Tana river red colobus and crested mangabey: Results of recent censuses

Censuses of the Tana River red colobus (Colobus badius rufomitratus) and crested mangabey (Cercocebus galeritus galeritus), primate species endemic to riverine forests of the lower Tana River in eastern Kenya, showed significant population declines for both species between the mid-1970s and 1980s. Red colobus declined in terms of group size and number of groups per forest; the mangabey population declined only in numbers of groups per forest. There was no significant change in mean group age/sex composition between the two time periods for either species. Differences in the feeding and ranging ecology of the two species may explain why the mangabey population suffered a less severe decline than the red colobus. Similarities in census results over 3 years in the late 1980s suggest that the primate populations are no longer decreasing.     

Population Sizes and Distribution of Primates in the Lower Tana River Forests,Kenya

We studied the population size and distribution of diurnal primates in the lower Tana River forests, Kenya. They are the only remaining habitats for 2 threatened primates: the Tana River red colobus (Procolobus rufomitratus) and the Tana River crested mangabey (Cercocebus galeritus galeritus). We conducted censuses in 73 forest patches from January through March 2001. We estimate population size of the red colobus to be 788 individuals in 82 groups and that of the crested mangabeys to be 2,070 individuals in 59 groups. The data suggest that over a 7-year period (1994-2001), there was an 18% increase in the crested mangabey population and a 5% decline in red colobus numbers. Further, the red colobus range has expanded both north and south, whereas that of crested mangabeys has only expanded south. Fifty-six percent of crested mangabeys and 46% of red colobus groups were inside the Tana River Primate National Reserve (TRPNR). Other primates encountered included 170 groups of Sykes' monkeys (Cercopithecus mitis), 70 groups of yellow baboons (Papio cynocephalus) and 4 groups of grivets [Chlorocebus (Cercopithecus) aethiops]. Mean group densities of the 2 endangered primates and of baboons were higher inside than outside the TRPNR, reinforcing the importance of TRPNR for their conservation. An intervention program is required to stem further decline in the red colobus population and to protect small isolated groups in forest patches outside TRPNR.     

Changes in forest fragment sizes and primate population trends along the River Tana floodplain, Kenya

The Tana River forest patches significantly decreased in total area by 1875 ha (34.5%), from 5439 to 3564 ha between 1979 and 2000. The area covered by forests outside the Tana River Primate National Reserve declined by 1246 ha (38%) from 3283 to 2037 ha. This loss was higher than that inside the reserve, where total forest area reduced by 629 ha (29%) from 2156 to 1527 ha. The numbers of Tana River red colobus (Procolobus rufomitratus) and Tana crested mangabey (Cercocebus galeritus galeritus) groups were significantly correlated with forest sizes suggesting that both red colobus and crested mangabeys are likely to be affected by forest loss and fragmentation. However, comparison of the 1974/75 and 2001 census data did not reveal any significant change in the number of groups of either the red colobus or crested mangabey. The two endangered primates may have developed strategies to cope with a shrinking habitat.     

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

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

Primate census and habitat evaluation in the Tana delta region, Kenya

Nineteen indigenous forest patches in the Tana River delta region, Kenya were surveyed between October and November 2000 for primates and habitat disturbance. Special emphasis was placed on the endangered Tana River red colobus (Procolobus rufomitratus Peters) and crested mangabeys (Cercocebus galeritus galeritus Peters), both of which are endemic to the region. Habitat disturbances evident in the forests included cutting of trees, harvesting of thatching material, firewood collection, dyke construction, cultivation, palm wine tapping and charcoal burning. A total of 85 groups of five primate species were counted. These comprised eighteen, ten, 22, 31 and four groups of red colobus, crested mangabey, baboons (Papio cynocephalus L.), sykes monkeys (Cercopithecus mitis Wolf) and vervet monkeys (Cercopithecus aethiops L.), respectively. A wider distribution of red colobus and crested mangabeys than was documented previously was noted, implying that they are probably more abundant than hitherto reported. It is hypothesized that extensive studies on some fauna considered endangered world‐wide would probably redefine their conservation status. Future studies in the lower Tana River region should cover the previously unsurveyed forests and focus on ways of curbing forest destruction.     

