Habitat connectivity and resident shared predators determine the impact of invasive bullfrogs on native frogs in farm ponds

Habitat connectivity is considered to have an important role on the persistence of populations in the face of habitat fragmentation, in particular, for species with conservation concern. However, it can also impose indirect negative effects on native species through the spread of invasive species. Here, we investigated direct and indirect effects of habitat connectivity on populations of invasive bullfrogs and native wrinkled frogs and how these effects are modified by the presence of common carp, a resident shared predator, in a farm pond system in Japan. The distribution pattern analysis using a hierarchical Bayesian modelling indicated that bullfrogs had negative effects on wrinkled frogs, and that these negative effects were enhanced with increasing habitat connectivity owing to the metapopulation structure of bullfrogs. The analysis also suggested that common carp mitigated these impacts, presumably owing to a top-down trophic cascade through preferential predation on bullfrog tadpoles. These presumed interspecific interactions were supported by evidence from laboratory experiments, i.e. predation by carp was more intense on bullfrog tadpoles than on wrinkled frog tadpoles owing to the difference in refuge use. Our results indicate that metacommunity perspectives could provide useful insights for establishing effective management strategies of invasive species living in patchy habitats.