How do differences in species and part consumption affect diet nutrient concentrations? A test with red colobus monkeys in Kibale National Park,Uganda

Within a primate species, diet can be highly variable in composition, even at small spatial scales within the same forest, or temporally, suggesting that primates use different plant species and parts to meet similar nutritional needs. To test whether such differences in the plant species and parts that primates eat affects the nutrient concentrations that they obtain, we observed feeding of seven groups of red colobus monkeys ( Procolobus rufomitratus) residing in Kibale National Park, Uganda. The different groups consumed mostly young leaves from many of the same plant species, but spent different amounts of time feeding on them. As protein and fibre are suggested to be important determinants of colobine food choice and abundance, we analysed multiple samples of 47 food species for protein and fibre. Despite the differences in the plant species and parts eaten, the protein and fibre concentrations for the seven red colobus groups were similar. Our results suggest that colobus monkeys eating diets with differing amounts of species and parts may ultimately receive similar concentrations of nutrients.     

Patterns of Frugivory in Three West African Colobine Monkeys

We studied the comparative feeding ecology of three species of colobus (Procolobus badius, Procolobus verus, and Colobus polykomos) on Tiwai Island, Sierra Leone. We collected dietary data on each species by scan-sampling habituated groups. Because these groups were observed in the same study area during overlapping time periods, the confounding effects of temporal and spatial variability in food availability were reduced. Our results show that the annual diets of the two larger species (Procolobus badius and Colobus polykomos) include roughly equal proportions of fruits (including seeds), young leaf parts, and mature leaf parts, although P. badius had a greater intake of floral parts. Procolobus verus consumed almost no mature leaf parts, few fruits and seeds, and many young leaf parts. Colobus polykomos commonly fed from lianas. Seeds were the dominant fruit item eaten by all three colobus, and the fruits they selected were generally dull and non-fleshy, in contrast to the brightly-colored, pulpy fruits eaten by guenons. Leguminous plants contributed substantially to the diets of both the larger species, but comparisons with other African forest sites indicate that colobine biomass is not closely correlated with the abundance of leguminous trees in the forest. (Deceased)     

Foraging challenges of red colobus monkeys: influence of nutrients and secondary compounds

The diet selection of two groups of red colobus monkeys (Procolobus badius) in Kibale National Park, Uganda are considered with respect to protein, fiber, digestibility, alkaloids, total phenolics, tannins, saponins, and cyanogenic glycosides. Both groups selected young leaves over mature leaves and young leaves had more protein, were more digestible, and had a higher protein to fiber ratio than mature leaves. Young and mature leaves did not differ with respect to secondary compounds. There were no differences in the phytochemical factors examined between frequently eaten foods and leaves that red colobus were never known to eat, but were relatively common in the environment. Regression analyses predicting foraging effort from the phytochemical components of the large group's diet revealed selection for only one factor, foods that are high in protein and low in fiber, when differences in food tree availability were taken into consideration. A similar analysis with the small group did not suggest selection or avoidance of foods with respect to any of the factors considered. Previous studies have found the biomass of folivorous primates to be related to the ratio of protein to fiber concentration of mature leaves in the environment. These investigations have considered variation in folivore biomass and forest composition among sites separated by hundreds of kilometers; however, large variation in folivore abundance occurs over much smaller spatial scales. In Kibale National Park the average protein to fiber ratio of the mature leaves of the 20 most abundant tree species predicted the biomass of red colobus among four neighboring sites. We examined the generality of this relationship by adding our biomass and leaf chemistry values to previously published values; 62% of the variance in colobine biomass was explained by variation in the protein to fiber ratios of mature leaves at the sites. There was no evidence that red colobus avoided plants with high levels of secondary compounds. In fact, one of the most preferred trees (Prunus africana) was the species with the highest levels of cyanogenic glycosides, and the highest saponin levels were found in the young leaves of Albizia grandibracteata, the sixth and fourth most preferred plant species for the large and small groups, respectively.     

