RESPONSES OF AN INSECT FOLIVORE AND ITS PARASITOIDS TO MULTIYEAR EXPERIMENTAL DEFOLIATION OF ASPEN

Abstract

Foliage quality may decline in deciduous trees following defoliation, thus affecting the insect generation responsible for the herbivory (rapid induced resistance, RIR), or future generations (delayed induced resistance, DIR). During outbreaks, trees often suffer partial or complete defoliation for two or more successive years, yet most studies have examined induced resistance following only one season of defoliation, which may not reveal its full impact on herbivores. In a field experiment, 40 trees from each of two clones of trembling aspen, Populus tremuloides, were severely defoliated for one, three, and four years in succession by experimentally manipulating densities of an outbreak folivore, the forest tent caterpillar, Malacosoma disstria. Treatments were applied such that the individual and combined effects of RIR and DIR on the fitness of the forest tent caterpillar could be assessed independently. In field assays, defoliation treatments did not affect larval development time and only marginally affected survival. However, fecundity in both clones was significantly reduced by a single season of defoliation concurrent with the bioassay (effects of RIR), by three consecutive years of defoliation prior to the year of the bioassay (effects of DIR), and by four consecutive years of defoliation (combined effects of RIR and DIR). There were no differences among the three defoliation treatments, indicating that the effects of DIR and RIR combined were not greater than each acting alone. Reductions in fecundity were less than half those observed during natural outbreaks, suggesting that other factors also must contribute to declining fecundity during the collapse of outbreaks. Short-term laboratory bioassays indicated that defoliation effects observed in long-term field assays were not due to changes in relative growth rate (RGR) of second instars or final-instar males, which were unaffected, possibly because of increased relative consumption rates (RCR). Defoliation treatments decreased RGR of final-instar females in laboratory bioassays, despite elevated RCR. Both defoliation treatment and aspen clone influenced parasitism by tachinid flies that detect hosts through volatiles released from leaves damaged by caterpillars. Parasitism was highest on trees defoliated concurrently with the larval bioassay. However, there were no differences between trees defoliated prior to the bioassay and control trees, indicating that effects on parasitism were not due to defoliation-induced changes in host quality per se. Thus, there were no additive interactions between DIR and parasitism that would amplify delayed density-dependent effects on population dynamics. Spatial responses of these tachinids to host density or to current defoliation rather than the defoliation history of the trees may enhance the stabilizing effect of RIR on population dynamics. Conversely, differences in parasitism among clones could contribute to spatial variation in tent caterpillar population density. Neither defoliation effect on host quality nor parasitism was sufficient to slow reproductive rates to levels observed in declining outbreaks in nature, suggesting that single-factor explanations for tent caterpillar population dynamics are unlikely.