Seed-spitting primates and the conservation and dispersion of large-seeded trees

Primate frugivory may reduce density-dependent predation on seeds and seedlings via effective seed dispersal. Accordingly, the tendency of cercopithecines to spit and scatter seeds > 4 mm wide could represent a prominent means of dispersal. However, the importance of seed-spitting may vary according to the life history adaptations of plants. Indeed, the actions of cercopithecines may be incongruent with the reproductive biology of plants that rely on large frugivores to swallow and defecate their seeds. This possibility raises conservation concerns because large frugivores are often susceptible to extirpation or extinction from hunting and habitat fragmentation. It is therefore important to determine if cercopithecines have a compensatory effect; that is, whether or not seed-spitting effectively conveys large seeds to recruitment sites. To test this concept, we used geospatial techniques to measure and analyze the dispersion of tree species dispersed by elephants, chimpanzees, and cercopithecines to different spatial extents. We studied adult trees of Balanites wilsoniana, Chrysophyllum gorungosanum, and Uvariopsis congensis in a 2.2-ha plot in Kibale National Park, Uganda. Despite the tendency of cercopithecines to spit the seeds of Uvariopsis congensis, adult trees were highly clumped, with a modal nearest-neighbor distance of < 5 m and a crown overlap of 1.5 m. Virtually identical results for Balanites wilsoniana and Chrysophyllum gorungosanum, the seeds of which are not spat, suggest that seed-spitting may be a poor mechanism of dispersal for some large-seeded plants.