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.     

Activity and Ranging Patterns of <Emphasis Type="Italic">Colobus angolensis ruwenzorii</Emphasis> in Nyungwe Forest,Rwanda: Possible Costs of Large Group Size

With group sizes sometimes >300 individuals, the Angolan black-and-white colobus (Colobus angolensis ruwenzorii) population in Nyungwe Forest, Rwanda is an intriguing exception to the tendency for folivores to live in smaller groups than expected relative to body size. Researchers have hypothesized that the unusually high quality of foliage at Nyungwe allows colobus there to avoid intragroup feeding competition, releasing constraints on the formation of large groups (Fimbel et al., 2001). We collected data on the activity and ranging patterns of a >300-member Nyungwe colobus group and compared our results to those from smaller groups in other black-and-white colobus (Colobus spp.) populations. Colobus at Nyungwe spent far more time feeding and moving (62%) and far less time resting (32%) than black-and-white colobus at any other site. The annual home range of the Nyungwe colobus was also many times larger (95% minimum convex polygon: 20.7 km ² ; 95% fixed kernel: 24.4 km ² ) than those for other populations. We terminated our research after the group engaged in an unprecedented migration among black-and-white colobus by moving 13 km south of their former range. Our results suggest that intragroup scramble competition may be more intense than originally believed within the large colobus groups at Nyungwe and that long periods of resource renewal may be necessary after a large colobus group passes through an area, thereby potentially helping to explain their wide ranging patterns. We discuss the socioecological convergence between the Nyungwe colobus and Chinese snub-nosed monkeys (Rhinopithecus spp.) and suggest directions for future research on the unique black-and-white colobus population at Nyungwe.

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.     

Group size in folivorous primates: ecological constraints and the possible influence of social factors

The ecological-constraints model assumes that food items occur in depletable patches and proposes that an increase in group size leads to increased day range due to more rapid patch depletion. Smaller groups become advantageous when an increase in travel costs is not repaid by an increase in energy gained or some other fitness advantage. On the other hand, we also know that group size can be influenced by social factors. Here we contrast the diet and group size of red colobus (Procolobus badius) and black-and-white colobus (Colobus guereza) in Kibale National Park, Uganda to consider how ecological and social factors are affecting their group sizes. Subsequently, we examine whether the insights gained from this detailed comparison can provide an understanding of why the social organization and group size of mantled howlers (Alouatta palliata) and black howlers (A. pigra) differ. Two groups of red colobus and two groups of black-and-white colobus were studied over 10 months. Red colobus groups were larger (48 and 24) than black-and-white colobus groups (9 and 6). The two groups of red colobus overlap home ranges with the two groups of black-and-white colobus; 75% and 95% of their home ranges were within red colobuss home range. There was a great deal of similarity in the plant parts eaten by the two species and both species fed primarily on young leaves (red colobus 70%, black-and-white colobus 76%). In terms of the actual species consumed, again there was a great deal of similarity between species. The average dietary overlap among months for the two neighboring groups of red colobus was 37.3%, while the dietary overlap between the red colobus and the black-and-white colobus group that had its home range almost entirely within the home range of the red colobus groups averaged 43.2% among months. If ecological conditions were responsible for the difference in group size between the two colobine species, one would expect the density of food trees to be lower in the home ranges of the black-and-white colobus monkeys, since they have the smaller group size. We found the opposite to be true. Both black-and-white colobus groups had more food trees and the cumulative size of those trees was greater than those in the red colobuss home ranges. We quantify how these differences parallel differences in mantled and black howlers. The average group size for mantled howlers was 12.9 individuals, and for black howlers it was 5.3 individuals. We explore possible social constraints, such as infanticide, that prevent black-and-white colobus and black howlers from living in large groups.This revised version was published online in April 2005 with corrections to the cover date of the issue.     

The status ofColobus polykomos dollmani Schwarz

The two West African black-and-white colobus taxa,Colobus polykomos andC. vellerosus, are distinct species. A supposedly intermediate subspecies,C. polykomos dollmani, is actually a hybrid swarm, in whichC. vellerosus genes greatly predominate. We propose a hypothesis to explain this situation.     

