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I .II' ;‘I:,',:': III: IIIIIIIIIIIIII :II :E’J- l lulu; lllljlflllll l l will will IL in; l LIB R A R Y Michigan State University This is to certify that the thesis entitled HABITAT RELATIONS OF THE SAMBAR (CERVUS UNICOLOR) IN KHAO-YAI NATIONAL PARK, THAILAND presented by Choompol Ngampongsai has been accepted towards fulfillment of the requirements for Ph.D. Department of Fisheries and Wildlife degree in //O 4 Z7/w ///6’/::2é4/ Major professor Date November 30, 1977 0-7639 I,-wr- , rn ~ IVA; fab in; " 3t ’7 'V‘V 4- u’hurmu HABITAT RELATIONS OF THE SAMBAR (CERVUS UNICOLOR) IN KHAO-YAI NATIONAL PARK, THAILAND BY Choompol Ngampongsai A DISSERTATION Submitted to Michigan State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Fisheries and Wildlife 1977 ABSTRACT HABITAT RELATIONS OF THE SAMBAR (CERVUS UNICOLOR) IN KHAO-YAI NATIONAL PARK, THAILAND BY Choompol Ngampongsai Habitat relations of sambar were studied between March 1976 and March 1977 in Khao-Yai National Park, Thailand. Species composition of both grassland and forest vegetation was determined. Seven grassland and 21 forest plants were determined to be preferred foods of sambar. Imperata cylindrica was by far both the most available forage and the most important food in the deer's diet. As a preferred species, it was among those plants which were consumed to a greater degree than indicated by their abundance. Vegetative analysis of the grassland range indicated that 52% of the 72 species present were pre- ferred foods, 11% were neglected forages, and 37% were avoided. No signs of range degradation were noted despite high deer density. A number of other species showed high preference rating values but their availabilities were Choompol Ngampongsai limited. Animals were judged to have only unimportant effects on forest production, in terms of their killing forest trees. The average daily food consumption by a captive three-year-old female sambar was 1084.21 grams dry weight. Assuming 40% greater food intake, the daily food consump- tion of a wild active animal was estimated to total 1517.89 grams. Energy used for maintenance and growth over a 12-day feeding trial was 29,694 kcal or 55.84% of the energy consumed. Daily water intake averaged 1.4 liters. The captive sambar spent more time resting, rumi- nating, and walking than it did feeding and standing. Feeding occurred in the early morning and late evening and also at night. Average numbers of wild deer per hectare over the period May through December were calculated to be 0.02 as determined by late-afternoon roadside counts, 0.11 from pre-midnight spotlight tallied, 3.68 from pellet-group censuses, and 3.15 on an experimentally burned food- removal study plot. The extremely high densities indicated by pellet- counts must be tested in future studies. Regardless of precise values, however, a very high grassland density seems certain to be characteristic of sambar populations in the park. This is especially true during certain seasons and especially sambar concentrations are due to the attractions of grassland foods. Choompol Ngampongsai The population consisted of 44% adults, 40% sub- adults, and 16% fawns. The sex ratio among adults was 6.17 females per male and there were 2.31 adult females per fawn. Recommendations for maintaining sambar populations in national parks are given. It is also suggested that the potential for ranching sambar for meat-production should be investigated. In view of the findings of this study, sambar may be more a productive source of meat than domestic livestock and will not require improved pasture- lands. ACKNOWLEDGMENTS Great appreciation is expressed to Dr. George A. Petrides, Department of Fisheries and Wildlife, committee chairman, for his direction, guidance, valuable counsel, criticisms, and for careful editing of the manuscript. I also thank my other committee members:' Drs. Leslie W. Gysel, also of that department, Peter G. Murphy of the Department of Botany, and Gary Schneider of the Department of Forestry for their support and enthusiastic encourage- ment throughout my graduate study. The visits of both Drs. William B. Drew, Chairman emeritus of the Department of Botany, and George A. Petrides during my data collection in Thailand are deeply appreciated. A warm expression of thanks goes to the Khao-Yai National Park staff for their cooperation and assistance which was essential to my study. My sincere appreciation and thanks also go to Dr. Tem Smitinand and Mr. Weerachai Nanakorn for plant identification, to Ph.D. candidate Thomas M. Butynski who helped me in preparing the manu- script, and to Ms. Pantipa Jantawat for her assistance in preparing drawings. ii Special thanks are extended to the Kasetsart University for educational scholarships and to the Faculty of Forestry there. In particular, I wish to express gratitude to the staff of the Conservation Department at Kasetsart University and to the Royal Thai Forest Department for providing facilities and study areas. Without financial support from the New York Zoological Society for field work, I would certainly never have had the opportunity to complete this study and for this I am most grateful. Unknown plants were identified through the kind- ness of the Division of Botany, Royal Thai Forest Depart- ment, Bangkok; the energy values were determined by the Department of Animal Husbandry, Kasetsart University, Bangkok, and the author wishes to express his sincere gratitude to them. Finally, special thanks go to my wife, NIPAPUN, whose constant inspiration, encouragement, and understand- ing helped me to complete the study. iii TABLE OF CONTENTS Page INTRODUCTION . . . . . . . . . . . . . . 1 Description of the Study Area. . . . . . . . 5 Topography . . . . . . . . . . . . . 6 Vegetation . . . . . . . . . . . . . 8 Fauna . . . . . . . . . . . . . . . 11 Climate O O O O O O O O O 0 O O O 0 1]- METHODS AND PROCEDURES. . . . . . . . . . . 14 Study Sites. . . . . . . . . . . . . . l4 Vegetation . . . . . . . . . . . . 14 Food Preference Study . . . . . . . . . . 20 Feeding Trial . . . . . . . . . . 22 Animal Effects on Forest Production. . . . . . 23 Population Study . . . . . . . . . . . . 24 RESULTS AND DISCUSSIONS . . . . . . . . . . 30 Grassland Community . . . . . . . . . . . 30 Preferred and Important Foods: Grassland Forages. . . . . . . . . . . . 36 Range Condition and Trend . . . . . . . . 40 Feeding Trial . . . . . . . . . . . . 43 Forage Production and Utilization. . . . . . 48 Sambar Populations . . . . . . . . . . . 53 Population Density. . . . . . . . . . . 53 Forage-Removal Studies . . . . . . . . . 53 Roadside Counts. . . . . . . . . . . . 54 Pellet-Group Counts . . . . . . . . . . 57 Population Structure . . . . . . . . . . 63 Daily Activity . . . . . . . . . . . . 65 Barking Deer. . . . . . . . . . . . . 68 Forest Community . . . . . . . . . . . . 68 Preferred and Important Foods: Forest Forages . 71 Animal Effects on Forest Production . . . . . 75 iv RECOMMENDATIONS . SUMMARY. . . . APPENDIX . . . LITERATURE CITED. Page 78 81 86 111 Table LIST OF TABLES Page Total and common forage species found in the grasslands, Khao-Yai National Park (July 1976) o o o o o o o o o o o o o 31 Sambar food preference ratings for grassland forages, Khao-Yai National Park, January to September 1976 . . . . . . . . . 34 Dry-weight forages eaten by a 3-year-old female sambar during a 12-day feeding trial (9 AM January 12 to 9 AM January 24, 1977) at Khao-Yai National Park, Thailand . 47 Dry weight standing crops, production, and utilization on square-meter fenced and unfenced plots burned on September 30, 1976, plus estimated numbers of sambar present on utilized plots, Nhong-King site, Khao-Yai National Park, October 1976 to January 1977 dry period . . . . 49 Combined data on sambar direct daylight counts, between 17:00 and 19:00 hours, Khao-Yai National Park, from May to December 1976 . . . . . . . . . . 55 Summary of spotlight counts made between 20:00 and 23:00 hours, from April 1976 to January 1977, Khao-Yai National Park. . . 56 Defecation rate of three-year-old captive female sambar, Khao-Yai National Park, Thailand, 1977 . . . . . . . . . . 58 Deer densities on grassland study sites as determined from fecal pellet-group counts, Khao-Yai National Park, Thailand, 1976— 1977 . . . . . . . . . . . . . 59 vi Table 10. 11. 12. l3. 14. 15. 16. 17. 18. Page Summary of calculations of average sambar den- sities on grasslands, Khao-Yai National Park, Thailand, 1976-77 . . . . . . . . 62 Proportions of size categories from 184 day- light counts of sambar between May and December 1976 on grassland sites, Khao-Yai National Park . . . . . . . . . . . 64 Number of minutes spent in various activities by a 3-year-old captive female sambar (recorded day-night every 5 minutes) over a 10-day period, Khao-Yai National Park, 1977 . . . . . . . . . . . . . . 66 Average barking deer densities as determined from fecal pellet-group counts between May 12, 1976, and March 4, 1977, Khao-Yai National Park (based on a defecation rate of 11.88 pellet-groups/day determined by Dr. Richard H. Yahner, personal correspondence. . 69 Sambar food preference ratings, forest forages, Khao-Yai National Park, January to December 1976 O I O O O O O O O O O O O O 72 Numbers of stems affected by sambar on two 400 m forest sites, Khao-Yai National Park. Numbers accumulated bimonthly, June 1976—February 1977. . . . . . . . . . 76 Relative density, frequency, and cover percen- tages plus importance index values for plant species on the Nhong-King grassland, Khao- Yai National Park, July 1976 . . . . . . 86 Relative density, frequency, and cover percen- tages plus importance index values for plant species on the Moor-Singh-Toe grassland, Khao-Yai National Park, July 1976. . . . . 89 Relative density, frequency, and cover percen- tages plus importance index values for plant species on the Nhong-Puck-Chee grassland, Khao-Yai National Park, July 1976. . . . . 92 Density and basal area of plant species, by diameter classes, Nhong-King forest site Khao-Yai National Park, June 1976. . . . . 95 vii Table 19. 20. 21. 22. 23. 24. 25. Density and basal area of plant species, by diameter classes, Moor-Singh-Toe forest site, Khao-Yai National Park, June 1976 . Density and basal area of plant species, by diameter classes, Nhong-Puck-Chee forest site, Khao-Yai National Park, June 1976 . Browsed and unbrowsed plant stems, Nhong- King forest site, Khao-Yai National Park, June 1976 . . . . . . . . . . . Browsed and unbrowsed plant stems, Moor- Singh-Toe forest site, Khao-Yai National Park, June 1976 . . . . . . . . . Browsed and unbrowsed plant stems, Nhong- Puck-Chee forest site, Khao-Yai National Park, June 1976 . . . . . . . . . Length and dry weight of leafy twig, energy values of leafy vegetative species and dropping, Khao-Yai National Park, January 1976-March 1977 . . . . . . . . . Moisture percentages and weights of fresh fecal pellets, dry at 100° C for 24 hours, Khao-Yai National Park, January 12-24, 1977. O I I O O I O O O O O 0 viii Page 97 99 101 104 106 108 110 Figure 1. 10. LIST OF FIGURES Page Sambar stag and hind showing typical body characteristics . . . . . . . . . . 4 Map of Khao—Yai National Park and its five major vegetation types (from Smitinand, 1968) o o o o o o o o o o o o o 7 Distribution of precipitation and temperature at Khao-Yai National Park, based on 1966- 1976 rainfall and 1970-1976 temperature data. . . . . . . . . . . . . . l3 Topographic map of the three study sites, Khao-Yai National Park. . . . . . . . 15 Species-area curve at Nhong-King grassland, Khao-Yai National Park, July 1976 . . . . 17 Vegetative canopy-cover at Nhong-King (open), Moor-Singh-Toe (solid), and Nhong-Puck-Chee (cross-hatched) grasslands, Khao-Yai National Park, July 1976 . . . . . . . 33 Percentages of forage categories available (open) and in the sambar's diet (solid) for (a) grasslands, January to September 1976, and (b) forests, January to December 1976, Khao-Yai National Park. . . . . . . . 37 Density, frequency, and cover percentage of preferred (open), neglected (solid), and avoided (cross-hatched) sambar foods on grasslands in Khao-Yai National Park, July 1976. . . . . . . . . . . . . . 44 Typical sambar habitat at Khao-Yai National Park, October 1976 . . . . . . . . . 45 A typical sambar grazing site, Khao-Yai National Park, October 1976 . . . . . . 45 ix Figure Page 11. Total dry-weight standing-crop production on fenced plot (solid), unutilized production on grazed area (dotted), and percentage utilization (bars) where fenced and un— fenced plots are compared, Khao-Yai National Park during October 1976-January 1977 dry period . . . . . . . . . . . . . 