Coping with Forest Fragmentation: The Primates of Kibale National Park,Uganda

A goal of conservation biology is to determine which types of species are most susceptible to habitat disturbance and which types of disturbed habitats can support particular species. We studied 20 forest fragments outside of Kibale National Park, Uganda, to address this question. At each patch, we determined the presence of primate species, tree species composition, patch size, and distance to nearest patch. We collected demographic, behavioral, and dietary data for Abyssinian black-and-white colobus (Colobus guereza). Black-and-white colobus and red-tailed guenons (Cercopithecus ascanius) were in almost all fragments; Pennant's red colobus (Procolobus pennantii) and chimpanzees (Pan troglodytes) were in some fragments; and blue monkeys (Cercopithecus mitis) and gray-cheeked mangabeys (Lophocebus albigena) were absent from all fragments. No species characteristics—home range, body size, group size, or degree of frugivory—predicted the ability of species to live in patches. No characteristics of patches—area, distance to the nearest patch, distance to Kibale, or number of food trees present—predicted the presence of a particular species in a patch, but distance to Kibale may have influenced presence of red colobus. Black-and-white colobus group size was significantly smaller in the forest patches than in the continuous forest of Kibale. For a group of black-and-white colobus in one patch, food plant species and home range size were very different from those of a group within Kibale. However, their activity budget and plant parts eaten were quite similar to those of the Kibale group. The lack of strong predictive variables as well as differences between other studies of fragmentation and ours caution against making generalizations about primate responses to fragmentation.     

Implications of small scale variation in ecological conditions for the diet and density of red colobus monkeys

In this 3-year investigation we documented patterns of density, diet, and activity of red colobus monkeys (Procolobus tephrosceles) in six areas in or near Kibale National Park, Uganda and related these patterns to availability of food resources. There were large differences in the density and behavior of the red colobus among the sites. For example, the red colobus at one site with a diverse plant community of more than 61 tree species, had a diet that included at least 42 species. In contrast, at a second site red colobus spent 92% of their feeding time eating from one species that dominated the tree community. The density of important red colobus food trees varied among sites from 32 trees/ha to 204 trees/ha, and red colobus density ranged from 0.70 groups/km² to 7.41 groups/km². Among sites, red colobus density was related to the cumulative DBH of important food trees, when one apparently anomalous site was excluded, and populations with more plant species in their diets tended to be those that were found at higher densities. Activity budgets of the red colobus populations varied markedly among sites. For example, feeding time ranged among sites from 29 to 55%, and traveling varied from 5 to 20%. When faced with increased foraging demands, red colobus reduced the time spent resting, while the time spent socializing remained fairly constant. Comparative socioecological studies typically contrast species separated by large geographical distances to ensure there is sufficient variation in the environment to detect behavioral responses. The marked differences in ecological conditions and red colobus behavior we documented over short geographical distances, suggests that small-scale contrasts are a useful tool to examine ecological determinants of behavior and community structure.     

The role of phytochemistry in dietary choices of Tana River red colobus monkeys(Procolobus badius rufomitratus)

We conducted a phytochemical survey of tree species growing within the riverine forests of the Tana River National Primate Reserve in Kenya to understand better the feeding ecology of an endangered resident primate, the Tana River red colobus monkey (Procolobus badius rufomitratus).Young leaves, which make up a large percentage of this monkey's diet, are significantly higher in nitrogen and lower in acid detergent fiber than more abundant mature leaves are. Phenolic chemistry had little inhibitory effect on feeding by P. b. rufomitratus.Choice among tree species by P. b. rufomitratusappears to be influenced largely by leaf availability,once an acceptable threshold of nitrogen and fiber is reached When mature leaves are eaten, they selected species that are high in nitrogen and low in fiber. A significantly higher nitrogen content was found for the mature leaves of all leguminous versus nonleguminous tree species. Consequently, the availability of certain types of mature leaf species during periods of preferred food scarcity may prove critical to groups of Tana River red colobus monkeys.     

