Quantity and quality of seed dispersal by a large arboreal frugivore in small and large Atlantic forest fragments

Seed dispersal is a key process driving the structure, composition, and regeneration of tropical forests. Larger frugivores play a crucial role in community structuring by dispersing large seeds not dispersed by smaller frugivores. We assessed the hypothesis that brown howler monkeys (Alouatta guariba clamitans) provide seed dispersal services for a wide assemblage of plant species in both small and large Atlantic forest fragments. Although fruit availability often decreases in small fragments compared with large ones, we predicted that brown howlers are efficient seed dispersers in quantitative and qualitative terms in both forest types given their high dietary flexibility. After a 36-month study period and 2,962 sampling hours, we found that howlers swallowed and defecated intact the vast majority of seeds (96%-100%) they handled in all study sites. Overall, they defecated ca. 315,600 seeds belonging to 98 species distributed in eight growth forms. We estimated that each individual howler dispersed an average of 143 (SD = 49) seeds >2 mm per day or 52,052 (SD = 17,782) seeds per year. They dispersed seeds of 58% to 93% of the local assemblages of fleshy-fruit trees. In most cases, the richness and abundance of seed species dispersed was similar between small and large fragments. However, groups inhabiting small fragments tended to disperse a higher diversity of seeds from rarely consumed fruits than those living in large fragments. We conclude that brown howlers are legitimate seed dispersers for most fleshy-fruit species of the angiosperm assemblages of their habitats, and that they might favor the regeneration of Atlantic forest fragments with the plentiful amount of intact seeds that they disperse each year.