Male takeover in Colobus vellerosus at Boabeng-Fiema Monkey Sanctuary, central Ghana

We describe a case of male takeover in the ursine black-and-white colobus (Colobus vellerosus). In April 2001, an all-male group attacked and eventually invaded our uni-male study group. Aggression increased following the takeover and the former resident male, severely wounded, became peripheral. The youngest immature received severe aggression from the new males but survived. The immatures mother intervened in most instances of this aggression. Eventually, the former resident male re-established relationships with some of the females and concurrently intervened to protect the immature. Defeated males that stay in their group can contribute to the protection of infants born during their tenure from infanticidal males. The females mated with the new males. Takeovers may be a means by which males acquire groups of females in C. vellerosus.     

Comparison of habitat quality and diet of Colobus vellerosus in forest fragments in Ghana

The forest fragments surrounding the Boabeng-Fiema Monkey Sanctuary (BFMS) in central Ghana shelter small populations of Colobus vellerosus. Little is known about these populations or the ability of the fragments to support them, despite the fact that these fragments represent potentially important habitat for the colobus in this region. We compared the diet of three groups of C. vellerosus in the fragments to two groups in BFMS. We also examined the differences in plant species composition and food abundance among fragments. The study took place from June to November 2003. Dietary data were collected using scan sampling. Plant species composition and food abundance were evaluated using tree plots and large tree surveys. As in BFMS groups, leaves constituted the highest proportion of the diet of fragment groups, yet the colobus in fragments fed on more lianas than did those in BFMS. Over 50% of all species observed eaten by colobus in the fragments were not consumed in BFMS groups during the same season. Food abundance was similar between fragments and BFMS, although species composition differed. There was no relationship between the density of colobus and the density of food trees or percentage of food species, suggesting that other factors may be influencing the number of colobus present. This study highlights the broad dietary range of C. vellerosus, which may be a factor allowing its survival in these fragments.     

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.     

Cross-species handling of colobine infants

At Tiwai Island, Sierra Leone, several instances of red colobus (Procolobus badius) adults handling white infants of black-and-white colobus monkeys (Colobus polykomos) were observed. These observations are described and their significance is discussed.     

Vigilance in ursine black and white colobus monkeys (Colobus vellerosus): an examination of the effects of conspecific threat and predation

Vigilance is thought to have evolved as an antipredator defense but, in primates, conspecific threat often better explains this behavior. We examined vigilance in one group of Colobus vellerosus inhabiting the Boabeng‐Fiema Monkey Sanctuary in Ghana. We aimed to (1) describe factors affecting vigilance in this population, and (2) examine the importance of predation avoidance and conspecific threat in explaining vigilance patterns. Because of a male takeover preceding the study, our focal group (B2) consisted of six adult males and three adult females. We collected 490 10‐min focal samples (82 hr) from all adults in the group (N=9) from July to November, 2001. To avoid predators, individuals should be more vigilant (i) with fewer neighbors, and (ii) in areas of the canopy with higher predation risks. Conspecific threats can be divided into extra‐ and intra‐group threats. Extra‐group threats should lead to higher vigilance levels (iii) during intergroup encounters, and (iv) in areas where the home range overlaps with other groups of colobus. Intra‐group threats should lead to greater vigilance (v) in the presence of neighbors and (vi) while feeding or occupying food patches (if resources are limiting). Our results best support the hypothesis that vigilance functions primarily to detect extra‐group, conspecific threats. Individuals were most vigilant during intergroup encounters and in areas of range overlap, and monthly vigilance rates were associated with monthly encounter rates. Individuals tended to scan less in proximity to familiar vs. unfamiliar group mates, suggesting that relationship quality may affect colobus vigilance. Finally, predation pressures or anthropogenic disturbances might have influenced vigilance, as individuals were more vigilant low in the canopy. However, this last result could also be due to the lower visibility because of dense foliage or to the fact that the monkeys have access to fewer escape routes when ranging lower in the canopy. Am. J. Primatol. 71:919–927, 2009. © 2009 Wiley‐Liss, Inc.     

