorded by Ale (2006). For both pasturelands, the low kid-to-female ratio is con- 467 sistent with what has been regularly reported for the area since 1992 (Lovari 1992, 468 Sandro Lovari, personal communication). In contrast, Schaler (1973) reports a kid- [AU3] 469 to-female ratio of 0.56 in Kang Chu, eastern Nepal, where tahr is hunted, and 0.57 470 in the Annapurna region of western Nepal, an area where there are no large preda- 471 tors of tahr (Gurung 1995). The low kid-to-female ratio in Sagarmatha may be due 472 to predation or disease. Snow leopard has recently re-colonized the Everest region. 473 In order toUncorrected determine, whether predation by snow leopards isProof responsible for the low 474 kid to female ratio of Himalayan tahr in Sagarmatha, the diet of snow leopards there 475 is currently being studied. 476 During this study only a small number of livestock (113 individuals) grazed the 477 pastures at Mongla and Phortse, compared to the 3169 at Namche and Khumjung 478 VDC (VDC is an administrative region in Nepal) in the Sagarmatha National Park 479 in 2003 (DNPWC/TRPAP 2006). This may be because and naks are moved to 480 high pastures in spring and during the monsoon months, and returned to settle- 481 ments at lower altitudes in summer and winter, and this study was done during the 482 summer season. 5 Interactions Between the Himalayan Tahr, Livestock and Snow Leopards...

5.5.2 Habitat Selection by Himalayan Tahr and Livestock 483

Himalayan tahr and livestock prefer to occupy similar habitats, with similar 484 aspects and to some extent slopes (Table 5.3). Both tahr and livestock were found 485 in rugged areas at middle altitudes with a southern aspect and slopes of 31°–60°. 486 Tahr, however, can be found in more rugged and very steep areas at low altitudes 487 and livestock in rolling, flat areas at high altitudes. Livestock is regularly observed 488 feeding in areas more than 150 m from escape terrain, whereas tahr is almost 489 consistently seen only in areas less than 100 m from escape terrain. A similar 490 overlap in habitat is recorded for ibex and livestock in other trans-Himalayan 491 protected areas, with ibex changing its grazing habitats during the season, which 492 reduces competition. 493

5.5.3 Food of Himalayan Tahr and Livestock 494

In terms of species composition, the diets of livestock and tahr are similar, but not 495 in the relative proportions of individual species. The proportion of woody plants 496 (Rhododendron and Cotoneaster) was higher, and that of grasses and sedges lower 497 in the diet of livestock than in that of tahr. This may be because livestock graze in 498 the vicinity of villages, where other species of plants are scarce due to harvesting or 499 overgrazing. 500 Parkes and Thompson (1995) found grass in the rumens of 48–65% of 253 tahr 501 shot in the Southern Alps, particularly snow tussocks; they claim that tahr more 502 often includes herbaceous than woody plants in its diet. In this study, the percentage 503 of woody plants, like Cotoneaster microphyllus and Rhododendron lepidotum, was 504 higher in the tahr’s diet, but nevertheless they still eat more soft and herbaceous 505 plants, compared to livestock, which often eat woody plants like C. microphyllus 506 and R. lepidotum. According to Forsyth and Tustin (2001) males of tahr prefer her- 507 baceous and woody plants to grasses and sedges in the period when the males and 508 females are segregated. This study was also done during this period (July-August) 509 and tahr grazed more on woody plants than is reported by Forsyth and Tustin (2001). 510 The percentage of woody plants may, however, be overestimated, as they are diffi- 511 cult to digestUncorrected and therefore more likely to be found in the faeces.Proof 512 In this study, the tahr’s diet consisted of 47% herbaceous plants and shrubs, 28% 513 grasses and 25% sedges, which is similar to the yearly averages reported by Green 514 (1979) for tahr in the Langtang valley (38% herbaceous plants and shrubs, 34% 515 grasses, 21% sedges, 4% ferns and 4% mosses). According to Green (1979), how- 516 ever, there are seasonal differences in these percentages, e.g., in winter tahr supple- 517 ments its diet with small amounts of mosses and ferns, presumably because other 518 food is less readily available. The previous results differ from those of Parkes and 519 Thompson (1995) who record that their diet is made up of 16.3% herbaceous plants, 520 55.7% grasses, 26.6% woody plants and 1.1% ferns. 521 B. Shrestha et al.

