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Habitat and feeding ecology of alpine (Moschus chrysogaster) in Kedarnath Wildlife Sanctuary, ,

Article in Production Science · January 2015 DOI: 10.1071/AN141028

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Zarreen Syed Orus Ilyas Wildlife Institute of India Aligarh Muslim University

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Habitat preference and feeding ecology of alpine (Moschus chrysogaster) in Kedarnath Wildlife Sanctuary, Uttarakhand, India

Zarreen Syed A and Orus Ilyas B,C

AWildlife Institute of India, Chandrabani, Dehradun, Uttarakhand, 248001, India. BDepartment of Wildlife Sciences, Aligarh Muslim University, Aligarh, Uttar Pradesh, 202002, India. CCorresponding author. Email: [email protected]

Abstract. The , Moschus chrysogaster, a small member of family , is a primitive deer threatened due to and habitat loss, and therefore classified as Endangered by IUCN and also listed in Appendix I of CITES. Although the is legally protected in India under Wildlife Protection Act 1972, conservation of the species requires better understanding of its distribution and resource-use pattern; therefore, a study on its feeding and habitat ecology was conducted from February 2011 to February 2014, at Kedarnath Wildlife Sanctuary. The Sanctuary is one of the known remaining habitats for musk deer in India. Four locations, namely, Shokhark I, Shokhark II, Tungnath and Chandrashila, were intensively monitored and further categorised on the basis of habitats and altitudinal gradient. The habitat overlap between musk deer and all other species of the area was calculated through Pianka’s overlap index. The results indicated a large overlap between musk deer and Himalayan . Species dietary spectra were studied using microhistological techniques for faecal pellet-group analysis coupled with Bonferroni approach. The dicotyledon to monocotyledon ratios were found to be 73.62–26.38% and 52.016– 47.984% in the pre- and post-monsoon seasons, respectively. The study showed that the musk deer is predominantly a browser. The most preferred species of the were found to be Gaultheria trichophylla, Ophiopogon intermedius., Cyperus sp. and Sibbaldia cuneata. During the field survey, opportunistic sightings of the species were also recorded. The species was found to be restricted to areas where the density of preferred vegetation was high. Therefore, it is recommended to provide strict protection to the areas such as Shokhark.

Additional keywords: feeding habit, habitat overlap, microhistology.

Received 26 December 2014, accepted 30 April 2015, published online 22 June 2015

Introduction unique niche, thus allowing their coexistence (Whitfield 2002; The alpine musk deer, Moschus chrysogaster, is a small member Bagchi et al. 2003a, 2003b). Ecological isolation in temporal of a primitive deer family named Moschidae. Being under high and spatial scales governs the co-existence of species and the pressure from centuries of poaching for musk pods and extensive sympatric species discriminate themselves at being in one place, habitat loss, this elegant species is rapidly disappearing from by showing isolation in terms of the use of various resources, its previous range of distribution (Green 1985). The species is which governs their habitat-use pattern (Dar et al. 2012). Several classified as Endangered in IUCN Red List Data (IUCN 2014) resource axes exist that could be partitioned by the species and and is also listed in Appendix I of CITES (2003). Although the if there is complete overlap between two species at one niche species is legally protected in India under Wildlife Protection Act axis, there must be partitioning along another unmeasured 1972, illegal poaching is still going on, more particularly in the axis (Schoener 1974; Bagchi et al. 2003a, 2003b). Hence, a border areas of the country. In the present scenario, the continuum habitat-overlap study is important in understanding community of its distribution is broken into small fragments that have organisation and depends on the measurement of how organisms subpopulations in isolated pockets of the (Green utilise and share their environment, and also plays an important 1986a; Sathyakumar 1991, 1992; Sathyakumar et al. 1993a). role in the efforts to generate sufficient knowledge for effective This situation further deepens the risk to its survival viability as wildlife management (Otis 1997; Dar et al. 2012). it increases the danger of genetic drift. It was realised that understanding the musk deer food In the present study, one of the objectives was to examine preferences and changes resulting from biotic influences are habitat overlap between musk deer and other ungulate species, also important in interpreting relationships between environment e.g. (Hemitragus jemlahicus), sambar ( and species (Leopold and Krausman 1987). In the case of musk unicolor), (Naemorhedus goral) and the livestock of the deer, observing while feeding was not possible in natural area. It was hypothesised that each species is adapted to utilise a conditions as the species is very shy. Therefore, pellet groups

