Intraguild predation and species exclusions in amphipods: The interaction of behaviour, physiology and environment

1. Data from field surveys, laboratory experiments and computer simulations of community dynamics revealed that a novel interaction among intraguild predation, physiological adaptation and environment may explain the complex distributions of two putatively competing aquatic amphipods. 2. Gammarus pulex and G. tigrinus both thrive in fresh and oligohaline waters in western Europe. However, the native European G. pulex excludes the invading North American G. tigrinus from freshwaters of relatively low conductivity, whereas the reverse occurs at higher conductivities. Additionally, there is much spatio-temporal fluctuation in the patterns of coexistence of these species. 3. Laboratory experiments in The Netherlands and Ireland revealed that mutual predation of moulting individuals occurred frequently between these species. However, predation frequencies were differentially in favour of G. pulex under the ionic conditions to which this species is physiologically adapted (freshwater). On the other hand, predation was not differential under the ionic conditions to which G. tigrinus is physiologically adapted (oligohaline water). 4. A mathematical model, which extends the logistic equation to include mutual intraguild predation, simulated interactions over a range of values of relevant population parameters. This indicated that G. pulex would be excluded when balanced instantaneous rates of mutual predation were combined with the known greater reproductive output of G. tigrinus. However, this reproductive advantage is overcome by any relatively small bias in the instantaneous rate of predation favouring G. pulex, leading to the exclusion of G. tigrinus. This occurs even when the reproductive advantage to G. tigrinus is relatively large. Moreover, the model generated 'switches' in species dominance that are determined by the relative values of reproductive rate and mutual predation. The time taken to 'switch' may explain the transient periods of apparent coexistence of these species observed in the field. 5. The complex community dynamics of such species may thus be understood in terms of variation in the intensity of species interactions mediated by behavioural, physiological and environmental factors.