522 5.5.4 Palatability

523 Herbivore diet depends not only on the abundance of vegetation, but also on the 524 palatability of individual species. There are conflicting opinions about plant palat- 525 ability in the literature. Species like Rhododendron, Cotoneaster, Anaphalis con- 526 torta, Carex sp. and Bistorta affinis are often considered to be unpalatable (Bauer 527 1990; Koirala and Shrestha 1997; Buffa et al. 1998). However, Schaler (1973) 528 reports that tahr eats small quantities of Rhododendron, Sharma (2000) that blue 529 include Anaphalis contorta and Cotoneaster microphyllus in their diet, 530 Awasthi et al. (2003) that ungulates in the Himalayas include Carex sp. in their diet 531 and Wangchuk (1995) that both blue sheep and yak eat Carex sp., Bistorta sp., 532 Anaphalis sp. and Cotoneaster microphyllus.

533 5.5.5 Niche Breadth and Food Overlap

534 Body size is the most important factor determining the metabolic rate and food 535 requirements. Large-bodied mammals have higher food requirements, since they 536 have higher cost of maintenance and production compared to small species (Geist 537 1974). Thus small-bodied ungulates tend to be limited by forage quality and large- 538 bodied ungulates by forage quantity (Hanley 1982). Small animals tend to be more 539 selective not only in terms of the number of plant species they eat, but also in terms 540 of their diversity and have a smaller niche breadth. 541 In this study, livestock, which is larger than tahr, grazed more plant species than 542 tahr and have a larger niche breadth. Thus, tahr is more selective feeder than live- 543 stock. There is also a large overlap in the diets of tahr and livestock expressed by the

544 Morisita index (CH = 0.83). However, the fieldwork was carried out during the 545 ­monsoon season, when net primary productivity is high and forage quality very 546 good. During winter, the yak herders interviewed reported that yak eats any plant 547 matter, including shrubs and tree bark. The same might hold for tahr. Consequently, 548 dietary overlap might be 100% during winter. 549 Shrestha (2006) and Buffa et al. (1998) suggest that the spatial overlap in the 550 habitats of tahr and livestock can lead to competition, which is not always the case 551 (Squires 1982) as is well illustrated by the interactions between ungulates in the 552 SerengetiUncorrected National Park in East Africa (Krebs 1994). The present Proof study also ­indicates 553 there is a partial, but not a complete spatial overlap between tahr and livestock. 554 However, there is a good availability of forage on both rangelands. In addition, tahr, 555 which is more agile than livestock, is able to reach the vegetation growing in steep 556 and rocky areas and therefore spatially separated from livestock. Therefore, strong 557 competition for food is unlikely. 558 There were more individuals of tahr at Phortse than at Mongla, where forage 559 diversity and availability are lower. Tahr is often observed grazing together with 560 livestock (Gurung 1995) and sometimes even at low altitudes. This might be how it 561 avoids predators like snow leopard. A similar phenomenon is reported by Basnet 5 Interactions Between the Himalayan Tahr, Livestock and Snow Leopards...

(2002) for blue sheep, which graze together with livestock that protect the blue 562 sheep from predators or attackers by chasing them away. 563 The worrying fact is, however, that the signs of overgrazing, such as bare and 564 eroded pastures, are becoming increasingly noticeable in the SNP. This is a warning 565 for the future, because if this trend continues, tahr will become an endangered spe- 566 cies in this area. 567

