Facilitation and predation structure a grassland detrital food web: the responses of soil nematodes to isopod processing of litter.

1. Detritus can support successive consumers, whose interactions may be structured by changes in the condition of their shared resource. One model of such species interactions is a processing chain, in which consumers feeding on the resource in a less processed state change the resource condition for subsequent consumers. 2. In a series of experiments, the hypothesis was tested that a common detritivore, the terrestrial isopod Porcellio scaber, affects soil nematodes through the processing of plant litter. Different detrital resources were added to soil from a California coastal prairie in order to simulate litter processing by the detritivore. Treatments that included only whole grass litter corresponded to detrital food webs lacking detritivores, while treatments that included mixtures of P. scaber faeces and grass litter corresponded to different densities or feeding rates of P. scaber. 3. Simulated litter processing by P. scaber increased the abundance of bacterivorous nematodes by between 32% and 202% after 24-44 days in laboratory experiments, but had no effect on fungivorous or predaceous nematodes. 4. In a subsequent field experiment, however, fungivorous nematodes were suppressed by isopod litter processing while bacterivores showed no response. Instead, P. scaber processing of litter increased the abundance of predaceous nematodes in the field experiment by 176%. 5. When simulated litter processing of litter was crossed in laboratory experiments with predaceous nematode addition (comparable to the response of predators in the field experiment), the abundance of bacterivores was increased by isopod processing of litter (by an average of 122%), but suppressed by elevated densities of predaceous nematodes (by an average of 41%). 6. This suggests that litter processing by P. scaber facilitates the bacterial channel of the soil food web, but that predaceous nematodes suppress the response of bacterivores in the field. Processing chain interactions may, therefore, be important in understanding the relative importance of bacterial and fungal channels in the soil food web, while top-down effects of predators determine the resulting changes in population abundance and biomass.