Factors influencing the stability of nutrient-enriched freshwater macrophyte communities: the role of sticklebacks Pungitius pungitius and freshwater snails

1. Enclosures, with the following characteristics, were used to investigate the effects of increased nutrient (nitrate and phosphate) loading on freshwater communities: (a) controls with normal densities of invertebrates, and including macrophytes, molluscs and fish (sticklebacks), and communities similar to the controls but with (b) much reduced snail densities and (c) increased fish densities. 2. The addition of nutrients resulted in increases in the biomass of phytoplankton and epiphyton, as well as zooplankton counts, in all the experimental enclosures. 3. A reduction in the snail population density resulted in a significant decrease in phytoplankton density and an increase, compared to controls, in the density of epiphytic algae growing on the leaves of the submerged macrophytes, Potamogeton crispus and Groenlandia densa. It also resulted in a significant increase (3.5 times that in the controls) in chlorophyll-a values of epilithic algae growing on glass slides, an increase in senescent tissue in macrophytes and a decline in the mass of Groenlandia densa by the end of the experiment. 4. An increase in the fish population resulted in a significant decline, compared to the controls, in the densities of zooplankton (including Chydorus sphaericus and copepods), Lymnaea peregra (from May to mid-June), Asellus meridianus, Crangonyx pseudagracilis, Culex (P.) molestus and Physa fontinalis. It also resulted in a change in the composition of the phytoplankton, with an increase in the large colonial Eudorina elegans and a decline in smaller planktonic algae, including Ankistrodesmus and Cryptomonas spp. 5. The possible mechanisms responsible for the effects are evaluated and their relevance to conservation and management of freshwater macrophyte communities are discussed.