5.5.6 Comparison of the Difficulties Associated with Direct 568 Observation and Micro Histological Techniques 569

Direct observation is a simple, cheap and easy way to determine the food habits of 570 both livestock and tahr, as neither is shy. Livestock is clearly used to the presence of 571 man; males of tahr escape very quickly when encountered, while females with juve- 572 niles do not, as they seem to be used to the presence of man. Thus tahr was often 573 observed from a distance of about 50 m. The problem with direct observation is in 574 defining and quantifying the species of plant being grazed, and how much of a plant 575 is consumed. It is difficult to differentiate between freshly and earlier eaten plants 576 and even those that have only been trampled (Holechek et al. 1982). 577 Histological analysis is the most commonly used method for evaluating herbi- 578 vore food habits (Holechek et al. 1982). The problem with faecal analyses is that the 579 material has to be examined under a microscope in order to identify the plant frag- 580 ments (Fitzgerald and Waddington 1979). The grass and sedge species are usually 581 overestimated and herbaceous plants underestimated (Vavra and Holechek 1980; 582 Gyawali 1986). In the present study, most of the fragments of grasses were very 583 characteristic: Avena sp. has a very distinct trichome and inter-stomatal cell ­structure, 584 which is clearly seen in faecal samples. The microstructure of other species, like 585 Carex sp., Cyperaceae sp., Trisetum spicatum is, however, very similar and difficult 586 to distinguish. Ephedra, Saxifraga brachypoda and Potentilla sp. have distinct char- 587 acters and are easy to recognize. 588

5.5.7 FloristicUncorrected Composition Proof 589

Species diversity was higher at Phortse than at Mongla, which may be explained by 590 their different altitudes. The range of most high altitude plants in the Northwest 591 Himalayas is 3,600–5,500 m, sometimes only 3,900–4,200 m (Mani 1978). The 592 rangeland at Mongla is at 3,400–3,800 m and at Phortse 3,600–4,200 m. Therefore, 593 that at Phortse is in the transition zone between shrubland and grassland, in which 594 vegetation diversity tends to be high. It is reported that the peak number of species 595 occurs at 4,250 m (Gurung 1995). The floristic composition however, is also affected 596 by slope and aspect. 597 B. Shrestha et al.

598 The dominant shrub species on both rangelands are Rhododendron lepidotum 599 and Cotoneaster microphyllus. Their dominance seems to be increasing (Bauer 600 1990) and they are rapidly colonizing new areas (Buffa et al. 1998). Bushes of 601 R. lepidotum and C. microphyllus sometimes form very dense growths, especially on 602 gentle slopes. Some grass and sedge species, like Avena sp. and Carex anomoea are 603 almost uniformly distributed on both rangelands. The species diversity was highest, 604 where the terrain is not rugged and the vegetation least disturbed on steep slopes.

605 5.5.8 Effect of Predation by Snow Leopard on Himalayan 606 Tahr and Livestock

607 The snow leopard has re-colonized the Everest region. An analysis of its scat indi- 608 cates that Himalayan tahr is its most important food, closely followed by livestock. 609 Our sample size is too small to reach a definitive conclusion, but certain indirect 610 factors indicate that snow leopards depend on tahr as food throughout the year. 611 These include the low kid-to-female ratio in tahr, for which snow leopard might be 612 responsible. 613 Ale (2006) reports that snow leopard started killing livestock in 2004. Our study 614 also indicates that because snow leopard is quite rare, it does not kill many live- 615 stock. In addition, most villagers have a spiritual belief that animals should be 616 respected. However, they may change their mind in the future, especially if the 617 numbers of snow leopard increase and those of tahr decrease, which will result in 618 snow leopard’s supplementing their diet with livestock, beyond the level tolerable to 619 local herders. This is typical of areas with depleted prey populations that are insuf- 620 ficient to sustain the local predators and can lead to a serious conflict interest.