Journal compilation CSIRO 2015 www.publish.csiro.au/journals/an B Animal Production Science Z. Syed and O. Ilyas were collected and analysed to assess the feeding habits of this receives 3000 mm of annual precipitation, of which ~60% falls species. Studies on feeding habits, using the microhistological during the monsoon season (June–August). The relative humidity technique for the identification of diet constituents, have appeared varies from 35% to 85% annually. There is moderate to heavy in the literature after Baumgartner and Martin (1939). Denham snowfall during December–February, even in low-altitude areas (1965) and Sparks and Malechek (1968)verified the technique by (Bhat et al. 2013). Temperatures are highest in May–June before hand-compounding mixtures of grasses and forbs. the arrival of monsoon, and are lowest in the first week of The work on conservation and management of wildlife is often January. The area is snow bound for 3 months in the year, hampered due to non-availability of data on species presence and following heavy snowfalls in December (Green 1986; Pande habitats occupied (Kushwaha and Roy 2002). Conservation of et al. 2001; Singh 2009). alpine musk deer requires a good understanding of its ecology, Owing to connection with timber line which support especially habitat and feeding ecology; therefore, a study on seasonal succession and distributional pattern, the alpine the habitat overlap and feeding ecology of the species was vegetation of this area has many characteristic features. Some conducted in the Kedarnath Wildlife Sanctuary, providing important that separate the alpine zone from timber line baseline information for managers to develop strategies to are Clematis montana, Berberis edgeworthiana, Hypericum conserve the species. hookerianum, campanulatum and R. arboreum in the south-facing slopes and Anemone spp., Skimmia spp., Materials and methods Geum spp. and Trillium govanianum in the north-west-facing Study site slopes. The plant species that appear soon after the snow melt are Primula denticulata, Oxygraphis polypetala and Gentiana spp., The Kedarnath Wildlife Sanctuary (KWLS) spreads over an area 2 immediately followed by Rannunculus spp., Anemone obtusiloba, of 975 km (Fig. 1) and is situated in the north-eastern part of Potentilla spp., Pedicularis spp., Senecio spp., Saussurea spp., 0– 0 0– 0 the Garhwal Himalayas, 30 25 30 41 N, 78 55 79 22 E. It is Polygonum spp. and others during July–August. In the alpine bordered by high mountain peaks Kedarnath (6940 m), zone, the plants observed in the late-flowering season are Mandani (6193 m) and Chaukhamba (7068 m) in the north, Delphinium spp., Selinum vaginatum, Tanacetum longifolium – – Gopeshwar Chopta Ukhimath motor road in the south, and Taraxacum officinale. Some plants such as Oxygraphis Urgam Reserve (RF) in the east and by the river polypetala, Rannunculus spp. and Gentiana spp. reflower at the – Mandakini and Chamoli Uttar Kashi district boundaries in the end of the season, during October–November (Semwal and west. The altitude varies from 1200 m (near Phata) to 7068 m 1981). (Chaukhumba peak) (Sathyakumar et al. 1994). This protected area was named after the famous 8th century Hindu Shrine at Data collection Kedarnath (3562 m) situated close to its northern boundary and which is one of the five ‘Kedars’. An intensive study area of Four sites, namely Shokhark I, Shokhark II, Tungnath and ~4 km2 was selected in the southern region of the sanctuary in Chandrashila, with known species presence were selected on and around the Tungnath area (30300N, 79150E). The study the basis of past studies (Green 1985; Sathyakumar 1994; Ilyas area included Shokhark, Tungnath and Chandrashila zones, 2007). These sites were located on different habitats and with an altitudinal range between 2800 and 4000 m. The area altitudes in the KWLS. These sites were further divided into

N Kedarnath Wildlife Sanctuary

Tungnath Chandrasila

To Okhimath Chopta Anusuya Devi Shokhark Bulkhana Ridge Motor road

Village/ Settlement Amrit Ganga Anusuya Ridge Anusuya Kanchula Kharakh Temple

River

Path Points

To Gopeshwar Path

Muskdeer Breeding Center

Fig. 1. Map of Kedarnath Wildlife Sanctuary, India. Habitat and feeding ecology of alpine musk deer Animal Production Science C