Endangered species in small habitat patches can possess high genetic diversity: the case of the Tana River red colobus and mangabey

We used mtDNA sequence data from the Tana River red colobus and mangabey to determine how their population genetic structure was influenced by dispersal and habitat fragmentation. The colobus and mangabey are critically endangered primates endemic to gallery forests in eastern Kenya. The forests are a Pliocene–Pleistocene refugium that has recently undergone significant habitat loss and fragmentation due to human activities. We expected both primates to exhibit low levels of genetic diversity due to elevated genetic drift in their small populations, and to show a strong correspondence between genetic and geographic distance due to disruption of gene flow between forests by habitat fragmentation. Additionally, because mangabey females are philopatric, we expected their mtDNA variation to be homogeneous within forest patches but to be heterogeneous between patches. In contrast, colobus have a female-biased dispersal and so we expected their mtDNA variation to be homogeneous within and between forest patches. We found high levels of haplotype and nucleotide diversity as well as high levels of sequence divergence between haplotype groups in both species. The red colobus had significantly higher genetic variation than the mangabey did. Most of the genetic variation in both primates was found within forest fragments. Although both species showed strong inter-forest patch genetic structure we found no correspondence between genetic and geographic distances for the two primates. We attributed the high genetic diversity to recent high effective population size, and high sequence divergence and strong genetic structures to long-term habitat changes in the landscape.     

Predation on mammals by chimpanzees in the montane forest of Kahuzi, Zaire

The rate of predation on mammals by chimpanzees was determined from carcasses and from fecal specimens found on fresh trails during a 16-month period in the montane forest of Kahuzi-Biega National Park, Zaire. A unit-group of semi-habituated chimpanzees, composed of 22 – 23 individuals including 8 adult or adolescent males, appeared to kill about 18 – 30 mammalian prey (16 – 28Cercopithecus monkeys) per year, if the multiple kills by chimpanzees were not considered. A juvenile l'Hoest's monkey was recorded for the first time as the prey of chimpanzees in this study. Predation occurred in the late dry and the early rainy seasons, when the diversity of ripe fruits was the highest during the year. The Kahuzi chimpanzees tended to kill mammals less frequently but to killCercopithecus monkeys more frequently than chimpanzees in other habitats. The absence of red colobus monkeys, which are the most frequent prey in Gombe, Mahale, and Tai, might be responsible for the low predation rate. However, the estimated rate of predation onCercopithecus monkeys is the highest record among various chimpanzee habitats. At least 11 – 18% of theCercopithecus population seemed to be lost annually as a result of being killed by chimpanzees. Chimpanzees may be the most important predators on these monkeys in the absence of leopards at Kahuzi. The examination of fecal samples and carcasses suggested that adult (probably male) or adolescent chimpanzees tended to eat juvenile or subadult monkeys most frequently, as is also seen for chimpanzees in Gombe, Mahale, and Tai.     

Rates of predation on mammals by gombe chimpanzees, 1972–1975

Rates of chimpanzee predation on mammals are calculated using data on 75 kills recorded during focal observation in Gombe National Park, Tanzania, from January 1972 to April 1975. The chimpanzees were members of two study communities (Kanyawara, or Northern, and Kahama, or Southern, community), and were observed as focal individuals for 14,583 hr by more than 30 researchers and field assistants working in pairs. The rate of predation by females was too low to allow reasonable estimates. For males, the mean rate of killing during the study period was 0.31 kills per male per 100 hr (N=17 males), or 4.65 kills per 100 hr in the two communities. In contrast to results from Mahale Mountains, there was no difference in predation rate between wet and dry seasons. However, predation rates varied over time, increasing by four times between the first three and last four seasons of the sample period. In an average year the 15 adult and subadult male chimpanzees are calculated to have killed 204 prey per year in an area of 16 km², varying between 99 and 420 prey per year in periods of low and high predation rate. Red colobus were the most frequent prey, followed by bushpig and bushbuck. Predation rates varied greatly on different prey species, and were not related to either the proportion of time spent within 200 m of male chimpanzees, or to their population densities. In relation to encounter rates and population density, baboons, blue monkeys, and redtail monkeys were killed at a fraction of the rate of red colobus monkeys, which suffered severe mortality from chimpanzee predation. Predation on bushpig and bushbuck also appears to have been high in relation to population density. The amount of food provided by predation is estimated to have averaged 600 kg per year for chimpanzees in the two communities (totalling 14–17 adult or subadult males, 18–20 adult of subadult females, and about 19 infants or juveniles). This suggests that adult males consumed around 25 kg of meat per year, although any average figure undoubtedly masks considerable individual variation. Present data suggest that chimpanzees in Gombe and Tai National Park, Ivory Coast, prey on mammals at rates higher than other populations.     

