Leaf surface specializations of birds and Arthropods in a Panamanian forest.

Most species of Panamanian lowland forest birds specialize on leaf undersurfaces when hunting foliage insects. The few species of leaf surface generalists and leaf upper surface specialists are omnivorous gleaners. We estimate that while over 90% of the avian understory insectivory is directed towards leaf undersurfaces, only 50% of canopy foliage insectivory is directed towards the undersides of leaves. In the low understory we found 70-80% of the arthropods on leaf undersides. The excess use of leaf-bottoms by understory birds may be a result of their greater visibility. It is hypothesized that less proficient insectivores are unable to take advantage of the greater effective density of underleaf insects because they can only efficiently attack the closest leaf surfaces; these closest surfaces will usually be the leaf tops from the branch on which the bird is perched. Alternatively, leaf-top specialists may have special foraging adaptations for overcoming the disadvantages of leaf-top foraging. These adaptations may involve attack behavior (Tachyphonus luctuosus) or searching behavior (Dacnis cayana). Dacnis often used leaf damage as a foraging cue; this may be the first report of a bird using leaf damage for searching for insects. The greater use of leaf upper surfaces by canopy birds may be influenced by four factors: greater seasonality of insects in the canopy favoring omnivores which may be less efficient insectivores; more insects on leaf tops; fewer planar leaf arrangements in canopy plants; or the greater visibility of leaf upper surfaces of the outer shell of foliage of massive trees. Based on the greater number of arthropods on leaf bottoms in the dry season, the higher abundance of smaller insects on leaf bottoms, as well as the greater proportion of insects on leaf tops at cooler higher elevations, we suggest that arthropods prefer leaf bottoms in tropical areas for physiological, not predator avoidance reasons.