Factors Influencing Interannual and Intersite Variability in the Diet of <Emphasis Type="Italic">Trachypithecus francoisi</Emphasis>

Comparative studies of the diet of the same species in different habitats, and over the long term, are essential to understanding a species’ behavioral and ecological plasticity. Moreover, such studies can help researchers and managers evaluate a species’ capacity to cope with changes in habitat quality resulting from natural processes or human disturbance, which is important for developing conservation strategies. We compared dietary data for François’ langur (Trachypithecus francoisi) collected at Nonggang and Fusui Nature Reserves, Guangxi Province, China, over 2 separate study periods to evaluate interannual and intersite variation in diet. Young leaves were the preferred staple foods for langurs, whereas mature leaves and seeds served as fallback foods in response to seasonal shortage in the abundance of young leaves. Species composition of the diets and the percentage of feeding records for plant species varied between the 2 study periods. The langurs at both study sites fed selectively, and they did not base their diet simply on the abundance of plant species in the habitat. However, the plant species eaten by langur groups inhabiting the 2 different reserves were markedly different, and the top 10 food species eaten by the Fusui group showed no overlap with those eaten by the Nonggang group. The variation may be related to differences in forest composition resulting from different level of human disturbance. In summary, our results indicate that François’ langurs exhibit a comparable dietary pattern both temporally and geographically, but there is marked interannual and intersite difference in species composition of the langur diet.     

Frequent water drinking by Zanzibar red colobus (Procolobus kirkii) in a mangrove forest refuge

Isolated populations of Procolobus kirkii on Uzi Island, Zanzibar, use Rhizophora mucronata-dominated mangrove forest for refuge. Three groups, observed over 14 months, spent up to 85% of total observation time in mangroves with brief excursions to adjacent upland coral rag forest, habitat degraded by human cutting. A large proportion of monkeys' diets consisted of plant parts of five mangrove species. Water drinking was common and 326 water-drinking events were recorded at a rate of up to 0.87 drinks hr(-1). Groups used different strategies to obtain water including licking dew, drinking from treeholes, licking rain off leaves and tree trunks, and drinking from coral rock crevices with Cercopithecus mitis albogularis. Drinking frequency increased with time spent in and consumption of mangroves. Strategies for obtaining water were group-specific and likely the result of learning. Drinking appeared to be an acquired behavior in movement-restricted groups living in a habitat with low plant species diversity and limited salty foods.     

Dietary Variability in Redtail Monkeys (<Emphasis Type="Italic">Cercopithecus ascanius schmidti</Emphasis>) of Kibale National Park,Uganda: the Role of Time,Space, and Hybridization

Studies of the diet of different groups of the same species allow us to understand intraspecific dietary variability. I collected dietary data from six neighboring groups of redtail monkeys (Cercopithecus ascanius schmidti) and three hybrid monkeys over 12 years at Ngogo and from one group at Kanyawara in Kibale National Park, Uganda and compared these results with previous studies of redtail diets elsewhere in Kibale and from the Kakamega Forest of Kenya. I scored feeding as a particular monkey ingesting a species-specific plant part, or catching insects from a species-specific substrate. A new feeding score was tallied for the same combination of parameters only after a 30-min interval or if the identity of one of the three parameters changed. I counted trees along transects in the home ranges of the two main study groups to calculate food selection ratios. I used chi-square tests to compare diets between groups and time periods and Spearman rank correlation coefficient tests for dietary correlates. These comparisons reveal considerable variation in plant parts and species eaten by redtails between months, years, and neighboring groups with overlapping ranges. Selection ratios show that some tree species are important sources of plant food, while others are more important as sources of invertebrates. The high incidence of insectivory by redtails demonstrates another ecological role they play in addition to seed dispersal. The intrademic variation in diets I describe for Kibale was often as great as and sometimes greater than the interdemic variation. The diets of the hybrid monkeys at Ngogo differed in some ways from their parental species, particularly in their greater consumption of invertebrates. Introgression may have led to some of these differences within and between redtail demes. The pronounced variability in redtail diets demonstrates why a typological perspective of species is unwarranted and that the validity of interspecific comparisons requires a thorough understanding of intraspecific variation.     

