Stand structure interacts with previous defoliation to influence herbivore fitness

The intensity and duration of Neodiprion abietis outbreaks have recently increased in forests of North America that were precommercially thinned more than a decade earlier. We tested the hypotheses that changes in stand structure following thinning increase the fitness (i.e., survival rate×fecundity) of N. abietis by either (a) increasing foliar availability and/or quality (i.e., increased availability of primary metabolites and/or reduced foliar defenses) or (b) by reducing any negative effects on foliar quality and/or availability resulting from herbivory that occurred during the preceding season(s). Effects of thinning and previous herbivory on N. abietis and its host plant (Abies balsamea) were determined through (i) a manipulative field experiment that evaluated the effects of experimental defoliation on N. abietis in a thinned stand, (ii) a manipulative field experiment that examined the effects of thinning on N. abietis in undamaged and naturally defoliated stands, and (iii) a field survey to estimate survival of N. abietis in natural populations. Defoliation caused reductions in the availability of different-aged foliage available to larvae and in the fitness of a subsequent N. abietis generation feeding on defoliated branches, but decreases in fitness were smaller in thinned than unthinned stands. In thinned stands, defoliation was associated with increases in foliage production and foliar contents of monoterpenes and nitrogen, as well as with a decrease in foliar contents of water. Conversely, only small changes in plant growth and foliar contents of nutrients and secondary chemicals were observed in defoliated unthinned stands. This suggests that deleterious effects of defoliation on sawfly fitness were offset by an increase in the foliar content of nitrogen, a primary compound known to improve larval growth in sawflies, which supports the hypothesis that thinning moderates negative effects of previous defoliation on sawfly fitness. The present study demonstrates that forestry practices that alter stand structure by reducing tree density may increase herbivory by affecting the way trees respond to insect attack, even after crown closure, with consequences on the buildup of herbivore populations in attacked trees.