50 12. New Imperata dry-period sprouts on a pilot fenced pIot, Khao-Yai National Park, October 1976 . . . . . . . . . . . 51 13. Sambar grazing signs on an unfenced plot, Khao-Yai National Park, October 1976 . . . 51 14. Seasonal changes in sambar abundance as indi- cated by fecal pellet-group counts (see Table 8), Nhong-King, Khao-Yai National Park, Thailand 1976-1977 a o o o o o o 60 15. Total and average sambar activities (observed over a 10-day period), Khao-Yai National Park, January 12-21, 1977. . . . . . . 67 16. Animal effects on forest tree stems at Nhong— King (a), Moor-Singh-Toe (b), and Nhong- Puck-Chee (c), June 1976-February 1977 . . 77 INTRODUCTION Thailand resembles other developing countries in having only recently undertaken the scientific management of its forest lands for purposes other than timber pro- duction. Attempts by the national government to establish and/or enforce regulations pertaining to forest reser- vations, national parks, and wildlife sanctuaries are often ineffective. This is mainly because of existing low levels of education and the low standard of living. The public is often unprepared to accept restrictions on lands that have traditionally been free of regulation. Villagers, especially those living within easy reach of the forests, national parks, and wildlife sanctuaries, often believe that trees, wildlife, scenic beauty, and other national resources are gifts of nature and belong to no one. They believe that they should have the right to occupy any lands, cut any trees, and hunt all wildlife freely (Banijbatana, 1966, 1967). Country people have always looked to the forest as a source of food, fuel, and shelter and do not now feel the necessity of having forest areas set aside as national parks. With rapid population growth and sub- ‘ sequent expansion of towns and cities, however, the need for recreational areas has increased. This is especially true to satisfy the need of people in urban areas (Banij- batana, 1966, 1967). Yet the people are apathetic to the necessity of establishing recreational areas or even for the protection of wildlife for their own enjoyment. Successful wildlife management requires basic understandings of animal habitat requirements, such as food, cover, water, and living space. Together, these essentials determine the ability of an area to support animal life. Due to effective fire control in Khao-Yai National Park, seedlings and saplings of woody species have become established in the grassland areas. While the present grasslands in this Park are important to wild herbivores, it is believed that secondary forests will eventually replace the meadows. No studies of wild hoofed animals and their relation to their habitats have been undertaken in all of Thailand. The objective of this study was to establish basic standards for the management of sambar (Cervus unicolor) in the park. The investigation was conducted between March 1976 and March 1977 in Khao-Yai National Park, Thailand. Specific study objectives were: (1) to compute food preference ratings for the sambar (2) to apply the ratings of forages used by the sambar in predicting range condition and trend (3) to appraise the effects of wild animals on forest regeneration and timber production (4) to provide sound management policies so as to insure the survival of sambar and the preservation of their habitats in the park and elsewhere. The sambar is related to the red deer or stag of EurOpe and Asia and to the elk of North America. It is a typical forest dweller of southeastern Asia. Sixteen subspecies of sambar (Whitehead, 1972; Grzimek, 1972) range from Ceylon through India to Burma, southern China, and Taiwan, south to Thailand, Malaysia, Sumatra, Borneo, and Celebes (Medway, 1969). Full-grown stags stand about 1.37 m at the shoulder and weigh about 227 kg. The color- ation in both sexes is uniform light brown, with under- parts paler. Stags are generally darker than hinds, approaching black or slaty grey in old males. The hair is long and coarse. It forms a heavy ruff around the throat of the mature stag (Thom, 1937; U Tun Yin, 1976). The males have antlers that are shed and replaced annually (Figure 1). The first set of antlers are grown at 3 years of age and are straight spikes. The second set has two tines, and the third and subsequent sets of antlers always have three tines (Medway, 1969). For most sambar, especially at Khao-Yai National Park, the Figure l. Sambar stag and hind showing typical body characteristics. antler-shedding period is from May to July. Antlers develop to the hard stage in December. During the rutting season, old stags stalk about with erect tail, outstretched muzzle, and everted face glands (Thom, 1937). At this time they are highly dangerous, especially in captivity. Each male fights for his territory and retains females which enter his area. After the mating season, stags leave the females and stay by themselves until the next rut. Most fawns are dropped during the rainy season. At Khao-Yai National Park, mating occurs in January and February, with most young being born during September and October. The ges- tation period is about eight months (Thom, 1937; Kenneth gt 31., 1953). The sambar is essentially an animal of the more open deciduous forests and does not favor dense tree growth. It is quite diurnal in its habits (Krishnan, 1972). When alarmed, sambar utter a loud, whistling call and raise the tail to reveal the white underside. Both sexes stamp their feet when suspicious of danger (Peacock, 1933; Bentley, 1967). Sambar in captivity have lived to a maximum of 26 years (Manville, 1957). Description of the StudyAArea Following a period of increased deforestation and land cultivation, Khao-Yai National Park was established as Thailand's first national park in September 1962 under the recommendation of Dr. George C. Ruhle (1964). This was done to preserve at least some natural areas for the future enjoyment of Thai citizens and visitors from other parts of the world. Geographically, the park is about 200 km northeast of Bangkok. It lies between 14° 5' and 14° 15' north latitude; and between 101° 5' and 101° 50' longitude. The park is rectangular in shape, 2168 km2 in size and extends into four political provinces: Pak Chong district of Nakorn Ratchasima Province in the north, Nakorn Ratcha- sima and Prachinburi Provinces in the east, Nakorn Nayok Province in the south, and Saraburi Province in the west (Figure 2). Topography Except for low undulating land in the east, the park is mountainous and extends from 250 to 1400 m above sea level. The western limb of the Dongrak Range includes two vital watersheds: Lam Ta Kong in the north and Mae Nam Nakorn Nayok to the south. In general, the northern and southern boundaries are steep escarpments while the western border is made up of irregular limestone peaks and outcroppings. The eastern slopes descend gradually from upland forest to agricultural areas outside the park (Enderlein and Maxwell, 1976). The Dongrak Range is composed of Permian and Jurassic limestones of the Ratchaburi and Kamawkala Eonwv mommp coflumummm> Momma o>Hm muH can xumm HMCOHum .Ammaa .cemeAuAEm z Hmwuomnx mo mm: (5.218.: 205.42 #1 3 mkéo; Mean minimum monthly temperatures Figure 3. JFMAM l 1 Month ASOND ‘3‘) 1") fix ‘3 O \a O a. 3 20"; h. 0’ EL 5 *- Distribution of precipitation and temperature at Khao-Yai National Park, based on 1966-1976 rainfall and 1970-1976 temperature data. METHODS AND PROCEDURES Studnyites As time and transportation facilities were limited and provisions for security were minimal, all study sites were necessarily located within 10 km of the Park Head- quarters (Figure 4). Intensive investigations were con- ducted on three grassland sites at Nhong-King, Moor-Singh- Toe and Nhong-Puck-Chee (Figure 4) and in the forests sur- rounding them. The attitude at these places was between 700-900 m above sea level. Vegetation Square-meter plots were used to sample the grass- land community. Plots were distributed regularly through- out the three sample grassland areas. It was assumed that this distribution provided a representative sample of the grassland community. The species-area curve was used to determine the number of plots needed in order to sample the forage species adequately. Braun-Blanquet (1932) considered that the sample was adequate when the species- area curve become approximately horizontal while Cain (1938) stated that the sample size is adequate when a 14 15 . . , ) 14 , a l I. ‘ '- I '1' ' l.’ ' .‘u, , 1' . I \I“ .‘ ~ ‘.IJ \ .4) . . f //4 . .' ' . ' ‘ 3 r k‘ - I ' \v " ’ 0 am an « t .-.. Figure 4. Topographic map of the three study sites, Khao-Yai National Park. 16 10% increase in the sample area results in a 10% increase in the number of species present. Braun-Blanquet's cri- terion was applied in this study. At Nhong-King (Figure 5), 35-40 plots were required to meet these requirements. To insure an even sample dis- tribution and to obtain additional information, vegetation data were gathered from 48 plots spaced at 100 m intervals along lines 100 m apart. Distance between plots were measured with a plastic rope of known length. Lines were kept equidistant using a hand compass. The index of similarity "S" (Odum, 1971) was used to compare the vegetative composition of paired study sites. With reference to areas A and B: S = 2C /(A + B) Where C is the number of species common to both areas, and A and B are the numbers of species found in areas A and B, respectively. 8 is the fraction of species on the two areas which is common to both areas. The closer the value of "S" to unity the more similar the samples. The index of dissimilarity, "D," is l-S. Clipping and weighing were used to estimate pro- duction and utilization (Brown, 1954; National Research Council, 1962). Herbs were clipped at the mean grazing height observed for each species and placed in a plastic bag. After weighing the fresh specimens, grasses and l7 .ohma mash .xumm HMQOHumz Hmwlomsm .Unmammmum mcHMImconz um m>uso nonalmmflommm .m musoflm «HG-m no awn-Ea: Aun- _Dnv .Unv nfln" awn" .me .Umu nun nun nu .U j H u L- . u d 1 fi u q q s {o Jaqulnu saload 18 herbs were dried at 75° C for 24 hours in a vacuum oven. Leafy twigs were treated similarly but at 100° C. After cooling to room temperature, specimens were weighed to the nearest 0.001 gm and moisture percentages computed. Production and utilization plots were established on the grassland at Nhong-King. Two 100 m2 plots were established approximately 20 m apart, one fenced to exclude large herbivores and the other not. On Septem- ber 30, 1976, before the plot markers and fencing were placed, all forage species within them were removed by burning. Each 100 m2 area was divided into 25 smaller plots, measuring 4 m2 in area. These were marked by wooden pegs extending 5 cm above ground level. Every 15 days, average production and utilization rates were determined by clipping from two randomly selected 4 m2 plots on each of the fenced and unfenced sites. Utili- zation was computed as the difference between forage weights on the protected and grazed plots. On each plot, all species of grasses and sedges, forbs, shrubs, and trees present were identified. For woody plants, the number of leafy twigs up to 2 m, the maximum browsing height of sambar, were tallied by the twig-count method (Shafer, 1963). Additionally, the line-intercept method was applied to determine the percentage of plant cover 19 (Canfield, 1941). The canopies of plants which inter- cepted the right-hand border of the wooden square-meter frame were measured as a straight-line distance and recorded by species. Similar procedures were used on 43 and 45 plots at the Moor—Singh-Toe and Nhong-Puck-Chee grasslands, respectively. In the forest community, a system of nested plots (Daubenmire, 1968; Gysel, unpublished) was adopted for study. Three rectangular plots, 6 x 15 m, l x 15 m, and 1 x 3 m, were established within each other and with a common center line. All trees over 1.5 cm diameter at breast height (d.b.h.) were measured throughout the 6 x 15 m plot. Woody plants taller than 0.5 m and having a d.b.h. greater than 1.3 cm were measured in the l x 15 m plot. Plants less than 0.5 m in height and smaller than 1.3 cm d.b.h. were measured in two 1 x 3 m plots located at both ends of the central line of the macro-plot. The percentage of crown cover was estimated along the central line of each 6 x 15 m plot (Cain and Castro, 1959). To assess the influence of sambar on forest trees, all plants rooted within 1 x 15 m and l x 3 m plots were recorded in terms of browsed and unbrowsed stems. Since sambar mainly utilize those portions of the forest in close proximity to the grasslands, 10 sets of nested plots were established in the forest in each locality. At each site, four parallel lines were extended 20 from the grassland at right angles to forest edge. The lines were 100 m apart and each set of plots was estab- lished on the lines no nearer than 100 m from the next one. The number of plots on each line varied between 2 and 4. A hand compass was used to keep the lines equi- distant. Plastic ropes marked at the desired lengths were used to delineate plot boundaries. The twig-count method (Shafer, 1963) was used on the ten 1 x 6 m forest plots to determine the amounts of browsed and unbrowsed twigs. The original lengths of browsed twigs were obtained by comparing their twig diameters with those of unbrowsed twig specimens (see Shafer, 1963); 30-50 specimens of unbrowsed twigs were selected randomly from different plants to determine mean dry weights. Food Preference Study Papageorgiou (1972) stated that there are two important observations in herbivore food habit studies: (1) the percentage of the animal's diet which a plant species contributes, and (2) the percentage of each forage which is cropped by the feeding animal. The dietary percentage indicates the principal foods con- sumed. The percentage cropped illustrates the degree to which that species is chosen from among those available to be eaten. 