When population growth intensifies intergroup competition, female colobus monkeys free-ride less.

Intergroup aggression often results in the production of public goods, such as a safe and stable social environment and a home range containing the resources required to survive and reproduce. We investigate temporal variation in intergroup aggression in a growing population of colobus monkeys (Colobus vellerosus) to ask a novel question: "Who stepped-up to produce these public goods when doing so became more difficult?". Both whole-group encounters and male incursions occurred more frequently as the population grew. Males and females were both more likely to participate in whole-group encounters when monopolizable food resources were available, indicating both sexes engaged in food defence. However, only females increasingly did so as the population grew, suggesting that it was females who increasingly produced the public good of home range defence as intergroup competition intensified. Females were also more active in male incursions at high population densities, suggesting they increasingly produced the public good of a safe and stable social environment. This is not to say that males were chronic free-riders when it came to maintaining public goods. Males consistently participated in the majority of intergroup interactions throughout the study period, indicating they may have lacked the capacity to invest more time and effort.

Serologic evidence for novel poxvirus in endangered red Colobus monkeys, Western Uganda.

Enzyme-linked immunosorbent assay, Western blot, and virus neutralization assays indicated that red colobus monkeys in Kibale National Park, western Uganda, had antibodies to a virus that was similar, but not identical, to known orthopoxviruses. The presence of a novel poxvirus in this endangered primate raises public health and conservation concerns.

Temporal dynamics of nutrition, parasitism, and stress in colobus monkeys: implications for population regulation and conservation.

The need to develop conservation plans calls for the ability to identify ecological factors that influence population density. Because stress is known to affect fecundity and mortality, increasing stress may provide a warning of potential population declines. We examined the effects of temporal variation in nutrition and parasitism on stress in endangered red colobus monkeys in Kibale National Park, Uganda. First, we tested the hypothesis that parasitism and nutrition would individually affect stress levels. We found that periods of poor-quality diet corresponded with an increase in cortisol. Similarly, increases in parasite infections were associated with increased cortisol. Next, we predicted that a poor-quality diet would facilitate increased parasite infections, and that together, they would lead to amplified stress. However, we found no support for such amplification, likely because the quality of the diet had little effect on parasite infections. Third, we tested whether individuals in a larger group were subject to food stress due to greater within-group competition, which would intensify nutritional stress and parasitism, and lead to reduced reproduction. Although we found no evidence to support a group size effect on parasites, cortisol levels in the large group tended to be higher than those in the small group, and the large group had fewer infants per female. The results suggest that parasitism and poor nutrition lead to increased stress which, because they are known to be associated with reduced fecundity and increased mortality, may lead to population declines.

Controversy over the application of current socioecological models to folivorous primates: Colobus vellerosus fits the predictions.

We explore the applicability of the current socio-ecological model to characterize the social structure of Colobus vellerosus, a folivorous primate. The current socio-ecological model predicts that female social relationships should respond in predictable ways to food abundance and distribution and associated competitive regimes. It appears to successfully explain variation in social structure in some primate species; however, recent research indicates that several folivorous or folivore-frugivorous species seem to be exceptions. We present data on social relationships and social structure in two groups of C. vellerosus over 15 months at Boabeng-Fiema, Ghana. As predicted by the model, our results indicate the co-occurrence of 1) low levels of grooming between female C. vellerosus when compared with other species, 2) an absence of female coalitions over food, and 3) female dispersal. Taken together, these traits suggest a "Dispersal-Egalitarian" species. However, our results also indicate female-female affiliation was higher than male-female affiliation, which was more indicative of a "female resident" species. Our data also suggests inter-sexual affiliation varied among groups. This variation in inter-sexual affiliation could be due to variation in the intensity of infanticidal threats between groups. The combination of these social characteristics lead us to conclude that C. vellerosus' social structure is largely congruent with the ecological indicators of food distribution and female competitive regime that we have previously documented, which indicated high quality foods were not monopolizable or usurpable and behavioral evidence of within-group contest competition (i.e. supplantations over food) was rare. But the combination of higher female-female affiliation (compared to male-female) and female dispersal is difficult to explain in light of predictions, unless future work reveals female residency is more predominant than female dispersal in our population. We also discuss reasons why some folivores do not appear to fit the predictions of the current socio-ecological model.

Do food availability, parasitism, and stress have synergistic effects on red colobus populations living in forest fragments?

Identifying factors that influence animal density is a fundamental goal in ecology that has taken on new importance with the need to develop informed management plans. This is particularly the case for primates as the tropical forest that supports many species is being rapidly converted. We use a system of forest fragments adjacent to Kibale National Park, Uganda, to examine if food availability and parasite infections have synergistic affects on red colobus (Piliocolobus tephrosceles) abundance. Given that the size of primate populations can often respond slowly to environmental changes, we also examined how these factors influenced cortisol levels. To meet these objectives, we monitored gastrointestinal parasites, evaluated fecal cortisol levels, and determined changes in food availability by conducting complete tree inventories in eight fragments in 2000 and 2003. Red colobus populations declined by an average of 21% among the fragments; however, population change ranged from a 25% increase to a 57% decline. The cumulative basal area of food trees declined by an average of 29.5%; however, forest change was highly variable (a 2% gain to a 71% decline). We found that nematode prevalence averaged 58% among fragments (range 29-83%). The change in colobus population size was correlated both with food availability and a number of indices of parasite infections. A path analysis suggests that change in food availability has a strong direct effect on population size, but it also has an indirect effect via parasite infections.

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.

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 immature's 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.