Diet of the red howler monkey (Alouatta seniculus) in French Guiana

We report the feeding behavior and food preferences of a troop of red howler monkeys (Alouatta seniculus) over two annual cycles in primary tropical rain forest in French Guiana. The monkeys used 195 plant species from 47 families as food. Major food categories were young leaves (54%), mature fruits (21.5%), and flowers (12.6%). Other food categories included old leaves, immature fruits, termitarium soil, bark, and moss. The monkeys were less selective than other howler groups, since 19 plant species contributed 1% to their diet and accounted for only 35.7% of their total diet. The Sapotaceae was the most frequently eaten plant family and represented >10% of the total diet.     

Estrogenic plant foods of red colobus monkeys and mountain gorillas in Uganda

Phytoestrogens, or naturally occurring estrogen-mimicking compounds, are found in many human plant foods, such as soybeans (Glycine max) and other legumes. Because the consumption of phytoestrogens may result in both health benefits of protecting against estrogen-dependent cancers and reproductive costs of disrupting the developing endocrine system, considerable biomedical research has been focused on the physiological and behavioral effects of these compounds. Despite this interest, little is known about the occurrence of phytoestrogens in the diets of wild primates, nor their likely evolutionary importance. We investigated the prevalence of estrogenic plant foods in the diets of two folivorous primate species, the red colobus monkey (Procolobus rufomitratus) of Kibale National Park and mountain gorilla (Gorilla beringei) of Bwindi Impenetrable National Park, both in Uganda. To examine plant foods for estrogenic activity, we screened 44 plant items (species and part) comprising 78.4% of the diet of red colobus monkeys and 53 plant items comprising 85.2% of the diet of mountain gorillas using transient transfection assays. At least 10.6% of the red colobus diet and 8.8% of the gorilla diet had estrogenic activity. This was mainly the result of the red colobus eating three estrogenic staple foods and the gorillas eating one estrogenic staple food. All estrogenic plants exhibited estrogen receptor (ER) subtype selectivity, as their phytoestrogens activated ERβ, but not ERα. These results demonstrate that estrogenic plant foods are routinely consumed by two folivorous primate species. Phytoestrogens in the wild plant foods of these two species and many other wild primates may have important implications for understanding primate reproductive ecology.     

Bioactive properties of plant species ingested by chimpanzees (Pan troglodytes schweinfurthii) in the Kibale National Park, Uganda

We measured the biological activities of a selected sample (84 crude extracts) of 24 species eaten by wild chimpanzees (Pan troglodytes schweinfurthii) in the Kibale National Park, western Uganda, to assess their potential chemotherapeutic values. Antibacterial, antimalarial, and/or antileishmania activities were observed in some crude extracts, and five of these extracts showed a significant cytotoxicity against human tumor cells. Active compounds isolated from three plant parts occasionally ingested by chimpanzees (Diospyros abyssinica (Ebenaceae) bark, Uvariopsis congensis (Annonaceae) leaves, and Trichilia rubescens (Meliaceae) leaves) showed highly significant medicinal properties. Two novel antiparasitic limonoids were isolated from Trichilia rubescens and their molecular structures were determined. In addition to elucidating the natural equilibrium maintained between hosts and pathogens, our investigation of the diet of wild chimpanzees may serve as a guideline to discovering plants with bioactive properties that should be preserved from destruction because of their health maintenance value for great ape populations.     