21 If all forage species were present and available in equal quantities, the composition of the animal's diet alone would reveal its food preferences. Such conditions, however, do not exist in nature. While forage preference may be expressed as the percentage of each species removed by the animal (Casebeer, 1948; Papageorgiou, 1972), it is usually more revealing to express forage preference ratings as the quotient which results when the species' percentage in diet is divided by its percentage availability (Petrides, 1975). The ratio of the percentage in the diet to the per- centage availability yields a value which if greater than 1.00 indicates the relative degree of preference, and if under 1.00 illustrates the degree of unattractiveness. A ratio of 1.00 demonstrates that a species is eaten as it is encountered, being neither sought out nor neglected. The ratio serves as a standard of relative comparison according to the degree of preference. The computed ratios are specific to the time that the data were collected and are restricted to the locality involved. Other food species present, the chemical com- position of both soil and plant, as well as the season of the year are important factors influencing preference ratings (Ivins, 1959; Guy, 1976). The reason why certain foods are preferred over others is not disclosed by either percentage utilization determinations or by food prefer- ence values. Relative forage preference ratings, however, 22 may be employed to appraise range condition and trend. Reductions in highly preferred species and increases in less-eaten species may be used to indicate increases in population levels with respect to range carrying capacity (Stoddart and Smith, 1955). The formula prOposed by Petrides (1975) was used to calculate a preference rating for each food plant. These values identified preferred foods as opposed to those which were neglected or avoided. The percentages used in the ratios were calculated from species' dry weights. Grasslands in the Khao-Yai are burned annually in late December or early January, and utilization by sambar begins in January. Food preference calculations were made from January to September in the grasslands and from January to December in the forest habitat. Feeding Trial Although only one captive sambar could be used, the effort was made to determine its daily food con- sumption and maintenance requirements. The tame 3-year- old female sambar used was held in a 10 x 10 m cage. The feeding trial was conducted for the 12 days between January 12-24, 1977, but the deer was penned for 15 days prior to the study to allow for acclimation to captive conditions. The sambar's weight was recorded before and after the trial. 23 Each day, pre~weighed foods were placed in a wooden feeder. Only preferred foods were offered to the animal. Since most preferred forage species were difficult to find during the time of study and a con- siderable amount was needed, only the relative abundant Imperata cylindrica and Neyraudia reynaudiana were util- ized. Food was provided at double the animal's expected daily intake to insure that foods were available to the animal. No other foods were given to the animal during the study period. Droppings were collected and weighed every morn- ing. Approximately 12-15% of the droppings were placed daily in a plastic bag. These were dried for 24 hours at 100° C. The energy values both of forage plants and sambar feces were determined using a bomb calorimeter. Drinking water was available ad libitum in a 5-1iter plastic container. Urine was not collected in this study. Animal Effects on Forest Production To determine the effects of deer on forest pro- duction, six 400 m2 plots were located at three forest sites. Two such plots were placed about 50 m apart at each site. Their boundaries marked with red paint. Only those plants affected by animals were tallied. Three damage categories were recognized: (l) rubbed: rubbed by deer antlers and the bark damaged; (2) overbrowsed: removal of the terminal shoot or over half of stems and 24 branches; (3) killed: dead stems which were clearly caused by deer. Stems killed by other causes were not counted. For simplicity, the 4 m2 plot system applied over 400 m2 sample plot as used in the grasslands surveys was adopted. Plastic ropes were used to delineate plot boundaries of the 4 m2 plots only when records were made. All trees were examined in each 4 m2 unit over all parts of each 400 m2 plot. The number of damaged trees was carefully tallied every two months between June 1976 and February 1977. Efforts were also made to count all tree seedlings browsed by animals taking care not to overlook any which might have been clipped close to the ground. Population Study In studies of wild herbivores, the relations between vegetation and animal population are of essential importance. Among census techniques, the fecal pellet- group count is a standard procedure. First described by Bennett gt El, (1940), the method has advantages in field plot sampling and statistical analysis. Numerous refine- ments have made the technique a useful research and man- agement tool (Harris, 1959; Neff, 1968). The pellet-group count technique is the one most universally recommended by wildlife biologists throughout the northern and western deer ranges of the United States (Ryel, 1971). The method assumes that deer defecate at a constant frequency, that the pellet-groups persist long 25 enough to be counted, that the groups can be found and counted correctly, that the deposition period can be delineated, that the age of the groups found can be related to the deposition period. Moen (1973) and Smith (1974) agreed that in applying this technique, knowledge concerning the defe- cation rate of the species under study is necessary. Neff (1968) summarized factors which are believed to cause higher defecation rates such as good range con- dition and relatively high feed intake (Rogers et 31., 1958); high moisture content in forage (Longhurst, 1954); change in diet such as from native range to concentrates (Smith, 1964); high percentage of fawns in the population (Smith, 1964); and psychological effects of captivity (Neff, 1964). Though defecation rates do vary, the average number of pellet-groups defecated per animal per day can be used to estimate the actual or relative numbers of animals in a given area. Pellet-group sampling involves plots and one of the most important considerations is the effect of plot size and shape on observer error. In general, the smaller the sample area and the more accurate its delin- eation, the smaller the error. Circular plots are easier to delineate accurately than are those of belt transects. Pellet-group sampling is more efficient in areas of moderate pellet-group density because high 26 density may cause overlapping groups. Preliminary surveys provide estimates of mean pellet-group density and variance as well as some idea of pellet-group distribution (Neff, 1968). In this study, permanent line-transects were set out at each study site, using wooden stakes to mark line directions. Directions of the lines varied, depending 2 (so ftz), upon site topography. Circular plots of 4.65 m recommended by Smith (1968) as being the most economic and precise, were used. The number of plots needed in each sample site was obtained from pre-sample survey plots applying the formula used by Grieb (1958): N = “0.10)2 52 (0.20 x §)2 where: N = number of plots needed 32 = variance of preliminary data R = mean of the preliminary data 0.20 = selected risk of error (e.g., the estimate here is expected to fall within 20% of the mean 95 times out of 100) t0.10 = tabular value of "t" for the selected level of probability 27 Based on preliminary pellet—group surveys, 102, 147, and 73 sample plots were calculated to be necessary at Nhong-King, Moor-Singh-Toe, and Nhong—Puck-Chee, respectively. Due to the scarcity of pellet-groups in the forests, counts there were omitted. Sample plots were distributed over the entire study areas at each locality. The distance between plots was 20 m. Plot centers were permanently marked with wooden pegs. Before data were collected, all plots were cleared of pellet- groups. After two months, the deposited pellet-groups were counted on each plot. Pellet-groups present on the plots were identified as belonging to sambar or to barking deer. Each plot was counted both clockwise and counter-clockwise. The investigation was first performed on May 12, 13, and 14, 1976, for Nhong-King, Moor-Singh-Toe, and Nhong-Puck-Chee, respectively. The standard value of 12 pellet-groups per deer-day was used except as described beyond. This value was obtained by the investigator during the study (Table 9) and agrees well with the 12.1 value established by Nootong (1970) for sambar in an area near Khao-Yai, and with the value of 11.88 derived for captive barking deer being studied in Virginia by Dr. Richard H. Yahner (personal letter) of The Smithsonian Institution. Daylight counts were obtained from a truck for animals in the meadows along a permanent 10 km road route 28 (Figure 4) as well as from towers located at two sites. For daylight counting, a 300 m average strip width was estimated for counts made every day between 17:00 and 19:00 hours, a time when sambar come out from the shade to graze in the open. Counts were not undertaken on Saturdays and Sundays because of tourist interruptions. The number, sex, and size of sambar seen were recorded. Sex was identified with the aid of binoculars (7 x 35), except that accuracy may have suffered during the season when males were without antlers. Size-classes tallied in this study were based on the characteristics of 4-tame sambar of known age. Experience gained by observing known-age and tame park sambar indicated that the spotted coats were replaced by unspotted pelage at 2 to 3 months of age and both males and females under 3 years of age were always accompanied by their mothers. The three age-classes recognized by size and the above characteristics were: fawns under 11 months, sub-adults under 3 years, and adults over 3 years of age. Spotlight counts were made from the same vehicle and over the same route as the daylight tallied. A sealed—beam spotlight attached to the truck battery revealed reflected eyes. Reflected eye—shines could be seen for approximately 150 m on each side of the road. There were difficulties in identifying sex and age at a distance in the dark, and only the number of sambar seen 29 were recorded at night. Counts were undertaken between 20:00 and 23:00 hours. A food-removal census was derived from the pro- duction and utilization plots as described previously. Food-removal was computed as the difference between forage weights on the protected and grazed plots. Obser- vations made every 15 days from October 15, 1976, to January 13, 1977. The average forage-removal per square meter per day was obtained by dividing the food-removal during each period by 15 (days). Based on the known daily food consumption of the sambar from the feeding trial, the average deer present per hectare could be estimated by dividing the dry-weight food removed per day per hectare by the known daily food consumption figure. RESULTS AND DISCUSSIONS Grassland Community At Nhong-King, Moor-Singh-Toe, and Nhong-Puck- Chee, 46, 46, and 42 plant species respectively were identified (Table 1, Tables 15-17 in Appendix). At the several sites, grass and sedge species varied between 10 and 13 and forbs between 22 and 25. There were from 3 to 6 kinds of shrubs and 4 to 6 tree species found on the three areas. Herbaceous plants provided most ground cover. Grasses and forbs together provided canopy-cover for 98.3%, 98.7%, and 99.98% of the respective areas (Tables 15-17 in Appendix, Figure 6). The coarse field—invading grass, Imperata cylin- drica, dominated all three sites, covering 49.49%, 41.30%, and 31.95% of the areas respectively (Tables 15—17 in Appendix) and comprised 69.77% dry weight of all vege- tation (Table 2). Following the dominant species Imperata cylin- drica, Eupatorium odoratum, was most common. While this forb did not cover a large percentage of the grasslands, it was distributed widely and may be an important compe- titor of Imperata. Shrubs and trees were uncommon 30 31 Table 1. Total and common forage species found in the grasslands, Khao-Yai National Park (July 1976). m- ____..._., ..