smaller blocks by using the toposheets of Survey of India, and in for all species (Pio = n/SN). Utilisation was significantly greater or each block, existing trails and straight transects used by humans smaller than expected when Pio lay outside the 95% confidence were monitored on a daily basis. The 10-m-radius circular plots limits constructed for proportional utilisation (Pie), which was were also laid at intervals of 100 m on each trail and transect for calculated by dividing number of plant fragments identified for sampling the vegetation and indirect evidence (pellets group) of each species (b) by the total number of identified browse fragment the animals. Pellet groups were identified on the basis of their size (B) of all species (Pie = b/SB). The statistical tests were conducted and shape. Musk deer pellet groups were collected during the pre- in Microsoft Excel following Zar (1999) and Fowler and Cohen monsoon as well as post-monsoon seasons from different (1986). surveyed sites to analyse the feeding habits of musk deer. Habitat overlap of musk deer with other ungulate species in the Results study area was analysed on the basis of the distribution of their On the basis of the pellet survey and Pianka’s habitat-overlap faecal pellets at the study site. Stratified random sampling was results, a large overlap of 0.76 (pre-monsoon) and 0.90 (post- conducted and pellets or dung of all were searched monsoon) between musk deer and Himalayan tahr was observed inside 10-m radius of every plot. To assess the resource (Table 1). The overlap between musk deer and sambar was partitioning in term of habitat overlap, between the musk deer estimated to be 0.57 during the pre-monsoon and 0.63 during and other ungulate species of the area, the Pianka’s niche overlap the post-monsoon season. Medium to higher overlap, i.e. 0.49 and index was used (Pianka 1973), as follows: 0.74, was found between the musk deer and the goral during pre- P PijPik and post-monsoon seasons, respectively, in comparison with Pianka index O ¼ pPffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiP ; jk 2 2 the overlap between musk deer and Himalayan tahr. In case of P ij P ik the overlap between musk deer and livestock (sheep and ), the index value showed decremental overlap from pre-monsoon where Pij is percentage of Utilised habitat i (through pellet-group (0.79) to post-monsoon (0.53) season. Overall, it was found density) by Species j and Pik is the percentage of Utilised habitat i (through pellet group density) by Species k. The index ranges that the musk deer has a large habitat overlap with Himalayan between values of 0 (no resource used in common between two tahr and livestock. The habitat-overlap value of musk deer with species) and 1 (complete overlap in resource use). livestock was highest, i.e. 0.7911, which is higher than the The collected pellet samples of musk deer were then preserved overlap among the wild species. under dry conditions and pellet-group analysis for feeding From the total of 164 plots, only 17 (from 13 plots) and four ecology was performed by using microhistology technique pellet groups of musk deer were found at Shokhark sites during following Vavra and Holechek (1980), Holechek et al.(1982), pre-monsoon 2011 and post-monsoon 2012, respectively, while Alipayo et al.(1992) and Ilyas et al.(2003). Plant species were from Tungnath and Chandrashila, no pellet group was found at collected from all sites for reference-slide preparation for all the any plot. The percentage count of the presence of pellet groups available plant species. Dicot plant fragments were identified to was calculated as 10.36% (pre-monsoon) and 2.43% (post- on the basis of micro-anatomical characters such as the size monsoon) from the total 164 plots. and shape of the cells, the pattern of the cell wall, and the In total, 5295 (pre-monsoon) and 1786 (post-monsoon) plant arrangement and size of stomata and trichomes, silica cells, fragments were evaluated to determine the diet of musk deer. In companion cells and crystals (Eames and MacDaniels 1947; total, two shrubs, 29 herbs and three grasses in the pre-monsoon Esau 1960; Green 1985). season and one tree, two shrubs, 17 herbs and four grass species For the pre-monsoon and post-monsoon seasons, the overall were identified during the post-monsoon season from the diet of browse (dicot) to grass (monocot) ratios for the musk deer were musk deer (Fig. 2). computed. The browse particles were analysed and identified to The analysis of pre-monsoon pellet groups showed the calculate the proportion of different trees, shrubs and herbs at the presence of a total 5295 fragments (3898 dicots and 1397 species level in musk deer diets, although grass particles could monocots), of which, 2084 plant fragments were identified to generally not be identified to species level. The proportion of species level, and included 1939 dicots and 145 monocots. The each plant species in the diet of musk deer was compared with total composition of vegetation in the analysed faecal pellets was the availability of food-plant species in the KWLS during pre- calculated as percentages and included 73.62% dicots and and post-monsoon seasons, by calculating 95% Bonferroni confidence intervals following Neu et al.(1974). The formula Table 1. Habitat overlap between the Himalayan musk deer and other for calculating 95% Bonferroni confidence interval is as ungulate species of Kedarnath Wildlife Sanctuary during pre-monsoon follows: and post-monsoon seasons pffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi Ojk, Pianka’s mesure of niche overlap P Z/= P ðÞ1 P =n P P ie 2k pieffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiie ie ie þ ðÞ = ; Z/=2k Pie 1 Pie n Overlapping species Ojk Pre-monsoon Post-monsoon where Pie is the observed proportional utilisation of each species season season and Z/=2k is the upper standard normal table value corresponding to a probability tail area of /=2k and n is the total number species Musk deer and Himalayan tahr 0.756636472 0.898688472 recorded in the sampling. For each species, its proportional Musk deer and sambar 0.573753356 0.62872558 availability (P ) in the sample was calculated by dividing its Musk deer and goral 0.494453434 0.735289237 io Musk deer and livestock 0.791106006 0.531560918 total number (n) by the total number of all individuals (N) sampled D Animal Production Science Z. Syed and O. Ilyas

35 during pre-monsoon were Dryopteris spp., Epilobium spp., Galium spp., Geum elatum, Hemiphragma spp., Ophiopogon 30 29 intermedius, Polygonum amplexicaula, Polystichum spp., Senecio spp., Sibbaldia cuneata, Stachys spp., Viola spp. and 25 Cyperus spp. The species that were utilised less than could be Pre-monsoon expected from their availability were Cotonaester acuminatus, Rubus spp., Anemone spp., Astragalus spp., Fragaria spp., 20 Post-monsoon 17 Gentiana argentina, Geranium spp., Nepeta spp., Oxygraphis spp., Plantago ovata, Potentilla spp., Primula denticulata, 15 Tanacetum longifolium, Chrysopogon spp. and Thamnocalamus spathiflora. Trachydium roylei was utilised in proportion to its 10 availability (Table 3). During the post-monsoon season, 10 plant species were

