Ecology of hoolock gibbon of Bangladesh

Of the nine known species of gibbon,Hylobates hoolock is found in the evergreen and semi-evergreen forests of Bangladesh. It is monogamous, living in groups of 2–5, each group maintaining its territory, in a home range of 30–35 ha. The day range varies from 600m to 1600m. Detailed study on this gibbon has not been conducted. Therefore, a study on the ecology and behaviour of this species was made between February 1990 and January 1991. The gibbon spends 39% of the daytime in feeding, 25% in moving, 26% in resting, 4% in calling, and 6% in other social activities. They are completely arboreal and are mainly frugivorous. The total number of this species is unlikely to exceed 200. They are in danger due to their habitat destruction.     

Lion-tailed macaques (Macaca silenus) in a disturbed forest fragment: Activity patterns and time budget

We describe the activity patterns and time budget of a feral group of lion-tailed macaques that were confined to a disturbed forest fragment of 65 ha and compare the results with those obtained for groups in protected forests. The degraded nature of the study site was reflected in low tree densities, low specific diversity, gaps in the girth distribution of trees, and frequent disturbance by humans. The study group of 43 subjects was twice as large as lion-tailed macaque groups in protected habitats. They spent the most time ranging (34.0%), followed by foraging (23.7%), feeding (17.9%), resting (16.0%), and other activities such as social interactions (8.4%). Monthly variations are significant for all activity categories except ranging. Times spent resting and foraging are negatively correlated (r = −0.5) and show significant seasonal differences. Foraging time was highest from September to November, when key food sources such asCullenia andArtocarpus were absent or marginally available. The study group spent most time (40.4%) at canopy levels between 21 and 30 m. They spent more time each day ranging than resting or feeding and more time terrestrially compared with groups in protected forests. Large group size, poor habitat quality, and seasonal variation in food availability were the major variables affecting their time budget, and these variables accounted for differences from the time budgets of groups in protected forests.     

On the ecology and behavior ofCebus albifrons in eastern Colombia: I. Ecology

A-503 contact-hr study of a 35-member group ofCebus albifrons was conducted in eastern Colombia in 1977 and 1978. The group had a female: male socionomic sex ratio of 2.5:1 and used a home range of 110–120 ha which overlapped the home range of another group ofC. albifrons about 20–30 ha. The animals spent about 80% of their foraging time eating plant material and about 20% of their foraging time eating animal materials. A birth peak at the end of the dry season extending into the wet season was indicated by data available. Attempted predation was recorded by the mustelidEira barbara and the black-and-white hawk-eagleSpizastur melanoleucus. Some association was observed with the red howler monkeyAlouatta seniculus. The group at times spent more than half the day foraging and traveling on the ground, exhibiting a level of terrestriality not reported for other New World primates.     

Composition and Nutritional Characteristics of Fungi Consumed by Callimico goeldii in Pando, Bolivia

Though ≥22 species of Primates consume fungi, most do so at low rates, comprising <5% of their feeding time. Goeldi's monkeys (Callimico goeldii), spend up to 29% of their feeding time year-round consuming fungal sporocarps, the fruiting bodies of fungi. We provide comprehensive data on the nutritional characteristics of 4 species of fungi consumed by Callimico goeldii (Ascopolyporus polyporoides, Ascopolyporus polychrous, Auricularia auricula, and Auricularia delicata). The composition of the fungi is similar to that of other fungi: predominantly fiber (66.2–83.0% dry matter) with small amounts of sugar (2.0–5.6% dry matter) and crude fat (0.9–1.6% dry matter). Though the crude protein content is substantial (5.5–13.4% dry matter), much of the nitrogen in the fungi is not likely to be available to Callimico goeldii because it is associated with indigestible food components or is in nonprotein form. The mineral content of the fungi are within the normal range for fungi generally and the calcium-to-phosphorus ratio is low (0.07–0.25). Fungi appear to be a low-quality food resource for Callimico goeldii and may contribute to their relatively large home ranges and low population density compared to other Callitrichinae. Research on the ability of Callimico goeldii to digest fungi is needed to understand fully the nutritional value of fungi to them. We discuss adaptations Callimico goeldii may have for improving their ability to obtain nutrients from fungi and potential ecological correlates of mycophagy.     

