Lophocebus Albigena Research Articles

Primates Take Weather into Account when Searching for Fruits

Temperature and solar radiation are known to influence maturation of fruits and insect larvae inside them . We investigated whether gray-cheeked mangabeys (Lophocebus albigena johnstonii) of Kibale Forest, Uganda, take these weather variables into account when searching for ripe figs or unripe figs containing insect larvae. We predicted that monkeys would be more likely to revisit a tree with fruit after several days of warm and sunny weather compared to a cooler and more cloudy period. We preselected 80 target fig trees and monitored whether they contained ripe, unripe, or no fruit. We followed one habituated monkey group from dawn to dusk for three continuous observation periods totalling 210 days. Whenever the group came within a 100 m circle of a previously visited target tree for a second time, we noted whether or not individuals proceeded to the trunk, i.e., whether they "revisited" or simply "bypassed" the tree. We found that average daily maximum temperature was significantly higher for days preceding revisits than bypasses. The probability of a revisit was additionally influenced by solar radiation experienced on the day of reapproach. These effects were found only for trees that carried fruit at the previous visit but not for trees that had carried none. We concluded that these nonhuman primates were capable of taking into account past weather conditions when searching for food. We discuss the implication of these findings for theories of primate cognitive evolution.

Dental microwear texture analysis: technical considerations

Dental microwear analysis is commonly used to infer aspects of diet in extinct primates. Conventional methods of microwear analysis have usually been limited to two-dimensional imaging studies using a scanning electron microscope and the identification of apparent individual features. These methods have proved time-consuming and prone to subjectivity and observer error. Here we describe a new methodological approach to microwear: dental microwear texture analysis, based on three-dimensional surface measurements taken using white-light confocal microscopy and scale-sensitive fractal analysis. Surface parameters for complexity, scale of maximum complexity, anisotropy, heterogeneity, and textural fill volume offer repeatable, quantitative characterizations of three-dimensional surfaces, free of observer measurement error. Some results are presented to illustrate how these parameters distinguish extant primates with different diets. In this case, microwear surfaces of Cebus apella and Lophocebus albigena, which consume some harder food items, have higher average values for complexity than do folivores or soft fruit eaters.

Spatial distribution of primates in a mosaic of colonizing and old growth forest at Ngogo, Kibale National Park, Uganda

Primate censuses were conducted in a mosaic of colonizing (two locations) and old-growth forests using line transect methods at the Ngogo study site, Kibale National Park, Uganda. Black and white colobus monkeys (Colobus guereza) were encountered more frequently in the colonizing forests than in the old growth forest, while chimpanzees (Pan troglodytes) were encountered more frequently in the old growth forest than in colonizing forests. Although not significant, results suggest that blue monkeys (Cercopithecus mitis) frequented colonizing forests more often than old growth forest. The encounter rates of mangabey (Lophocebus albigena), and redtail (Cercopithecus ascanius) groups were ambiguous with their density being higher in some colonizing forests but not others as compared to old-growth forest. No significant differences were detected for baboons (Papio anubis), L'hoest's (Cercopithecus lhoesti), and red colobus monkeys (Piliocolobus tephroscales). The conversion of forests to farmland is one of the major problems encountered in primate conservation. This study shows that secondary forests replacing anthropogenic grasslands have the potential of supporting some primate species such as black and white colobus, redtail monkeys, and possibly blue monkeys. Therefore, such areas should not be given up but should be conserved for the benefit of primates that can survive in secondary forests; as the forests mature further, primate species that are adapted to old growth forest will colonize the area provided there is a nearby source.

