Most forest birds include arthropods in their diet, sometimes specializing on arthropods that consume plant foliage. Experimental tests of whether bird predation on arthropods can reduce plant damage, however, are few and restricted to relatively low-diversity systems. Here, we describe an experimental test in a diverse tropical forest of whether birds indirectly defend foliage from arthropod herbivores. We also compare how the indirect effects of bird predation vary with different levels of foliage productivity in the canopy vs. the understory. For three Neotropical tree species, we observed that birds decreased local arthropod densities on canopy branches and reduced consequent damage to leaves. In contrast, we observed no evidence of bird-arthropod limitation on conspecific saplings in the less productive understory of the same forest. Our results support theory that predicts trophic cascades where productivity is high and suggest that birds play an important role in Neotropical communities by means of their indirect defense of some canopy tree species.F or decades, ecologists have debated the circumstances under which a predator limits its prey's consumption of organisms in lower trophic levels (i.e., a predator-driven trophic cascade) (1-4). One body of theory predicts such cascades will occur in terrestrial systems with high plant productivity (5, 6). Opposing theory predicts that cascades will not occur in terrestrial systems and only in low-diversity aquatic or marine systems (7). Proponents of the latter theory suggest that higher diversity in terrestrial systems leads to diffuse food webs, rendering trophic levels nonexistent (7). Field experiments demonstrate that insectivorous birds can limit arthropod abundances and decrease damage to plants, but these tests have been conducted in settings with relatively low tree species diversity such as temperate forests (8-11) or agricultural systems (12, 13). Along with high tree species diversity, tropical forests support a high diversity and biomass of leaf-chewing arthropods (14) as well as a high biomass of birds that consume them (15). In a lowland forest of Panama, we used canopy crane access (16) to test the hypotheses that (i) birds limit arthropod densities and consequent herbivore damage, and (ii) the effects of bird predation are strongest where foliage production rates are high.For one year, we observed how the local density and taxonomic composition of the arthropod community responded to the absence of bird predation and also assessed changes in herbivore damage. We estimated and compared these quantities on control branchesÍsaplings where birds had access to foliage and on branchesÍsaplings in experimental exclosures where foliage was inaccessible to birds. A within-site comparison of canopy branches and conspecific understoryÍedge saplings allowed us to investigate the effects of bird predation across a 3-fold vertical gradient of foliage production.
MethodsA canopy crane provided forest canopy access in a dry, semideciduous lowland tropical for...