Scramble Competition Among <Emphasis Type="Italic">Colobus vellerosus</Emphasis> at Boabeng-Fiema,Ghana

We investigated the occurrence of scramble competition among Colobus vellerosus at Boabeng-Fiema, Ghana. If scramble competition had an impact on feeding efficiency among females, we expected a positive relationship between group size and the proportion of time spent feeding, day journey length, or home range size assuming resource availability is similar among the groups compared. We collected focal data on the feeding behavior of adult females and males over 11 mo (September 2000–August 2001) on 2 study groups: WW (n = 31–33 individuals) and B (n = 8–16 individuals). We also collected ranging data on group movements at half-hour intervals. The large group (WW1) had a significantly longer day journey length than the small group (B1), and females in the large group spent a significantly greater proportion of time feeding in the wet season, a period of low food availability, which suggests it may be a bottleneck period when food resources are scarce and Colobus vellerosus is close to being energy limited. The proximity data suggested females may be able to reduce or adjust for competition by having fewer neighbors when they feed and by spreading out when in a larger group. However, we found no relationship between home range size and group size or that females spent a greater proportion of time feeding than adult males did. Our results highlight the need to factor in differences in food availability when investigating scramble competition. Though equivocal, our results suggest scramble competition occurs among Colobus vellerosus, leading us to suggest there was a match with the potential competitive regime, i.e., food distribution.     

Foraging strategies of yakushima macaques (Macaca fuscata yakui )

I investigated the activity budget and diet of Yakushima macaques (Macaca fuscata yakui,)in warm temperate broad- leaved forest of Yakushima, Japan. Both time spent feeding and time spent moving varied considerably between half- months. However, total time spent in active behaviors— feeding time plus moving time— was stable. The composition of the diet also showed considerable variation between half- months. The macaques fed mainly on fruits, seeds,mature leaves, fallen seeds, flowers, and young leaves, each of which accounted for more than 30% of feeding time in at least 1 half- month. They also ate insects and fungi, but each of them comprised ≤ 25 and ≤ 8% of feeding time in any half- month, respectively. Time spent feeding on mature leaves, young leaves, flowers, or fallen seeds is positively correlated with total time feeding and is negatively correlated with time moving. In contrast, time feeding on fruits, seeds, insects or fungi is negatively correlated with time feeding and is positively correlated with time moving. Foraging on foods that have a low energy content, a high density, and a relatively even distribution— mature leaves— or that need much manipulation to be processed— flowers and fallen seeds— increased feeding time, while foraging on foods for which monkeys must search intensively in the forest— fruits, seeds, insects, and fungi— led to increased moving time. I examined foraging strategies of Yakushima macaques in terms of moving costs and the quality of food items. Regarding time feeding on fruits, which have more energy and may need less manipulation than other foods, as a benefit, and moving time as a cost, they seemed to employ a strategy that balanced the costs and benefits of foraging.     

Studying <Emphasis Type="Italic">Daphnia</Emphasis> feeding behavior as a black box: a novel electrochemical approach

We present a novel approach for examining the complex feeding behavior of a filter feeder at a previously unexploited scale. A Daphnia lives in a viscous environment and thus creates a feeding current with a distinct laminar inflow and a repetitive pulsed outflow. We propose that by treating the feeding apparatus as a black box, and using the pulsed outflow current as a surrogate to the inside working of the apparatus, we can calculate feeding rate in near real time. The structure of the outflow is interpreted as a direct representation of the organism’s response to its environment. Therefore, we examine how the work performed by an organism’s feeding apparatus is altered according to environmental factors and metabolic demands. Our approach is an integration of optical (Schlieren system) and electrochemical (chronoamperometry) techniques that allow for real time visualization and temporal analysis of flow systems, respectively. As electrochemistry requires a tracer chemical, we employed low dopamine concentrations (≤ 1mM), and tested the effect of dopamine on the heart rate and swimming of Daphnia. It appears that dopamine free in solution at concentrations below 10 mM has no adverse effects on the organism, and all observed differences in Daphnia feeding behavior were due to environmental or metabolic factors. The feeding nature of daphnids in the presence or absence of food, and differences between the sexes is reported. Our results indicate that in the absence of food a Daphnia has a strict and repetitive feeding behavior with short delays between pumping actions. However, in the presence of food this behavior becomes complex, with increased delays between pumps, perhaps designed to maximize feeding efficiency. Our observations demonstrate that males have a higher appendage beat frequency than females under identical conditions. We hypothesize that the difference may be dictated by metabolic demand, as a male spends more time actively seeking a mate. The application of electrochemistry to the study of Daphnia feeding behavior is an improvement over current methods for its near real time quantification of behavioral response, its versatile application under varying environmental conditions and its extreme sensitivity to changes in the organism’s feeding behavior. This technique is a valuable addition to the current tools available for studying Daphnia feeding behavior and will allow us to learn more about the interactions of an organism with its environment. Guest editor: Piet Spaak Cladocera: Proceedings of the 7th International Symposium on Cladocera     