621 5.6 Conservation Recommendations

622 Based on this study, it is recommended:

623 1. The tahr populations should be monitored regularly and any changes in popula- 624 tion structureUncorrected (mortality, fecundity), prevalence of diseases, Proof reproduction, gen- 625 eral health and other factors associated with the well being of populations 626 recorded. 627 2. Trends in the productivity and carrying capacity of the tahr’s habitat should be 628 closely monitored and action taken if the decline in productivity continues. 629 3. An in-depth predator-prey population study based on monitoring the abundance 630 of snow leopard, tahr, livestock and vegetation is needed. 631 4. As both snow leopard and tahr need protection and there is currently no serious 632 competition between livestock and tahr for food, the main threat now comes 633 from the decline in plant productivity in the region due to overgrazing. This trend 5 Interactions Between the Himalayan Tahr, Livestock and Snow Leopards...

could seriously change the situation, as tahr and livestock would then compete 634 for food. 635 5. The most effective way of reversing this trend is to introduce measures that limit 636 the amount of grass that is harvested for feeding livestock during winter, which 637 is rapidly increasing. 638 6. Delivery of hay from low altitude areas for feeding livestock during winter 639 should be subsidized, so that it becomes economically attractive for the farmers 640 not to use the overgrazed areas for hay production. 641

Acknowledgements The research was supported by the grant No. LC06073 of the MSMT, by the 642 Nepal Academy of Science and Technology (NAST), Nepal, and by the Department of National 643 Park and Wildlife Conservation (DNPWC) Nepal. We thank NAST and DNPWC for granting us 644 permission to work in the study area. We also thank Ranjit Pandey for help with fieldwork and 645 micro-histological lab analyses. 646

Annexes 647

Annex 5.1 Floristic composition of the vegetation at Mongla (see text for notation) t11.1 Plant species D f RF RD A RA IVI t11.2 Anaphalis contorta 4.6 52.4 4.5 3.5 8.8 2.1 10.1 t11.3 Anaphalis triplinervis 0.6 9.5 0.8 0.5 6.5 1.5 2.8 t11.4 Androsace sarmentosa 5.4 52.4 4.5 4.2 10.4 2.5 11.1 t11.5 Avena sp. 19.4 90.5 7.7 14.9 21.4 5.1 27.7 t11.6 Bistorta affinis 2.1 19.1 1.6 1.6 10.7 2.5 5.7 t11.7 Briza media 1.0 4.8 0.4 0.7 20.0 4.7 5.9 t11.8 Buplerium sp. 0.6 4.8 0.4 0.5 13.0 3.1 4.0 t11.9 Carex anomoea 11.5 66.7 5.7 8.8 17.3 4.1 18.6 t11.10 Cheilanthus sp. 0.3 4.8 0.4 0.3 7.0 1.7 2.3 t11.11 Compositeae 0.1 4.8 0.4 0.0 1.0 0.2 0.7 t11.12 Cotoneaster microphyllus 6.2 81.0 6.9 4.8 7.7 1.8 13.5 t11.13 Cyperaceae sp. I 0.5 4.8 0.4 0.4 10.0 2.4 3.1 t11.14 Cyperaceae sp. II 1.2 19.1 1.6 0.9 6.2 1.5 4.0 t11.15 Cyananthus hookeri 8.4 38.1 3.3 6.4 22.0 5.2 14.9 t11.16 Cyananthus microphyllus 9.2 57.1 4.9 7.1 16.2 3.8 15.8 t11.17 Cypripedium himalaicum 0.8 14.3 1.2 0.6 5.7 1.3 3.2 t11.18 Drosera peltataUncorrected3.1 47.6 4.1 2.4 Proof6.6 1.6 8.1 t11.19 Festuca sp. 0.3 9.5 0.8 0.2 3.0 0.7 1.7 t11.20 Fragaria daltoniana 0.3 14.3 1.2 0.3 2.3 0.6 2.0 t11.21 Gerbera gossypina 6.4 47.6 4.1 4.9 13.4 3.2 12.1 t11.22 Gueldenstaedtia himalaica 4.1 33.3 2.9 3.1 12.3 2.9 8.9 t11.23 Habenaria aitchisonii 0.1 9.5 0.8 0.1 1.5 0.4 1.3 t11.24 Herminium josephii 3.6 38.1 3.3 2.7 9.4 2.2 8.2 t11.25 Hieracium sp. 0.1 4.8 0.4 0.1 3.0 0.7 1.2 t11.26 Iris sp. 0.3 4.8 0.4 0.3 7.0 1.6 2.3 t11.27 Juniperus sp. 0.0 4.8 0.4 0.0 1.0 0.2 0.7 t11.28 (continued) B. Shrestha et al.

