Nearby large islands diminish biodiversity of the focal island by a negative target effect

The Equilibrium Theory of Island Biogeography postulates that larger and closer islands support higher biodiversity through the dynamic balance of colonization and extinction processes. The negative diversity-isolation (i.e. the distance to the mainland) relationship is derived based on the assumption that the mainland is the only source pool for island biotas. However, nearby islands could also act as species sources for focal islands via a source effect. In this study, we move a further step and hypothesize that nearby islands may reduce bird colonizers of the focal island and diminish its biodiversity, resulting in a negative target effect. To test our hypothesis, we assessed the effects of island area and isolation (metrics considering both the mainland and nearby islands) on taxonomic (i.e. species richness), functional and phylogenetic diversity of terrestrial breeding birds on 42 islands in the largest archipelago of China, the Zhoushan Archipelago. Furthermore, we compared the predictive power of the distance to the large island under a set of relative area thresholds and the relative area of nearby islands on species richness under a set of distance thresholds to explore the role of nearby islands as a source and/or target island. We found that island area had a positive effect on species richness, phylogenetic diversity and functional diversity, while the distance to the mainland had a negative effect only on species richness. Species richness on the focal island increased with increasing distance to the nearest larger island, indicating the negative target effect. Furthermore, the negative target effect depended on the area of nearby islands relative to the area of the focal island. Our finding of the negative target effect suggests islands located between the mainland and the focal island can be not only sources or stepping stones, but also colonization targets. This result demonstrates the importance of considering multiple geographical attributes of islands in island biogeographic studies, especially the characteristics related to source and/or target effects.