Dissecting the drivers of population cycles: Interactions between parasites and mountain hare demography

There is a growing awareness that cyclic population dynamics in vertebrate species are driven by a complex set of interactions rather than a single causal factor. While theory suggests that direct host-parasite interactions may destabilise population dynamics, the interaction between host and parasite may also influence population dynamics through indirect effects that result in delayed responses to either density or to life-history traits. Using empirical data on mountain hares (Lepus timidus) infected with a nematode parasite (Trichostrongylus retortaeformis), we developed an individual-based model (IBM) that incorporated direct effects and delayed life-history effects (DLHEs) of a macroparasite, alternative transmission mechanisms and seasonality in host population dynamics. The full model describes mean characteristics of observed mountain hare time series and parasite abundance, but by systematically removing model structure we dissect out dynamic influences of DLHEs. The DLHEs were weakly destabilising, increasing the propensity for cyclic dynamics and suggesting DLHEs could be important processes in host-parasite systems. Further, by modifying model structure we identify a strong influence of parasite transmission mechanism on host population stability, and discuss the implications for parasite aggregation mechanisms, host movement and natural geographical variation in host population dynamics. The effect of T. retortaeformis on mountain hares likely forms part of a complex set of interactions that lead to population cycles.