Gastrointestinal parasites of critically endangered primates endemic to Tana River, Kenya: Tana River red colobus (Procolobus rufomitratus) and crested mangabey (Cercocebus galeritus)

We conducted fecal egg counts of gastrointestinal parasites of 2 critically endangered primates endemic to the forest of Tana River, Kenya. We aimed to use the fecal egg counts as proxies to quantify the prevalence of gastrointestinal parasites between the 2 primates. The Tana River red colobus (Procolobus rufomitratus) and crested mangabey (Cercocebus galeritus) are of similar body size, but their behavioral ecology is very different. We predicted that mangabeys would have a higher prevalence of parasites because they are mostly terrestrial omnivores, live in larger social groups, and therefore range widely. We detected 10 nematodes and 3 protozoans in mangabeys and 7 nematodes and 2 protozoans in colobus. We detected a higher number of different parasite species in individual mangabeys, and 4 of the 5 nematodes requiring intermediate hosts were found in mangabeys. The overall prevalence of parasites was higher for mangabeys, but this difference was not statistically significant. For colobus, we found a trend whereby the number of different parasite species in individual monkeys was higher in males and in lactating females. However, there was no difference in the prevalence of parasites between the sexes or between lactating and nonlactating females.     

Population Declines of Colobus in Western Uganda and Conservation Value of Forest Fragments

The processes of habitat loss and fragmentation are probably the most important threats to biodiversity. It is critical that we understand the conservation value of fragments, because they may represent opportunities to make important conservation gains, particularly for species whose ranges are not in a protected area. However, our ability to understand the value of fragments for primates is limited by the fact that researchers have conducted many studies in protected areas, which do not represent most fragments, and studies are typically short term. Here we determine the long-term survival probability of red (Procolobus pennantii) and black-and-white colobus (Colobus guereza) inhabiting forest fragments outside of Kibale National Park, Uganda. Local communities use the fragments primarily for subsistence agriculture and fuelwood. We surveyed primate populations 3 times over 8 yr, made a total inventory of all trees 2 times, contrasted behavior of groups inhabiting 1 fragment with groups in the continuous forest, and judged the conservation value of the fragments by quantifying patterns of forest use by local people. Of the 20 fragments surveyed, 16 supported resident populations of colobus in 1995, 2 were cleared in 2000, and an additional 2 fragments were cleared by 2003. In 1995 we counted 165 black-and-white colobus, whereas in 2000 and 2003, we counted 119 and 75 individuals, respectively. Seven fragments supported red colobus in 1995, 11 in 2000, and 9 in 2003. In 2000 we counted 159 red colobus, while in 2003, we saw 145 individuals. For both species, activity patterns in continuous forest were similar to those in a fragment, with the exception that individuals in the fragment rested more. Colobus in the fragment ate more mature leaves than colobus in the continuous forest did. Fragments supported all the fuelwood needs of an average of 32 people who lived immediately adjacent to them, and partially supported families up to 3 farms away (ca. 400 m), representing 576 people. Intensive harvesting for fuelwood occurred when neighboring households engaged in beer brewing (an average of 9.6% of the households), gin distilling (8.8%), or charcoal production (14.5%). Overall, between 2000 and 2003, the average density of trees declined by 14 trees/ha (range = 0–60 trees/ha). If current rates of clearing continue, the probability that the fragments will continue to support colobus populations is low.     

