Nutrient fluxes from insect herbivory increase during ecosystem retrogression in boreal forest

Ecological theory, developed largely from ungulates and grassland systems, predicts that herbivory accelerates nutrient cycling more in productive than unproductive systems. This prediction may be important for understanding patterns of ecosystem change over time and space, but its applicability to other ecosystems and types of herbivore remain uncertain. We estimated fluxes of nitrogen (N) and phosphorus (P) from herbivory of a common tree species (Betula pubescens) by a common species of herbivorous insect along a -5000-yr boreal chronosequence. Contrary to established theory, fluxes of N and P via herbivory increased along the chronosequence despite a decline in plant productivity. The herbivore-mediated N and P fluxes to the soil are comparable to the main alternative pathway for these nutrients via tree leaf litterfall. We conclude that insect herbivores can make large contributions to nutrient cycling even in unproductive systems, and influence the rate and pattern of ecosystem development, particularly in systems with low external nutrient inputs.