___-.—_ - —_ ..— Nhong- Moor-Singh Nhong-Puck Forage Spec1es King Toe Chee Grass and sedges: Imperata cylindricaa Carex indica Carex cruciatab Ischaemum muticumb Coelorachis glandulosa Cyperus sp.b Neyraudia reynaudiana Panicum notatum Chrysopogon aciculatus Scirpus grossus Eragrostis capensisb Cyanodon dactylon Paspalum conjucatumC Cyperus digitatus Fimbristylis trichoides Forbs: Eupatorium odoratumb Portulaca quadrifida Erechtites hieracifolia Pteris sp. Hedyotis sp. Adiantum sp. Helicteres obtusab Utricularia aurea Ipomoea aquatica Ipomoea sp.b Hygrophila erecta Portulaca sp. Portulaca oleacea Dioscorea stemonoidesb Vernonia ellipticaa Scoparia dulcisb Spilanthes ocmellab Alpinia sp.a Polygonum chinense Costus speciosus Mosses Jussiaca suffruticosa Amaranthus gracilis Stachytarpheta indica Euphorbia hirta Hygrophila minor Phyllanthus urinaria Commellina nudifolia x b XXXXX XXXXXXX a XX >4><><><><><><><><><><><>< X >4 X XX XXXXXXXXX XXXXXXX XXXXXKX XX X X X XXXXX xxxxxxx X xxxxxxxxxxxxxx XX Table 1. Continued 32 Forage Species Moor-Singh Nhong-Puck Toe Chee Nhong- King Forbs (continued): Centella asiatica Parameria bartata Arisaema sp. Crotalaria elliptica Vernonia parishii Alpinia oxymytra Hedyotis coronaria Hedyotis corymbosa Eryngium foetidum Blumea napifolia Emilia sonchifolia Ageratum conyzoides Merremia gemella Shrubs: Desmodium biarticulatab Desmodium cephalotoides Prismatomeris albidifolia Melastoma malabathricum Ixora sp. Pandanus sp. Trees: Schima wallichii Cratoxylon formosuma Wrightia tomentosaa Bridelia sp. Sapium baccatum Choerospondias axillaris Altingia excelsa Trema orientalis Hibicus macrophylla Oroxylum indicum Total: Common: species species Grazing intensities deer/ha/day (Table 10) XXXXXXXX XXXX X XX XXXXX XXXX XXXX XX X XX 42 29 46 46 31 4.68 4.14 2.23 aPreferred (p b Neglected (p CHighly preferred (p (blank) avoided < 1. 1.00 to 1.99) 00) = > 1.99) 33 so - 74.7 70 . so - 55. 59.8 b so - '5 40 . \2 r" a: ==:= 3 A B \ 0 so % § 2: ‘h.. .2 § § \ no - § \ \ § \ IO - § \ \ § § 0 .J S 1'0 tr. W Grass—Sedges Forbs Shrubs Trees Figure 6. Vegetative canopy-cover at Nhong-King (open), Moor—Singh-Toe (solid), and Nhong-Puck-Chee (cross-hatched) grasslands, Khao-Yai National Park, July 1976. 34 00.0 AH.0H 000.0 00.0 0H0.0 ANH.0 .mm msummao 00.0 m0.ma 0H.0 Hm.0 mm0.0 050.m mmsuno mmumuoflamm m0.0 00.0H 0m.0 0N.H mnm.~ m0H.0H mammoum msmuflom m0.0 mm.0H mm.0 H>.0 ~00.H mma.0 moflncfl xmumo 00.0 NA.AH 00.0 ~H.0 05~.0 00m.a mflmcmmmo mflumoummum 0m.0 ~0.0H m0.0 0H.0 nnm.0 m0~.~ mmoasccmam manomuoamoo 00.0 0A.0H 00.~ 00.m 0m0.0 505.00 mumnosuo xmnmo A0.0 NN.~N «0.0 00.0 000.0 00m.0 mmeflocosmum mmuoomofla A0.0 00.mm 0000.0 0000.0 «00.0 000.0 asoflumom asflmcmnm 00.0 ma.mm mN.H 00.H m0~.m N00.~H asofluse aasmmsomH 0H.H 00.~0 mm.0 0m.0 H50.H ~m0.m nesmosu0m coamxoumuo -.H A0.mv H0.00 55.00 H00.000 AH0.00m MOHuecflamo mnmummaH 0m.a 00.00 m0.H 50.0 005.0 000.0 monumflaam «Haocnm> N0.H 0m.vm 0H.m 00.~ 00~.0H MAH.0N mamaesmcmmn mfiwsmummz ~0.H 00.nm Hm.0 ma.0 sm0.0 0m0.a .mm mflcflmaa ~0.H 0m.0m 0m.0 H~.0 0mm.a 000.~ ammoucwaou mannmflnz 00.m 50.00 00.0 0~.0 00~.0 ~0~.0 saummsflcoo adamammm in: we: m2: m3 2: 20 Hm>OEwm uOHD OHQMHHMNSN HM>OE®~H OHQmHHm>¢ mmflommm mwmuom mwwmwwmwum mommucmoumm unmflmbmmmmxwmmuom .mhma HmQEoummm on wumscmn .xumm HMQOfluwz chlomzm .mommuow cacammmum How mmcflumu wocmummoum poow “828mm .N mHQMB .mdomomnumn mum mmHOmdm Hmnuo 000 “madman pCMHmmmuon mm _MM 35 s.“ 0 a s s 0 .5 " onl H o I .III 0 x H o x u 0000000 0 0 0 0 000 x 0 0 000 0 0 000 0 00 00.0 . 00.000 00.000 000.000 000.0000 0 u 00000 00.0 00.0 00.0 00.00 000.0 000.000 00000000 0000000000 00.0 00.0 000.0 00.0 000.0 000.0 .00 0000000 00.0 00.0 00.0 00.0 000.0 000.0 00000000 0000000> 00.0 00.0 00.0 00.0 000.0 000.00 000000000000 E00005000 00.0 00.00 00.0 00.0 000.0 000.0 000000 00000000 00.0 00.0 000.0 00.0 000.0 000.0 000000000 000000 00.0 00.00 00.0 00.0 000.0 000.00 0000000 0000000000 00.0 00.00 000.0 00.0 000.0 000.0 .00 00000000 0000 0000 0000 0000 000 000 Hm>OEOm #OHQ OHQMHHM>< HM>OEOM OHQMHHMNSN mmflommm mmmuom 0000000 100\000 0000000000 00000000000 030003 >00 000000 pmssflucoo .m wanna 36 (Figure 6), probably because of burning-frequency. Only plants having Special fire resistant characteristics can normally exist on annually burned areas. Both Imperata and Eupatorium sprout from the roots after the above- ground parts have been killed by fire. Based on the formula 8 = 2C/(A + B), the simi- larity index "S" between Nhong-King and Moor-Singh-Toe, and Nhong-King and Nhong-Puck-Chee grasslands were 0.674 and 0.659 respectively, and between Moor-Singh-Toe and Nhong-Puck-Chee was 0.591. There were differences in the vegetation present on each of the three areas but they were not great. These differences did not seem clearly to be correlated with sambar grazing intensities (Table 1). Preferred and Important Foods: Grassland Forages In calculating food preference ratings for grass- land species, data for the three grassland sites were combined. Of the 72 plants found in the grasslands (Table 1), only 25 were eaten by sambar. By dry weight, species available in the field were 79.37% grasses and sedges, 17.12% forbs, 2.99% shrubs, and 0.50% trees. Consumption of these respective categories was 96.07%, 2.41%, 0.59%, and 0.66% (Figure 7, [a]). It is evident that the sambar is both a grazer and browser but that grasses are by far predominate in the diet. In order of preference (Table 2), seven species were found to be eaten to a great extent than indicated 37 .xnmm Hmcowumz Hmwlomnm .mhma “anamomo ou >um:cwn .mUmmuom Anv cam .mhma Hwnsmummw ou mnmscwn .mncmammmum Adv Mom AUflHOmV umwc m.HmnEMm wnu ca can “ammov mandawm>m mmfluommumo meHOM mo mmmmucmoumm 2.. . a. nuJTufl umfimfln flung-... uflaflzm an COL 81—59 mum—u... mun-31¢ ”DID“- mmflflu ill .5 musmflm 38 by their abundance. These were: Paspalum conjugatum, Wrightia tomentosa, Alpinia sp., Neyraudia reynaudiana, Vernonia elliptica, Imperata cylindrica, and Cratoxylon formosum. Up to 80.97% of Paspalum was utilized and over 50% of each of the next four species were consumed. Eighteen species (Table 2) were eaten less often than their abundance would indicate. Twenty-two other plant species (Table 1) showed no utilization by sambar. Only a few forage species were quantitatively important in the sambar's diet (Table 2). Imperata cylindrica, Neyraudia reynaudiana, Ischaemum muticum, Carex cruciata, and Vernonia elliptica comprised 96.07% of the total consumption with Imperata alone comprising 88.81% of the sambar's diet. Known locally as ya-kar, Imperata was an abundant grass. It made up 69.77% of the available forage and was utilized heavily all year round. Eupatorium odoratum comprised 13.25% of the available forage and was the second most abundant grass- land species. Yet it constituted only 0.27% of the sambar's diet and was the most neglected of all dietary items. It appeared to compete with Imperata in some areas but whether this is an important matter in a national park depends upon the degree to which habitat management may be desirable there. In areas where sambar production is given priority (sambar ranching is thought 39 to warrant investigation, see beyond), increases in Eupatorium may serve as an indication of heavy grazing pressure. Paspalum conjugatum, with a food preference rating of 2.00, had the highest preference rank of the grassland forages. While it was heavily utilized by sambar, its availability was very low (0.02%). It appears to be a species which is sought out by the sambar and any reduction in its abundance might indicate range overuse. Wrightia tomentosa and Cratoxylon formosum were tree species having preferred ratings. They were scarce both in the grasslands and forests, however, and were relatively unimportant in the sambar's diet. It is evident from data on the seven preferred species (Table 2) that five of these were lower both in percentages of availability and dietary contribution than either Imperata cylindrica or Neyraudia reynaudiana. It can be said with confidence that these two species are the most important foods for sambar on this rangeland. The fact they not only comprised most of the forage eaten but also were sought out by sambar as preferred items indicates their especial importance to sambar. The close dependence of the sambar on Imperata cylindrica would seem to be a matter of particular ecological and possible economic significance. Ya-kar 40 is a coarse grass and an invader of open fields through- out south and southeast Asia. Because domesticated sambar are not wild sambar, in no way could the practice of sambar-ranching be con- sidered to be wildlife conservation (Petrides, 1977). Yet investigations should be undertaken in the interests of national and international economics to determine whether meat-production would be more efficient by ranch- ing tamed or domesticated sambar on coarse grasslands than by raising beef cattle on improved pastures. Alter- natively, sambar might be useful in some area to control ya-kar, perhaps being ranched along with cattle or other domestic stock (see also feeding trial data, beyond). Range Condition and Trend Species composition and the degree of forage utilization are both useful in appraising the current condition of a range area and in evaluating whether it is becoming degraded. Forage species with the highest and lowest pref- erence ratings can be used as indicators of animal stock- ing density with respect to carrying capacity, that is as indicators of range condition. When animals remove vege- tation faster than it can be replaced, the most-preferred food species tend to be removed first and to the greatest extent. Other forage species are consumed more or less severely depending largely on their preference ratings 41 but also on animal densities and the duration of grazing. Unless grazing pressures are too severe to allow any species to reproduce or grow, the avoided species and neglected forages tend to increase under heavy grazing pressures. Range condition, or the current status of the vegetative habitat, is a reflection of current and past stocking rates of the grazing-animal populations using the area. The degree of past range overuse and the evaluation of whether overgrazing continues can be deter- mined by observing the performance of both the preferred and neglected food species. On areas where preferred foods are few, species which normally are neglected or avoided may be heavily cropped. Such a range is in poor condition, and lack of food may prevent herbivore pOpu- lation growth. The density of any species is the number of animals occupying a defined area. A given density may be lower or higher than the carrying capacity of the range. If a high stocking density prevails, over- utilization of the range may occur which ultimately leads to range degradation and habitat destruction. Malnutrition, reduction in animal production, and even starvation cannot be avoided on seriously degraded ranges unless range improvement and rehabilitation are undertaken. The term "overstock" does not refer only to the absolute number of animals present but to animal density 42 with respect to the range carrying capacity for that species. The number of malnourished animals that survive on an area may still exceed that which the range can support on a long-term basis. Any stocking density which exceeds the range carrying capacity will prevent habitat recovery and lead to continuing range depletion. The current condition and trend of any rangeland can be assessed easily once food preference ratings for the area and season are determined (Petrides, 1975). To appraise range condition and trend in this study, two indicators were used: (1) the degree of util- ization of highly preferred forage species, and (2) the extent to which the more heavily utilized range species held dominance in the plant community. Plant species having preference ratings over 1.00 often are termed "decreasers" because of the effects of heavy grazing. Species rated below 1.00 are "increasers." Those plant species which are not utilized as foods can be classified as "avoided-increasers." Forage use which removes over 50% of either the vegetative or reproductive parts of preferred food plants will normally result in harm to the range (Stoddart and Smith, 1943; Sampson, 1952; Heady, 1960; Bell, 1973). On the study areas, there were four grassland species (Table 2) and seven forest plants (Table 13) which were consumed in excess of the 50% level, and these 11 species also were scarce plants in the community. 