Densities of Two Frugivorous Primates with Respect to Forest and Fragment Tree Species Composition and Fruit Availability

Conservation of wildlife populations requires extensive knowledge of their habitat requirements, efficient methods to evaluate habitat quality, and an understanding of the value of fragments and edges. Kibale National Park, Uganda has areas that differ in the densities of 2 species of frugivorous monkeys—Cercopithecus mitis and Lophocebus albigena—including one on an edge and forest fragments outside the park that lack both species. We compared the basal area densities of important food trees with primate densities. The density of Cercopithecus mitis correlates most strongly with the basal area density of all types of food trees combined. The density of Lophocebus albigena does not correlate with the basal area densities of any category of food trees or with fruit availability. An index of their density—number of groups seen per km walked—correlates to fruit availability but with marginal significance. Lack of a relationship between the basal area densities of food trees and density of Lophocebus albigena may be the result of a mismatch in scale between the forest area measured and their large home ranges. We compared the unused area of forest to the other areas of the forest and the fragments and found it had higher basal area densities in all food tree categories for both species than the fragments and lower basal area densities of most categories than the other parts of the forest, indicating that the fragments are poor quality and would probably be unused even if dispersal were likely.

Seasonal variation in the quality of a tropical ripe fruit and the response of three frugivores

Seasonality in tropical forests can be more subtle than that of temperate forests but still affects the resources available to wildlife. Much work has been done describing changes in fruit availability and dietary composition but the nutritional quality of any particular food item is assumed to be relatively constant. We investigated seasonal changes in the quality of the ripe fruit of Celtis durandii, a common tree that produces fruit year-round and is important in the diets of many species. The lipid content of the ripe fruit was found to be highly variable (0.3–30.8% dry matter) among months and this variation was positively correlated with the summed daily rainfalls of the previous and concurrent months. The amount of this fruit in the diets of three frugivorous primate species (Cercopithecus mitis, Cercopithecus ascanius and Lophocebus albigena) was positively related to measured or estimated lipid levels in the fruit. Such predictable changes in the quality of a constantly available fruit have not been previously reported and suggest that the resources provided by tropical forests may be more seasonal than shown by common measures of fruit availability.

Dental microwear and diets of African early Homo

Conventional wisdom ties the origin and early evolution of the genus Homo to environmental changes that occurred near the end of the Pliocene. The basic idea is that changing habitats led to new diets emphasizing savanna resources, such as herd mammals or underground storage organs. Fossil teeth provide the most direct evidence available for evaluating this theory. In this paper, we present a comprehensive study of dental microwear in Plio-Pleistocene Homo from Africa. We examined all available cheek teeth from Ethiopia, Kenya, Tanzania, Malawi, and South Africa and found 18 that preserved antemortem microwear. Microwear features were measured and compared for these specimens and a baseline series of five extant primate species ( Cebus apella, Gorilla gorilla, Lophocebus albigena, Pan troglodytes, and Papio ursinus) and two protohistoric human foraging groups (Aleut and Arikara) with documented differences in diet and subsistence strategies. Results confirmed that dental microwear reflects diet, such that hard-object specialists tend to have more large microwear pits, whereas tough food eaters usually have more striations and smaller microwear features. Early Homo specimens clustered with baseline groups that do not prefer fracture resistant foods. Still, Homo erectus and individuals from Swartkrans Member 1 had more small pits than Homo habilis and specimens from Sterkfontein Member 5C. These results suggest that none of the early Homo groups specialized on very hard or tough foods, but that H. erectus and Swartkrans Member 1 individuals ate, at least occasionally, more brittle or tough items than other fossil hominins studied.