Number of plant species identified 4 5 3 preferred for food, including two shrubs, five herbs and three 2 2 1 0 grasses. While 12 species (1 tree, 10 herbs and 1 grass) were 0 avoided during the post-monsoon season; however, it is Tree Shrub Herb Grass possible that these plant species were consumed by musk deer but their fragments were not identified. The post-monsoon fi Fig. 2. Composition of identi ed plants in the diet of musk deer, through results showed differences from the pre-monsoon preferred microhistology during pre-monsoon and post-monsoon seasons. food plants as the preferred plant species in this season were Cotonaester acuminatus, Rubus spp., Gaultheria spp., 26.38% monocots (with a dicot : monocot ratio of 2.8 : 1), which Ophiopogon intermedius, Primula denticulata, Saussurea sp., were further analysed to species level; 49.74% (1939 · 100/3898) Selinum vaginatum, Cyperus sp., Danthonia cachemeriana of dicot plant species were identified among the total dicot and Thamnocalamus spathiflora. The plant species that were fragments analysed and 10.38% monocot plant species used less than could be expected from their availability were were identified among the total analysed monocot fragments. , Anemone obtusiloba, Corydalis Moreover, the total composition was 36.62% herbs, 2.74% govaniana, Fragaria spp., Geranium spp., Geum elatum, Geum grasses, 36.99% unidentified herbs and 23.64% unidentified spp., Gnaphalium spp., Polygonum amplexicaula, Potentilla spp., grass species. The proportion of Anemone spp. in the diet of Saxifraga spp. and Chrysopogon spp. Gentiana argentina and musk deer during the pre-monsoon season was found to be the Sibbaldia cuneata were utilised in proportion to their availability. greatest (19.90%), followed by Polygonum amplexicaula and The forb species diversity was found to be higher; however, Potentilla spp. (15.83% and 13.48%, respectively; Table 2). grasses were the major diet ingredient during the post-monsoon The post-monsoon data collection showed the presence of a season (Table 3). total of 1786 fragments of vegetation in the pellet-group samples, Some plant species were found to be preferred in both of which, 755 vegetation fragments were identified to species seasons, e.g. Gaultheria trichophylla, Ophiopogon intermedius, level and included 401 dicot and 354 monocot species fragments. Cyperus spp. and Sibbaldia cuneata. Among these, Gaultheria The total composition of vegetation of the analysed faecal trichophylla was found to be the most preferred plant species, but pellets was calculated in percentages and included of 52.02% was not even recorded in sampling during pre-monsoon season of dicots and 47.98% of monocots (dicot : monocot ratio of although fragments of the species were recorded in the musk deer 1.08 : 1), which were further analysed to species level and diet in both the season. included 43.16% of dicot plant species and 41.31% of monocot plant species. The total composition was 22.45% herbs, 19.82% grass, 29.56% unidentified herbs and 28.16% Discussion unidentified grass species. During the post-monsoon season, the The coexisting animal species segregate primarily by their highest proportion of consumed plant fragments was recorded habitats and, subsequently, by the dietary and temporal for Danthonia cachemeriana (21.32%) followed by Cyperus specialisations (Schoener 1974, 1983; Toft 1985). A variety of spp. and Thamnocalamus spathiflora (12.98% and 11.92%, indices can be used to measure the ecological niche, such as respectively; Table 2). dietary components, microhabitat, or temporal or spatial activity The feeding analysis showed that during pre-monsoon season (Dar et al. 2012). Pianka’s overlap index is one of such the musk deer prefer browse items to grass, because of the 13 measures that is used to estimate the extent of the overlap, on consumed species, 12 species were herbs and only one species the basis of the usage of a particular resource. was grass. Fifteen species (2 shrubs, 11 herbs and 2 grasses) were The present study showed a high habitat overlap between avoided according to the plant fragments identified; however, it is musk deer and Himalayan tahr, with respect to their habitat possible that some of these species were included in the diet of utilisation. This may be because of the fact that the Himalayan musk deer but their fragments were among the unidentified plant tahr is utilising all habitats in the study area and is found to prefer fragments. Preference and avoidance of food-plant species by the habitats such as alpine meadows and alpine scrub, which are the musk deer with respect to the available composition of vegetation preferred habitat for musk deer as well. Also, the value of the was calculated using the Bonferroni approach. The Bonferroni overlap index showed that the overlap between these two species results showed that the preferred plant species of the musk deer was highest during the post-monsoon season. Habitat and feeding ecology of alpine musk deer Animal Production Science E

Table 2. Percentage occurrence of fragments of tree, shrub and herb species within the identified browse fragments in faecal pellets of musk deer during pre-monsoon season in 2011 and post-monsoon season in 2012 in Kedarnath Wildlife Sanctuary N, number of individuals of a species sampled; n, number of plant fragment recorded in diet composition; %P, percentage occurrence