Activity and Ranging Patterns of Guerezas in the Kakamega Forest: Intergroup Variation and Implications for Intragroup Feeding Competition

From March 1997 to February 1998, I investigated the activity patterns of 2 groups and the ranging patterns of 5 groups of eastern black-and-white colobus (Colobus guereza), aka guerezas, in the Kakamega Forest, Kenya. Guerezas at Kakamega spent more of their time resting than any other population of colobine monkeys studied to date. In addition, I recorded not one instance of intragroup aggression in 16,710 activity scan samples, providing preliminary evidence that intragroup contest competition may be rare or absent among guerezas at Kakamega. Mean daily path lengths ranged from 450 to 734 m, and home range area ranged from 12 to 20 ha, though home range area may have been underestimated for several of the study groups. Home range overlap was extensive with 49–83% of each group's range overlapped by the ranges of other groups. Despite the high level of home range overlap, the frequently entered areas (quadrats entered on 30% of a group's total study days) of any one group were not frequently entered by any other study group. Mean daily path length is not significantly correlated with levels of availability or consumption of any plant part item. Mean daily path length is also not significantly correlated with group size, though the largest group did have the longest mean daily path length. This finding suggests that intragroup scramble competition may have been rare or absent among guerezas at Kakamega except perhaps in the largest group, which was unusually large.     

Test of the ecological-constraints model on ursine colobus monkeys (Colobus vellerosus) in Ghana

For group-living mammals, the ecological-constraints model predicts that within-group feeding competition will increase as group size increases, necessitating more daily travel to find food and thereby constraining group size. It provides a useful tool for detecting scramble competition any time it is difficult to determine whether or not food is limiting. We tested the ecological-constraints model on highly folivorous ursine colobus monkeys (Colobus vellerosus) at the Boabeng-Fiema Monkey Sanctuary in Ghana. Three differently sized groups were followed for 13 months and two others were followed for 6 months each in 2004-2005 using focal-animal sampling and ranging scans; ecological plots and phenology surveys were used to determine home-range quality and food availability. There was relatively little difference in home-range quality, monthly food availability, diet, adult female ingestion rates, and rate of travel within food patches between the groups. However, home-range size, day-range length, and percent of time spent feeding all increased with group size. We performed a single large test of the ecological-constraints model by combining several separate Spearman correlations, each testing different predictions under the model, using Fisher's log-likelihood method. It showed that the ecological-constraints model was supported in this study; scramble competition in this population is manifesting in increased ranging and time spent feeding. How costly this increased energy expenditure is for individuals in larger groups remains to be determined.     

Effects of habitat disturbance on the behavioral ecology and demographics of the Tana river red colobus (Colobus badius rufomitratus)

Annual surveys in 1985–1987 revealed that, since 1975, the total population of the Tana River red colobus (Colobus badius rufomitratus)declined by approximately 80%. An intensive study in 1986–1988 of two groups of colobus in the Tana River Primate National Reserve indicated that habitat disturbance from the changing river course and shifting agricultural practices were primarily responsible for the decline. Clearcutting around Mchelelo forest in the late 1960s compressed colobus populations to levels probably above the carrying capacity. Between 1975 and 1986 primate population density declined dramatically, the number of red colobus groups in Mchelelo forest decreased by half, and the size of the remaining group was greatly reduced. In 1986, there were fewer solitary colobus and small parties in the forest, harem male takeovers did not occur, infant survivorship increased, and demographic parameters generally had improved. The colobus groups in Mchelelo in 1973–1975, living at higher densities, showed different feeding and ranging behaviors than 1986–1988 groups. Range size was smaller in 1975, range overlap occurred, and a greater portion of the forest was used per day and per month. Mature leaves accounted for a much higher proportion of the diet. Time spent feeding and resting was the same in both studies. Social organization in predominantly one- male groups was maintained and adult and subadult females transferred between groups.     