43 Vegetative analysis of the grassland range in Khao-Yai revealed that preferred food species or decreasers covered 52% of the range while 11% of the plants present were increasers and 37% were avoided-increasers (Figure 8). No scarcity or overuse of preferred foods was noted. Density and frequency percentages for these categories were: 76.5% and 18.2% for decreasers; 16.4% and 49.4% for increasers, and 7.1% and 31.9% for avoided—increasers respectively (Figure 8). There were no obvious signs of excessive utilization of forage species leading to range degradation. In further support of these findings: (1) there are only 26.9% nonpreferred food species on the range as compared with 73.1% preferred food species available; (2) the most abundant plant on the range (Imperata cylindrica) was nevertheless a preferred food; (3) only slight signs of soil erosion were seen; (4) even during the dry period, few bare-soil areas occurred on the range (Figures 9 and 10). Feeding Trial A three-year-old tame female sambar was made available by park authorities and was penned for a feed- ing trial. This was done over the 12 days between 9 AM January 12 and 9 AM January 24, 1977. 13" b Percentage n o 0 O 0 12¢! h 44 Figure 8. Density Frequency Cover Density, frequency, and cover percentage of preferred (open), neglected (solid), and avoided (cross-hatched) sambar foods on grasslands in Khao-Yai National Park, July 1976. Figure 9. Typical sambar habitat at Khao-Yai National Park, October 1976. Figure 10. A typical sambar_grazing site, Khao-Yai National Park, October 1976. 46 The captive sambar preferred Neyraudia reynaudiana over Imperata cylindrica (Table 3). Though comparable preference ratings cannot be calculated because other forage species were lacking, the feeding trial response supported the relative food preference values determined on the wild grasslands. During the investigation, 6,754.79 gm dry weight of Imperata and 6,255.75 gm of Neyraudia were utilized by captive sambar over the 12-day period. Daily food consumption was 3,329 gm wet weight or 1,084.21 gm dry weight. By the end of the study the sambar had gained 0.5 kg over its original weight of 118.5 kg. Feces col- lected over the trial period weighed 5,588.17 grams, averaging 465.68 gm dry weight per day (Table 25 in Appendix). The energy content of Imperata cylindrica, Neyraudia reynaudiana and droppings were 4.2043, 3.9604, and 4.2016 kcal/gm reSpectively (Table 24 in Appendix). Over the 12-day period, the food-energy consumed, energy defecated, and energy used in body maintenance and growth were computed to be 52,173, 23,479, and 29,694 kcal, respectively. Of the energy digested, 55.84% thus was used for maintenance and growth and 44.16% was returned to the ecosystem. Daily water consumption was about 1.4 liters. 47 Table 3. Dry-weight forages eaten by a 3-year-old female sambar during a 12-day feeding trial (9 AM January 12 to 9 AM January 24, 1977) at Khao- Yai National Park, Thailand. . Grams Grams Grass Spec1es Offered Consumed Percentages Neyraudia reynaudiana 6,863.06 6,255.75 91.15 Imperata cylindrica 7,632.02 6,754.79 88.51 Totals 14,495.08 13,010.54 Note: Feces collected over the trial period weighed 5,588.17 gm dry-weight. 48 For grazing sheep, Langlands 33 31. (1963), Lambourne and Readon (1963), Graham (1964) computed respectively that 33%, 30%, and 40% more energy is required for maintenance needs than is true for penned animals. Devendra (1967) found that free-ranging Malayan goats consumed about 44% more food than when confined. For the sambar, a 40% higher food consumption for wild deer would mean a daily food intake of 1,517.98 gm dry weight. Forage Production and Utilization Basic assumptions made in estimating forage pro- duction and utilization were that the forage growth rates on each plot were the same after burning and that the utilization measured was due to grazing by sambar alone. It must be emphasized that the heavy grazing on the study plot was a result of the local experimental burn which induced the growth of new grass shoots which were not available elsewhere. Forage production (Table 4, Figure 11) continued throughout the September 30-January 13 study although the increment rate varied between study-period segments. Almost full growth was attained on the protected plot 45 days after the burn. After that, the rate of plant growth gradually decreased. Imperata cylindrica con- tinued to grow, however, even during most of the December—January dry period (Figures 11 and 12). It 49 .maco E ooa u uon pmocmwco ”muoz .voaumm smcumoa wan um>o xuflmcmp Hmnfimm mmmum>m n mumuomn you How . I mm.hama I mm.nHmH I Auxmu mmmv xmp\HMQEmm pafi3 xn cmumm p00w U£mmm3 >HU manna p ma m «use I o cocoa I mumuownwwmp\chn ocean pm>oEoH magnum ma.m mH.H mm.am vnne.o ovma.om mv>H.o mvaw.~ onao.aa oava.am ma .cmn moa mv.m I mm.om mnmm.o mmom.nv mmmm.OI mmmm.mI mmmv.aa maom.mm mm .owo om mv.¢ ~v.H mm.mm Namm.o ammo.am mvam.o ovmm.m momh.w mmam.hm «a Home mu mm.m om.m ma.vm mnmn.o Hmmm.hw voom.o mnom.h hmmm.m mmmm.mm mm .>oz ow Hm.m mm.m mo.mn mmmm.o mamm.ov vooo.H owoo.ma mmmm.m~ Homh.mm ¢H .>oz mv hm.m mn.v oa.a> mmvm.o mmmm.mm mm~>.o mhnm.oa omom.oa mamm.mm om .uoo om mm.m mm.m mm.an Nmmm.o hnhv.va Nmom.o nhhv.va mnvm.m mmmm.om ma .uoo ma Ammlfich $553 $5.53 mausm m m ooo.oa m\m 00H «\m xoumv Ama\ov coflumm scam poam monam MWMWMM Guam mocflm woo mom Hmuoe xmo mom mom pmocwmco condom pofiumm ammo” on non wmp\mc\ummp pw>oawm Guam mocam mo pcm wmm hoop m>< mmmum>m mommucmouwm mE\Um>oEmu “mew NE\om>OEmu mew mmouu mcHUcmum Abe AHV Amy on Amy Ame ADV on Amy Aums¢mo on whma Honouoo .xumm HMCOflumz Hmeomnx .0UHm mafia Imconz .muon pmuaawus co ucwmmum umnfimm mo mumnEdc omumEHumm moan .mhma..om uwnfimummm so common muon poocwmcs paw pmocmm HmuoEImumdqm co downwaflawuo can .COHu0960um .mmouo mcfipcmum unmflm3 >Ho .v manna 50 100 100 - 4 I_I 80 - »‘F"‘ ‘IOO fl «A _I E p _. Ir» ‘ g ( V60 P A“ 1 6o .— 3 r’”) g a ‘ I: 3 o I «.3 ‘5 2 P N 5 ‘ 2 III I- a 3 ..20 E P~~ 4”““- p‘ ‘ “‘rfirv" O 60 75 90 105 DAYS Figure 11. Total dry-weight standing-crop production on fenced plot (solid), unutilized production on grazed area (dotted), and percentage utili- zation (bars) where fenced and unfenced plots are compared, Khao-Yai National Park during October 1976-January 1977 dry period. 51 Figure 12. New Imperata dry-period sprouts on a pilot fenced pIoE, Khao-Yai National Park, October 1976. Figure 13. Sambar grazing signs on an unfenced plot, Khao—Yai National Park, October 1976. 52 was impossible to ascertain when growth stopped com- pletely because both plots were destroyed by fires which swept throughout the park grasslands after seven records had been collected. In contrast to forage growth rates, the amount consumed per day on the unfenced plot decreased even though the grasses were kept short by grazing. Evidently as time went on, the plants became tough or lost nutri- tional value (Figure 13). Percentage utilization ranged from 71% to 88% (Table 4, Figure 11) on the unfenced plot during the study period, but it must be considered that this plot was the only burned (and therefore desirable green) spot in the entire region and attracted concen- trations of deer (see beyond). Comparisons between standing vegetation on the fenced and unfenced plots indicated the amounts of food removed for each period of the study (Table 4, Column D). The food utilized per day was highest during the first 45 days after the controlled burn of September 30. Even Imperata shoots quickly followed the fire and deer were attracted to this, the only green spot in the entire region, in large numbers as the vegetation dried out, in the November-December, deer use of the forage declined though animal densities nevertheless were high (Table 4, Column I). 53 Sambar Populations Population Density Data on sambar abundance were collected in four ways: (1) measurements of forage use, (2) daylight road- side counts, (3) spotlight roadside tallies, and (4) fecal pellet-group analyses. Forage-Removal Studies When computed on a per-hectare basis over a 105- day period (Table 4), an average of 4,777 grams dry weight of vegetation disappeared daily and was presumed to have been eaten by the sambar population. Since each deer consumed 1,517.89 grams dry weight per day (Table 4, p. 49), it may be calculated that the study plot was grazed with an intensity of 330.75 deer days per hectare. Sambar grazed there, that is, at an average density of 3.15 animals per hectare per day. An average density of 315 sambar per km2 is unbelievable over a large area. But the evidence indi- cates that this condition did prevail on the tiny (100 m2) plot after it was burned on September 30. This area soon thereafter sprouted green grass from the burned stubble clumps, and this was at a time when such forage was less available elsewhere. As is commonly recognized by vil- lagers in the region and as demonstrated by this experi- ment, green growth on Imperata grasslands induces high- density utilization by sambar. 54 ‘ That sambar occur at high densities on Khao-Yai grasslands was evident throughout the study (Tables 5, 6, 8, 9). During the same period that the food-removal study was conducted, a lower but still high average density of 135 sambar per km2 was calculated (see beyond) over the grassland as a whole. The fact that forest openings are few (grasslands comprise only 3.9% of the 2,168 km2 national park) un- doubtedly results in deer moving out of the forests to feed whenever new grasses become available. Seasonal levels of sambar abundance in the grasslands must be closely correlated with (and probably can serve as direct indicators of) the availability of green growth there. Roadside Counts Tallies made from a moving truck both before dark and prior to midnight revealed low sambar densities, at least when compared with pellet-group counts. The evening census conducted between 17:00 and 19:00 hours over the 10 km route from May to December yielded 0.02 sambar per hectare (Table 5) while during the same period and over the same route the 20:00-23:00 hours spotlight enumeration estimated an average of 0.10 per hectare (Table 6). Though indicated even more strongly by pellet- group data (see beyond), the roadside results also 55 Table 5. Combined data on sambar direct daylight counts, between 17:00 and 19:00 hours, Khao-Yai National Park, from May to December 1976. Month Mo. of No. of Avg. per Deer pera Sightings Counts Count Hectare May 168 19 8.84 0.03 June 360 54 6.67 0.02 July 416 49 8.49 0.03 August 356 78 4.56 0.02 September 280 60 4.67 0.02 October 218 86 2.53 0.01 November 157 90 1.74 0.006 December 213 85 2.51 0.01 Average 0.02 a Deer per hectare = Avg. no. deer seen = Strip width x length Deer seen _ Deer seen 0}3km x 10km " 300 ha 56 Table 6. Summary of spotlight counts made between 20:00 and 23:00 hours, from April 1976 to January 1977, Khao-Yai National Park. Month No. of No. Deer Avg. Deer Deer pera Counts Seen per Count Hectare April 1976 2 123 61.50 0.21 May 8 354 44.25 0.15 June 9 408 45.33 0.15 July 11 349 31.73 0.11 August 7 232 33.14 0.11 September 7 123 17.57 0.06 October 10 188 18.80 0.06 November 3 43 14.33 0.05 December 8 243 31.13 0.10 January 1977 7 267 38.14 0.13 Average 0.11 Deer per hectare Avg. deer per count = Strip width x length Deer seen _ Deer seen (7.3km x70km _ 300 ha 57 disclose that deer tend to congregate most after dark. During daylight hours, the deer are much less frequently seen. They are believed then to be mostly in the forest. Even though the sambar densities determined by these visual counts are very much lower than those derived beyond from pellet-group counts, at least the spotlight- determined average density of 10 per km2 indicates that sambar concentrations on grasslands in late evening near the roads are not low. The effect of road traffic and tourist activities near the road, too, may have resulted in reduced deer numbers there. The roadside counts also supplement the pellet- group censuses in revealing seasonal changes in abundance. Deer densities were highest on grasslands during the early rains when new and tender grass-shoots were most available. As the year progressed and, presumably, as the grasses became taller and more coarse and also more dry, sambar became fewer. Pellet-Group Counts Estimates of average deer densities on the several grassland sites over the period from May to March (Table 8) revealed that high-intensity sambar use was characteristic over wide areas, all of which were outside the boundaries of the roadside surveys. There was a close relationship between rainfall and deer density (Figure 14). Even during the driest season, however, 58 NH mommm>< was Hmuoa NH om as ma OH «N Ha as ma 4H ma mm HH mg «a ma OH mm NH RH NH NH ma Hm sumacms ma SH sumacmn ma Ha sumssms mmo mom moo mom mmo mom mmsoumnumaamm mama masouanumaamm muma mmsoumuumaamm memo .nama .ocmagmze .xumm Hmcoflumz Hmeomnm .HMQEMm mHmEmw m>Hummo UHOIHmmmImwunu mo mumn coaumomwmo .n magma 59 Table 8. Deer densities on grassland study sites as determined from fecal pellet-group counts, Khao-Yai National Park, Thailand, 1976-1977. Moor-Singh Nhong-Puck Site: Nhong-King Toe Chee Average Number of plots: 102 147 73 Number of deer per hectarea (and number of days since plots cleared of pellets) July 9.14 9.95 2.57 (69) (67) (67) September 6.90 3.29 1.82 (54) (57) (54) November 1.82 2.21 1.81 (60) (57) (57) January 0.80 0.88 0.96 (57) (29) (59) February 4.16 1.71 4.22 (38) (62) (39) March 3.24 5.18 3.78 (13) (20) (13) Average 4.68 4.14 2.32 3.68 (291) (272) (289) Number of pellet-groups/hectare Number of days since plots cleared x 12 groups/day/deer aAverage/deer/hectare = 6O .shmanmnaa ccmagmne .xumm Hmcogumz gmwuomns .mch Imconz .Am manna mmmv mucsoo maoumlumaamm Hmowm an kumoapcfi mm uncapcsnm HmnEMm CH mmmcmno Hmcommmm .va musmam augecc . ¢< .