Species Family Pre-monsoon Post-monsoon Nn%P Nn%P Trees Acer sp. Aceraceae 1.000 0.000 0.000 1.000 0.000 0.000 Abies sp. Pinaceae 0.000 0.000 0.000 6.000 0.000 0.000 Quercus semecarpifolia Fagaceae 344.000 0.000 0.000 306.000 0.000 0.000 Rhododendron arboreum 184.000 0.000 0.000 218.000 9.000 1.192 Rhododendron barbatum Ericaceae 115.000 0.000 0.000 80.000 0.000 0.000 Taxus baccata Taxaceaae 19.000 0.000 0.000 14.000 0.000 0.000 Shrubs Berberis jaeschkeana Berberidaceae 4.000 0.000 0.000 1.000 0.000 0.000 Cotonaester acuminata Rosaceae 21.000 4.000 0.192 41.000 15.000 1.987 Indigofera sp. Leguminosae 10.000 0.000 0.000 10.000 0.000 0.000 Lonicera myrtillus Caprifoliaceae 83.000 0.000 0.000 17.000 0.000 0.000 Myrsine sp. Myrsinaceae 1.000 0.000 0.000 0.000 0.000 0.000 Piptenthus nepalensis Leguminosae 4.000 0.000 0.000 4.000 0.000 0.000 Pyracantha sp. (Kilmoda) Rosaceae 21.000 0.000 0.000 39.000 0.000 0.000 Rhododendron anthopogon Ericaceae 1202.000 0.000 0.000 1373.000 0.000 0.000 Rhododendron barbatum Ericaceae 42.000 0.000 0.000 2.000 0.000 0.000 Rhododendron campanulatum Ericaceae 64.000 0.000 0.000 86.000 0.000 0.000 Rosa sericea Rosaceae 25.000 0.000 0.000 50.000 0.000 0.000 Rubus sp. Rosaceae 62.000 5.000 0.240 10.000 18.000 2.384 Salix denticulata Salicaceae 1.000 0.000 0.000 0.000 0.000 0.000 Vibernum grandiflorum Sambucaceae 12.000 0.000 0.000 50.000 0.000 0.000 Herbs Anaphalis roylei Compositae 78.000 0.000 0.000 9.000 0.000 0.000 Anemone sp. Ranunculaceae 1484.000 415.000 19.914 381.000 26.000 3.444 Astragalus sp. Leguminosae 13.000 1.000 0.048 0.000 0.000 0.000 Boeninghausenia albiflora Rutaceae 5.000 0.000 0.000 0.000 0.000 0.000 Campanula latifolia Leguminosae 0.000 0.000 0.000 2.000 0.000 0.000 Cirsium falconeri Compositae 0.000 0.000 0.000 8.000 0.000 0.000 Clematis sp. Ranunculaceae 0.000 1.000 0.048 0.000 0.000 0.000 Corydalis govaniana Papaveraceae 259.000 0.000 0.000 1437.000 9.000 1.192 Delphinium sp. Ranunculaceae 0.000 0.000 0.000 6.000 0.000 0.000 Dryopteris sp. – 4.000 5.000 0.240 0.000 0.000 0.000 Epilobium sp. Onagraceae 109.000 118.000 5.662 0.000 0.000 0.000 Fragaria sp. Rosaceae 1253.000 98.000 4.702 669.000 29.000 3.841 Galium sp. Rubiaceae 8.000 3.000 0.144 11.000 0.000 0.000 Gaultheria trichophylla Ericaceae 0.000 23.000 1.104 27.000 23.000 3.046 Gentiana argentina Gentianaceae 928.000 109.000 5.230 161.000 23.000 3.046 Geranium sp. Geraniaceae 647.000 50.000 2.399 418.000 52.000 6.887 Geranium sp. 2 Geraniaceae 24.000 0.000 0.000 0.000 0.000 0.000 Geum elatum Rosaceae 138.000 100.000 4.798 318.000 24.000 3.179 Geum sp. Rosaceae 0.000 0.000 0.000 251.000 16.000 2.119 Gnaphalium sp. Compositae 92.000 0.000 0.000 604.000 15.000 1.987 Hemiphragma sp. Scrophulariaceae 7.000 20.000 0.960 0.000 0.000 0.000 Lysimachya prolifera – 37.000 0.000 0.000 0.000 0.000 0.000 Micromeria sp. Labiateae 89.000 0.000 0.000 0.000 0.000 0.000 Morina sp. Dipsacaceae 11.000 0.000 0.000 158.000 0.000 0.000 Nepeta sp. Labiateae 144.000 3.000 0.144 0.000 0.000 0.000 Ophiopogon intermedius Liliaceae 117.000 49.000 2.351 36.000 22.000 2.914 Oxalis sp. Oxalidaceae 1.000 0.000 0.000 0.000 0.000 0.000 Oxygraphis polypetala Ranunculaceae 36.000 5.000 0.240 8.000 0.000 0.000 Pedicularis sp. Scrophulariaceae 0.000 0.000 0.000 25.000 0.000 0.000 Pedicularis pectinata (Bhootkes) Scrophulariaceae 16.000 0.000 0.000 0.000 0.000 0.000 Plantago ovata Plantaginaceae 41.000 5.000 0.240 0.000 0.000 0.000 Polygonum amplexicaula Polygonaceae 587.000 330.000 15.835 641.000 7.000 0.927

(continued next page) F Animal Production Science Z. Syed and O. Ilyas

Table 2. (continued )

