Solar Radiation Determines Site Occupancy of Coexisting Tropical and Temperate Deer Species Introduced to New Zealand Forests

Assemblages of introduced taxa provide an opportunity to understand how abiotic and biotic factors shape habitat use by coexisting species. We tested hypotheses about habitat selection by two deer species recently introduced to New Zealand’s temperate rainforests. We hypothesised that, due to different thermoregulatory abilities, rusa deer (Cervus timorensis; a tropical species) would prefer warmer locations in winter than red deer (Cervus elaphus scoticus; a temperate species). Since adult male rusa deer are aggressive in winter (the rut), we also hypothesised that rusa deer and red deer would not use the same winter locations. Finally, we hypothesised that in summer both species would prefer locations with fertile soils that supported more plant species preferred as food. We used a 250 × 250 m grid of 25 remote cameras to collect images in a 100-ha montane study area over two winters and summers. Plant composition, solar radiation, and soil fertility were also determined for each camera location. Multiseason occupancy models revealed that direct solar radiation was the best predictor of occupancy and detection probabilities for rusa deer in winter. Multistate, multiseason occupancy models provided strong evidence that the detection probability of adult male rusa deer was greater in winter and when other rusa deer were present at a location. Red deer mostly vacated the study area in winter. For the one season that had sufficient camera images of both species (summer 2011) to allow two-species occupancy models to be fitted, the detection probability of rusa deer also increased with solar radiation. Detection probability also varied with plant composition for both deer species. We conclude that habitat use by coexisting tropical and temperate deer species in New Zealand likely depends on the interplay between the thermoregulatory and behavioural traits of the deer and the abiotic and biotic features of the habitat.     

A comparison of food habits of two sympatric ruminants of Mt. Yatsugatake,central Japan: sika deer and Japanese serow

Comparison of food habits of sympatric animals provides understanding of interspecific relations. Previous studies of food habits of the two ruminants of Japan, sika deer (Cervus nippon) and Japanese serows (Capricornis crispus), have shown that sika deer are variable, and they are gazers in northern Japan, but browsers in southern Japan, whereas Japanese serows are browsers. However, these studies described the food habits of each species separately, and no study has compared the food habits of these species living in sympatry. Therefore, we examined these species on Mt. Yatsugatake, central Japan, using microhistological analyses and nutritional analyses of feces. Sasa nipponica, a dwarf bamboo, predominated in the fecal composition of sika deer, whereas both S. nipponica and dicotyledonous plants were found in the feces of Japanese serows. Crude protein levels of serow feces were higher than those of deer. The particle sizes of plant fragments in serow feces were smaller than those in deer feces, suggesting that serow fed more selectively on digestible plants than deer did. These results support the suggestions of previous studies conducted in different habitats and show that sika deer are less selective grazers, and Japanese serows are more selective browsers. Thus, food differences are likely explained by the feeding ecophysiology of the animals and not habitat differences.     

Summer dietary compositions of sika deer on Yakushima Island,southern Japan

Summer food habits of Sika deer (Cervus nippon yakushimae) on Yakushima Island, southern Japan, were studied focusing in particular onPseudosasa owatarii, a dwarf bamboo native to the island. Fecal analyses showed that the food habit of the Sika deer on Yakushima island was plastic: the dietary composition was dominated by graminoids (ca. 90%), particularlyP. owatarii (50%–60%) in the bamboo grassland of theP. owatarii zone above the timber line, whereas leaves and non-leafy parts of woody plants were important in the crytomeria forest and evergreen broad-leaved forest below the timber line. The maximum culm age ofP. owatarii was 8.5 yr, but the majority of leaves (97.8%) were younger than 2.5 yr. Since the weight contribution of the leaves is great (27.7%) and the turnover rate is high, theP. owatarii grassland affords a good foraging place for the Sika deer on Yakushima Island.     