‘ ¢< u q .u _. a. mi .t q o o o o O X o o u r. x . one ...u mu 0 o x MW “w e u x o .L can mu w m m m mm a 1. o _1 can \I mm ‘ O O m a .. o _. one 5:338... a x 535.. .8: x or . can 61 averages of 80 to 96 deer per km2 occupied the study sites. And during the June-July time of highest sambar utilization, between 257 and 995 deer per km2 occupied the openings. Since these high densities were not observed during either the early or late evening counts or during more general daytime observations, concentrated sambar grazing must have occurred between midnight and dawn. (Also since all pellet-count areas were further from the road than the 150 m half-strip width as seen from the vehicle and may have received more intensive grazing pressure.) Because sambar did not occupy the feeding areas throughout the day, many of the 12 fecal groups which they defecate daily (Table 7) must have been deposited in the forest, or at least away from the study sites. Hence, populations on the grasslands during the major late-night feeding hours must have been very great indeed. During the lOS-day period when the food-removal study indicated an average density of 3.15 sambar per hectare on the small previously burned plot (Table 4), pellet-group counts over the more widespread study sites indicated an average density of 1.35/ha (Table 9). The very high densities of sambar found here by the pellet-count method must be confirmed in later 62 Table 9. Summary of calculations of average sambar den- sities on grasslands, Khao-Yai National Park, Thailand, 1976-77. Deer/ha Roadside Counts: Evening tallies, May-December Spotlight surveys, May-December Censuses September 14-January 9: Pellet group counts2 Food removal, 100 m area only Pellet—group counts, May-Marcha 1.35 3.15 3.68 aAverages for all three sites and believed to be representative of the forest clearings near park head- quarters. 63 studies. It seems certain, however, that by any stan- dards deer utilization of the grasslands must be at a very high level. It would be most interesting to know the size of the area from which sambar are drawn to forest open- ings, and also to what degree sambar require woody vegetation to be present. Population Structure During the 184 daylight roadside counts, attempts were made to identify the sex and age-class of each sam- bar seen. Though care was taken to avoid errors, it is possible that some males with shed antlers were tallied as females. Fawns, too, because of their small size and likelihood to be hidden by vegetation, may have been undercounted. That the counts showed almost identical results on the three study areas (Table 10) may either indicate that errors in identification were not serious or that they were made in a uniform manner. Based on the data collected, males comprised only 21% of the population. Possibly, sex segregation occurred and male herds were not encountered. Mr. Weer- achai Nanakorn, an educated and experienced park officer, had never observed such segregation, however, and felt that a ratio of one male to six or seven females was normal for adult sambar. 64 .mmHmEmm can moans maamsqm coon w>mn ou pmfidmmum mum pun xwm wn pmflmaucmpw nos wum3 mn3mm a .mmuwm mwsonosmImQOSZIu can .moelmchIHoozIm .mcHMImGOSZIasm «Hangman: em.o mh.o~ av.o ms.o mom.o ~mm.o mamummoqmum mmusom 4m.o m~.- hm.o sm.o G¢H.o mmm.o .mm nuances so.a ow.m~ nv.o q¢.o mho.o sm~.o magaomggumc mamumoonom 4H.H Ho.m~ Go.a mm.o nna.o ~mm.o .mm xmusum mm.H oo.vm mm.o mm.o va.o mm¢.o .mm mgum>a ~4.H mo.m~ ma.m ms.~ Hmm.o amm.a maoowmgu machmsm vm.a so.mm o~.o «H.o mmo.o mmo.o mumHsoacmm mamuusz m¢.a sv.mm nm.o m~.o ~mo.o opa.o .mm mflcmmsm mm.a mm.am sv.o Hm.o mso.o mo~.o maamsoson «Humans ~m.a om.hm ma.> 54.4 Ho~.H o-.m maummuumagesom mummocmummo mm.a mm.hm mm.o sm.o mmH.o vm¢.o mflHoMflumsHmosm mamumoonugq mowa m~.os om.H~ mm.~H omm.m sma.m mamumoocmso manucmHmmq os.a No.~¢ mm.a HH.H mam.o ~mn.o .mm mguosuom mh.a mm.mv mo.o H¢.m mHo.H mam.~ asumaamusom monoamamz m~.H mm.m¢ mm.o om.o omH.o Hem.o mush maum>o mm.a mm.m¢ hm.~ m4.a om¢.o mmm.o mahnmuss engamnmmz «m.~ ms.hm sm.v va.H mmh.o Hmm.a .mm muons mv.~ oa.am ~m.m mm.H mmm.o «no.a mflmcmEMAm «gunman mm.~ sm.mo mm.H m~.o mmm.o mmm.o .mm whammam on.~ mH.mG hm.o oa.o mvo.o mmo.o mogcmasmu ammugaomz mn.~ n~.~m vv.o ma.o «so.o oaa.o mscmgummcou msmumuonugq 5H.m om.mh HH.H mm.o Gma.o av~.o mcHHsMH mamas mm.m om.am ma.a mm.o th.o mmm.o mumgnomun agaflmumu ME m3 .13 m3 9: a: Hm>OEmm umfla mangaflm>¢ Hm>oEwm mammaflm>¢ mocwumm mmflommm mmmuom mocmummmum mommucwoumm Amn\mxv unmflmz who mmmuom .mhma umnEmomo ou NHMDGMH .xumm HocOHumz Hmeomcm .mmmmuow ummHOM .mmcflpmn mocmummmnm GOOM Hmnfimm .mH magma 73 .m I m I mm I .mm I 000008500 I 0 .0 I 0 “000 x m I 0 “000 x m I 0 “000 x 0 I 00 00.000 00.000. 000.00 000.00 0 I 00009 00.0 00.0 00.0 00.0 000.0 000.0 00000000 000000 00.0 00.0 00.0 00.0 000.0 000.0 .00 50000000 00.0 00.0 00.0 00.0 000.0 000.0 .00 2000s000 00.0 00.0 00.0 00.0 000.0 000.0 000002000 0000000000 00.0 00.0 00.0 00.0 000.0 000.0 0000000 000000 00.0 00.0 00.0 00.0 000.0 000.0 00000000 0000>m 00.0 00.0 00.0 00.0 000.0 000.0 00000 2020200000 00.0 00.00 00.0 00.0 000.0 000.0 00000000 00000500 00.0 00.00 00.0 000.0 000.0 000.0 00000000000> 000000 00.0 00.00 00.0 00.0 000.0 000.0 0000000000 0000x000 00.0 00.00 00.0 00.0 000.0 000.0 .00 0000000 00.0 00.00 00.0 00.0 000.0 000.0 200000000 2000000000 00.0 00.00 00.0 00.0 000.0 000.0 00000000000 0000000 00.0 00.00 00.0 00.0 000.0 000.0 asu0>0 000000202 00.0 00.00 00.0 00.0 000.0 000.0 .00 008000 00.0 00.00 00.0 00.0 000.0 000.0 5000>0ns0 2050200000 00.0 00.00 00.0 00.0 000.0 «00.0 00000000 050000000 00.0 00.00 00.0 00.0 000.0 000.0 0uusesx0 0000000 00.0 00.00 00.0 00.0 000.0 000.0 000000000 000000000 00.0 00.00 00.0 00.0 000.0 000.0 000>00xm 00000000 00.0 00.00 00.0 00.0 000.0 000.0 0000000 000000200 00.0 00.00 00.0 00.0 000.0 000.0 00000000 x0000 0000 0100 0100 0x00 000 A00 am>oEmm umHQ mHQMHHm>< Hm>oamm magmaflm>< mmcflumm mmaommm ommnom mocmummmum mommucmoumm Amn\mxv unmflmz who mmmnom omscflucou .ma mHQMB 74 Lasianthus cyanocarpus, Ixora sp., Castanopsis acuminatissima, and Melaleuca seutellatum were the most important food species in the forests. They constituted 21.30%, 9.97%, 7.19%, and 6.08% to the sambar's diet, respectively. The first three of these were preferred forages. Collectively, they comprised about a third of the forest vegetation available to sambar. Twenty-six species were rated as neglected forages. These formed 62.83% of the available food and 38.99% of the sambar's diet. Only Lasianthus, Ixora, Castanopsis, Melaleuca, Alpinia, Clausena, Memecylon, and Ardisia constituted significantly to the sambar's available food (53.66%) and diet (57.60%). Comparing the preferred foods of grassland and forest, the forage available per hectare in the grasslands was far greater. Food preference rating values for the forest species showed that sambar preferred forest species, at least where grassland species were absent. Observa- tions, however, indicated that sambar utilized forest species heavily and during the period when the grasslands were dry. In Khao-Yai, the sambar's ideal habitat is a mixture of forest and grassland. Whether the sambar could survive on Imperata grasslands alone has not been tested. 75 Animal Effects on Forest Production Forest damage caused by rubbing and overbrowsing were greatest at the Nhong-Puck-Chee forest site (Table 14, Figure 16). The number of tree stems killed by animals, however, were similar on all sites. Killed stems caused by rubbing and overbrowsing were high on some sites but in general, the effects of animals in causing tree deaths were slight. Animals had but little effect on forest production on the study areas. Enderlein and Maxwell (1976) also reported no serious destruction of forest trees by wild animals at Khao-Yai. Park authorities and local people living near the park boundary, too, agreed with this finding. They reported further that wild animals are seldom seen outside the park. 76 .mumsunmm n m “nonsmomo D “Honouoo u o “umsms< n m mash u h “00:0Ixosmlmconz I U “moalcmcwmluooz I m “mcflxlmconz I £0 0 m m 0 N mN 0N m0 00 0 mm mm Nm 0N NN o 0 0 0 m N 00 0 v m N 00 00 00 00 m m 0 m m N 0 m m m 0 H 00 m0 NH OH 0 d m o o 4 h m o 0 ¢ o m a 0 d o pwaaflx Ummzounum>o pmnnsm mouam mEmum no 003652 .0000 00000000 I0000 mash .>0£uGOEHQ UmumasEdoom mmmnesz .xumm HMGOHumz 00» I003& .mmuflw ummHOM E oov 030 so HMQEMm an Umuommmm mEmum wo 0003552 .00 manme N 77 .0000 00000000 Imhma mash .on mmnolxosmImconz can .Anv moaInmcwmIHooz .Amv mc0MImconz um mEmum 000» ummMOM co muommmm HMEHcd +2.20: .00 000000 q O l O .— l l 8 8 ( (W 003 ) cams" sums :IO 'ou mmnwnaav O? 93.. .x .awmgouncw>0 Oman—z. RECOMMENDATIONS Procedures for the management of the sambar should vary on areas devoted to different forms of land use: National Parks. With the primary objective of maintaining natural conditions, it is suggested that: (1) Several exclosers be built, each perhaps 0.2 to 0.5 ha in area. These would exclude all large grazing animals so that comparisons could be made between grazed and ungrazed vegetation. Such exclosures would enable early detection of range overuse. Care should be taken to measure the effects of rodents and other small herbi- vores that might be attracted to the protected vegetation. A portion of the area could exclude all mammals including small one, while major part could exclude the ungulate. Exclosures should be sited in hopes of avoiding damage by elephants and should be protected by firebreaks against the danger of protected vegetation being burned. (2) Population counts of the major grazing animals should be made at least annually. Censuses should be attempted if suitable methods are available 78 79 but at least roadside counts should be made under stan- dardized conditions to establish indices to ungulate abundance. (3) Long-term studies of sambar population char- acteristics, behavior, movements, reproductive patterns, and relations to predators, diseases, and competitors are often best accomplished in national parks. The main- tenance of research program in each national park is essential to receiving full benefits from it. (4) Studies of burning practices should be designed to reveal their effects on the natural flora and fauna. Fire and fire control both have effects on nature preservation which must be evaluated. (5) As many forest species provide fruits as foods for sambar, the importance of these species as related to the deer should be studied. (6) Since extremely high deer density figures were derived from pellet-group counts, it is recommended that this type of census should be replicated and reviewed. The basic procedures should be tested and results confirmed. After-midnight deer densities also should be investigated by other means. (7) The area from which sambar are attracted to grassland openings should be determined, using telemetry or other marking methods. 