The objectives of this study were to determine responses by red mangrove (Rhizophora mangle) and its primary consumers to nutrient enrichment, to determine if nutrient limitation is responsible for the dwarf plant form of red mangrove, and to evaluate four competing hypotheses for the relation of nutrient status and invertebrate herbivory. In a factorial-designed experiment, I fertilized 48 dwarf trees along gradients of tidal elevation and water depth at Twin Cays, an intertidal mangrove island in Belize, Central America, and measured plant growth responses and herbivory for 2 yr. At the end of year- 2, I compared biomass accumulation and analyzed plant tissue for chemical and structural composition. Dwarf red mangrove trees on this tidal island responded quickly and grew vigorously when treated with P and NPK fertilizers. Leaf number, leaf area, branching, shoot length, and aerial root production increased dramatically over 2 yr. N-fertilized trees grew very slowly and their responses were not different from Control trees. N-fertilized and Control trees changed little over 2 yr. Some responses to phosphorus-containing fer- tilizers vary by water depth and tidal elevation, but physicochemical factors do not explain the differences in growth responses. I conclude that phosphorus availability is a major factor limiting red mangrove growth at my study site in the interior of Twin Cays. Herbivory by two specialized, endophytic insect species (Ecdytolopha sp., which feeds in apical buds, and Marmara sp., which mines stem periderm) increased in P- and NPK-fertilized trees compared to N-fertilized and Control trees. Twice as many apical buds were damaged or destroyed and the frequency of mines increased by 6-8 fold. However, fertilization had no effect on feeding rates and standing damage by a leaf-feeding guild of generalist herbivores or on the frequency of shoots killed by stem borers. Herbivory by the two specialists was not related to C:N ratios, but it was inversely related to concentrations of phenolic com- pounds. Neither nutrient ratios nor concentrations of phenolics affected rates of herbivory by the generalist folivores. Although sclerophylly of red mangrove leaves decreased in P- and NPK-fertilized trees but not in N-fertilized and Control trees, there was no relationship between leaf toughness and herbivory by generalist folivores. These data suggest that sclerophylly in oligotrophic ecosystems may be an adaptive mechanism related to nutrient conservation, and that it is associated with red mangrove survival in phosphorus-deficient soil rather than an adaptation to herbivory.
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