Metapopulation responses to patch connectivity and quality are masked by successional habitat dynamics.

Many species have to track changes in the spatial distribution of suitable habitat from generation to generation. Understanding the dynamics of such species will likely require spatially explicit models, and patch-based metapopulation models are potentially appropriate. However, relatively little attention has been paid to developing metapopulation models that include habitat dynamics, and very little to testing the predictions of these models. We tested three predictions from theory about the differences between dynamic habitat metapopulations and their static counterparts using long-term survey data from two metapopulations of the butterfly Plebejus argus. As predicted, we showed first that the metapopulation inhabiting dynamic habitat had a lower level of habitat occupancy, which could not be accounted for by other differences between the metapopulations. Secondly, we found that patch occupancy did not significantly increase with increasing patch connectivity in dynamic habitat, whereas there was a strong positive connectivity-occupancy relationship in static habitat. Thirdly, we found no significant relationship between patch occupancy and patch quality in dynamic habitat, whereas there was a strong, positive quality-occupancy relationship in static habitat. Modeling confirmed that the differences in mean patch occupancy and connectivity-occupancy slope could arise without changing the species' metapopulation parameters-importantly, without changing the dependence of colonization upon connectivity. We found that, for a range of landscape scenarios, successional simulations always produced a lower connectivity-occupancy slope than comparable simulations with static patches, whether compared like-for-like or controlling for mean occupancy. We conclude that landscape-scale studies may often underestimate the importance of connectivity for species occurrence and persistence because habitat turnover can obscure the connectivity-occupancy relationship in commonly available snapshot data.