Nitrogen effects on an interaction chain in a salt marsh community.

Nutrients can structure communities by influencing both plant interactions and plant herbivore interactions, though rarely do studies integrate these processes. In this study we examined how nitrogen fertilization influenced (1) the positive interaction between the marsh elder, Iva frutescens, and the black rush, Juncusgerardi, and (2) the quality of Iva as a host plant for the aphid, Uroleuconambrosiae. Previous studies have shown that by mitigating soil salt accumulation and hypoxia, Juncus is essential to the survival of Iva and its aphid herbivore at mid-marsh elevations. To address the effects of nitrogen on this interaction, we compared fertilized and unfertilized Iva plants subject to Juncus removal and control treatments in the field. Additionally, we measured the monthly population growth rates of aphids transplanted onto these Iva plants. Iva leaf biomass and flower number results indicated that fertilizing Iva eliminated its dependence upon Juncus, such that fertilized plants grown without Juncus were not different from unmanipulated plants. Aphid monthly population growth rates through mid-summer revealed that fertilization also eliminated the indirect dependency of aphids on Juncus, so that aphid growth rates on fertilized Iva without Juncus neighbors were similar to rates on unmanipulated Iva. Results also indicated that fertilizing Iva grown with Juncus increased Iva size, potentially enabling these plants to support larger aphid populations. Our results suggest that only under conditions of nitrogen limitation are the positive effects of Juncus essential to the mid-marsh persistence of Iva and its aphid herbivore. Furthermore, we found that nitrogen effects on aphid populations may arise not only from a direct effect of nutrients on Iva size but also through the indirect effects of nitrogen on the interaction between Juncus and Iva. We argue that studies integrating processes occurring both within and between trophic levels, are important to fully understanding the community-wide effects of nutrients.