穆棱林区野生东北马鹿种群冬季食物组成模式与植物代谢产物的关系

高纬度地带,在冬季食物资源有限的环境中,野生大型有蹄类动物满足营养需求的同时,需要对植物中次生代谢产物进行平衡,回避有害物质并选择对机体有益的成分,从而形成特定的食物组成模式。以东北马鹿（Cervus elaphus xanthopygus）为研究对象,于2020年11月,在黑龙江穆棱东北红豆杉国家级自然保护区境内,采集东北马鹿粪便和植物样本。通过粪便显微分析法确定保护区内马鹿冬季食物组成,采用k-means聚类分析揭示马鹿冬季食物组成模式。应用广泛靶向代谢组技术对部分植物中次生代谢产物的含量进行全覆盖定性和相对定量检测,将食物组成与次生代谢产物数据整合,进行曼特尔检验（Mantel test）分析,以探究植物次生代谢产物对马鹿种群内食物组成模式的影响。结果表明,林区内马鹿种群冬季共采食30种植物,其中木本植物占92.48%;且种群内分别呈现出以东北红豆杉（Taxus cuspidata）,簇毛槭（Acer barbinerve）,毛榛子（Corylus mandshurica）为主要食物的不同食物组成模式。广泛靶向代谢组技术共检测出638种次生代谢产物,有25种代谢物与马鹿采食频率显著相关,其中多数萜类物质抑制马鹿采食,而鞣质类物质对马鹿的采食选择有一定的正向作用;Mantel test结果显示,上述25种物质中黄酮、鞣质、萜类化合物相对含量与不同马鹿个体食物组成显著相关,说明这些代谢物相对含量和性质的差异会造成种群内不同个体食物组成的差异,是种群内形成不同食物组成模式的原因之一。从植物次生代谢产物角度揭示了该地区东北马鹿种群冬季食物组成模式呈现差异的可能因素,为进一步研究大型有蹄类营养策略和植物化学防御关系提供基础依据。     

Use of mires and food habits of sika deer in the Oze Area,central Japan

Sika deer Cervus nippon entered the Oze Area, a snowy area in central Japan, in the 1990s. The use of mires and the food habits of this species were studied in 1999 and 2000. Deer used the mires immediately after snow melted and did not use the mires in mid summer, although they did use them lightly in autumn. The summer food habits differed from those of the deer population living in Nikko in the lower area: non-Sasa graminoids and other monocotyledonous leaves occupied more (approximately 30%) of the feces in the former population, whereas Sasa nipponica, a dwarf bamboo, was the dominant food (60%) of the latter population. Seasonal changes in fecal composition in the Oze deer were that mire graminoids occupied a considerable portion of their food in spring (11.6%) and summer (17.0%), whereas the amount of dwarf bamboo increased in fall (26.0%). Despite the small size of the mires, Oze deer appear to prefer mires to forests. Nitrogen concentrations of the mire plants did not differ from those of the forest plants; however, foraging efficiency would be greater in the mires and this may explain the preference observed. Despite the small total biomass, leaf biomass of the mire was equivalent to that of the adjacent forest floor. In addition, biomass was concentrated near the ground, or the biomass concentration was greater than that of the forest floor.     

Dry season diets of sympatric ungulates in lowland Nepal: competition and facilitation in alluvial tall grasslands

Based on microhistological analyses of faecal material, we compared the early dry season diets of greater one-horned rhinoceros Rhinoceros unicornis, swamp deer Cervus duvauceli and hog deer Axis porcinus, which inhabit the same alluvial grassland habitat complex in lowland Nepal. Their diets were quite similar, both at the forage category level and within subcategories of graminoids and woody plants. Early successional tall grasses, especially Saccharum spontaneum, were the dominant food of all three species, underlining the key role of the threatened alluvial floodplains in large mammal conservation in South Asia. The two deer species ate significantly more graminoids (>66.5%) than did rhino (45.5%), and although they did not differ in proportions of graminoids, swamp deer ate significantly more late successional tall grasses (Narenga porphyrocoma and Themeda spp.) and short grasses (mainly Imperata cylindrica) than hog deer. The two deer consumed almost equal proportions of woody browse (ca. 10%), significantly less than that of rhino (33.0%). The prediction of the Jarman–Bell hypothesis, that large-bodied herbivores are less selective and subsist on lower quality graminoids than smaller ruminants, was not supported by the data. Based on this and previous studies in the same area we propose a conceptual model where the larger megaherbivores (rhino and elephant Elephas maximus) facilitate the smaller swamp deer and hog deer during the monsoonal growing season, while the smaller and more selective deer species outcompete the larger during the dry season when food is more limited. Owing to the all-year sprouting ability of S. spontaneum, facilitation may occur also in the dry season with low numbers of megaherbivores, thus accentuating competitive exclusion at higher deer densities.     