Some observations on the ecology ofColobus badius temmincki,Abuko Nature Reserve,the Gambia,West Africa

Three groups of red colobus (Colobus badius temmincki) were observed at the Abuko Nature Reserve in the Gambia, West Africa between December 1973 and August 1974. The population of the three groups ranged from 24–40 animals with home ranges measuring 4.3–12.8 hectares. The red colobus fed on a wide variety of flora and their diet included fruit, flowers, shoots, and bark, as well as new and mature leaves. There was a change in diurnal activity patterns over the two seasons (dry and rainy), with more definitive activity peaks occurring during the dry season.     

Ecology of the Zanzibar Red Colobus Monkey: Demographic Variability and Habitat Stability

We examined the Zanzibar red colobus' (Procolobus kirkii) social structure and population dynamics in relation to the density, diversity and dispersion of food resources in ground-water forest and agricultural land, which we characterized in terms of red colobus food species density, diversity, basal area and dispersion. We used transect sampling and group follows to describe population dynamics and social systems. Two agricultural areas, SJF Shamba and Pete Village, had higher densities and more uniformly dispersed red colobus food tree species than those of the ground-water forest. Red colobus at these two sites had greater population densities and natality, and smaller home ranges than red colobus in the ground-water forest. However, these findings apply to a very small area of agricultural land (approximately 18 ha) that is contiguous with an area of the forest reserve having a high density of red colobus. It is not representative of agricultural areas elsewhere on Zanzibar which support much lower densities or no red colobus. Although agricultural areas contiguous with the forest reserve had high densities of red colobus, they appear to be very unstable. Within the agricultural areas, we observed higher intergroup variation in group size and composition, study groups that decreased dramatically in size and disappeared from the study site, significantly lower levels of juvenile recruitment, and red colobus food trees that exhibited definite signs of overbrowsing. This apparent instability in the subpopulation of red colobus utilizing agricultural systems probably reflects the lower basal area of food trees and the greater fragmentation of suitable habitat and floristic dynamics due to human activities in these areas. A fusion-fission social system occurred only in the ground-water forest subpopulation, which we hypothesize to be due to highly clumped food resources.     

Population dynamics of Japanese serow in relation to social organization and habitat conditions. II. Effects of clear-cutting and planted tree growth on Japanese serow populations

Population responses of Japanese serow (Capricornis crispus) to clear-cut logging and planted tree growth were studied by a direct count of the number observed in three areas, Takiyama (305 ha), Gentouziro (270 ha), and Tanokashira (324 ha), near Wakinosawa Village, Aomori Prefecture, Japan, during 1976–89. At Takiyama, a mature mixed forest ofThujopsis dolabrata var.hondai andFagus crenata decreased from 73 to 39% coverage of the area due to clear-cutting during 1978–83. The serow population maintained a relatively low stable density (3–6 individuals per km²) until 1985–87, and increased 5–10 years after the clear-cutting, reaching 9–10 km⁻² in 1988. At Gentouziro and Tanokashira, young plantations of Japanese cedar (Cryptomeria japonica) covered about half of each area. The population densities between 1980 and 1983 were relatively high (14–19 km⁻²), but declined thereafter, reaching 10–14 km⁻² in 1988–89. These results indicated that the density increase resulted from an improvement of food supply due to growth of scrub following the clear-cutting, and that the density decline resulted from a habitat change due to growth of planted Japanese cedar and a decrease in the food supply. In Wakinosawa Village, serow density began to increase 5–10 years after forest cutting, and the high population density, about three- or six-fold larger than that in mature forest, is expected to be maintained for about 20 years after logging.     