Diet of chimpanzees (Pan troglodytes schweinfurthii) at Ngogo, Kibale National Park, Uganda, 2. Temporal variation and fallback foods

Highly frugivorous primates like chimpanzees (Pan trogolodytes) must contend with temporal variation in food abundance and quality by tracking fruit crops and relying more on alternative foods, some of them fallbacks, when fruit is scarce. We used behavioral data from 122 months between 1995 and 2009 plus 12 years of phenology records to investigate temporal dietary variation and use of fallback foods by chimpanzees at Ngogo, Kibale National Park, Uganda. Fruit, including figs, comprised most of the diet. Fruit and fig availability varied seasonally, but the exact timing of fruit production and the amount of fruit produced varied extensively from year to year, both overall and within and among species. Feeding time devoted to all major fruit and fig species was positively associated with availability, reinforcing the argument that chimpanzees are ripe fruit specialists. Feeding time devoted to figs-particularly Ficus mucuso (the top food)--varied inversely with the abundance of nonfig fruits and with foraging effort devoted to such fruit. However, figs contributed much of the diet for most of the year and are best seen as staples available most of the time and eaten in proportion to availability. Leaves also contributed much of the diet and served as fallbacks when nonfig fruits were scarce. In contrast to the nearby Kanywara study site in Kibale, pith and stems contributed little of the diet and were not fallbacks. Fruit seasons (periods of at least 2 months when nonfig fruits account for at least 40% of feeding time; Gilby & Wrangham., Behavioral Ecology and Sociobiology 61:1771-1779, 2007) were more common at Ngogo than Kanyawara, consistent with an earlier report that fruit availability varies less at Ngogo [Chapman et al., African Journal of Ecology 35:287-302, 1997]. F. mucuso is absent at Kanyawara; its high density at Ngogo, combined with lower variation in fruit availability, probably helps to explain why chimpanzee population density is much higher at Ngogo.     

Mechanical defenses in leaves eaten by Costa Rican howling monkeys (Alouatta palliata)

Primate species often eat foods of different physical properties. This may have implications for tooth structure and wear in those species. The purpose of this study was to examine the mechanical defenses of leaves eaten by Alouatta palliata from different social groups at Hacienda La Pacifica in Costa Rica. Leaves were sampled from the home-ranges of groups living in different microhabitats. Specimens were collected during the wet and dry seasons from the same tree, same plant part, and same degree of development as those eaten by the monkeys. The toughness of over 300 leaves was estimated using a scissors test on a Darvell mechanical tester. Toughness values were compared between social groups, seasons, and locations on the leaves using ANOVA. Representative samples of leaves were also sun-dried for subsequent scanning electron microscopy and energy dispersive x-ray (EDX) analyses in an attempt to locate silica on the leaves. Both forms of mechanical defense (toughness and silica) were found to be at work in the plants at La Pacifica. Fracture toughness varied significantly by location within single leaves, indicating that measures of fracture toughness must be standardized by location on food items. Monkeys made some food choices based on fracture toughness by avoiding the toughest parts of leaves and consuming the least tough portions. Intergroup and seasonal differences in the toughness of foods suggest that subtle differences in resource availability can have a significant impact on diet and feeding in Alouatta palliata. Intergroup differences in the incidence of silica on leaves raise the possibility of matching differences in the rates and patterns of tooth wear.     