Life history of the predatory mites <Emphasis Type="Italic">Neoseiulus paspalivorus</Emphasis> and <Emphasis Type="Italic">Proctolaelaps bickleyi</Emphasis>, candidates for biological control of <Emphasis Type="Italic">Aceria guerreronis</Emphasis>

The eriophyoid mite Aceria guerreronis Keifer (Eriophyidae), commonly called the coconut mite, is a key pest of coconut fruits. Surveys conducted on coconut palms in Brazil revealed the predatory mites Neoseiulus paspalivorus DeLeon (Phytoseiidae) and Proctolaelaps bickleyi Bram (Ascidae) as the most commonly associated natural enemies of A. guerreronis on coconut fruits. However, virtually nothing is known about the life history of these two predators. We conducted laboratory experiments at 25 ± 0.1°C, 70–90% RH and 12:12 h L:D photoperiod to determine the life history characteristics of the two predatory mites when feeding on A. guerreronis and other potential food sources present on coconut fruits such as Steneotarsonemus furcatus DeLeon (Tarsonemidae), coconut pollen and the fungus Rhizopus cf. stolonifer Lind (Mucoraceae). In addition, the two-spotted spider mite Tetranychus urticae Koch (Tetranychidae) was tested for its suitability as prey. Both predators, N. paspalivorus and P. bickleyi, thrived on A. guerreronis as primary food source resulting in shorter developmental time (5.6 and 4.4 days, respectively), higher oviposition rate (1.7 and 7.0 eggs/female/day, respectively) and higher intrinsic rate of increase (0.232 and 0.489 per female/day, respectively) than on any other diet but were unable to develop or lay eggs when fed T. urticae. Coconut pollen and S. furcatus were adequate alternative food sources for N. paspalivorus and Rhizopus for P. bickleyi. We discuss the relevance of our findings for natural and biological control of the coconut mite A. guerreronis.     

Taxonomy of theAlouatta seniculus group: Biochemical and chromosome data

Wild populations ofAlouatta belzebul andA. seniculus from Brazil were surveyed in relation to 20 protein loci. Estimates of genetic variability demonstrated thatA. seniculus presents the highest level of heterozygosity among several New World monkey species studied for the same loci. Additional information from DNA and chromosomes suggest thatA. seniculus andA. belzebul are not the closest species in theHershkovitz’sAlouatta seniculus group.     

Behavior, Diet, and Movements of the Sulawesi Crested Black Macaque (Macaca nigra)

We present the first field study of activity budgets, diet and ranging patterns of the Sulawesi crested black macaque, Macaca nigra, one of seven macaque species endemic to Sulawesi, Indonesia. We studied three crested macaque groups, ranging in size from 50 to 97 individuals, for 18 months in the Tangkoko-DuaSudara Nature Reserve, North Sulawesi. They spent 59% of the day moving and procuring food, especially fruits, and 41% of the day resting and socializing. Their diet is composed of more than 145 species of fruit (66% of observed feeding bouts), vegetative material (2.5%), invertebrates (31.5%), and occasional vertebrate prey. Group differences were more pronounced than seasonal or diurnal differences. Specifically, the largest group moved farther during the day, moved at a faster and more uniform rate, ate less fruit, rested more, and socialized less than the smaller groups did. The largest group had the largest home range, but it included less primary forest and more disturbed habitat than the ranges of smaller groups. There are individual differences in activity budgets of adult males and females in time spent moving, resting, feeding, and socializing that may reflect differences in reproductive strategies of males versus females. The behavior of large juveniles is more similar to that of adults than to that of small juveniles. Daily movements and use of home range are correlated with diet. Macaques moved shorter distances as the proportion of time spent feeding on fruit increased, and the top four dietary items accounted for most of the variance in entry into hectare blocks of home range.     