80 (8) Studies should be attempted to determine the extent to which sambar require forest cover or other woody vegetation. Ranching Areas. Sambar were found to feed largely on Imperata grass and to prefer this to many other available species. Imperata (ya-kar) is considered to be a troublesome weed in many areas of Asia and Africa. Because of this fact, it seems reasonable to undertake further studies to test whether the sambar may be more suitable for meat-production than domestic stock, espe- cially on unimproved rangelands. (9) It is recommended that research be undertaken on Imperata grasslands to determine how those lands should be managed for maximum useful forage production and how sambar react to being tamed or domesticated on such sites. (10) Since it was the new growth prevailing for 45-60 days after burning which sambar ate most during this study, the effects of mowing and controlled burning in stimulating new production should be determined. The frequency with which burning or mowing should be repeated needs to be ascertained. SUMMARY Sambar were studied between March 1976 and March 1977 at Khao-Yai National Park, Thailand. Three grass- land and three adjacent forest sites were selected for investigation. Study objectives were: (1) to calculate food preference ratings, (2) to apply these ratings in an appraisal of range condition and trend, (3) to appraise the effects of wild animals on forest production, (4) to learn as much as possible concerning sambar population status, and (5) to develop management policies to insure survival of sambar in the park and on other areas having suitable habitats. In light of the research findings in this study, it is recommended that sambar may be more productive than domestic livestock as producers of meat on unimproved pasturelands. There were 46, 46, and 42 plant species identified on the grasslands at Nhong-King, Moor-Singh-Toe, and Nhong-Puck-Chee respectively, yet Imperata cylindrica dominated these areas covering 50% of the ground area at Nhong-King, 41% at Moor-Singh-Toe, and 37% at Nhong-Puck- Chee. The similarity index values indicated only minor 81 82 differences between the vegetation on the three grassland areas and these differences do not seem to be correlated with sambar grazing intensities. Among 72 plant species found on the grasslands, only 25 were eaten by sambar. Their percentage availa- bilities were 79% grasses and sedges, 17% forbs, 3% shrubs, and 0.5% trees. Corresponding percentages of food con- sumed were 96%, 2%, 0.6%, and 0.7%. Of the 25 species eaten, Paspalum, Wrightia, Alpinia, Neyraudia, Vernonia, Imperata, and Cratoxylon were preferred foods. Paspalum was utilized up to 81%. Forty-two percent to 58% of available forage was eaten in the other preferred plants; 18 species were eaten to a lesser degree than their abun- dance would indicate. The remaining 47 species were not eaten by sambar. Imperata was the most abundant plant and most important food in the diet of the sambar, forming 67% of available forage and 88% of all food consumed. Imperata, Neyraudia, Ischaemum, Carex, and Vernonia were the impor- tant sambar foods on the grasslands, forming 96% of the sambar's diet. Forage production and utilization by sambar on study plots during the dry period indicated that the production of grass forages increased up to about 45 days after burning and then gradually decreased. Forage removal from the plots ranged from 71% to 88%, though 83 this heavy use doubtless was because this was the only area of green grass available; sambar heavily utilized_ all grasslands. The species composition, density, and basal area of trees over 1.5 cm d.b.h. in all forest sites were similar. The stem numbers ranged from 2965 to 4596 per ha and with basal areas between 27.1 to 31.5 mZ/ha. Common tree species were: Aquilaria crassna, Aphanamyxis polystachya, Cinnamomum iners, Evodia gracilis, Eugenia siamensis, and Gonocaryum lobbianum. Crown cover was 85% to 90% at all sites. The species composition and the number of browsed and unbrowsed stems of forest ground species were similar all on sites. The percentage of browsed stems was small compared to the total stems available. Animals did not significantly affect forest production. Among 42 plant species in the forest habitat, 21 were preferred foods. Seven were highly preferred, with Carallia and Knema being eaten 3 times as much as their abundance would indicate. Lithocarpus, Neolitsea, and Ficus had food preference ratings over 2.00. These highly preferred foods, which constituted only 5.5% of the forage available in the forest, comprised 14% of the sambar's diet, 57% to 82% of forages with preference ratings over 2.00. All 14 preferred species formed 30% of the food available and 48% of the diet. Lasianthus, 84 Castanopsis, Melaleuca, and Eugenia were most important in terms of bulk contribution of sambar forest foods. The 26 neglected species formed 63% of the food available and only 39% of the diet. Daily food consumption by a captive 3-year-old female sambar (118.5 kg) was 1,084.21 gm dry-weight. Energy needed for maintenance for 12 days period was 29,694 kcal or 55.48% of the total energy consumed. Daily water intake was about 1.4 liters. The captive sambar spent more time in resting, ruminating, and walking than it did feeding and standing. Feeding occurred in the early morning, late evening, and also at night. A 40% greater food intake was assumed for the wild active animal, so that daily consumption was estimated as 1,517.89 gm. Animal effects on forest production were heavy at Nhong-Puck-Chee forest site where considerable rubbing and overbrowsing occurred. The number of killed stems, however, was similar at all sites and animals were judged to have but unimportant total effects on forest production. The average population density (deer/ha) was 0.02 from direct daylight counts, 0.11 from spotlight counts, 3.68 from pellet-group counts, and 3.15 from a food-removal census. Population densities of up to 995 per km2 were calculated for certain seasons from pellet 85 group counts. Even the lowest densities were for 80 deer per km2. These high concentration levels remain to be confirmed by later studies; yet by any standards, sambar densities on the grasslands must be very high. The population consisted of 44% adults, 40% sub- adults, and 16% fawns. The fawn to adult female ratio was 1:2.31, and there were 2.31 adult females per fawn. Recommendations are made for further studies in the national park to answer research questions still outstanding and to benefit prOper park management. In particular, however, it is suggested that the potential be investigated for ranching sambar in meat—production enterprises. The coarse Imperata cylindrica (ya-kar) grass comprises 88% of the sambar's diet and it prefers that species over other available forages. This grass is an abundant and vigorous invader of open fields. It seems likely that sambar might be tamed and raised on ya-kar at less cost and more efficiently than domestic livestock can be fed on improved pastures. APPENDIX 86 00.0 00.0 00.0 .00 000000000 0m.N N0.N 0o.o 000055 000mmoom mm.v mo.m mN.0 0N.o 0000000 00505000mm 00.0 00.0 Nv.N mm.o .mm 000000 00.0 00.0 00.0 .00 00000004 00.00 00.0 00.0 0000000000 000000000 00.00 N0.0 00.N0 om.N 25000050 E500oumm5m 00.0m 00.Nm 00.m 000002 "0&000 0N.mm 000500 U50 0000000 00009 0N.o 0N.o 0o.o 0500050000 5omom000050 0N.o 0N.o 0o.o 8500005 5500500 mn.o 0m.o NH.o .mm 0500520 0o.0 0N.o om.o 50000000 5opo5>0 00.0 00.0 00.0 00.0 0000000000 00000000 00.0 0N.0 mv.o 00050500 0000000005 00.0 0m.0 mN.o 0500000 m5m000m 0o.N 00.0 mm.o 0000555000 00500000000 Nm.m mo.o oa.m 0m.o 000550 x0000 0N.0 0m.m no.o 05005505000 005500002 mo.o0 0m.o oo.0 00.N 5500055 55500500H 0o.0N 0m.m 00.00 00.0 00000500 00000 00.000 00.00 00.00 00.00 0000000000 00000000 "000500 550 0000000 0 w m 00500> x0c5H 00>oo 005055000 0000505 005000OQEH 0>00000m 0>Huma0m 0>00000m 00000mm U+m+< n O U m < .0000 0000 .0000 00500002 Hmwlomnm .550000000 @502Im5052 050 50 0000050 05005 000 005Hm> x0050 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cam mwmmmum Hmuoe mw.o h~.o ~«.o Eduwuoc Esowcmm mH.H mm.o m~.o mmoasvcmam manomuonoo mm.H hm.o mo.a mflam>wummm mflahumwunfifim Hm.¢ nm.m nm.o nN.H mSHMHDOHom comom0m>uno Ha.n mm.o mm.m vm.~ Mudflusmammu macdmuhmz nm.n Nh.v mm.H mo.a mwmcmmmo mflumoummum mo.m mo.~ Hm.v N¢.o moaned xmumo om.mm om.m mo.ma HN.HH mumaosuo xmumu mo.mm ~v.ma om.m mm.ma Esoflune EdammsomH mm.~aa om.H¢ mm.ma mp.hm mofiuccflawo mumummeH ”mmmcmm cam mmmmmuw w w w manam> waGH um>ou mocmskum muflmama mocmguomEH m>Humamm m>wumHmm m>fiumamm mmwommm U+m+4 n D U m < .mnma mash .xumm Hmcowumz Hmwlomnx .ccmHmmmnm moelsmcwmuuooz may no mmfiommm #:mam How wwsam> xmvcfl mocmunomafl mdam mmomucmoumm Hw>oo cam .mocmdwmum .wuflmcmw w>Humamm .oH magma 90 mm.o madnnm Hmuoe mm.o >~.o Ho.o .mm muoxH m~.o >~.o Ho.o mmwflouoamsmmo EdvaEmma m~.o b~.o Ho.o .mm mscmvcmm mm.o hm.o mo.o Edowunumnmame mfioummamz vo.a no.0 mm.o vo.o mmduno mmumuofiamm ma.h mm.o vm.m mo.a EdumasuwuHMfln EdadOEmmo "mnaunm vo.vm mnHOM Hmuoa mm.o h~.o ao.o 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h~.o Ho.o .mm mwamkum m~.o h~.o Ho.o asumoomn ssflmmm mm.o Hm.o h~.o Ho.o asmosuom coamxoumuu hm.o om.o na.o «mamoxm mflmcaua4 "mmwna w w w mmsam> mecH um>ou wocmsvmum muflmcmo mocmuuomEH m>Humamm m>HumHmm m>aumamm mmfiommm U+m+¢ .I. a U m d vmscfiucoo .ma manme 92 Ho.o oo.o m¢.o mo.o «HHoMflmmc «madam ~H.H no.H mo.o mfloasw mwnmmoom mm.a wm.H mo.o mmoowusummnm momwmmsb mm.a mo.o mm.a mm.o mofiumflaao masocum> mo.m mm.~ o~.o .mm nowasunom -.¢ ~m.m om.o .mm asucmflc¢ mm.m mm.h om.~ muflmwuvmnv momasuuom mm.ma >¢.o vm.aa nm.a Esumuoco Edfluoummsm m~.H~ om.m mo.HH mm.m «Hameoo mmguamaflmm mm.om mm.mm o~.m mmmmoz "mnuom ma.ov mmmwmm can mmmmmum Hmuoa hv.m mo.o mm.h on.H moavcfl xmumu m~.o «~.o mo.o msumufluflc msuomao m~.o v~.o mo.o mmsccmam mwaomuoamou mm.o v~.o mo.o mwam>wummm mflamumfiunefim m¢.o mv.o mo.o .mm manhoN oa.m om.H om.o .mm mnummxu Hm.m mm.v m~.H Esoauzfi EdfimmnomH m~.b ~m.m mv.H mcmwvnmcaou wwwnmummz oo.m mm.~ ma.m mammouo mnmufiom nv.ma mH.m mm.~ mm.m wumwosuo xmnmv mm.o~a mm.nm mh.aa oo.Hn moflucaflamo mumummaH "mmmcmm can mmmmmuu m w w mmsam> mecH Hm>oo mocmnvmum muflmcmo mocmuuomEH m>Humamm m>Humamm m>fiumamm mwflommm o+m+¢ u o u m 4 .whma mash .xumm Hmcoaumz Hmeomnx .wcmammmum manIxosmImaonz mng co mmflommm pamHm How mwsam> xmvcfl mocmuuomfifl mfiam mmmmucmouwm Hm>oo vcm .xocmsvwnm .wuflmcmc m>aumamm .ha magma 93 Hm.0 0v.0 m0.0 amounmfiou maunmfiuz mo.a no.H No.0 .mm mflamoflum mm.a Hm.H No.0 magmaaflxm mMflvcommoumono "mmmua mnzunm Hmuoa m~.0 v~.0 H0.0 mmvwouoamnmmo Edfivosmma 00.0 m0.0 m0.0 Edowunumnmamfi mfioummamz ~0.v mm.m 00.0 mumHsofiuumwn EdwUOEmmo ”manusm mau0m Hmuoa m~.0 v~.0 H0.0 mmummao momasunom mN.0 vm.0 H0.0 mmsuno mmumuoflamm mm.0 vm.0 H0.0 MflHOMfino:0m mwaflEm m~.0 vm.0 H0.0 maamfiwm mafimuuwz mm.0 ¢~.0 a0.0 mmwflocoEmUm mmuoomoflo o~.o ¢~.o no.0 .mm mflcwmad m~.o v~.o No.0 asofiumom asflmcflwum mm.0 vm.0 m0.0 MAHOmwomumwa mmuaunomum a~.o v~.o mo.o moflncfl mumnmumuasomum s¢.o o¢.o Ho.o .mm mfluoaomm mv.0 00.0 «0.0 mmUHoncoo Edumummd mm.0 0v.0 00.0 .mm mmoEomH @mscHuGOU "mQHom m m m mmSHm> xwvcH Hm>ou zocmdvmnm mufimcmn mocmuuomEH m>fiumamm m>flumHmm w>Humamm mmflommm U+m+< u D U m .a vmscflucou .na magma 94 mwmommm HHW mo mammmm>flvcwimo Hm>ooIkuom .onma .xoo ommm 00H x x mmflommm mo mamocw>flwcfl mo Hm>ou mmmommm Ham mo mmsam> >ocmsvmum mo Edm u U 00H x 00H x mmwommm HHN m0 Hmswfi>mmmw no “mass: Hafimm x mmflommm mo mamsnw>flocfl mo Hmnfisz x mmfiommwlwocmDWmum u m bbhboa 00.00H oo.ooa mmflommm unmam Ham Hmuos m0.0 mmmuu Hmuoe mm.0 vm.0 H0.0 maahnmouome msoflnflm mm.0 vm.0 H0.0 ESOflccH Edawxono mv.0 0v.0 No.0 mflamucmfluo mEmHB cmSCflucou "mmmue w w w wwaam> xmvcH um>ou mucwswmum muflmcmo mocmuuomEH m>aumamm m>flumem m>Humamm mmflommm U+m+¢ n Q o m m wmscflucou .FH manme 95 N N H Hv ONHSHQOWHDHNHHHHODMHHMOHIDMI‘NH (OH H r-l r-i N HHN HN H mm FIqu LDHMNv-l HNHV‘V‘LO Nr-I ... 