The Influence of Multiple Introduced Predators on a Littoral Pond Community

In a replicated field experiment we studied the effects of natural densities of two exotic consumers, the predatory and herbivorous signal crayfish (Pacifastacus leniusculus) and the predatory rainbow trout (Oncorhynchus mykiss), on multiple trophic levels of a pond community. The goals were to: (1) determine the individual and combined effects of predators on macroinvertebrates, macrophytes, and periphytic algae; (2) evaluate the strength of direct and indirect interactions in a food web influenced by omnivores; and (3) evaluate the relative importance of direct and indirect predator effects on mortality and growth of a native frog species, Rana temporaria. The experiment showed that both signal crayfish and rainbow trout had strong effects on multitrophic levels of a littoral pond community, through direct consumption and indirect effects on lower trophic levels. Crayfish had weak but significant negative effects on the biomass of predatory invertebrates and greatly reduced the biomass of snails, the most abundant invertebrate grazers. Although the number of active herbivorous tadpoles tended to be higher in crayfish cages relative to control cages, the proportion of surviving froglets was lower in crayfish cages than in control cages, possibly due to crayfish predation on injured tadpoles. The size of surviving froglets did not differ from controls, but tadpoles in crayfish cages often suffered tail injuries. Macrophyte coverage decreased as a result of crayfish consumption and nonconsumptive fragmentation. However, the biomass of periphyton increased in crayfish cages relative to controls, probably due to reduced grazing from snails. In contrast, trout had strong negative effects on the biomass of both predatory invertebrates and insect grazers, whereas trout had less effect on snail biomass than did crayfish. Also, in contrast to crayfish cages, the number of active tadpoles in trout cages was lower than in controls, probably due to a combination of trout predation and troutinduced reduced tadpole activity. Trout had a strong negative impact on froglet survival, and froglets in trout cages metamorphosed at a smaller size and had reduced growth rates compared to froglets in crayfish and control cages. As with crayfish, the biomass of periphyton increased in trout cages relative to controls, which may be due to a combination of both density and trait-mediated trout effects on tadpole grazing. In treatments with multiple predators the effects of crayfish and trout on caged communities were independent, and there were few interactions. Mostly effects of combined predators reflected those in single predator cages. Our results demonstrate that noninteracting, introduced multiple predators can have strong direct and indirect effects on multiple trophic levels in pond communities. Trophic cascades may develop in aquatic food webs even with omnivores such as crayfish, and in complex habitats with trout. These strong indirect effects are mediated through both predation on important grazers (i.e., the crayfish– snail–periphyton link) and a combination of density and behavioral responses of grazers to predators (i.e., the trout–tadpole–periphyton link). When two noninteracting predators have strong but different effects on prey survival or activity, their combined effects on intermediate trophic levels reflect responses to the more dangerous predator.