Species Family Pre-monsoon Post-monsoon Nn%P Nn%P Polystichum sp. – 27.000 34.000 1.631 0.000 0.000 0.000 Potentilla sp. Rosaceae 2134.000 281.000 13.484 631.000 39.000 5.165 Potentilla peduncularis Rosaceae 0.000 0.000 0.000 250.000 0.000 0.000 Primula denticulata Primulaceae 542.000 60.000 2.879 85.000 14.000 1.854 Rubus sp. Rosaceae 0.000 0.000 0.000 62.000 0.000 0.000 Rumex nepalensis Polygonaceae 108.000 0.000 0.000 0.000 0.000 0.000 Senecio sp. Compositae 185.000 128.000 6.142 0.000 0.000 0.000 Saussurea sp. Compositae 0.000 0.000 0.000 37.000 13.000 1.722 Saxifraga sp. Saxifragaceae 0.000 0.000 0.000 233.000 11.000 1.457 Selinum vaginatum Umbelliferae 0.000 0.000 0.000 55.000 14.000 1.854 Sibbaldia cuneata Rosaceae 65.000 37.000 1.775 155.000 22.000 2.914 Skimmia anquetilia Rutaceae 0.000 10.000 0.480 0.000 0.000 0.000 Smilax sp. Liliaceae 1.000 0.000 0.000 0.000 0.000 0.000 Spirae bella Rosaceae 1.000 0.000 0.000 0.000 0.000 0.000 Stachys sp. Labiateae 17.000 12.000 0.576 0.000 0.000 0.000 Tanacetum longifolium Asteraceae 18.000 3.000 0.144 0.000 0.000 0.000 Thalictrum sp. Ranunculaceae 0.000 2.000 0.096 0.000 0.000 0.000 Trachydium roylei Apiaceae 27.000 7.000 0.336 0.000 0.000 0.000 Vaccinium nummularia Ericaceae 0.000 0.000 0.000 8.000 0.000 0.000 Valeriana sp. Valerianaceae 0.000 2.000 0.096 0.000 0.000 0.000 Veronica cana Scrophulariaceae 3.000 0.000 0.000 0.000 0.000 0.000 Viola biflora Violaceae 32.000 19.000 0.912 15.000 0.000 0.000 Zanthoxylum nepalense Rutaceae 0.000 0.000 0.000 52.000 0.000 0.000 Grass Arundinaria sp. Poaceae 2.000 0.000 0.000 0.000 0.000 0.000 Chrysopogon sp. Gramineae 293.000 48.000 2.303 93.000 5.000 0.662 Cynodon sp. Gramineae 147.000 0.000 0.000 5.000 0.000 0.000 Cyperus sp. Gramineae 109.000 90.000 4.319 262.000 98.000 12.980 Danthonia cachemeriana Poaceae 881.000 0.000 0.000 406.000 161.000 21.325 Danthonia sp. Poaceae 736.000 0.000 0.000 45.000 0.000 0.000 Thamnocalamus spathiflora Poaceae 138.000 7.000 0.336 96.000 90.000 11.921 Subtotal 13 809 2084 39.358 9968 755 42.273 Browse unidentified 1065 1959 36.997 2303 528 29.563 Grass unidentified 2125 1252 23.645 759 503 28.163 Total 16 999 5295 100 13 030 1786 100

On the basis of the presence of indirect evidence, overlap the total 12 wild ungulate species inhabiting the subalpine and between musk deer and livestock was found to be highest; alpine zones of the Himalayas, only four have been studied in however, a major portion of the pellet groups of livestock was terms of their feeding ecology. The majority of the work on the reported from the Tungnath and Chandrashila regions, while no feeding ecology of ungulates has been restricted to temperate sign of musk deer presence in these regions was recorded. If we and subalpine regions, while a few studies have been conducted consider Shokhark (preferred musk deer habitat) only, then a in the trans-Himalaya (Awasthi et al. 2003). small overlap between the musk deer and livestock of the area was Duetotheshynatureofthemuskdeer,itisdifficult to observed. Therefore, it can be concluded that to conserve the make direct observations to study their feeding ecology in the musk deer, human-induced disturbance (e.g. livestock grazing) Himalayan environment. Moreover, animals cannot be should be reduced because, according to the results, the musk deer sacrificed to apply the oesophageal and rumen fistula is not utilising the area having such disturbance. technique or for collecting rumen content. Therefore, we Conservation of a species requires a good understanding of its mostly rely on information from pellet groups to study the ecology (especially habitat and feeding ecology). According to feeding ecology of this species. The microhistology Hanley (1996) and Ilyas and Khan (2004), deer species are the technique was used to investigate the feeding ecology of best indicator of the health of the forest and understanding their musk deer. It is undoubtedly the only technique available to food preferences and changes resulting from biotic influences are study the feeding ecology of small secretive species such as important in interpreting relationships between the environment musk deer. However, microhistology techniques for and the consumer, i.e. deer (Leopold and Krausman 1987). determining food habits of small herbivores can be biased by The Himalayas harbour a vast diversity of wildlife and the differential digestibility of ingested plant species (Holechek literature review on ungulates (Ilyas 2007) suggested that of et al. 1982). Forbs are usually highly digestible and, as a result, aia n edn clg fapn ukdeer musk alpine of ecology feeding and Habitat

Table 3. 95% Bonferroni confidence limits (C.L.) for available (Pio), and utilised (Pie) proportion of different grass and browse species in musk deer diet in pre-monsoon and post-monsoon seasons at Kedarnath Wildlife Sanctuary Pio, proportion of total sampling plots; Pie, proportion of plots with indirect evidence of utilisation; a, species neither recorded in sampling nor recorded in diet composition; b, species not recorded in sampling but recorded in diet composition; c, species available but not recorded in diet composition; +, utilised significantly more than the availability; 0, utilised in proportion to availability; , utilised significantly less than availability

