Frog community responses to recent American bullfrog invasions

Native species may decline quickly when confronted with an exotic species to which they are not adapted. The extent of decline may depend on the abundance of an invader and the length of time since it first arrived in the community （residence time）, and the interaction between these two variables. We tested these effects using data on the effects of American bullfrog Lithobates catesbeianus invasion on native frog communities in 65 permanent lentic waters on islands in the Zhoushan Archipel- ago, China. We examined variation in native frog abundance and species richness in relation to features of the American bullfrog invasion, habitat disturbance, characteristics of the water body and fish communities and the presence of red swamp crayfish. Bullfrog invaded sites had lower native frog density and species richness, higher submerged vegetation cover and greater fre- quency of repairs to the water body than did non-invaded sites. The minimum adequate general linear mixed models showed that both native frog density and species richness were negatively related to post-metamorphosis bullfrog density, and that native frog species richness was also positively related to the vegetation cover. There was no effect on either native frog density or species richness of residence time or its interaction with bullfrog density, or of the abundance of bullfrog tadpoles. The results suggested that post-metamorphosis bullfrogs had impacts on native frog communities in the islands, and that the extents of these impacts are proportional to post-metamorphosis bullfrog density