SEASONAL FOOD LIMITATION OF DETRITIVORES IN A MONTANE STREAM - AN EXPERIMENTAL TEST

Many species of detritivorous invertebrates in small streams depend almost entirely on inputs of leaf litter for their nutritional requirements, however the concentration of this resource varies considerably seasonally. An experiment designed to test the hypothesis that productivity of coarse particle detritus feeders (shredders) is seasonally food limited was performed using replicate streamside channels that received one of three input rates of whole leaf detritus (rates equal to those naturally falling into montane streams, and two levels of increased inputs). Seven of the nine common shredder species attained significantly higher adult mass, higher densities, or both when food was supplemented. Larval densities of Malenka spp. and Brillia retifinis were significantly higher when additional food was added. Brillia retifinis densities were more than 10 x greater in high food additions than in the "natural" treatment; B. retifinis apparently fills a role as a fugitive in this system and is better able to track shifts in resource abundance by virtue of its short generation time. Zapada cinctipes and Z. haysi were more dense in high and intermediate food input treatments during the last half of the summer. Other taxa had nonsignificant trends to higher densities when extra food was provided. Most of the change in benthic densities can be attributed to decreased rates of emigration with increased food supply. Six of the eight taxa for which adult mass at emergence was measured were significantly more massive (4-46%) when food was added. Both sexes exhibited this increase in mass, but females gained proportionally more in most species. There were no detectable changes in the timing of adult emergence due to food manipulations. The biomass of most taxa increased in proportion to the overall increase in biomass of the common coarse-detritus consumers. The main exceptions to that pattern were a disproportionate increase in the percentage of the biomass represented by Brillia retifinis and a decrease in the representation of Zapada cinctipes. The responses of this community to food supplementation demonstrate food limitation of detritivores. The exponential increase in benthic biomass under food addition shows the scope for productivity if food were not limiting. These coexisting species benefitted from enhanced food supply on a spatial scale that is relevant for population level processes. The large seasonal variation in resource abundance under natural conditions creates bottlenecks during periods of low food supply, which constrain subsequent production even during periods when food is abundant.