Vertical stratification of feeding by Japanese beetles within linden tree canopies: selective foraging or height per se?

Adult Japanese beetles, Popillia japonica Newman (Coleoptera: Scarabaeidae), tend to aggregate and feed most heavily in the upper canopy of host plants, defoliating them from the top down. We examined characteristics of linden (Tilia cordata L.) foliage from different canopy zones and tested two hypotheses that might account for vertical stratification of feeding by this vagile, polyphagous folivore. In the field, P. japonica caused 4-12 times more damage to upper canopy leaves in full sunlight than to lower canopy leaves in sun or shade. However, this within-tree pattern apparently cannot be explained by differences in nutritional parameters (protein, water, and sugar content) or "defensive" properties (toughness, tannins) of leaves. Furthermore, beetles did not discriminate between foliage from different canopy zones in laboratory choice tests, nor were fecundity or longevity higher for beetles fed upper canopy, sun-exposed leaves. Clonal grape plants suspended from vertical posts in the field at 3.65, 1.83 or 0 m above ground showed a top-down defoliation pattern identical to that seen in linden trees. This suggests that the height of foliage per se strongly affects initial orientation and attack by P. japonica. Recent related studies have shown that both sexes of P. japonica are strongly attracted to host volatiles induced by feeding of conspecific beetles. We suggest that Japanese beetles begin to feed in the upper canopy for reasons unrelated to host nutritional variation (e.g., behavioral thermoregulation, visual orientation to the host silhouette), and that top-down defoliation follows as additional beetles are attracted to feeding-induced volatiles acting as aggregation kairomones.