Feeding ecology of mandrills (Mandrillus sphinx) in campo animal reserve,Cameroon

A field study on the ecology of mandrills (Mandrillus sphinx) was carried out for 28 months in Cameroon. Fresh food remnants and large quantities of fresh feces were collected by following the groups. Analyses of these products indicated that fruit (including seeds), monocotyledonous plant leaves and insects (especially ants and termites), were frequently eaten. Mandrills mostly ate the plant and animal foods in the lower forest stratum and on the ground. Fallen seeds and monocotyledonous plant leaves were eaten more frequently in the minor fruiting season than in the major fruiting season presumably to compensate for the shortage of fresh fruit during the former. Daily travel distances were shorter during the minor fruiting season than during the major fruiting season, because in the minor fruiting season mandrills forage for small food items, such as the new leaves and piths of monocotyledons and fallen seeds which are sparsely distributed on the ground, while in the major fruiting season they search for widely distributed food such as fruit. The daily pattern of group movement and a food intake experiment suggest that mandrills move and feed continuously throughout the day. Use of fallen seeds and monocotyledonous plant leaves appears to enable mandrills to maintain a terrestrial life in the tropical rain forest. The feeding and ranging characteristics of mandrills are basically similar to those of other baboon species in open land, though their environments differ extremely.     

Fate of heavy metals in vertical subsurface flow constructed wetlands treating secondary treated petroleum refinery wastewater in Kaduna,Nigeria

This study examined the performance of pilot-scale vertical subsurface flow constructed wetlands (VSF–CWs) planted with three indigenous plants, i.e. Typha latifolia, Cyperus alternifolius, and Cynodon dactylon, in removing heavy metals from secondary treated refinery wastewater under tropical conditions. The T. latifolia-planted VSF–CW had the best heavy metal removal performance, followed by the Cyperus alternifolius-planted VSF–CW and then the Cynodon dactylon-planted VSF–CW. The data indicated that Cu, Cr, Zn, Pb, Cd, and Fe were accumulated in the plants at all the three VSF–CWs. However, the accumulation of the heavy metals in the plants accounted for only a rather small fraction (0.09–16%) of the overall heavy metal removal by the wetlands. The plant roots accumulated the highest amount of heavy metals, followed by the leaves, and then the stem. Cr and Fe were mainly retained in the roots of T. latifolia, Cyperus alternifolius, and Cynodon dactylon (TF < 1), meaning that Cr and Fe were only partially transported to the leaves of these plants. This study showed that VSF–CWs planted with T. latifolia, Cyperus Alternifolius, and Cynodon dactylon can be used for the large-scale removal of heavy metals from secondary refinery wastewater.     

Preliminary Evidence of the Importance of ENSO in Modifying Food Availability for White‐tailed Deer in a Mexican Tropical Dry Forest1

The influence of El Niño/Southern Oscillation (ENSO) on rainfall and its possible effect on availability of food for white‐tailed deer (Odocoileus virginianus) in a tropical dry forest in the Pacific coast of Mexico was studied. From 1977 to 2003 there were three significant El Niño and La Niña events. During El Niño years rainfall decreased during the wet season ( June to October) and increased during the dry season (November to May), with the opposite effect during La Niña years. Plant diversity was monitored in permanent plots during the wet and dry seasons of 1989–1993. The results provide evidence that ENSO events affect deer food availability, particularly in the dry season.     

Food habits of Sika deer on Mt. Goyo,northern Honshu

The food habits of Sika deer (Cervus nippon) on Mt. Goyo, northern Honshu (the main island of Japan), were studied. Among 88 plant species found in surveys, 36 species (40.9%) were eaten by the deer. The percentage frequency of feeding (F) was highest for graminoids (59.6%), followed by browses (16.6%) and ferns (7.7%) and lowest for forbs (5.4%). Though various kinds of plants were eaten, only limited species formed the bulk of the deer dietary composition.Sasa nipponica, in particular, was the most important constituent, ranking highest inF value (80.3%) and being the dominant component of feces. Fecal analyses revealed that, 1) in areas lower than 900 m in altitude,Sasa nipponica accounted for between 20 and 60% of the content and that pasture grasses were also important, and 2)S. nipponica occupied more than 50% in montane areas higher than 900 m, and even more than 80% in winter. The utilization ofS. nipponica as a forage for Sika deer was discussed. This plant can be a key forage species because it meets all the necessary conditions: abundance and stability of supply, palatability, nutritive quality and tolerance to grazing.     

