The effects of Daphnia on nutrient stoichiometry and filamentous cyanobacteria:: a mesocosm experiment in a eutrophic lake

1. Stoichiometric theory predicts that the nitrogen : phosphorus (N : P) ratio of recycled nutrients should increase when P-rich zooplankton such as Daphnia become dominant. We used an enclosure study to test the hypothesis that an increased biomass of Daphnia will increase the relative availability of N versus P sufficiently to decrease the abundance of filamentous cyanobacteria. The experiment was conducted in artificially enriched Lake 227 (L227) in the Experimental Lakes Area (ELA), north-western Ontario, Canada. Previous studies in L227 have shown that the dominance of filamentous, N-fixing cyanobacteria is strongly affected by changes in the relative loading rates of N and P. 2. We used a 2 x 2 factorial design with the addition or absence of D. pulicaria and high or low relative loading rates of N and P (+NH4 , -NH4 ) in small enclosures as treatment variables. If Daphnia can strongly affect filamentous cyanobacteria by altering N and P availability, these impacts should be greatest with low external N : P loading rates. The phytoplankton community of L227 was predominantly composed of filamentous Aphanizomenon spp. at the start of the experiment. 3. Daphnia strongly reduced filamentous cyanobacterial density in all enclosures to which they were added. The addition of NH4 had only a small impact on algal community composition. Hence, we conclude that Daphnia did not cause reductions in cyanobacteria by altering the N : P ratio of available nutrients. 4. Despite the lack of evidence that Daphnia affected filamentous cyanobacteria by altering the relative availability of N and P, we found changes in nutrient cycling consistent with other aspects of stoichiometric theory. In the presence of Daphnia , total P in the water column decreased because of an increase in P sedimentation. In contrast to P, a decrease in suspended particulate N was offset by an increase in dissolved N (especially NH4 ). Hence, dissolved and total N : P ratios in the water column increased with Daphnia as a result of differences in the fate of suspended particulate N versus P. There was minimal accumulation and storage of P in Daphnia biomass in the enclosures. 5. Our experiment demonstrated that Daphnia can strongly limit filamentous cyanobacterial abundance and affect the biogeochemical cycling of nutrients. In our study, changes in nutrient cycling were apparently insufficient to cause the changes in phytoplankton community composition that we observed. Daphnia therefore limited filamentous cyanobacteria by other mechanisms.