REGULAR ARTICLE Season mediates herbivore effects on litter and soil microbial abundance and activity in a semi-arid woodland

Abstract Herbivores can directly impact ecosystem function by altering litter quality of an ecosystem or indirectly by shifting the composition of microbial communities that mediate nutrient processes. We examined the effects of tree susceptibility and resis-tance to herbivory on litter microarthropod and soil microbial communities to test the general hypothesis that herbivore driven changes in litter inputs and soil microclimate will feedback to the microbial commu-nity. Our study population consisted of individual piñon pine trees that were either susceptible or resistant to the stem-boring moth (Dioryctria albovit-tella) and susceptible piñon pine trees from which the moth herbivores have been manually removed since 1982. Moth herbivory increased piñon litter nitrogen concentrations (16%) and decreased canopy precipi-tation interception (28%), both potentially significant factors influencing litter and soil microbial commu-nities. Our research resulted in three major findings: (1) In spite of an apparent increase in litter quality, herbivory did not change litter microarthropod abun-dance or species richness. (2) However, susceptibility to herbivores strongly influenced bulk soil microbial communities (i.e., 52 % greater abundance beneath herbivore-resistant and herbivore-removal trees than susceptible trees) and alkaline phosphatase activity (i.e., 412 % increase beneath susceptible trees relative to other groups). (3) Season had a strong influence on microbial communities (i.e., microbial biomass and alkaline phosphatase activity increased after the summer rains), and their response to herbivore inputs, in this semi-arid ecosystem. Thus, during the dry season plant resistance and susceptibility to a common insect herbivore had little or no observable effects on the belowground organisms and processes we studied, but after the rains, some pronounced effects emerged