Comprehensive Conservation Profile of Tana Mangabeys

Forests along 60 kilometers of the lower Tana River, Kenya, provide habitat for one of the world’s top 25 most endangered primates, the Tana mangabey (Cercocebus galeritus). There is no current accurate estimate of the mangabey population, but a 1994 census estimated the population at 1,000–1,200. Their habitat has been severely degraded since then: visual estimates indicated that 30% of the forest area has been cleared and product use has increased in > 80% of forests surveyed. As the mean number of mangabey groups per forest is positively correlated with forest area and density of trees, this loss is damaging to the mangabey population. There has also been an increase in mangabey-human conflict, e.g., crop raiding, set traps, mangabeys chased by dogs. Mangabeys exhibit ecological flexibility, but behavioral data come from only a few mangabey groups. A new conservation approach is needed because past approaches, particularly the Tana River Primate National Reserve and a World Bank/Global Environment Facility Project, failed to protect the forests. The failure was mainly due to a disregard of the land-tenure issue within the Reserve, exclusion of local people from decision-making, and neglect of forests outside the reserve. Future actions must include community conservation programs and forest and corridor restoration. Research should focus on traditional management, status of primate groups in severely degraded forests, ecology of additional groups, and a population estimate to inform management as they implement more specific conservation strategies for the species.     

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

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

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.     

Demographic Structure of Zanzibar Red Colobus Populations in Unprotected Coral Rag and Mangrove Forests

More than half of the global population of the endangered Zanzibar red colobus (Procolobus kirkii) live outside the single major protected area on Zanzibar Island. We present data on the 2 largest, discrete subpopulations living in unprotected areas at extremes of the species’ range. We compare the size and structure of 11 groups, specifically 6 core groups inhabiting interior, mature forest with 5 peripheral groups living in disturbed/degraded edge habitats. Groups living in southern mangrove forest—a species-poor but more productive and less seasonal habitat than coral rag thicket—had larger group sizes and more heterogeneous age structure, were more stable, and had higher rates of infant survival than did groups in northern coral rag. Group size ranged from 5.5 ± 1.6 SD (the smallest reported for this species) in edge coral rag to 31.2 + 1.9 SD in core mangroves. Edge groups were significantly smaller than core groups in northern coral rag while in the south, where all groups had access to mangroves, we found no significant difference in mean group size between edge and core areas. Groups using mangroves exhibited frequent social play, an indicator of habitat quality, and had a higher ratio of births per female per year. We suggest that mangroves are an important refuge and possibly source habitat for Zanzibar red colobus. We urge the conservation of mangrove and remaining coral rag in the unprotected areas described here in an effort to sustain this endemic species throughout its range.     

Conservation and Survival Adaptations of Temminck’s Red Colobus <Emphasis Type="Italic">(Procolobus badius temmincki)</Emphasis>, in Senegal

In the Fathala Forest, Saloum Delta National Park, Senegal, in 1974–1976 and 1988–2002, we studied the northernmost populations of the endangered Temminck’s red colobus (Procolobus badius temmincki). Drastic habitat changes in the last 30 yr have reduced the forests by > 50% (75% in the gallery forests used by the red colobus). The woody species diversity decreased by > 30%. Despite these changes, the red colobus population only decreased from ca. 600 to ca. 500 individuals. We examined the ecological conditions and behavioral adaptations that may have influenced survival of the population. We found that 5 major adaptations have emerged in the last 30 yr that may have influenced the survival of red colobus at Fathala: (1) frugivory, (2) terrestriality, (3) tendency to form polyspecific associations with green monkeys, (4) tendency to frequent more open habitats, and (5) use of mangrove swamps for refuge and forage. By developing these adaptations, red colobus demonstrated their ability to persist in degraded habitats of the Fathala Forest. Preserving the forest should ensure the conservation of the red colobus population there.     

Interspecific interactions between wild pygmy chimpanzees (Pan paniscus) and red colobus (Colobus badius)

Interspecific interactions accompanied by physical contacts between wild pygmy chimpanzees (Pan paniscus) and red colobus (Colobus badius) were observed on three occasions at Wamba, Republic of Zaire. In all cases, the red colobus initiated the interactions by approaching the pygmy chimpanzees. Most of the pygmy chimpanzees, which were within 5 m of the red colobus, were juveniles or infants but the adult male pygmy chimpanzees never showed any interest in the red colobus. The red colobus groomed the chimpanzees in two cases, but the latter never groomed the former. No true aggressive interactions were observed between the two species. The lack of any evidence of hunting of red colobus through longitudinal studies of the pygmy chimpanzees of Wamba, together with the present observations, suggests that red colobus are probably not targets of hunting by the pygmy chimpanzees.     

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.