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S'TV'9'68 S'6C-9'LE S'LE-9'SE S'IE-9'62 S'62-9'LZ S'LZ‘Q'SZ S'SZ-9'EZ S'EZ-9'IZ S'IZ-Q'GI S'6T-9'LT S'LT-9'ST S'ST-9'ET S'ET-9'II S'TI-9'6 -9°L “9'9 -9’8 -S°T 5'6 S'L 9'9 9’8 mmHommm A500 mommeo HmumEMHo 1 .ohmH mash .xumm HacOHumz Hmwuomsm .mUHm ummu0m moaIsmchIuooz .mmmmmHo umumfimHv an .mmHommm uGMHm mo noun Human can huHmcwo .mH mHnma 98 6V0'LZ 9OS'I OZV°T EBZ'T fisI'T 908’0 VOL'O 609'0 T9S'I 9SL'I GZL'O 0I8°T VT6°T Z99°T I88'Z E9€°I LEE'I BSL'T L68'T 668'0 Hmn\~60 mmum Hummm mmmv HH HH HH NHv H H H MNkDLnHNo-{I-IN mt-iNl‘ HH NN HH N H HH mm «v mm 50 mm 00H mmm m m HN mmH VH mmw HN NNm 0v hmHH mOH Fit-{MN mmmH me bra HNfl‘l‘r-lc-l H .mn umm mamum mo .oz me 000 CH mEoum mo .02 .mm mHonmwnom mammmHmE mHumeoumEmHum .mm mHnuHmaHom mHHHomum vao>m "mndunm mmoucmfiou mHuanuz MHHMEOEos MHHMUGD mHHOMH>Hmm chHmHsa HHnOHHHm3 mEHnum .Qm macsum muMHowocmH unmonm MOHum>Ham mummHmndz mwomucwemu wmmmz Edum>o cloomemz UmscHucoo ”mmmua IPQOL S'Ev-9'Iv S’IV—S'GE 9°6£-9'L£ S'LE-9'SE S'TE-9'6Z S'6Z-9'LZ S'LZ-9'SZ S'SZ-9'EZ S'EZ-9'TZ S'TZ-9'6I S'6I-9'LT S'LT-9'SI S'ST-9'EI S'ET-9'TI S'TI-9'6 5'6 ‘9'L S'L -9°S 9'9 '9'8 S'E ‘S'I HEOV mmmwMHo uwquMHo mmHoomm omscHucou .mH oHnms 99 Inl‘HNHI-INHHHNN OHM oomo MIDI-4 HmHNNmHMI—Ilfi N V‘ N HWCD VLDH HMN HI" H OHM Ho-Il HH V'LOOWHV‘ Fir-l aunm msmhmoocmmo manucmHmmq dendn mOEmma mmomunmno mHHMman Hanmaumn mauuonmuud "unaunm mmoucwsou MHusmHuz .mm MHHM>D HH£0HHHM3 wEHnom HHuwx MHcon mmmonEH mooonEhm MCHHDMH mOUOHmE>m MOHGMHamN mwmuHHomz Edum>o conomewz muMHsOHucmn mmcmumom: mduMHQOHcmm msuoHHmz .mm mHEmmUouon mHmcmfidccm msmumoo:UHH mcHHSMH «Ewan munnmum MHnHonmom mammoEHOm MHOHHmm EdCMHnnoH Eshumoocow mmo>uwc muchcouHo mHmaoEmHm MHcomsm mzumsnou monumoomMHm .mm Mchmsm mHHHomum msmumooumumHo mcoou MHmucmo mHmcwEmHm ESEOEMGQHU mumcH EdeosmccHu manomumeom waaemcmnmd "mmmua Iago; S'LV-9'SV 5'99-9’Eb S'LE-9'SE S'SE-9‘EE S’EE-9'IE S'6Z‘9'LZ S'LZ-9'SZ S'SZ-9'EZ S'EZ-9'TZ S'IZ-9'6I S'6T-9'LT S'LT-9'SI S'SI-9'ET S'EI-9'TI S'TI-9'6 '9'L 9'6 '9'5 S'L '9'8 9'9 -S'I S'E H800 mmmmMHU HGUGEMHQ mMHowmm .on0H mean .xumm HmcoHumz Hmwuomnx .0UHm umeOM manIxodmImconz .mmmmMHo HmumEMHU >2 .meommm ucmHm mo mwnm Human can wuHmcma .0N mHnma Continued Table 20. Diameter classes (cm) Species IPQOL S'LV-9'SV S’SV-9'EV S'L€-9'S€ S'Si-9'ES S'EE-9'TE S'62-9’LZ S'LZ-9'SZ S’SZ-9'EZ S'EZ-9'TZ S'TZ-9'6I S'61-9'LI S'LT-9'SI S'ST-9'ET S'EI-9'TI S'TT-9'6 9'6 '9'L S'L -9'5 9'9 '9'8 9'8 -S'T Continued Lasianthus sp. Pinanga sp. Shrubs: 100 I-IInN Hmv-O Salacia prinoides l 1 267 l l 54 28 20 104 No. of stems in 900 m2 122 67 67 44 44 33 33 56 11 11 11 11 11 2965 67 311 222 100 1155 589 No. of stems per ha. VOL'LZ TL8'I VIL'T VST'T TEO'I STG'O EBS'E 988'1 T9S'T S9L°T 697'I 0T8'T Tfif'T LZO'Z 888°C VL8'0 VLZ'T LVO'T LS6'0 L9S'0 Basal area (mz/ha) 101 nuanmm 800.0 H0625 mgcmHm muanmm 500.0 Hm>o mucMHm am.mmm no.00H 00.00N >0.00 .mm mnemoamm 00.000 00.00 EsOHusquMHmE maoumMHmz 00.00H 00.000 00.000 mamumoocmmo manuGMHmMH v0.00H .mm MHoHumab mo.oom~ mm.mm0 Ho.oo~ .mm muoxH 00.00H mHHHmmmndm mHEmoowHu v0.000~ 00.000 00.00 mHHHomum MHUo>m 00.00H 00.000 MHHomesoH Maoomusm v0.0000 00.000 Estm>mn5m EdEOEMcGHU "mnaunm 00.000H 00.000 mHmummoamum mmusom 00.00HH muMHnomm mEHmumaxo 00.000H HHHmuonu mEoumocaHH H0.000 mmUHoaoEmum mmuoomOHa 00.000H mumMHann mwnoomoHa 00.0000 00.00HH mHHmsHOHmmo manucmuoHnu H0.000H .mm MHchHd 00.000H .mm mssucmod 00.00HH .mm memmmHud 00.0000 00.000 muumfihxo MHGHQH¢ H0.00H~ 00.00H .mm EducmHU< "mnuom H0.0000H 00.000 Eduwuo: EdOHcmm 00.000 00.000 mGMHsno xmumu 00.000H 00.000H MOHUCH xwumu "mmmvmm can mmmuw cwm3ouncb cmmzoum Umm3ounss 0mmzoum Hm: ummv Am: Hmmv mmHowmm .0hmH mash .xumm Hm20Humz Hmwnomnx .muHm ummnom mcHxImaonz .mEmum ucmHm 0mmsonncs 0cm 0mm3oum .HN GHQMB 102 nuanmm 200.0 HmmvCD muhflam musmem 500.0 uw>o muGMHm 00.000 00.000 00.000 «0000000 manna 00.000 00.000 00.000 00.00 .mm chwmsm 00.00H 00.000 00.00 mHmcwEMHm «Hammsm H0.000 00.00H EfiumHo EdHcmuoma 00.000H muMHsomun MHHHmumo 00.0000 EdHH>SQOCH EsHHwnm0Hmu 00.00H mmo>umc muchconHU 00.00H mHmamEMHm ESEOEMGGHU 00.0000H 00.00H0 mamommccsun EaHsucmo v0.000 00.000 00.00 mum>moxm mamszHo 00.000 00.000H 00.000 mEHmmHumcHEdom mHmmocmummo 00.000H H0.000 00.000 H0.H00~ msmuou mnEmHmu 00.000H 00.0000 mumcH EsfiosmccHU 00.00 msHuouomu anMHoouumHoc¢ 00.000H 00.000 mammmuo MHHMHHsvd 00.00H 00.00H 00.00H manumummHom memsmcmnma HOA000 H0.00~ mumuwo MHMHO¢ 00.000 00.00 mumHSOHumu muon< 00.000 H0.000 00.000 00.00 mcwommuonum MHmHvud «0.000 00.000 00.000 mmHmoxm MHmcHuH¢ "mmmue 00.000 .Qm mnmwmmnom 00.000H 00.000 mcHusmH mooonshm 00.00H . .mm mEoumMHmz H0.000 00.00H MHMH:OHQMQ mmmuuaz vmscHucoo "mnsunm Umm3ounca 0mm3oum 0mm3ounco 0mm30um Hm: mev Am: me0 mmHoQO UmssHucou .Hm mHnma 103 .n.n.v Eu 0.H “was: mEmum was uanmn E 0.0 Hmong mcmEHommm n .£.Q.v EU 0.H Hm>o mEmum can uanms E 0.0 Hm>o mamEHommmm 0H.00 H0.0H 00.00 00.0H mmmucwoumm 00.0H000 00.00HOH 0.00v0H 0v.000m mfimum Hmuoa 00.000 .mm MHHM>D 00.000 MHHMEOEon MHHMUGD 00.000 mHmammon mccmuma «0.0000 .mm mumomuums 00.000 0H00H>Hmm MHaHmume H0.000 MUHcOQMn MHEmouumcumB 00.00H MHMHomosmH mnmonm 00.00H 00.00 HH:0HHHM3 MEHsom v0.000 H0.000 mdumoHunfiH msmumoooom 00.000 MUHsmHmmN mmmuHHomz» 00.00H MHMHaoHGMQ maHOHHmz 00.000 00.00 Edum>o cloumEmz 00.00 00Hum>H>m mumMHmcmz 00.00Hv 00.000H ESMMHHmusmm noanmeS H0.000 00.000 Edammuummdv Edeummonmoq 00.00H mumMHnwm ammuHH 00.000H mumHHmoHuum> ammuHH H0.000 00.000 mSGMHHmmvou msmumoonUHH H0.000 00.000 mHmcmEmcsm msmumoosuHH 00.000 msHHOMHummHmosm msmumoonUHH 00.00H 00.000 ESGMHAQOH Esmumoocow 00.000 00.000 00.00 mccanuOHm mscmeum UmscHucoo "mmmue 0mm3ounca 0mmzonm 0mm30unc3 wmmsoum Am: me0 Hm: Hmmv nusmHmm 500.0 mugmem 500.0 meommm Hmvca mMCMHm um>o mucmHm UmscHucou .Hm mHnma 104 H0.000 mumuowo MHMHmd 00.00H mmHmoxm MHmQHUHd 00.00H mazomummHom meaamcmnmd 00.000H 00.00 HHccmaumn mhuuonmuud H0.000 00.00HH mammmuo MHHMHHDU¢ «0.0000 00.00H H0.000 00.000 mcmummuonum MHmeud ”mmmua H0.00H0 00.000H .mm masmccmm 00.00H mdmnmoocmmu manuGM0mMH 00.00H .mm manu5m0mmq «H.0000 00.000 .mm MHoxH 00.00HH 00.000 00.00 mHHHomum mfluo>m 00.00H 00.00 .mm moEmmo 00.0000 5505m>mnnm EfiEOEmccflo "mnaunm 00.000 MHmuomocmum mmusom H0.00H0 HHHmHozu mfioumocHH «0.000 «0.00H mmUHososmum mmuoomoHa 00.0000 muwMHann mmuoomoHn 00.000H mHHmcHOHmmo manusmHoHsu 00.000H .mm Educmflvfi 0H.0000 00.00H mnuhexxo MHchH< "mnuom H0.0000 H0.000 Edumuos ESOHcmm _ 00.00H MHMHUSHU xmnmu 00.000 mowucH xmumo «mmmvmm 05m mmmuw cmmBOHQCD @mm3oum Umm3onnca mezohm Am: Hmmv Hm: Hmmv nuanmm 500.0 muanmm 500.0 mmHommm HmUCD mucmHm Hm>o mucmHm .000H wasw .xumm HmsoHumz Hmwuomsm .muHm ammuom moeuzmchunooz .m5mpm ucMHm ummzouncs cam 0mmzoum .NN mHnma 105 .n.n.© EU 0.H Amos: mamum 05m unmflmn E 0.0 Hmwca mcmfiflommmn .£.n.v Eu 0.H Hm>o mEoum 05m uanmn E 0.0 Hm>o mamEHommmm H0.00 00.0 00.00 H0.0H mwmuawouom 00.H0000 H0.0000 00.0000 00.000 mfimum Hmuoa 00.00H meomw>Hmm mwcwmnna 00.000 .00 mummmwsom 00.000H 00.000 00.00H m5055MH mooon500 00.000 00.00H «0.00H muMHomOGMH mnmonm H0.000 00.000 mowum>Hmm mumMHmamz 00.00H Edum>o 50H0005mz 00.0000 mummflnmm mmmuHH 00.0000 00.000 msGMHHmmcon msmnmoonuflq 00.000 mumHHmowuHm> mmmuHH 00.00HH 00.00 mHmcoEmccm msmumoonuHH 00.0000 00.00 MHHOMHumeHmonm mamumoonuHH H0.000 00.000 00.00 50:0HnnoH 5swumoocou 00.00H 05HHUMH mfimcx 00.000 mocanfluon mscmeHm 00.000 MHomecoH mfiooausm 00.000H 00.000 acacommn manna H0.000 00.00 00.00 .mm wflnmmsm 00.000H 00.00H 00.000 H0.000 mHmcmEMHm aflcmmsm 00.00H mHHHomum msmumooumuHo 00.000 00.00H mcmuou msEmHmu 00.00H H0.000 mumnfl EfiEOEmccHu 0H.0000 00.00H 00.000 MEHmmHumcflfidom mHmmocmummo cmscHuaou "mmmue 00030595: 0mmsoum 0mmsouncs cmmzoum Am: me0 Am: ummv nuanmm 500.0 musmHmm 500.0 mmHommm Hmcco muCMHm Hm>o mucmHm vmssHusou .00 anme 106 00.000 00.00H H0.000 H0.000 mHmcm5mHm «Hammsm H0.000 00.000 mHHflomum mdmumooumumHo 00.00H mcmuou mSEmHmu 00.000H 00.000 00.000 H0.000 mumcw EdEOEmccHU «0.00H manomuthom mehemcmnmd 00.00HH 00.000 00.00 HHccm5umn m05uonmuu< 00.000 00.00H H0.000 mumuoco mHmHmd "mmmus 00.00H 00.00 .mm «EcummHmz 00.00HH 00.00H 00.00 .mm mdnucmflmmq 00.000 00.00H 00.000 00.000 msmumoocmmo manucmfimma 00.000H H0.000 .mm muoxH H0.000 00.00H mHHHmmmndm mHEmoomHm 00.00H 00.000H mHHwomum mHuo>m 00.00H moandv moEme 00.00H0 H0.000 55H5m>mndm EUEoemccHu 00.000H 00.00 mmumuumno MHHmmmno "madnnm 00.0000 H0.000 mmUHocme maomHmm «0.000 mflmcmcHno EscomxHom 00.000 mmUHocoamum mmuoom00n 00.00H mHHMCHOHmmo manucmuoHsu 00.000 00.00HH mmOEEoo MHomo< 00.000 .mm msnucmod 00.0000H H0.00H0 .mm ESHGMHU¢ 00.0000H «0.000 muu050xo MHQHQH< "mnuom 00.000H 00.000 Esumuoc Edoficwm “mmmomm 05m mmmuu Ummzounco 0meOHm 0mmBOHQQD 0mm3oum Hm: ummv Am: ummv nuanmm 500.0 mmHummm Hmvco mucmHm muanmm 500.0 Hm>o muGMHm .000H $550 .xumm HMGOHumz Hmeomnx .muHm ummuom meUIxosmImcosz .mEmum uGMHm wmm3ouncs 65m cmm3oum .00 mHQMB 107 .n.n.© E0 0.H Hmwcs mEmum can unmflmn E 0.0 “muss mcmanwmm n .£.n.U E0 0.H Hw>o mEmum van #5000: E 0.0 nm>o mcmeommmm 00.00 H0.0H 00.00 00.0H mmmucmouwm 00.H0000 H0.0000 00.0000 H0.000H mEmum Hmuoe 00.00H0 00.000 H0.000 mCHHSMH mooonEhm 00.00 muMHsoHucmc mmqmumomHmz 00.000 00.00H mumHoncmm msuoHHmz 00.000H mumwanmm mmmpHH 00.000 00.00 mflmcmEmcsm mamumoonuwH 00.00H 00.000 m50HSMH mEmCM 00.000H 00.000 00.000 munanm mfluHmfimmuom 00.00H 00.000 mum>woxm mammDMHu 00.00H 00.00H mmo>umc mumficcouflw 00.000 H0.000 00.000 adamannoH 5:0umoocoo 00.00 .mm chmmsm wmsnfiusou “mmmue cmmzounca 0003000 UmmBOHnss 0mm3onm Hm: nmmv Hm: Hmmv munmflmm 500.0 mmflommm nunmflmm 500.0 Hm>o muGMHm Hmvas muGMHm cmscHucou .mm mHnme 108 Table 24. Length and dry weight of leafy twig, energy values of leafy vegetative species and dropping, Khao-Yai National Park, January 1976-March 1977. Avg. Avg. . Twig Dry Energy Forage 596C185 Length Wt. (kcal/gm) (cm) (gm) Paspalum conjugatum 3.5 0.0907 4.1201 Wrightia tomentosa 5.5 0.1511 4.4622 Alpinia sp. 7.0 0.0309 3.8716 Neyraudia reynaudiana 6.5 0.3803 3.9604 Vernonia elliptica 9.5 0.1737 4.0086 Imperata cylindrica 12.0 0.0958 4.2043 Crotoxylon formosum 5.5 0.1111 4.3561 Dioscorea stemonoides 3.5 0.0272 4.0936 Ischaemum muticum 3.5 0.0692 4.1243 Carex cruciata 14.5 0.1052 3.7984 Eragrostis capensis 9.5 0.0709 4.1594 Coelorachis glandulosa 8.5 0.2137 4.1156 Helicteres obtusa 7.5 0.1691 3.9507 Carex indica 5.5 0.1301 4.0490 Scirpus grossus 10.0 0.2015 3.8329 Spilanthes ocmella 8.0 0.0781 2.4962 Scoparia dulcis 4.0 0.0366 3.9450 Desmodium biarticulata 5.5 0.1371 4.2970 Vernonia parishii 3.5 0.0354 3.0588 Ipomoea sp. 4.5 0.0513 3.1489 Eupatorium odoratum 5.0 0.2477 4.4611 Costus speciosus 4.5 0.0234 4.0644 Bridelia sp. 5.0 0.0676 4.4202 Eryngium foetidum 8.0 0.0256 3.4543 Cyperus sp. 10.5 0.0276 3.7892 Carallia brachiata 2.5 0.1958 3.4935 Knema laurina 12.5 0.3340 4.0125 Lithocarpus rodgerianus 8.5 0.1985 1.9201 Neolitsea zeylanica 3.5 0.1224 4.1278 Eugenia sp. 2.0 0.1437 4.1834 Eugenia siamensis 2.5 0.6430 3.9744 Ficus sp. 12.5 0.3566 4.0105 Nephelium mutabile 3.0 0.2468 4.6963 Melaleuca seutellatum 5.0 0.0535 4.1387 Uvaria rufa 4.0 0.1398 3.4919 Hedyotis sp. 3.5 0.1424 4.3077 Lasianthus cyanocarpus 4.5 0.6008 3.1676 Lithocarpus eucalyptifolia 10.5 0.1951 1.8114 Castanopsis acuminatissima 9.5 0.1730 1.8849 Uncaria homomalla 5.5 0.1464 4.1055 Eugenia sp. 2.5 0.0511 4.2423 Murrya paniculata 2.0 0.1848 3.2861 109 Table 24. Continued Avg. Avg. . Twig Dry Energy Forage SpeCLes Length Wt. (kcal/gm) ( cm) (cm) Eugenia ripicola 2.5 0.1173 3.4555 Uvaria sp. 3.0 0.1473 4.2060 Podocarpus neriifolius 2.0 0.2141 4.0476 Linostoma thorelii 2.5 0.0354 3.5701 Rurea stenopetala 8.0 0.0618 3.9664 Curcuma parviflora 7.0 0.0309 3.8709 Styrax sp. 3.5 0.2731 4.1306 Dioscorea bulbifera 8.5 0.4116 4.0582 Cinnamomum subavinum 2.5 0.1222 4.6104 Litsea verticellata 2.0 0.2143 4.3154 Mangifera sylvatica 2.5 0.7115 4.0707 Ixora sp. 2.0 0.6430 3.8733 Ardisia arborescens 2.0 0.3916 4.0125 Desmos sp. 2.0 0.1547 4.2316 Gonocaryum lobbianum 5.0 0.2976 4.6425 Evodia gracilis 6.5 0.3217 4.2970 Clausena excavata 3.0 0.4634 4.0828 Symplocos laurina 4.5 0.1101 3.9280 Jasminum sp. 4.0 0.1685 3.8970 Camellia oleifera 2.5 0.1008 3.8466 Memecylon ovatum 3.0 0.1848 4.3624 Adiantum sp. 7.0 0.0983 1.9439 Uncaria sp. 6.5 0.1575 3.9857 Cinnamomum iners 3.5 0.1977 4.1420 Fraxinus floribunda 2.0 0.1013 4.3719 Aglaia odorata 3.5 0.1766 4.1311 Artabotrys harmandii 5.5 0.1543 3.8719 Litsea sebifrea 2.5 0.4108 4.1852 Alpinia oxymytra 10.5 1.5404 3.8382 Dropping (ave.) - - 4.2016 110 Table 25. Moisture percentages and weights of fresh fecal pellets, dry at 100° C for 24 hours, Khao-Yai National Park, January 12-24, 1977. Wet Weight Dry Weight Weight Loss Percentage sample (gm) (gm) (gm) Loss 1 160.0 60.40 99.60 62.25 2 90.0 35.66 54.34 60.38 3 210.0 73.38 136.62 65.06 4 130.0 76.33 55.67 41.29 5 130.0 61.67 68.33 52.56 6 150.0 59.78 90.22 60.15 7 230.0 78.79 151.21 65.73 8 225.0 87.38 137.62 61.14 9 290.0 118.84 171.16 59.02 10 200.0 82.49 117.51 58.76 Total 1815.0 734.72 1080.23 Average 181.5 73.47 108.02 59.52 LITERATURE CITED LITERATURE CITED Banijbatana, D. 1966. Management of forest lands for recreation. Royal For. Dept., Bangkok, Thailand. 9 pp. . 1967. The forest problems and forest programs in Thailand. Royal For. Dept., Bangkok, Thailand. 31 pp. Bell, H. M. 1973. Rangeland management for livestock pro- duction. Univ. Oklahoma Press. 303 pp. Bennett, L. T., P. F. English and R. McCain. 1940. The study of deer populations by use of pellet-group counts. J. Wildl. 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