Patterns of male residency and intergroup transfer in gray‐cheeked mangabeys (Lophocebus albigena)

We describe the movements and fates of 36 collared gray-cheeked male mangabeys (Lophocebus albigena) that resided in seven social groups in Kibale National Park, Uganda. The male mangabeys were captured, radiotagged, and then contacted regularly over a period of up to 8 years. Individuals varied considerably in how tightly they were associated with their groups. "Central" males were usually found in close spatial association with their own groups, but they sometimes visited others. "Peripheral" males were often found several hundred meters away from their own groups. Most adult males were characterized by repeated intergroup transfers, manifested as "visits" and "dispersals." Visits were transfers that did not result in mating. The visits were brief, and ranged from a few minutes to a few days before the male returned to his previous group, moved on to another group, or returned to solitary life. The term "dispersal" refers to a transfer that results in mating, after which the male remains in the new group. Young males began to drift away from their natal groups in early subadulthood, but the timing of first associations with estrous females in other groups was highly variable. Natal dispersers were generally solitary for a month or more, and at least half moved into nonadjacent groups. Secondary dispersal was common. After they immigrated, adult males remained in a group for a median of 19 months before they emigrated again, with the probability of departure being approximately constant in time. Secondary dispersal was usually made into an adjacent group and lasted less than 7 days. Emigration and immigration were not seasonal, and males emigrated singly and apparently independently; however, immigrations were clumped in time, and male dispersal contributed to considerable heterogeneity of group composition. L. albigena dispersal patterns are intermediate between those described for Papio baboons and forest Cercopithecus, and models that predict demographic consequences of dispersal for baboons can be refined to apply to mangabeys.

Distribution, population density, and status of sympatric cercopithecids in the Campo-Ma'an area, Southwestern Cameroon.

A study on species composition, distribution, and population density of cercopithecids in the Campo-Ma'an area, Southwestern Cameroon, was undertaken from December 1997 until August 2000. A total of 665.5 km of line transects was used for the census. Thirteen diurnal primate species including five endangered species (Gorilla g. gorilla, Pan troglodytes, Mandrillus sphinx, Colobus satanas, Cercocebus torquatus) were recorded in the Campo Forest, the greatest part of which is a logging concession. Cercopithecus nictitans (1.43 groups/km2), C. cephus (1.13 groups/km2), C. pogonias (0.81 groups/km2), and C. torquatus (0.51 groups/km2) occurred at medium frequencies compared to figures from other Central African study sites. Mandrill densities estimated (0.27 groups/km2) show that the area is very important for the conservation of this rare species. Guenon densities found inside areas with a high level of human activities did not differ significantly from densities estimated in areas with a lower level of human activities. C. torquatus densities were significantly higher in the areas with a low level of human disturbance and encounter rates with Lophocebus albigena also indicate a preference of less disturbed areas. Mangabeys are thus likely to be adversely affected by human activities in the area. In the Ma'an Forest, which has not been logged yet, ten species were confirmed. The population densities of two guenons (C. nictitans and C. cephus) were estimated to be twice as high in the unlogged forest area as compared to the logged forest of Campo. Other species are rarer in the Ma'an Forest than in the Campo Forest. Although mangabeys are adversely affected by human activities, the results still indicate that selective logging may be compatible with the conservation of cercopithecids, if a reduced damage logging concept and antipoaching measures are implemented. Increased hunting activities following logging operations will definitely have a negative longterm impact on primate populations in the Campo-Ma'an area if no further, more effective conservation measures will regulate wildlife use in future.