Effects of Forest Fragmentation on the Abundance of <Emphasis Type="Italic">Colobus angolensis palliatus</Emphasis> in Kenya’s Coastal Forests

We documented the occurrence and abundance patterns of Angola black-and-white colobus (Colobus angolensis palliatus) in 46 coastal forest fragments ranging from 1 ha to >1400 ha in the Kwale District, Kenya. In field surveys conducted in 2001, we also recorded forest spatial, structural, resource, and disturbance characteristics to determine the effects of habitat quality and fragmentation and the factors most critical to the continued survival of the little-known species. We tested 13 hypotheses to explain variation in patch occupancy and abundance patterns of Colobus angolensis palliatus in relation to habitat attributes. Minimal adequate models indicated that the occurrence of colobus in forest fragments is positively associated with fragment area and canopy cover, whereas the density of colobus in occupied fragments is attributable to forest area, the proportion of forest change over the previous 12 yr, and the basal area of 14 major food trees. Large-scale illegal extraction of major colobus food trees in the District for human resource use, in both protected and unprotected forests, together with ongoing forest clearance and modification, are the major threats to Colobus angolensis palliatus in Kenya.     

Proximity and grooming patterns reveal opposite-sex bonding in Rwenzori Angolan colobus monkeys (<Emphasis Type="Italic">Colobus angolensis ruwenzorii</Emphasis>)

Close proximity and social grooming are important bonding mechanisms in primates. These behaviors show the social structure of a species and many studies have found positive correlations between the degree of kinship and grooming and proximity. We used 1 year of data collected via instantaneous scan sampling on a large “supertroop” of Colobus angolensis ruwenzorii at Lake Nabugabo, Uganda, to examine partner preferences for grooming and nearest neighbors in each age-sex class. Little is known about this species, so we based our hypotheses on congeners. Of the five species of black-and-white colobus, data on sex-biased dispersal patterns are available for three (C. guereza, C. vellerosus, and C. polykomos), all of which show male-biased dispersal with occasional female dispersal. We thus predicted that female C. a. ruwenzorii would be more strongly bonded than males, showing greater proximity and grooming. We did not expect bonding between the sexes since congeners do not show this pattern. We found that among adult dyads, males and females were more likely to be found in loose proximity, and to groom, than would be expected given group composition. Conversely, both males and females had relatively weak same-sex relationships. Between the sexes, adult males had higher proximity and grooming indices with adult females without infants than with females with infants. These observations indicate that this subspecies is cross-bonded and that both sexes may disperse. Furthermore, our findings suggest that the social organization and social structure of C. a. ruwenzorii differ greatly from other black-and-white colobus species.     

Monkey Abundance and Social Structure in Two High-Elevation Forest Reserves in the Udzungwa Mountains of Tanzania

The effects of human activity on population and social structure are a pantropical concern for primate conservation. We compare census data and social group counts from two forests in the Udzungwa Mountains, Tanzania. The main aim is to relate differences within and between the forests to current theory on the effect of human disturbance on primate abundance and group size. The survey reveals the presence of the restricted-range red colobus, Procolobus gordonorum, in New Dabaga/Ulangambi Forest Reserve (NDUFR). The primate community of NDUFR is impoverished compared to that in Ndundulu forest. Red colobus and black-and-white colobus (Colobus angolensis palliatus) abundance and group size are lowest in NDUFR. Fission-fusion of red colobus social groups may be occurring in previously logged areas of both forests. Our observations are consistent with current theory on the effect of habitat degradation and hunting on primates, but the relative effects of the 2 factors could not be differentiated. We pooled the results with previous data to show that abundance of red colobus in the Udzungwa Mountains is lowest at high elevations. Low red colobus group sizes appear to be related to human activity rather than elevation. Black-and-white colobus and Sykes monkeys (Cercopithecus mitis) show no relationship with elevation. Future studies will require more detailed information on vegetation, diet and ranging patterns to interpret fully intraspecific variation in population demography and social structure in the Udzungwa Mountains.     