Diets of some French guianan primates: Food composition and food choices

We studied food choices of black spider monkeys (Ateles paniscus) and red howlers (Alouatta seniculus) in an undisturbed tropical forest of French Guiana for 6 months in the rainy season. We made additional observations on tufted capuchins (Cebus apella) and examined the differences and similarities in feeding behavior with respect to the plant specific composition of the habitat and the biochemical characteristics of their food. Capuchins and spider monkeys mainly fed on ripe fruit pulp, to which they added invertebrates (capuchins) or young leaves (spider monkeys); their plant diet was more varied than that of howlers, which contained approximately equal proportions of ripe fruits and young leaves of many species, with large monthly variations in these food categories. BothAteles andAlouatta tended to feed preferentially on abundant plant species, with a large overlap in their fruit choices. The former species included a high proportion of soluble sugars and a low proportion of protein in its diet compared, to howlers. Leaves of several species selected byAlouatta reacted positively when screened for alkaloids and phenolic compounds. Besides specific metabolic requirements and adaptations to deriving nutrients from distinct food types, we hypothesize that the ability to taste sugars, which varies among primates, affects the range of foods appearing to be palatable and, consequently, contributes to the differentiation of feeding niches.     

Food habits and home range use of Japanese macaques on an island inhabited by deer

Food habits and home range use of a troop of Japanese macaque Macaca fuscata Blyth on Kinkazan Island (9.6km²), northern Japan, were studied from the spring of 2000 to the winter of 2002. The home range of this troop covers the area where vegetation is modified by foraging of sika deer Cervus nippon Temminck. The core areas of the home ranges of the macaque troop corresponded closely to the distributions of the staple food plants in every season. For example, leaves of Berberis thunbergii DC. and leaves of Zelkova serrata Makino in spring, berries of Berchemia racemosa Sieb. & Zucc. in summer, nuts of Zelkova serrata and Carpinus spp. in the fall of 2000 and winter of 2001, seeds of Torreya nucifera Sieb. & Zucc. in the fall of 2001, and bark of Zanthoxylum piperitum DC. in winter of 2002. Among the staple food plants, Berberis thunbergii and Zanthoxylum piperitum are more abundant on Kinkazan Island than other macaque habitats in northern Japan because they are spiny and unpalatable to sika deer, hence survive under the heavy foraging by the deer. Further, another staple food plant, Berchemia racemosa, a liana, grows abundantly at the edges of Illicium forests. Illicium anisatur is toxic and is not consumed by sika deer. Monkeys of this troop using the unique vegetation induced by sika deer grazing, fed on several plants which are not consumed by monkeys in other areas, and the home range was heavily affected by the distributions of these plants.     

Seasonal change in browsing by sika deer on hinoki cypress trees on Mount Takahara,central Japan

Browsing by sika deer (Cervus nippon Temminck) on hinoki cypress (Chamaecyparis obtusa (Sieb. et Zucc.) Endl.) was monitored for approximately 3years on 300 trees at three sites on Mount Takahara, central Japan. We investigated the seasonal change in browsing on hinoki cypress in relation to the availability of other food plants (dominated by Sasa nipponica Makino et Shibata) and preferred food plants (deciduous broad leaves). Browsing occurred mostly from November to April, while no browsing damage occurred from June to October. Browsing was observed in November, even when the amount of other food plants did not decrease. Therefore, the cause of browsing cannot be fully explained by a decreased availability of other food plants. Comparison of fecal composition and food availability showed that deer had a preference for deciduous broad leaves. The availability of these preferred plants decreased in November and increased in May. Chemical analysis of the hinoki cypress and deciduous broad leaves (crude protein and neutral detergent fiber) showed that the quality of deciduous broad leaves decreased in November, and as a result the quality of hinoki became higher than that of the deciduous broad leaves. Browsing on hinoki cypress occurred when the availability of deciduous broad leaves decreased, and vice versa. These results suggest that the availability of preferred food plants influences browsing on hinoki cypress by sika deer.     

