Quantification of invertebrate predation and herbivory in food chains of low complexity.

Zooplankton grazing impact on algae, heterotrophic flagellates and bacteria, as well as invertebrate predation on herbivorous zooplankton, were investigated in two sub-Antarctic lakes with extremely simple food chains. The two species of herbivorous zooplankton present in the lakes (the copepods boeckella michaelseni and Pseudoboeckella poppei) exerted substantial grazing pressure on algae. However, the dominant algal species exhibited properties that enabled them to avoid (large size or extruding spines, e.g. Staurastrum sp., Tribonema sp.) or compensate (recruitment from the sediment, Mallomonas sp.) grazing. There are only two potential invertebrate predators on the herbivorous copepods in the two lakes: the copepod Parabroteas sarsi and the diving beetle Lancetes claussi. Vertebrate predators are entirely abscent from sub-Antarctic lakes. Based on our experiments, we estimated that the predators would remove at most about 0.4% of the herbivorous copepods per day, whereas planktivorous fish, if present in the lakes, would have removed 5-17% of the zooplankton each day. Consequently, the invertebrate predators in these high-latitude lakes had only a marginal predation impact compared to the predation pressure on zooplankton in the presence of vertebrate predators in temperate lakes. The study of these simple systems with only two quantitatively functionally important trophic links, suggests that high grazing pressure foreces the algal community towards forms with grazer resistant adaptations such as large size, recruitment from another habitat, and grazer avoidance spines. We propose that due to such adaptations, predictions from food web theory are only partly corroborated, i.e. algal biomass actually increases with increasing productivity, although the grazer community is released from predation. In more species-rich and complex systems, e.g temperate lakes with three functionally important links, such adaptations are likely to be even more important, and, consequently, the observable effects of trophic interactions from top predators on lower trophic levels even more obscured.