Annex 5.1 (continued) Plant species D f RF RD A RA IVI Leontopodium stracheyi 0.1 4.8 0.4 0.1 2.0 0.1 1.0 Microula pustulosa 0.4 4.8 0.4 0.3 8.0 1.9 2.6 Notholirion macrophyllum 1.6 19.0 1.6 1.2 8.5 2.0 4.9 Parnassia nubicola 0.1 4.8 0.4 0.1 2.0 0.5 1.0 Pedicularis siphonantha 0.0 4.8 0.4 0.0 1.0 0.2 0.7 Persicaria capitata 1.8 14.3 1.2 1.4 12.7 3.0 5.6 Poa sp. 1.1 23.8 2.0 0.9 4.8 1.1 4.1 Polygonatum hookeri 8.3 42.9 3.7 6.4 19.33 4.6 14.6 Potentilla sp. 7.0 19.0 1.6 5.3 36.5 8.7 15.6 Rhododendron lepidotum 5.4 71.4 6.1 4.1 7.5 1.8 12.0 Salvia hians 1.0 19.0 1.6 0.8 5.5 1.3 3.7 Satyrium nepalense 4.7 57.1 4.9 3.6 8.2 1.9 10.4 Saxifraga brachypoda 3.1 28.6 2.4 2.4 11.0 2.6 7.5 Saxifraga parnassifolia 2.7 57.1 4.9 2.1 4.7 1.1 8.1 Sedum sp. 1.9 9.6 0.8 1.5 20.0 4.7 7.0 Sedum sp. 0.0 4.8 0.4 0.0 1.0 0.2 0.7 Silene sp. 0.1 9.6 0.8 0.1 1.5 0.3 1.3 Trisetum spicatum 0.3 4.8 0.4 0.3 7.0 1.6 2.3 Unidentified graminae (15) 0.2 4.8 0.4 0.2 5.0 1.2 1.8

t12.1 Annex 5.2 Floristic composition of the vegetation at Phortse (see text for notation) t12.2 Plant species D f RF RD A RA IVI t12.3 Allium wallichii 0.0 3.3 0.2 0.0 1.0 0.2 0.4 t12.4 Anaphalis contorta 0.2 6.7 0.4 0.1 2.5 0.5 1.04 t12.5 Avena sp. 9.0 96.7 6.4 5.8 9.3 1.8 14.1 t12.6 Bistorta affinis 2.8 46.7 3.1 1.8 6.1 1.2 6.1 t12.7 Campanula pallida 2.0 10.0 0.7 1.3 20.0 3.9 5.9 t12.8 Carex anomoea 12.2 100.0 6.7 7.8 12.2 2.4 16.9 t12.9 Cassiope fastigiata 1.7 10.0 0.7 1.1 17.3 3.4 5.2 t12.10 Cheilanthus spp. 0.2 10.0 0.7 0.1 2.3 0.5 1.3 t12.11 Compositeae 0.0 3.3 0.2 0.0 1.0 0.2 0.4 t12.12 Cotoneaster microphyllus 1.9 43.3 2.9 1.2 4.5 0.9 5.0 t12.13 Cyperaceae sp. 0.1 3.3 0.2 0.1 5.0 1.0 1.3 t12.14 Cyananthus hookeri 8.4 63.3 4.2 5.4 13.3 2.6 12.2 t12.15 CyananthusUncorrected microphyllus 5.2 50.0 3.3 3.3 Proof10.3 2.0 8.7 t12.16 Cyperus sp. 4.0 43.3 2.9 2.6 9.2 1.8 7.3 t12.17 Cypripedium himalaicum 0.6 6.7 0.4 0.4 9 1.8 2.6 t12.18 Dactylorhiza hatagirea 0.1 3.3 0.2 0.1 3.0 0.6 0.9 t12.19 Drosera peltata 5.4 46.7 3.1 3.5 11.6 2.3 8.9 t12.20 Dubyaea hispida 1.2 6.7 0.4 0.8 18.5 3.6 4.9 t12.21 Ephedra gerardiana 0.2 6.7 0.4 0.1 2.5 0.5 1.0 t12.22 Euphrasia himalayica 2.3 13.3 0.9 1.5 17.5 3.4 5.8 t12.23 Fragaria daltoniana 2.9 56.7 3.8 1.8 5.1 1.0 6.6 t12.24 Gentiana depressa 6.6 23.3 1.6 4.2 28.1 5.5 11.3 (continued) 5 Interactions Between the Himalayan Tahr, Livestock and Snow Leopards...