Fallback Foods of Red Leaf Monkeys (Presbytis rubicunda) in Danum Valley, Borneo

Animals in Southeast Asia must cope with long periods of fruit scarcity of unpredictable duration between irregular mast fruiting events. Long-term data are necessary to examine the effect of mast fruiting on diet, and particularly on the selection of fallback foods during periods of fruit scarcity. No such data is available for colobine monkeys, which may consume substantial amounts of fruits and seeds when available. We studied the diet of red leaf monkeys (Presbytis rubicunda, Colobinae) in Danum Valley, Sabah, northern Borneo, using 25 mo of behavioral observation, phenology and vegetation surveys, and chemical analysis to compare leaves eaten with nonfood leaves. The monkeys spent 46% of their feeding time on young leaves, 38% on seeds, 12% on whole fruits, 2.0% on flowers, 1.0% on bark, and 1.2% on pith. They spent more time feeding on seeds and whole fruit when fruit availability was high and fed on young leaves of Spatholobus macropterus (liana, Leguminosae) as fallback foods. This species was by far the most important food, constituting 27.9% of the total feeding time, and the feeding time on this species negatively correlated with fruit availability. Consumed leaves contained more protein than nonconsumed leaves, and variation in time spent feeding on different leaves was explained by their abundance. These results suggest that red leaf monkeys show essentially the same response to the supra-annual increase in fruit availability as sympatric monogastric primates, increasing their seed and whole-fruit consumption. However, they depended more on young leaves, in particular Spatholobus macropterus, as fallback foods during fruit-scarce periods than did gibbons or orangutans. Their selection of fallback food appeared to be due to both nutrition and abundance.     

Increasing Group Size Alters Behavior of a Folivorous Primate

Group size influences many aspects of mammalian social life, including stress levels, disease transmission, reproductive rates, and behavior. However, much of what is known about the effects of group size on behavioral ecology has come from comparisons across multiple groups of different sizes. These findings may be biased because behavioral differences across groups may be more indicative of how environmental variation influences animal behavior, rather than group size itself. To partially circumvent this limitation, we used longitudinal data to examine how changes in group size across time affect the behavior of folivorous red colobus monkeys (Procolobus rufomitratus) of Kibale National Park, Uganda. Controlling for food availability, we demonstrated that increasing group size resulted in altered activity budgets, based on 6 yr of data on a group that increased from 57 to 98 members. Specifically, as group size increased, individuals spent less time feeding and socializing, more time traveling, and increased the diversity of their diet. These changes appear to allow the monkeys to compensate for greater scramble competition apparent at larger group sizes, as increasing group size did not show the predicted relationship with lower female fecundity. Our results support recent findings documenting feeding competition in folivorous primates. Our results also document behavioral flexibility, an important trait that allows many social mammals to maximize the benefits of sociality (e.g., increased vigilance), while minimizing the costs (e.g., increased feeding competition).     

Baboon diet: a five-year study of stability and variability in the plant feeding and habitat of the yellow baboons (Papio cynocephalus) of Mikumi National Park, Tanzania

The habitat and plant feeding of 64 well-habituated, individually identified adult male and female yellow baboons was studied for 5 years at Mikumi National Park, Tanzania. Variation across the years showed that a study of only one or two years would have been incomplete and misleading. The list of baboon food species obtained from Mikumi is considerably larger and more diverse than any previously reported. One to six plant parts were eaten from each of more than 180 species. The 25 most common tree genera all contained species used for food. Of the 50 most common grass, shrub and herb genera, 93% included plant foods. Using months in which a species was eaten during at least one year of the study, 21 staple species were eaten during a mean of 8.86 months and 7 were eaten in all 12 months. Although many foods were from commonly available plant species, 15 such species were only rarely eaten. The number of parts of a species eaten per month and an estimate of the amounts eaten per month both varied with temperature and rainfall, being lowest near the end of the cool, dry season. There were substantial differences from year to year in the timing and amount of food production of many species; nevertheless, the same broad feeding pattern was repeated in each of the 5 years of the study. Despite yearly variation in food availability, 14 or more staples and other common foods were eaten in any given month. If crops of many of these foods were to fail, a large number of less commonly eaten species could be substituted. Baboons are eclectic feeders that appear to be optimizing their diet by selective feeding from among a wide array of available foods in an ever-changing floristic environment.     

