Tree leaf chemical characters: selective pressures by folivorous primates and invertebrates

Plants have evolved a variety of chemical means to deter herbivory. Several studies have documented that secondary compounds are strong deterrents to certain herbivores, while others have demonstrated that some herbivores ingest large quantities of these compounds without exhibiting deleterious effects. This inconsistent response suggests that plants have evolved compounds to deter specific herbivores. Based on a study in Kibale National Park, Uganda, we explored how two major groups of herbivores, invertebrates and colobus monkeys, respond to chemical characteristics of leaves: protein, attractive from a nutritional perspective, and alkaloids, saponins and cyanogenic glycosides, which are all plant defences, detering herbivory. The intensity that colobus monkeys fed on leaves of different tree species was determined by observations (1300 h), and invertebrate herbivory was indexed by collecting leaves from 20 species and digitizing tracings to quantifying invertebrate damage. Invertebrate damage to leaves varied among species (1.5–22.5%), but showed no relationship with saponin or protein content, or the presence or absence of alkaloids. Colobine foraging effort did not relate to the saponin and protein of leaf species, nor to the presence or absence of alkaloids. Prunus africana, the only species to test positive for cyanogenic glycosides, was fed on by colobus monkeys for 8.1% of their foraging time, but, as it occurred at low densities, it was the most preferred species. These results can be interpreted in different ways. First, it is possible that inactive compounds are retained because they increase the probability of producing new active compounds. Secondly, the indices used to evaluate compound effects may be inappropriate. For example, monkeys may only be able to tolerate a toxin to a specific threshold in a single feeding session, but our index of foraging effort was averaged over the year. Thirdly, it may be that these compounds play an active role with organisms not considered (e.g. prevent fungal attack). Finally, these compounds may serve some unknown function and selection may operate for that purpose.