DIFFERENTIAL RESOURCE USE BY PRIMATES AND HORNBILLS: IMPLICATIONS FOR SEED DISPERSAL

Arboreal frugivores, such as primates and hornbills, are important seed dis- persers for many tropical plant species, yet the degree to which they use the same resources is unknown. If primates and hornbills consume the same fruit species, they may be redundant in their roles as seed dispersers, and the loss of one of these taxa may be compensated for by the other. To examine resource use by tropical frugivores, we quantified the feeding habits of two hornbill species, Ceratogymna atrata and C. cylindricus, and five primate species, Colobus guereza, Lophocebus albigena, Cercopithecus pogonias, C. cephus, and C. nictitans, in the lowland rainforest of south-central Cameroon. Based on over 2200 feeding observations recorded between January and December 1998, we characterized the diets and estimated dietary overlap among frugivore species. Previous studies have cal- culated dietary overlap by counting the number of diet species that two animals share, often leading to inflated estimates of overlap. Our method incorporated the proportional use of diet species and fruit availability into randomization procedures, allowing a clearer as- sessment of the actual degree of overlap. This added complexity of analysis revealed that, although the diets of a hornbill and a primate species may have as many as 36 plant species in common, actual dietary overlap is low. These results suggested that there are distinct hornbill and primate feeding assemblages, with primates consuming a greater diversity of plant species and higher levels of nonfruit items like leaves and seeds. Using Correspon- dence Analysis, we also identified two primate assemblages, separated largely by degree of frugivory and folivory. In addition, we found that hornbills feed at significantly higher strata in the forest canopy and eat fruits of different colors than primates. Averaged across the year, overlap between groups (hornbill-primate) was significantly lower than combined within-group overlap (primate-primate and hornbill-hornbill), showing that primates and hornbills have dissimilar diets and are not redundant as seed dispersers. In equatorial Africa, primate populations face greater declines than hornbill populations because of hunting. It is unlikely that seed dispersal by hornbills will compensate for the loss of primates in maintaining forest structure.

Differential Resource Use by Primates and Hornbills: Implications for Seed Dispersal

Arboreal frugivores, such as primates and hornbills, are important seed dispersers for many tropical plant species, yet the degree to which they use the same resources is unknown. If primates and hornbills consume the same fruit species, they may be redundant in their roles as seed dispersers, and the loss of one of these taxa may be compensated for by the other. To examine resource use by tropical frugivores, we quantified the feeding habits of two hornbill species, Ceratogymna atrata and C. cylindricus, and five primate species, Colobus guereza, Lophocebus albigena, Cercopithecus pogonias, C. cephus, and C. nictitans, in the lowland rainforest of south-central Cameroon. Based on over 2200 feeding observations recorded between January and December 1998, we characterized the diets and estimated dietary overlap among frugivore species. Previous studies have calculated dietary overlap by counting the number of diet species that two animals share, often leading to inflated estimates of overlap. Our method incorporated the proportional use of diet species and fruit availability into randomization procedures, allowing a clearer assessment of the actual degree of overlap. This added complexity of analysis revealed that, although the diets of a hornbill and a primate species may have as many as 36 plant species in common, actual dietary overlap is low. These results suggested that there are distinct hornbill and primate feeding assemblages, with primates consuming a greater diversity of plant species and higher levels of nonfruit items like leaves and seeds. Using Correspondence Analysis, we also identified two primate assemblages, separated largely by degree of frugivory and folivory. In addition, we found that hornbills feed at significantly higher strata in the forest canopy and eat fruits of different colors than primates. Averaged across the year, overlap between groups (hornbill–primate) was significantly lower than combined within-group overlap (primate–primate and hornbill–hornbill), showing that primates and hornbills have dissimilar diets and are not redundant as seed dispersers. In equatorial Africa, primate populations face greater declines than hornbill populations because of hunting. It is unlikely that seed dispersal by hornbills will compensate for the loss of primates in maintaining forest structure.

Bone geometry in cercopithecoid mandibles

This study explores the relation between cortical bone geometry in the mandibular corpus and in vivo masticatory stress patterns and dietary specialization in cercopithecoid primates. Cortical bone distribution in the mandibles of three species of Old World monkeys ( Macaca fascicularis, Procolobus badius, Lophocebus albigena) was measured by computed tomography. The arrangement of bone within sections was quantified as (1) the ratio of cortical area to the enclosed (total) subperiosteal area; (2) the ratio of orthogonal second moments of area; and (3) size-adjusted measures of cortical area and regional thickness. Cross-sectional geometry differed among samples, but consistent patterns of cortical thinning and bone area were found within individual sections. This consistency was despite the marked differences in diet and feeding behavior that distinguish the three taxa. Lingually thin and basally thick cortical bone was found in the three monkeys; previously published data suggest that this pattern may be stereotypical among anthropoid primates. It is hypothesized that the interactive effects of shear, bending and torsion produce eccentric loads in corpus sections, which are mirrored by this asymmetrical arrangement of cortical bone. When interpreted against existing data for other primate groups, these results are consistent with the hypothesis that masticatory-loading profiles are broadly similar across anthropoids despite the distinctive occlusions found among the suborder. Understanding of the impact of diet on jaw morphology is, therefore, not improved by considerations of cortical bone distribution, i.e. the inference of diet from jaw form is best predicated on considerations of relative corpus size rather than cross-sectional geometry.