Understanding sitatunga (Tragelaphus spekii) habitats through diet analysis in Rushebeya‐Kanyabaha wetland,Uganda

Between January and December 2008, we assessed the diet and habitat selection of sitatunga, a highly endangered tropical wetland antelope threatened by habitat loss in Rushebeya‐Kanyabaha wetland. Microhistological faecal analysis and vegetative sampling methods were used to assess plant forms, species and seasonal changes in the sitatunga diet. Habitat use was indirectly determined by assessing feeding patterns, distribution of dung and trails within the wetland. Sitatunga fed mainly on herbs, sedges, grasses and shrubs. A total of 34 plant species were recorded as eaten by sitatunga. The most eaten plant species was Cyperus papyrus L. (22%). Malenthera scandens Schum. & Thonn., Polygonum senegalense Meisu (12%) and Polygonum pulchrum Blume (5%) were the most eaten herbs. Zea mays L. was the most eaten agricultural crop (58% of domestic crops), mainly during the wet season. Sitatunga feeding was mainly concentrated on the wetland edge habitat (46%) where most of its food (53%) was located. The other preferred habitat was the tall closed papyrus. We conclude that the long‐term survival of sitatunga requires a management plan focussing on the conservation of the most preferred plant species and habitats.     

Facilitative interactions between an exotic mammal and native and exotic plants: hog deer (<Emphasis Type="Italic">Axis porcinus</Emphasis>) as seed dispersers in south-eastern Australia

Endozoochory by exotic mammalian herbivores could modify vegetation composition by facilitating the dispersal and establishment of exotic and native plant species. We examined the potential for endozoochoric dispersal of native and exotic plants by exotic hog deer (Axis porcinus) in south-eastern Australia. We quantified the germinable seed content of hog deer faecal pellets collected in five vegetation types within a 10,500-ha study area that was representative of their Australian range. Twenty exotic and 22 native species germinated from hog deer faecal pellets and significantly more native species germinated compared to exotic species. Seedlings of the encroaching native shrub Acacia longifolia var. sophorae emerged, but no native trees emerged and the percentage of grasses that germinated was low (11%). The species composition of germinants was similar among the five vegetation types. We estimated that the hog deer population in our study area could potentially disperse >130,000 viable seeds daily. Our study shows how an exotic mammal can disperse seeds from both native and invasive plants and highlights the need for endozoochory to be considered more widely in studies assessing the impacts of exotic mammals on plant communities.     

贺兰山同域分布高山麝和阿拉善马鹿秋季食性的比较研究

对贺兰山同域分布的高山麝和阿拉善马鹿秋季的粪便进行采集,利用粪便显微分析法对二者秋季食性进行了研究。分别采集高山麝和阿拉善马鹿活动区域内粪样和植物样本,分别获得两种动物40个复合粪便样本,采用频率转换法对数据处理计算,得到贺兰山同域分布的高山麝和阿拉善马鹿秋季的食性组成及组成比例。结果显示,高山麝秋季共取食植物20科30种(属),其中蓝锭果忍冬(11.96%)、秦氏黄芪(10.17%)、密齿柳(8.32%)、折枝绣线菊(6.82%)、内蒙野丁香(6.18%)组成了高山麝的主要食物;阿拉善马鹿秋季共取食植物12科29种(属),其中针茅(22.15%)、灰榆(21.14%)、早熟禾(16.90%)、小叶金露梅(11.62%)、山杨(10.00%)为阿拉善马鹿秋季的主要食物。高山麝和阿拉善马鹿共有9种食物重叠,生态位重叠指数为83.75%。秋季高山麝取食的Shannon-Wiener指数、Pieou均匀性指数、食物生态位宽度指数和最大的物种多样性指数均高于阿拉善马鹿。     

Food habits of sika deer and nutritional value of sika deer diets in eastern Hokkaido, Japan

Rumen content analysis and field observations were used to investigate the food habits and diet quality of sika deer (Cervus nippon yesoensis Heude) from 1991 to 1993 in eastern Hokkaido, Japan. Diets varied seasonally, with deer consuming graminoids and browse in winter, forbs and agricultural crops in spring and summer and all of these plant foods in autumn. Eighty-four plant species with sika deer bite marks were identified and their use also varied seasonally. The diversity of food resources available provided both critical protein and digestible energy, allowing for physiological maintenance and seasonal growth. With these high-quality diets, deer maintained good body condition in eastern Hokkaido, where the population density was relatively low.     

