Resolving the paradox of clumped seed dispersal: positive density and distance dependence in a bat-dispersed species.

One of the hypothesized benefits of seed dispersal is to escape density- and distance-responsive, host-specific, natural enemies near maternal plants where conspecific seed and seedling densities are high. Such high conspecific neighbor densities typically result in lower offspring growth and survival (i.e., negative density-dependent effects), yet many dispersal modes result in clumped seed distributions. New World leaf-nosed bats transport fruits to their feeding roosts and deposit seeds, thereby creating high-density seed/seedling patches beneath feeding roosts in heterospecific trees away from maternal trees, which seemingly nullifies a key benefit of seed dispersal. Such dispersal may still be adaptive if negative density-dependent effects are reduced under feeding roosts or if the benefit of being dispersed away from maternal trees outweighs negative effects of conspecific seed/seedling density below roosts. We mapped the entire post-germination population of a bat-dispersed tree species Calophyllum longifolium (Calophyllaceae) in a 50-ha plot on Barro Colorado Island, Panama in each of three successive years. We tested two hypotheses: (1) distance-dependent effects are stronger than density-dependent effects on seedling performance because seedlings far from conspecific adults are more likely to escape natural enemies even when at high densities and (2) negative density-dependent effects will be reduced far from vs. near conspecific adults. Density and distance were naturally decoupled, as expected. However, in contrast to our expectation, we found positive density effects on seedling survival and density-dependent effects did not differ with distance from conspecific adults. Both density and distance had positive effects on seedling survival when considered together, while only year had a significant effect on seedling growth. Thus, both being dispersed under bat feeding roosts and escaping the vicinity of conspecific adults were beneficial for C. longifolium seedling survival, supporting the directed dispersal and escape hypotheses, respectively. Despite resulting in high densities of conspecific seedlings, favorable habitat under bat feeding roosts and lack of negative density-dependent effects appear to provide evolutionary advantages in C. longifolium.