Population characteristics of howlers: Ecological conditions or group history

We examined the relative importance of ecological parameters-habitat productivity and seasonality-and group history-episodic predation, disease, and sudden habitat deterioration-to explain variation in the density and group structure of howlers (Alouatta spp.). We use data from a census of Guanocaste National Park, Costa Pica, and a literature review characterizing 80 howler populations. In Guanocaste National Park both habitat type and degree of protection affect howler density and group structure. Howlers were found at the highest density and in the largest groups in areas of semievergreen forest, which ecological sampling indicates have the most consistent level of food production. Differences in density between the sector of the park that first received protected status and more recently protected areas may be due partially to the degree of protection the areas received. We test the prediction that howler density and group structure would be influenced by habitat productivity as indexed by rainfall. Average group size and sex ratios differ among species, but female-to-immature ratios do not. Considering all censuses at one site to be independent, there are significant interspecific differences in density, with Alouatta pigra occurring at lower densities than the other species. In spite of such variability there is no relationship between annual rainfall and howler density and rainfall had a variable effect on group size depending on the level of independence that was considered. While such ecological comparisons are unrefined, e.g., rainfall must be used as a surrogate for habitat production, the fact that so few relationships were documented suggests that factors other than the ecological factors considered here are responsible for the observed differences in population characteristics. We suggest that much of the variability in howler population characteristics is related to events occurring in the recent history of the groups, such as habitat alteration, hunting, food tree crop failure, and disease.