Constraints on Group Size in Red Colobus and Red-tailed Guenons: Examining the Generality of the Ecological Constraints Model

The ecological constraints model proposes that an increase in group size will increase intragroup feeding competition and thereby constrain group size. Although this model has received wide acceptance, tests of it are based only on a few studies of species that have similar ecological requirements and social organizations, and there are reasons to question the widespread acceptance of the assumptions underpinning it. Via a 2-year study, we explored determinants of group size in species that feed on markedly different types of foods: the folivorous red colobus (Procolobus pennantii) and the frugivorous/insectivorous red-tailed guenon (Cercopithecus ascanius). We established 4 study sites approximately 15 km apart in Kibale National Park, Uganda, to examine the relationship between average group size and food availability. In both species, we quantified interdemic variation in diet, density of food trees, rate of travel, and group size. Red colobus at all sites relied heavily on leaf resources (75.5%–86.9%), but fruit (6.4%–13.9%) and flowers (2.0%–13.9%) were important in some populations. In general, red-tailed guenons fed on fruit (35.7%–59.7%), insects (14.5%–17.6%), and young leaves (12.2%–32.8%), but the amount of time allocated to these foods varied among sites. Average monthly density of trees bearing food items ranged among sites from 45 to 79 trees/ha for red colobus and from 19.6 to 67.3 trees/ha for red-tailed guenons. For both species, rate of travel was similar among sites, with one exception for red colobus. Average red colobus group size varied among sites from 14 to 40 (28 groups counted). Red-tailed guenon group size varied among sites from 11 to 24 (16 groups counted). As predicted by the ecological constraints model, group size increased with food tree density across sites for both species.     

Incisor microwear and diet in three species of Colobus

Examination of incisor microwear in three species of Colobus revealed that the predominantly folivorous C. badius more closely resembles C. satanas, a seed predator/folivore, than C. guereza, another predominantly folivorous species. This demonstrates that species of the same broad dietary category can have very different patterns of incisor microwear, indicative of differences in food procurement behavior and/or the physical properties of dietary items for some portion of the diet. Conversely, species of different categories can have microwear patterns that, superficially at least, are quite similar. The dissimilarity in incisor microwear between C. badius and C. guereza is mirrored to a certain extent in molar microwear, although the differences are not nearly so great on the molars. The differences between C. badius and C. guereza may involve different food items in the major, folivorous portions of their diets, or they may relate to differences in the very minor fruit and bark components. The similar microwear patterns of C. badius and C. satanas demonstrate that incisor microwear by itself is unreliable for assigning fossil species to broad dietary categories. Incisor microwear can be used to infer finer dietary distinctions in fossil species for which dietary category has been determined by other means.     

The diet and feeding behavior of the silver leaf monkey (Trachypithecus auratus sondaicus) in Indonesia

I describe the diet and feeding behavior of silver leaf monkeys (Trachypithecus auratus sondaicus) in the Pangandaran Nature Reserve, West Java, Indonesia, and compare a group living in old secondary rain forest with a group living in mixed plantation/secondary forest to determine intraspecific variation in feeding behavior and the importance of the plantation species in the diet of the monkeys. Young leaves and leaf buds made up slightly less than half of their diets, with both groups showing a preference for a few species when seasonally available. Fruits and flowers of a few species were also preferentially selected when available. These included sweet, fleshy fruits, which most other colobines tend to avoid. Young leaf intake was greatest in months when fruit intake was low. Mature leaves were rarely eaten. Both groups spent approximately 20% of feeding time foraging on Moraceae species. Differences in the diet of the two groups were related largely to differences in vegetational composition and the availability and abundance of food items for the species common to both sites. Teak (Tectona grandis) was the top food species of the group living in mixed plantation/secondary forest, with the midribs of young leaves preferentially selected. Young leaves ofT. grandis, available throughout the study, provided a staple food and were eaten when preferred foods were scarce. More favored food items were available to the group living in old secondary forest, though none was a staple food.