Global patterns of insect herbivory in gap and understorey environments, and their implications for woody plant carbon storage

Abstract

Insect herbivory is thought to favour carbon allocation to storage in juveniles of shade-tolerant trees. This argument assumes that insect herbivory in the understorey is sufficiently intense as to select for storage; however, understoreys might be less attractive to insect herbivores than canopy gaps, because of low resource availability and - at temperate latitudes - low temperatures. Although empirical studies show that shade-tolerant species in tropical forests do allocate more photosynthate to storage than their light-demanding associates, the same pattern has not been consistently observed in temperate forests. Does this reflect a latitudinal trend in the relative activity of insect herbivory in gap versus understorey environments? To date there has been no global review of the effect of light environment on insect herbivory in forests. We postulated that if temperature is the primary factor limiting insect herbivory, the effect of gaps on rates of insect herbivory should be more evident in temperate than in tropical forests; due to low growing season temperatures in the oceanic temperate forests of the Southern Hemisphere, the effect of gaps on insect herbivory rates should in turn be stronger there than in the more continental temperate climates of the Northern Hemisphere. We examined global patterns of insect herbivory in gaps versus understories through meta-analysis of 87 conspecific comparisons of leaf damage in contrasting light environments. Overall, insect herbivory in gaps was significantly higher than in the understorey; insect herbivory was 50% higher in gaps than in understoreys of tropical forests but did not differ significantly between gaps and understories in temperate forests of either hemisphere. Results are consistent with the idea that low resource availability - and not temperature - limits insect herbivore activity in forest understoreys, especially in the tropics, and suggest the selective influence of insect herbivory on late-successional tree species may have been over-estimated.