Functional response comparisons among freshwater amphipods: ratio-dependence and higher predation for Gammarus pulex compared to the non-natives Dikerogammarus villosus and Echinogammarus berilloni

Abstract

The invasive Ponto–Caspian amphipod Dikerogammarus villosus strongly impairs the structure of recipient freshwater communities, mostly through predation of a wide range of invertebrates. Useful insights on the ecological impact of an invader can be provided by comparing its functional response (FR)—namely the relationship between resource use and resource availability—to that of trophic analogs from the recipient communities. We applied this method and derived the FR of D. villosus, the native Gammarus pulex, and Echinogammarus berilloni, another gammarid that is also undergoing range expansion, feeding on live dipteran larvae. We tested a range of predator densities to account for mutual interference among predators, assuming that predators do not forage alone in their natural environment. We also analysed the predators’ spatial distribution to investigate whether spatial behaviour might be at the origin of interference. For the three gammarid species, the per capita predation rate was a function of the prey-to-predator ratio and showed a decelerating rise to an asymptote. There was no difference in searching efficiency between the three species while prey handling time was significantly lower in G. pulex than in D. villosus and E. berilloni, leading to a higher food intake for the native at high prey densities. Differences in morphology and behaviour between the three gammarids might explain our results. Indeed, D. villosus tended to aggregate more than the other two gammarid species, which promotes conspecific interactions. Our findings suggest that mutual interference between conspecifics is a fundamental shaper of gammarid predation. Time spent interacting with conspecifics is likely to decrease the biotic pressure of the Killer shrimp D. villosus. From a broader perspective, in addition to higher predation risk, parasitism, habitat complexity or climatic variables, the presence of conspecifics of the predator (i.e. predator density) is another factor that has the potential to alter FR comparisons.