Species Family Pre-monsoon Post-monsoon Pio Pie 95% Bonferroni Conclusion Pio Pie 95% Bonferroni Conclusion C.L. for Pie C.L. for Pie (Min Pi Max) (Min Pi Max) Trees Acer sp. Aceraceae 0.00005 0.0000 0.00 Pi 0.00 c 0.0001 0.0000 0.00 Pi 0.00 c Abies sp. Pinaceae –– – a 0.0005 0.0000 0.00 Pi 0.00 c Quercus semecarpifolia Fagaceae 0.0202 0.0000 0.00 Pi 0.00 c 0.0235 0.0000 0.00 Pi 0.00 c Rhododendron arboreum Ericaceae 0.0108 0.0000 0.00 Pi 0.00 c 0.0167 0.0050 0.002 Pi 0.007 Rhododendron barbatum Ericaceae 0.0067 0.0000 0.00 Pi 0.00 c 0.0061 0.0000 0.00 Pi 0.00 c Taxus baccata Taxaceaae 0.0011 0.0000 0.00 Pi 0.00 c 0.0011 0.0000 0.00 Pi 0.00 c Shrubs Berberis jaeschkeana Berberidaceae 0.0002 0.0000 0.00 Pi 0.00 c 0.0001 0.0000 0.00 Pi 0.00 c Cotonaester acuminatus Rosaceae 0.0012 0.0007 0.00 Pi 0.001 0.0031 0.0084 0.005 Pi 0.011 + Indigofera sp. Leguminosae 0.0005 0.0000 0.00 Pi 0.00 c 0.0008 0.0000 0.00 Pi 0.00 c Lonicera myrtillus Caprifoliaceae 0.0048 0.0000 0.00 Pi 0.00 c 0.0013 0.0000 0.00 Pi 0.00 c Myrsine sp. Myrsinaceae 0.00005 0.0000 0.00 Pi 0.00 c 0.0000 0.0000 0.00 Pi 0.00 c Piptenthus nepalensis Leguminosae 0.0002 0.0000 0.00 Pi 0.00 c 0.0003 0.0000 0.00 Pi 0.00 c Pyracantha crenulata Rosaceae 0.0012 0.0000 0.00 Pi 0.00 c 0.0030 0.0000 0.00 Pi 0.00 c Rhododendron anthopogon Ericaceae 0.0707 0.0000 0.00 Pi 0.00 c 0.1054 0.0000 0.00 Pi 0.00 c Rhododendron barbatum Ericaceae 0.0024 0.0000 0.00 Pi 0.00 c 0.0002 0.0000 0.00 Pi 0.00 c Rhododendron campanulatum Ericaceae 0.0037 0.0000 0.00 Pi 0.00 c 0.0066 0.0000 0.00 Pi 0.00 c Rosa sericea Rosaceae 0.0014 0.0000 0.00 Pi 0.00 c 0.0038 0.0000 0.00 Pi 0.00 c Rubus elipticus Rosaceae 0.0036 0.0009 0.000 Pi 0.0001 0.0008 0.0101 0.071 Pi 0.012 + Salix denticulata Salicaceae 0.00005 0.0000 0.00 Pi 0.00 c –– – a Viburnum grandiflorum Sambucaceae 0.0007 0.0000 0.00 Pi 0.00 c 0.0038 0.0000 0.00 Pi 0.00 c Herbs Anaphalis roylei Compositae 0.0045 0.0000 0.00 Pi 0.00 c 0.0007 0.0000 0.00 Pi 0.00 c Anemone obtusiloba Ranunculaceae 0.0873 0.078 0.07 Pi 0.09 0.0292 0.0146 0.011 Pi 0.018 Astragalus sp. Leguminosae 0.0007 0.0002 0.00 Pi 0.0005 –– – a Boeninghausenia albiflora Rutaceae 0.0003 0.0000 0.00 Pi 0.00 c –– – a Science Production Animal Campanula latifolia Leguminosae –– – a 0.0002 0.0000 0.00 Pi 0.00 c Cirsium sp. Compositae –– – a 0.0006 0.0000 0.00 Pi 0.00 c Clematis sp. Ranunculaceae 0.0000 0.0002 0.0002 Pi 0.0005 c –– – a Corydalis govaniana Papaveraceae 0.0152 0.0000 0.00 Pi 0.00 c 0.1103 0.0050 0.003 Pi 0.007 Delphinium sp. Ranunculaceae –– – a 0.0005 0.0000 0.00 Pi 0.00 c Dryopteris sp. – 0.0002 0.001 0.00 Pi 0.00 + –– – a Epilobium sp. Onagraceae 0.0064 0.022 0.02 Pi 0.03 + –– – a Fragaria sp. Rosaceae 0.0737 0.019 0.01 Pi 0.02 0.0513 0.0162 0.01 Pi 0.02 Galium sp. Rubiaceae 0.0004 0.0005 0.000 Pi 0.001 + 0.0008 0.0000 0.00 Pi 0.00 c Gaultheria trichophylla Ericaceae 0.0000 0.004 0.003 Pi 0.006 b 0.0021 0.0129 0.009 Pi 0.016 + G H Gentiana argentina Gentianaceae 0.0546 0.021 0.016 Pi 0.024 0.0124 0.0129 0.009 Pi 0.016 0 Geranium sp. Geraniaceae 0.0381 0.0094 0.006 Pi 0.01 0.0321 0.0291 0.02 Pi 0.03 Geranium sp. 2 Geraniaceae 0.0014 0.0000 0.00 Pi 0.00 c –– – a Science Production Animal Geum elatum Rosaceae 0.0081 0.019 0.015 Pi 0.022 + 0.0244 0.0134 0.006 Pi 0.011 Geum species Rosaceae –– – a 0.0193 0.0090 0.010 Pi 0.016 Gnaphalium sp. Compositae 0.0054 0.0000 0.00 Pi 0.00 c 0.0464 0.0084 0.005 Pi 0.011 Hemiphragma sp. Scrophulariaceae 0.0004 0.004 0.00 Pi 0.01 + –– – a Lysimachya prolifera ———- 0.0021 0.0000 0.00 Pi 0.00 c –– – a Micromeria sp. Labiateae 0.0052 0.0000 0.00 Pi 0.00 c –– – a Morina sp. Dipsacaceae 0.0006 0.0000 