Prey distribution,foliage structure,and foraging behavior of insectivorous birds in two oak species (<Emphasis Type="Italic">Quercus serrata</Emphasis> and <Emphasis Type="Italic">Q. variabilis</Emphasis>)

We examined foraging behaviors of four insectivorous bird species (Great Tit Parus major, Varied Tit P. varius, Long-tailed Tit Aegithalos caudatus and Japanese White-eye Zosterops japonica) on the two oak species (Q. serrata and Q. variabilis) in relation to fine-scale foliage structures, abundance, and distribution of folivorous insects and physical and chemical characteristics of leaves. There was no difference in the mean number of insects for each oak species, but there was a significant difference in the mean biomass. The distance from twigs or branches to leaf tips or blade-petiole junctions showed that leaves of Q. variabilis were more dispersed than those of Q. serrata. The different size distribution of insects influenced insect sizes selected and foraging maneuvers used by each bird species. Parus major and P. varius fed on beetles (low energy gain) with perch-gleaning (the most energy-saving maneuver) from the upper side of leaves on Q. variabilis, whereas they fed on large lepidopteran larvae (high energy gain) from the lower side of leaves on Q. serrata using a variety of maneuvers (including energy-consuming ones). Both A. caudatus and Z. japonica fed on insects from the lower side of leaves by hang-gleaning and reach-gleaning, respectively. These species appeared to be morphologically adapted to the energy-consuming but more effective maneuvers used to take preys from the undersides of leaves. We hypothesize that differences in foliage structures between closely related species of trees have evolved as a balance between defense against folivorous insects and attraction of insectivorous birds.