Social, state-dependent and environmental modulation of faecal corticosteroid levels in free-ranging female spotted hyenas.

Little is known about to what extent the sensitivity of the hypothalamic-pituitary-adrenal (HPA) axis may be state dependent and vary in the same species between environments. Here we tested whether the faecal corticosteroid concentrations of matrilineal adult female spotted hyenas are influenced by social and reproductive status in adjacent ecosystems and whether they vary between periods with and without social stress. Females in the Serengeti National Park frequently become socially subordinate intruders in other hyena territories by undertaking long-distance foraging trips to migratory herds, whereas in the Ngorongoro Crater they usually forage inside their own small territories on resident prey. The faecal corticosteroid concentrations in Serengeti females were significantly higher than in Ngorongoro females. Energy expenditure by lactation is exceptionally high in spotted hyenas and this may be reflected in their corticosteroid levels. The faecal corticosteroid levels in both populations were higher in lactating than in non-lactating females. During periods of social stability, faecal corticosteroid concentrations increased in non-lactating females but not in lactating females as social status declined. Lactating Serengeti females had significantly higher faecal corticosteroid concentrations during periods with acute severe social stress than during periods without, indicating that the HPA axis is sensitive to social stimuli even in lactating females. So far few studies have used non-invasive monitoring methods for assessing social stress in freeranging animals. This study demonstrates for the first time, to the authors' knowledge, that corticosteroid concentrations may differ between periods with and without social stress for a free-ranging female mammal and that the modulating effect of social status may depend on reproductive status.

Predation on mammals by the grey-cheeked mangabeyLophocebus albigena

Behavioral observations were made on a habituated group of grey-cheeked mangabeys (Lophocebus albigena) in the Dja Reserve, Cameroon, using 5-min scan sampling periods at 15-min intervals. On two occasions male mangabeys captured and fed on mammalian prey, a squirrel and a bushbaby. Although grey-cheeked mangabeys have been studied at several sites, these are the first observations of carnivory by them.

Activity patterns, habitat use and mortality risks of mangabey males living outside social groups

Isolated individuals of most species are thought to be at a higher risk of mortality from starvation and predation than those in groups. However, supporting evidence remains scant. As part of a broader study of grey-cheeked mangabey,Lophocebus albigena , dispersal in Kibale National Park, Uganda between October 1996 to February 1999, we investigated starvation and mortality risks within a sample of 27 radiotagged mangabey males. Each male was contacted on average three times a week to collect data on spatial relations, habitat use, activity patterns and fates. We assessed risks by comparing data from ‘isolated’ (≥200m from the group) males with data from ‘within-group’ males. We assessed starvation risks by comparing proportions of time spent in feeding, foraging and moving, comparing rates of movement during focal samples, and comparing proportions of time that males spent in different habitat types and canopy levels. We assessed mortality risks by comparing scanning rates and mortality events of within-group and isolated males. Isolated males spent significantly more time scanning, but we could detect no other significant differences in time allocated to different activities by the two categories of males. Nor could we detect differences between within-group and isolated males in the use of habitat types and canopy levels. At least three of five established mortality events involved males known to be isolated from groups. Given that males were found isolated only 11% of the time, it is estimated that isolated males were at least 12 times as likely to die as within-group males. These results suggest that dispersing males may not face increased risk of starvation, but are at higher risk of mortality (primarily through predation by eagles) than nondispersing males.