0.00 Pi 0.00 c –0.0121 0.0000 0.00 Pi 0.00 c Nepeta sp. Labiateae 0.0084 0.001 0.00 Pi 0.001 –– – a Ophiopogon intermedius Liliaceae 0.0069 0.009 0.01 Pi 0.01 + 0.0028 0.0123 0.009 Pi 0.015 + Oxalis sp. Oxalidaceae 0.00006 0.0000 0.00 Pi 0.00 c –– – a Oxygraphis polypetala Ranunculaceae 0.0021 0.001 0.00 Pi 0.00 0.0006 0.0000 0.00 Pi 0.00 c Pedicularis sp. Scrophulariaceae –– – a 0.0019 0.0000 0.00 Pi 0.00 c Pedicularis pectinata (Bhootkes) Scrophulariaceae 0.0009 0.0000 0.00 Pi 0.00 c 0.0000 0.0000 0.00 Pi 0.00 c Plantago ovata Plantaginaceae 0.0024 0.001 0.00 Pi 0.00 –– – a Polygonum aamplexicaula Polygonaceae 0.0345 0.062 0.06 Pi 0.07 + 0.0492 0.0039 0.002 Pi 0.005 Polystichum sp. ———- 0.0016 0.006 0.00 Pi 0.01 + –– – a Potentila sp. Rosaceae 0.1255 0.053 0.05 Pi 0.06 0.0484 0.0218 0.017 Pi 0.026 Potentilla peduncularis Rosaceae –– – a 0.0192 0.0000 0.00 Pi 0.00 c Primula denticulata Primulaceae 0.0319 0.011 0.01 Pi 0.01 0.0065 0.0078 0.005 Pi 0.010 + Rubus sp. Rosaceae –– – a 0.0048 0.0000 0.00 Pi 0.00 c Rumex nepalensis Polygonaceae 0.0063 0.0000 0.00 Pi 0.00 c –– – a Senecio sp. Compositae 0.0109 0.024 0.02 Pi 0.03 + –– – a Saussurea sp. Compositae –– – a 0.0028 0.0073 0.004 Pi 0.009 + Saxifraga sp. Saxifragaceae –– – a 0.0179 0.0062 0.003 Pi 0.008 Selinum vaginatum Umbelliferae –– – a 0.0042 0.0078 0.005 Pi 0.010 + Sibbaldia cuneata Rosaceae 0.0038 0.007 0.00 Pi 0.01 + 0.0119 0.0123 0.009 Pi 0.015 0 Skimmia anquetilia Rutaceae 0.0000 0.002 0.00 Pi 0.003 b –– – a Smilax sp. Liliaceae 0.00005 0.0000 0.00 Pi 0.00 c –– – a Spirae bella Rosaceae 0.00005 0.0000 0.00 Pi 0.00 c –– – a Stachys sp. Labiateae 0.0010 0.0022 0.00 Pi 0.00 + –– – a Tanacetum longifolium Asteraceae 0.0011 0.0005 0.00 Pi 0.00 –– – a Thalictrum sp. Ranunculaceae 0.0000 0.0004 0.0001 Pi 0.0009 b –– – a Trachydium roylei Apiaceae 0.0016 0.001 0.00 Pi 0.00 0 –– – a Vaccinium nummularia Ericaceae –– – a 0.0006 0.0000 0.00 Pi 0.00 c Valeriana sp. Valerianaceae 0.0000 0.0003 0.0003 Pi 0.002 b –– – a Veronica cana Scrophulariaceae 0.0002 0.0000 0.00 Pi 0.00 c –– – a Viola biflora Violaceae 0.0019 0.004 0.00 Pi 0.01 + 0.0012 0.0000 0.00 Pi 0.00 c Zanthoxylum nepalense Rutaceae –– – a 0.0040 0.0000 0.00 Pi 0.00 c Grass –– – Arundinaria sp. Poaceae 0.0001 0.0000 0.00 Pi 0.00 c a Ilyas O. and Syed Z. Chrysopogon sp. Gramineae 0.0172 0.009 0.01 Pi 0.01 0.0071 0.0028 0.001 Pi 0.004 Cynodon sp. Gramineae 0.0086 0.000 0.00 Pi 0.00 c 0.0004 0.0000 0.00 Pi 0.00 c Cyperus sp. Gramineae 0.0064 0.017 0.01 Pi 0.02 + 0.0201 0.0549 0.048 Pi 0.061 + Danthonia cachemeriana Poaceae 0.0518 0.0000 0.00 Pi 0.00 c 0.0312 0.0901 0.081 Pi 0.098 + Danthonia sp. Poaceae 0.0433 0.0000 0.00 Pi 0.00 c 0.0035 0.0000 0.00 Pi 0.00 c Thamnocalamus spathiflora Poaceae 0.0081 0.001 0.00 Pi 0.00 0.0074 0.0504 0.044 Pi 0.056 + Habitat and feeding ecology of alpine musk deer Animal Production Science I underestimated by faecal analyses (Vavra et al. 1978; Vavra and his valuable comments on the manuscript. Special thanks go to the fields Holechek 1980;McInniset al. 1983). Some grass and browse assistants Satyendra Singh Negi, Anil Negi and Jitendra Singh Negi. species are overestimated by faecal analyses, while others are underestimated (Dearden et al. 1975;Leslieet al. 1983). References fi Bonferroni test results with a 95% con dence interval showed Alipayo D, Valdez R, Holechek JL, Cardenas M (1992) Evaluation of that the musk deer preferred dicot over monocot plant species microhistological analysis for determining diet botanical as food. This may be to avoid competition with a sympatric composition. 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