INVITED REVIEW: PART OF AN INVITED ISSUE ON TREE NUTRITION Nutrition of mangroves

Mangrove forests dominate the world's tropical and subtropical coastlines. Similar to other plant communities, nutrient availability is one of the major factors influencing mangrove forest structure and productivity. Many mangrove soils have extremely low nutrient availability, although nutrient availability can vary greatly among and within mangrove forests. Nutrient-conserving processes in mangroves are well developed and include evergreeness, resorption of nutrients prior to leaf fall, the immobilization of nutrients in leaf litter during decomposition, high root/ shoot ratios and the repeated use of old root channels. Both nitrogen-use efficiency and nutrient resorption efficiencies in mangroves are amongst the highest recorded for angiosperms. A complex range of interacting abiotic and biotic factors controls the availability of nutrients to mangrove trees, and mangroves are characteristically plastic in their ability to opportunistically utilize nutrients when these become available. Nitrogen and phosphorus have been implicated as the nutrients most likely to limit growth in mangroves. Ammonium is the primary form of nitrogen in mangrove soils, in part as a result of anoxic soil conditions, and tree growth is supported mainly by ammonium uptake. Nutrient enrichment is a major threat to marine ecosystems. Although mangroves have been proposed to protect the marine environment from land-derived nutrient pollution, nutrient enrichment can have negative consequences for mangrove forests and their capacity for retention of nutrients may be limited.