The effects of planktivorous fish on lower trophic levels through predation on zooplankton and nutrient excretion were experimentally separated and their relative importance quantified in a eutrophic humic lake. The experiment was performed in 12 enclosures (3 m 3 ), which initially were identical with respect to all components except fish. At the start of the experiment, caged fish not able to feed on zooplankton were added to four of the enclosures (excretion treatment), and free swimming fish to four enclosures (excretion plus predation treatment). Four enclosures were left as controls. Samples for nutrients and all major groups of organisms from bacteria to zooplankton were taken after 14 and 28 d. The effect sizes of fish excretion and predation were calculated for each variable. Our results suggest that in eutrophic lakes fish predation on zooplankton may be more important than nutrient excretion by fish for the structure and dynamics of planktonic communities. Fish predation on zooplankton was the most important mechanism accounting for fish effects on nutrient concentrations in the water, on phytoplankton biovolume, on rotifers and total zooplankton biomass, as well as on protozoan densities. However, our results suggest that nutrient excretion by fish may have important indirect effects on zooplankton. Hence, the effects of planktivorous fish through both predation on zooplankton and nutrient excretion act in concert and may be quantitatively important to shape the structure and dynamics of planktonic communities.Although the effects of planktivorous fish on planktonic communities have been well demonstrated in many aquatic systems, the mechanisms accounting for these effects are still not clear. The classical interpretation is that these effects result from size-selective predation processes throughout the food chain. Planktivorous fish selectively consume large herbivores and often shift zooplankton structure toward dominance by rotifers, small cladocerans, and copepods (reviewed by Gliwicz and Pijanowska 1989), which have a lower grazing impact on phytoplankton and higher massspecific rates of nutrient recycling (reviewed by Sterner 1989). The increase in phytoplankton biomass arising from 1 Corresponding author.
AcknowledgmentsThe Brazilian Government, through CAPES (Post-Graduate Federal Agency), has awarded J.L.A. with a Ph.D. grant and has financed this research. Funding was also received from the Swedish Natural Science Research Council (NFR) to L.A.H. and from the Swedish Foundation for International Cooperation in Research and Higher Education (STINT) to J.L.A. We thank Marie Svensson, Ola Svensson, Renata Panosso, Alexandre Anesio, Maur铆cio Petr煤cio, and Ramunas Stepanauskas for their assistance in the field or lab. We also thank Renata Panosso, Christer Br枚nmark, Wilhelm Gran茅li, Gary Mittelbach, and two anonymous reviewers for valuable comments and suggestions to improve the paper.