Habitat,diet, and activity patterns of free-ranging woolly spider monkeys (Brachyteles arachnoides E. Geoffroy 1806)

Results of a 10-month study of the ecology and behavior of free- ranging woolly spider monkeys (Brachyteles arachnoides)in Brazil show that these animals are strongly folivorous. Leaf-eating accounted for more than 50% of the total feeding time in all samples but one and accounted for more than 80% of the total feeding time in three samples. Mature foliage was routinely eaten. Woolly spider monkeys consistently spend more than 50% of each day quietly resting and sleeping. Animals travel little except when actively feeding and show low levels of social interaction. Such an activity profile suggests that woolly spider monkeys may often be living near the limits of their energetic resources. The social organization of the species is unusual for a folivorous primate in that small groups of females and associated immature animals confine their activities to discrete home-range areas, whereas males are itinerant, traveling over the home ranges of various female groups. Animals sharing a common home-range area show no permanent daily pattern of association other than that of mother-dependent offspring. Foraging alone or with few conspecifics should maximize each individual’s returns from foraging by minimizing the day range that must be traveled each day to locate foods while simultaneously lowering interference competition for higher-quality dietary resources.     

Feeding behavior ofLemur catta andLemur fulvus

The feeding behavior of two sympatric species of lemurs, Lemur cattaand Lemur fulvus,was studied in an enclosure simulating a natural habitat at the Duke University Primate Center. L. fulvusspent less time feeding during the day than L. catta.But the former species ate more fruit and had longer feeding bouts on preferred food items than L. catta.They also had a shorter food passage time than L. cattaand their choice of resting places was more influenced by food distribution. Furthermore, the two lemur species ate parts of different plant species and showed different reactions to chemical plant components. According to these results, L. fulvusis a more conservative feeder than L. catta.These interspecific differences in feeding behavior may be one of a number of differences that allow the two species to coexist. In allopatry, however, L. fulvusmay also adopt feeding patterns similar to those of L. catta.But L. cattawas never found to change its feeding strategies in different areas. It may be this option of L. fulvusto adopt different feeding strategies in different situations that allows this species to have the widest range of all Malagasy lemurs. Duke University Primate Center Publication No. 259.     

Energy budget of the lesser flamingo (Phoeniconaias minor Geoifroy)

A lesser flamingo's energy surplus, which is defined as the net rate at which it gains chemical energy (averaged over 1 or more whole days) is calculated by estimating the rate at which it filters algae from the water, and then subtracting the energy needed to pump water through the filter, and that expended on general metabolism. From the assumptions made, it follows that a flamingo should be able to make a positive energy surplus if the food concentration exceeds about 0–12 kg dry matter per m³ of water, and if it spends 80% of its time feeding. During incubation, however, less than half the total time can be spent feeding, and in this case the food concentration would need to be at least 0–25 kg/m³. About 2 days would be required to produce an egg at this concentration, and less than 1 day at the highest concentrations observed.     

In situ feeding tactics of short-nosed fruit bat (Cynopterus sphinx) on mango fruits: evidence of extractive foraging in a flying mammal

We report a sequence of behaviors exhibited by the short-nosed fruit bat Cynopterus sphinx while feeding on fruits of Mangifera indica. They peel off the outer skin to form a feeding area of about 3–6 cm diameter. Such food preparatory behaviors were more pronounced on larger mangoes. Bats competed among themselves to feed on the mangoes that had such feeding areas exposed. Individuals that spent a considerable amount of time on food preparatory behaviors actively secured the fruits. Altogether, these behaviors indicate that Cynopterus bats might have learnt, over evolutionary time, and developed behaviors that facilitate efficient processing and feeding of fruits such as mangoes. It appears that actions exhibited by C. sphinx in peeling off the outer skin of mangoes exemplify “extractive foraging”, a behavior that is prominently known in large-brained mammals. Thus, our findings will have implications on the distribution and evolution of extractive foraging and “technical intelligence” among mammalian lineages.     