Aspects of Male Competition in <Emphasis Type="Italic">Colobus vellerosus</Emphasis>: Preliminary Data on Male and Female Loud Calling,and Infant Deaths After a Takeover

Male Colobus vellerosus are the main participants in intergroup encounters, and lead incursions in neighboring groups during which they attack infants. Extragroup copulations, all-male groups, and male takeover occur in the species. Here, we provide additional information on behaviors associated with male reproductive competition in Colobus vellerosus. We examined 1 resident male loud calling and participation in intergroup encounters in relation to a takeover. We also report a second case of takeover that led to the death of the former resident male and the death of 2 male infants, presumably as a result of aggression from the all-male group. The new resident male wounded the third infant of the group, which apparently died after its mother abandoned it. During the period characterized by the attacks on the infant and after its disappearance, females initiated and participated in loud call bouts with the new resident male. We examine the possible functions of female loud calling, and suggest that in this context, it might force the resident male to call along to indicate his presence.     

Primate surveys in Côte d'Ivoire's Sassandra–Bandama interfluvial region with notes on a remnant population of black‐and‐white colobus

We conducted surveys in Côte d'Ivoire's Bandama–Sassandra interfluvial region to assess the abundance of primates and investigate further a population of black‐and‐white colobus monkey whose taxonomic position has been debated. Primates were encountered in five of eight forests surveyed. The most common primates were Campbell's guenons and lesser spot nosed monkeys. Roloway monkeys were located in one reserve (Dassioko FR), sooty mangabeys in two (Dassioko and Port Gauthier FR) and olive colobus in three reserves (Dassioko, Port Gauthier, Bakadou) as well as one forest patch was considered a sacred grove (Grébouo I FG). Red colobus and chimpanzees were never observed. The only black‐and‐white colobus encountered were located in a forest adjacent to the Grébouo I village. The coat of this population is similar to that of Colobus vellerosus; however, vocal data are more complex. In one vocal attribute, the Grébouo I colobus is similar to Colobus polykomos while in another it is similar to C. vellerosus. We hope that by investigating the taxonomy of the Grébouo 1 colobus population, highlighting the conservation value of sacred forests and advocating for more active law enforcement in protected reserves, we can prevent the extirpation of all primates within the Bandama–Sassandra interfluvial zone region.     

Pattern, Distribution, and Function of Greeting Behavior Among Black-and-White Colobus

Various species of primates engage in greeting, a ritualized pattern of nonaggressive behavior that usually occurs during a reunion. Black-and-white colobus (Colobus guereza) perform overhead mounting, mounting, and embracing behavior soon after an aggressive act and in nonagonistic situations. We studied the pattern, distribution, and function of the greeting behavior in 2 captive groups of black-and-white colobus. Overhead mounting was the most frequent pattern, accounting for >60% of all greetings (N=333). In nonagonistic situations, younger subordinate individuals greeted an older dominant individual more frequently than vice versa. A dominant male in a small multimale group frequently initiated contacts with adult females, though he was the oldest in the group. Conversely, the dominant male in a large 1-male group never greeted group members. Immediately after performing the greeting, the greeter groomed the recipient in more than half the cases. In the large group, greeting frequency correlates positively with the age difference between the pair; however, there is no correlation between the relatedness, affiliation frequency, or aggression frequency of a pair in either the small or large group. The results are consistent with the hypothesis that greeting behavior functions as a tension-reducing mechanism in nonagonistic situations. There is no evidence, however, that greeting functions to express social status or to attract a mate. In addition, the frequent greeting by the adult male in the small multimale group may indicate that individuals affirm social bonds via greeting behavior.     

Group Size and Composition of Colobus guereza in Kyambura Gorge, Southwest Uganda, in Relation to Chimpanzee Activity

We documented and assessed the influence of chimpanzee activity on group size and composition of Colobus guereza in Kyambura Gorge, southwest Uganda, from July to September, 1994 and in February and March 1996. The population density of colobus is very high: 347 individuals per km ². Density differed outside activity centers of chimpanzees (525 individuals per km ² ) and within the centers (186 per km ² ). We identified a total of 24 colobus groups, ranging between 3 and 13 individuals. Of the 24 groups, 22 were one-male groups. Groups were smaller and the percentage of subadults and juveniles was lower in groups within chimpanzee activity centers. Estimates of home ranges are between 1.7 and 6.2 ha, but neighboring home ranges overlapped 80%. Only a territory of approximately 0.5 ha was defended by the alpha male.