Visual categorization of natural and abstract items in forest monkeys and humans

A preference for novelty paradigm was used to investigate whether mangabeys ( Lophocebus albigena), an arboreal non-human primate species, were able to discriminate and to categorize different visual stimuli belonging to natural (food items) and abstract (non-food items) categories. In a comparative perspective human subjects were tested with the same procedure and the same stimuli. Two out of four mangabeys and three out of the four humans showed significant preference for novelty when comparing food versus non-food items. Hence they discriminated between these two sets of items. The two mangabeys and one non-adult human subject sorted the food items in one category, showing no preference for novelty when comparing known and unknown food-items and different views of the same food items. In contrast the two adult human subjects who showed preference for novelty in the between-category, did not show preference for novelty when comparing known and unknown food-items but did show such a preference when comparing different views of the same food items. Compared to human performances, the results suggest that mangabeys are able to form at least a perceptual category of natural, ecologically relevant stimuli.

Correlates of Intergroup Transfer in Male Grey-cheeked Mangabeys

We studied factors influencing intergroup transfer in male mangabeys (Lophocebus albigena) inhabiting 7 social groups in Kibale National Park over a 2-year period. The sample consisted of 40 males including 36 that we captured and marked during the study. Intergroup transfers are movements between groups that culminate in either long-term (dispersal) or short-term (visits) residency by males in the new groups. Both dispersers and visitors had a greater tendency to move into groups that contained higher numbers of estrous females than their prior group. Using averages of weekly group counts, we found a significant positive relationship between the number of estrous females in a group and the number of adult males in a group, but not between the number of nonestrous females and the number of adult males in the 7 study groups. There is no evidence that dispersal events were released by aggression. For a sample of males followed ≥2 mo before emigration, aggression did not increase just before emigration. Results suggest that spatiotemporal availability of estrous females is a major proximate factor influencing intergroup transfer in mangabeys. The results also suggest that dispersing males are more sensitive to relative numbers of estrous females than to measures of female availability such as operational sex ratio, socionomic ratio, and the number of excess females that take into account the potential for male-male competition.

Catarrhine photopigments are optimized for detecting targets against a foliage background.

The colour vision of many primates is trichromatic, whereas that of all other mammals is thought to be dichromatic or monochromatic. Moreover, the triplets of cone pigments in different catarrhines (Old World apes and monkeys) are strikingly similar in their spectral positions. We ask whether the selective advantage of trichromacy lies in an enhanced ability to find edible leaves or fruit. Further, we ask whether any factor in these two search tasks has constrained the particular set of cone spectral sensitivities observed in all catarrhines. We measured the spectral properties of the natural environments of six primate species in Uganda: Pan troglodytes, Cercopithecus mitis, Cercopithecus ascanius, Lophocebus albigena, Colobus guereza and Colobus badius. We concentrated on the fruit and leaves in their diets and the leaves of the trees that make up the background against which these diet items must be found. We plotted these measured stimuli in colour spaces appropriate for each primate species, and found that both frugivory and folivory are facilitated by the extra dimension of colour vision found in catarrhines but lacking in most other mammals. Furthermore, by treating the task of searching for food as a signal-detection task, we show that, of all possible combinations of cone sensitivities, the spectral positions of the actual primate pigments are optimal for finding fruit or young leaves against the background of mature leaves. This is because the variance of the chromaticities of the mature leaves is minimised in one channel of the primate's colour vision, so allowing anything that is not a mature leaf to stand out.