Use of space, activity patterns, and foraging behavior of red howler monkeys (Alouatta seniculus) in an Andean forest fragment in Colombia

Howler monkeys are among the most studied primates in the Neotropics, however, behavioral studies including estimation of food availability in Andean forests are scarce. During 12 months we studied habitat use, behavior, and feeding ecology of two groups of red howler monkeys (Alouatta seniculus) in an isolated fragment in the Colombian Andes. We used a combination of focal animal and instantaneous sampling. We estimated fruit production (FP) using phenology transects, and calculated young leaf abundance by observing marked trees. The home range area used by each group was 10.5 and 16.7 ha and daily distances traveled were 431 ± 228 and 458 ± 259 m, respectively. We found that both groups spent most of their time resting (62–64%). Resting time did not increase with leaf consumption as expected using a strategy of energy minimization. We did not find a relationship between daily distances traveled and leaf consumption. However, howlers consumed fruits according to their availability, and the production of young leaves did not predict feeding time on this resource. Overall, our results are similar to those found on other forest types. We found that despite limited FP in Andean forests, this did not lead to a higher intake of leaves, longer resting periods, or shorter traveling distances for red howlers. Am. J. Primatol. 73:1062–1071, 2011. © 2011 Wiley‐Liss, Inc.     

Ranging Behavior and Possible Correlates of Pair-Living in Southeastern Avahis (Madagascar)

Researchers have proposed several hypotheses to explain pair-living in primates. In particular, when males are not involved in direct parental care, pair-living may be related to female dispersal, infanticide prevention, or male mate/resource defense. We aimed to evaluate, through a better understanding of the ranging patterns of avahis, which hypotheses may best account for pair-living in these nocturnal lemurs. We collected focal observations over 26 nights, June–September 2004, in a littoral forest (Sainte Luce, southeastern Madagascar) on 4 adult radiocollared avahis [Avahi laniger (Tattersall, I. (1982). The Primates of Madagascar. Columbia University Press, New York.) or Avahi meridionalis (Zaramody in Primate Reports 74:9–22, 2006)]. We followed 2 males and 2 females from 2 groups: B, a male-female couple, and A, comprising the parental pair and 1-yr-old female offspring. The adult females birthed in August. We recorded resting and feeding tree points (via global positioning system) for home range calculation through minimum convex polygon and kernel methods. We provide the first quantitative information on the ranging behavior of 2 male and 2 female avahis. Home range/daily path length values (means) are higher than the ones previously reported for the same and other folivorous pair-living lemur species. On average, the 2 females spent more time feeding and traveled shorter distances than the 2 males did. Male–female cohesion (mean values), possibly enhanced by offspring presence, was higher in A and after births than in B and before births, respectively. Although male avahis may be forced into pair-living owing to energy constraints related to size, locomotion, and diet, females might accept pair-living in exchange for indirect territory defense and mate guarding.     

Influence of Human Food Consumption on the Time Budget of Vervets

In nonhuman primate groups that utilize crop and food-raiding strategies, the inclusion of human food in the diet can have a pervasive impact on the activity budget. Human food is usually of higher quality and energy per unit than wild food. Thus, metabolic demands may be reached sooner on a human food diet, which in turn leads to improved foraging efficiency. In particular, a reduction in time spent feeding is associated with an increase in time spent resting. We investigated changes in time budget in a group of vervets (Chlorocebus aethiops pygerthrus) living in a tourist and cultivated area of Entebbe, Uganda (Zoo group). Saj collected 123 hr of focal observations on 17 individuals. Human food constituted half of the diet of Zoo group. Access to human food resulted in a high proportion of time resting and a low proportion of time feeding. The group had a small average daily range and a small home range in relation to other reports on unprovisioned groups. The time budget differences suggest that the juveniles spent more time moving and feeding and less time resting than adults did, and that adult females were more social than adult males were. These patterns have been observed in both provisioned and unprovisioned vervet groups. In contrast to a report from an unprovisioned group (Harrison, 1983), lactating females in this provisioned situation were able to maintain feeding and activity levels similar to those of other adults. However, adult females spent significantly less time consuming human food than juveniles did. We suggest that the risk associated with human food consumption may explain the low consumption of human food by females with dependent offspring. The differences may also reflect the different nutritional requirements of lactating females or the tendency for juveniles to be more exploratory than other group members.     