Food and feeding ecology of the rock fowl Picatthartes oreas in Old Oyo National Park, Nigeria

The rock fowl is an endangered species and its food and feeding ecology was studied in the forest zone of Old Oyo National Park, Nigeria. The methods used were direct observation, and fragmentation analysis of faecal and crop content of samples collected during both the wet and dry seasons. The results showed that rock fowl feed on a wide range of species of insects, fruits and flower buds. In the wet season, animal resources represented 54 and 62% of the adult and chick diet, respectively, whereas in the dry season, animal food resources represented about 60% of both adult and the chick diets. There was a significant difference (P < 0.05) in the food composition of the birds between the dry and wet seasons. There was also a significant (P < 0.05) difference in food preference between adults and chicks. Food sharing was not common, although it was observed between rock fowl cockerels and hens. Food search and selection indices were directly related to season and food availability. It was concluded that food scarcity may not be a serious contributory factor in the threat to the rock fowl.     

Food habits of sika deer on the Boso Peninsula,central japan

The rumen contents of sika deer (Cervus nippon Temminck) on the Boso Peninsula, central Japan, were analyzed to identify local, sexual and age-specific differences in food habits. Graminoids and woody plants were the primary foods throughout the year. In winter, the use of evergreen broad leaves increased. The food habits of sika deer on Boso Peninsula were intermediate between those of populations inhabiting northern and southern Japan. Acorns, mainlyLithocarpus edulis Nakai, were consumed in fall and winter with a peak in October. Since the availability of acorns is not influenced by foraging in previous years, it can be regarded as a stable food supply and hence may be important for deer on the Boso Peninsula. The local difference between the Amatsukominato (AT) area, having a large plantation ofLithocarpus producing acrons, and the Kamogawa-Katsuura (KK) area, having a small plantation ofLithocarpus, was recognized; seeds and fruit were consumed more in AT than in KK in fall and winter. Males consumed more seeds and fruit than females at both sites in fall. This can be attributed to sexual differences in nutritional requirement.     

ELEPHANT ECOLOGY IN THE QUEEN ELIZABETH NATIONAL PARK, UGANDA

1. Aerial and ground counts in the Queen Elizabeth National Park from 1963 to 1969 indicate that the number of elephants has more than doubled. The increase appears to be the result of immigration rather than reproduction. 2. Elephant food habits were studied for 10 months in two areas of the Park. In the short-grass/thicket area, browse intake rose markedly whenever the rainfall fell below 50 mm/month. In the tall-grass area without thicket, herbs were eaten frequently during the rains and Cymbopogon grass bases in the dry season. 3. Food availability had an important influence on the diet of elephants. Sometimes, however, elephant were very selective, e.g. for Panicum maximum, Azima tetracantha, Securinega virosa, and Tribulus terrestris. Certain grass inflorescences were selected during the rains, and bases in the dry season. 4. Most browse was consumed in the thicket area during the dry season and most grass was eaten in the tall-grass area of the Park. 5. Stomach and faecal samples gave a similar result and indicated that the diet in the tall-grass area was comparable to that of elephant in Murchison Falls National Park. 6. Analyses of important plant species showed that, in general, tall-grass had a lower crude protein content than short-grass, herbs and browse. Browse leaves had a relatively high crude-protein content in the dry season. Differences were less marked in the rains. 7. Ether extract, or fat content, was high in browse leaves and in Cymbopogon, in particular its bases. These may have been selected in the dry season for this reason. 8. Grass and browse stems were very fibrous and this may have contributed to their being avoided. 9. High carbohydrate levels probably enhanced the palatability of grass bases and browse leaves. 10. Grasses were very siliceous, in particular their bases when contaminated with soil. Elephant took considerable trouble to remove soil. Browse leaves had little silica but were rich in other minerals, and this may have contributed to their greater palatability. 11. Studies of woodland dynamics show a decline in large trees which corresponds to the increase in elephants. In some cases trees have been almost eliminated. Damage to trees indicates that a most important factor in their decline is the elephant. There is some evidence that the trees are eliminated selectively. 12. In the tall-grass area there has been an increase of 46.3% in elephant numbers, but a decline of 80.3 % in buffalo, over the past 15 years. An overall decline in combined biomass of 36.9 % is calculated. Estimates of energy utilization of the annual vegetation production by elephant and buffalo indicates a decline from 6.1 % to 3.9% over this period.