Mass Differences among Male Mangabey Monkeys Inhabiting Logged and Unlogged Forest Compartments

Abstract: Remedies for reduction in primate numbers following rainforest habitat disturbance must incorporate understanding of how disturbance affects population biology. Between July 1996 and July 1998, I captured 31 adult male grey cheeked mangabeys ( Lophocebus albigena) from groups occupying logged and unlogged forest in Kibale National Park, Uganda. Captured males were weighed, measured, fitted with radiocollars, and radiotracked three to five times a week. Males from groups in the unlogged forest were significantly heavier than males from the logged forest. Males that immigrated into the study groups were significantly lighter than resident males in unlogged forest, but they did not differ in mass from males in logged forest. I tested hypotheses that these differences were due to differences in age, skeletal size, and tissue mass. Results support the hypothesis that differences are due to differences in tissue mass. Results of further tests showed that differences in tissue mass were explained more by differences in habitat type than by rates of male turnover. There was no evidence to suggest that differences in tissue mass were related to differences in costs of travel, crowding, or seasonal variation in habitat quality. My results suggest that selective logging may affect primate biomass, although these changes may or may not be reflected in estimates of primate density.

More on the meanderings of mangabeys: how to test whether bounded walks are random

Abstract1. Barrett & Lowen (1998)andWaser (1976)attempted to explain the net monthly and yearly displacements of Grey‐Cheeked Mangabeys using observed short‐term step lengths and assuming a random walk, with and without boundaries. This paper reanalyses their data.2. Analytic approaches require the root‐mean‐square step length, not the mean. However, a more flexible approach to making and testing predictions is Monte‐Carlo simulation. With a random walk long‐term displacements have a large variance, so a single observation is unlikely to disprove this null hypothesis.3. Restricting movement to a square lattice is a reasonable approximation even when rectangular boundaries are incorporated. Describing the boundary configuration accurately is more important.4. The observed non‐uniformity in turning angles should have been incorporated as it has a large effect on predicted net displacements, unless the arena is tightly constricted. Randomness of movement within a day can be distinguished from that between days. For Waser's population it makes sense to predict long‐term displacements using only long‐distance daily displacements.5. There are better approaches to establish both whether boundaries exist and whether movements follow a random walk.

Interdemic variation in mixed-species association patterns: common diurnal primates of Kibale National Park, Uganda

We used interdemic variation in the tendency to form mixed-species groups to examine the costs and benefits of association among the primates of Kibale National Park, Uganda. A year-long survey of six sites revealed that the amount of time that the five common diurnal primates [red colobus (Procolobus tephrosceles), black-and-white colobus (Colobus guereza), redtail monkeys (Cercopithecus ascanius), blue monkeys (Cercopithecus mitis), and grey-cheeked mangabeys (Lophocebus albigena)] spent in mixed-species groups varied dramatically among sites. In many cases, the proportion of time that species associated was positively related to their densities. By using detailed behavioral observations of redtail monkeys and red colobus made over 4 years (2660 h) at four sites, we were able to reject the null hypothesis that associations occur by chance for only one of four sites. However, a correlative approach exploring the costs and benefits of association suggests that ecological variables do influence association patterns. We found that redtail monkeys and red colobus overlapped in diet (19.2% of their foraging effort) and traveled further when in mixed-species groups than when alone. Having demonstrated this, we examined the applicability of the ecological constraints model for predicting the proportion of the time spent in mixed-species groups based on food availability. For this analysis we concentrated on red colobus from the site with 35 months of observation and demonstrated that their tendency to be in mixed- species groups was related to food availability. We used two methods to examine if mixed-species associations function to decrease predation risk. First, chimpanzees are known to prey heavily on red colobus, but rarely kill other primates. The time red colobus spent in mixed-species groups was correlated to chimpanzee density, but it was not for the other monkey species, suggesting that mixed-species groups serve to decrease predation risk. Second, when red colobus groups contain more infants and are presumably at the greatest risk of predation, they form mixed-species groups most often. These results demonstrate that the costs and benefits of mixed-species associations vary dramatically over small spatial and temporal scales. If such variation is generally the case, then studies conducted at different locations or different times could easily highlight the importance of difference selective agents in favoring mixed-species associations.