Estimation of food intake and ingested energy in Daubenton’s bats (Myotis daubentonii) during pregnancy and spermatogenesis

We studied food intake of and estimated ingested energy in female and male Myotis daubentonii during the periods of pregnancy (period 1, 8 May–4 June) and of intense spermatogenetic activity (period 2, 24 July–22 August) over 8 years (1996–2003) in central Germany. We used radiotelemetry to determine the time spent foraging and marked animals with chemiluminescent light-sticks to determine prey attack rates. Body length, body mass, moisture content, and caloric content of chironomids, the main prey of Daubenton’s bats, were measured to estimate the nightly food intake and, in consequence, energy intake. Pregnant females spent significantly more time foraging than males during period 1 and females during the post-lactation period. In contrast, male foraged longer during the period of highest spermatogenetic activity than during late spring and also significantly longer than post-lactating females. Based on a mean number of 8.3 prey attacks per minute, the time spent foraging, and a capture success rate of either 50 or 92%, calculated intake values with a feeding rate of 7.6 insects per minute (=92% capture success) were more consistent with literature data for other insectivorous bats than that of values calculated on the basis of a capture success rate of 50%. In the high capture-success model, calculated insect intake of female bats was 8.0 g during pregnancy and 4.9 g per day during post-lactation, providing 5.0 and 3.0 kJ of ingested energy per gram body mass per day. Calculated intake of male bats was 3.6 g insects per day during late spring and 8.0 g during period of intensive spermatogenesis, providing 2.6 and 5.7 kJ of ingested energy per gram body mass.     

Effect of Group Size on Activity Budgets of Colobus vellerosus in Ghana

Group size influences foraging efficiency in several primates. We examined the activity budgets of 3 groups of Geoffroy's pied colobus (Colobus vellerosus) at the Boabeng-Fiema Monkey Sanctuary in Ghana to determine whether larger group size induces scramble competition. We studied 2 groups (B1; N = 7-8 and WW; N = 31-33) occupying slightly overlapping home ranges from August to November 2000. We observed the third group, B2 (N = 15-16), comprising B1 and 7 male invaders in the same home range as B1 from August to November 2001. By comparing groups belonging to the same population and occupying sligthly overlapping or similar home ranges, we were able to control, to a certain extent, for differences in food distribution. We recorded a total of 3353 scans, yielding 14,886 activity records, over 73 days. As with other black-and-white colobus, resting was their most common activity (59%). Intergroup comparisons suggest that time spent feeding, resting and moving did not vary in relation to group size. However, intragroup comparisons between the sexes show that females in the large group spent more time feeding than males did, whereas this was not the case in the small group, which suggests that scramble competition may be occurring among female Colobus vellerosus at BFMS. It is also possible that this may be due to greater nutritional requirements because of a higher proportion of infants in the large group. In fact, the proportion is quite similar between the two groups, lending support to the idea that females in the two groups had comparable nutritional demands due to lactation. This suggests that increased feeding in females in the large group was partly an effect of scramble competition. Group size and group composition also influenced the frequency of social behavior. There was more grooming in the large group, and it was performed mostly by females. The distribution of activities throughout the day was similar to the pattern reported for other black-and-white colobus.     

Predicting the Competitive Regime of Female <Emphasis Type="Italic">Colobus vellerosus</Emphasis> from the Distribution of Food Resources

We examined the spatial and temporal distribution of the foods of ursine colobus (Colobus vellerosus) at Boabeng-Fiema, Ghana as a means to predict the monopolizablity and usurpability of their food resources. Recent evidence suggests that food may not be limiting for folivorous primates, and that male sexual coercion may be a more important influence on folivore social organization. To address the question, we collected focal data on the feeding behavior of adult females and males over 11 mo (September 2000-August 2001) on 2 groups: WW (n = 31–33 individuals) and B (n = 8–16 individuals). We also conducted phenological monitoring and a tree survey of the two-group home ranges to establish food availability and distribution. We used 2 behavioral or organism-defined indicators of feeding behavior to assess potential resource contestability: food site residence time and distance moved between food sites. The colobus fed on a high diversity of species, most of their food trees were not clumped in distribution, within-tree interfood distances were short, and food trees were large. The only condition associated with the potential for monopolization was low food tree density. However, low food tree density may be offset by the colobus’ use of large trees. Taken together, the ecological and behavioral indicators suggest the food resources of Colobus vellerosus had a low potential for monopolization. Our results also indicate mature leaves had the longest food site residence time, which may suggest they should be the most usurpable plant part, though their presumed low quality and high abundance probably counteracted the effect. The pattern implied the potential for direct feeding competition among Colobus vellerosus at Boabeng-Fiema was low and agonistic interactions over food are not expected. Instead, a group size effect on feeding efficiency should be a more predominant influence on feeding efficiency, if food is limiting for the species.