Annex 5.1 (continued) Plant species D f RF RD A RA IVI Gentiana sp. 1.0 23.3 1.6 0.7 4.4 0.9 3.1 Gerbera gossypina 7.2 73.3 4.9 4.7 9.9 1.9 11.5 Gueldenstaedtia himalaica 2.7 46.7 3.1 1.8 5.9 1.2 6.0 Halenia elliptica 1.5 20.0 1.3 1.0 7.5 1.5 3.8 Herminium josephii 0.3 6.7 0.4 0.2 5.0 1.0 1.6 Hieracium sp. 3.3 43.3 2.9 2.1 7.6 1.5 6.5 Leontopodium jacotianum 3.0 20.0 1.3 1.9 14.8 2.9 6.2 Morina nepalensis 3.5 36.7 2.4 2.3 9.6 1.9 6.6 Neottianthe calcicola 1.6 23.3 1.6 1.0 6.9 1.3 3.9 Notholirion macrophyllum 1.7 20.0 1.3 1.1 8.5 1.7 4.1 Parnassia nubicola 1.9 33.3 2.2 1.3 5.9 1.2 4.6 Pedicularis siphonantha 3.4 66.7 4.4 2.2 5.0 1.0 7.6 Persicaria capitata 5.6 36.7 2.4 3.6 15.2 3.0 9.0 Poa sp. 0.6 20.0 1.3 0.4 2.8 0.6 2.3 Polygonatum cirrhifolium 0.2 3.3 0.2 0.1 6.0 1.2 1.5 Polygonatum hookeri 29.7 56.7 3.8 19. 52.4 10.3 33.2 Polygonum sp. 1.0 16.7 1.1 0.6 5.8 1.1 2.9 Polygonum sp. 0.4 3.3 0.2 0.3 12.0 2.4 2.8 Potentilla sp. 105.0 26.7 1.8 1.0 5.6 1.1 4.5 Primula sp. 2.2 6.7 0.4 1.4 33.5 6.6 8.5 Rhododendron lepidotum 3.1 60.0 4.0 2.0 5.2 1.0 7.0 Salvia hians 0.7 20.0 1.3 0.5 3.5 0.7 2.5 Satyrium nepalense 0.4 10.0 0.7 0.3 4.0 0.8 1.7 Saxifraga brachypoda 2.4 40.0 2.7 1.6 6.1 1.2 5.4 Saxifraga parnassifolia 1.4 36.7 2.4 0.9 3.7 0.7 4.1 Sedum sp. 0.2 13.3 0.9 0.1 1.7 0.3 1.4 Sedum sp. 0.6 6.7 0.4 0.4 9.0 1.8 2.6 Silene sp. 0.1 3.3 0.2 0.0 2.0 0.4 0.7 Thermopsis barbata 4.9 26.7 1.8 3.2 18.5 3.6 8.6 Thesium emodi 0.4 6.7 0.4 0.2 5.5 1.1 1.8 Trachyspermum ammi 0.7 13.3 0.9 0.5 5.2 1.0 2.4 Woodfordia sp. 0.1 6.7 0.4 0.1 1.5 0.3 0.8

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