Escapes from herbivory in relation to the structure of mangrove island macroalgal communities.

Some shallow habitats that surround mangrove islands exhibit abruptly discontinuous macrophyte boundaries; in other regions, plant distributional patterns are less defined. Where distinct boundaries do occur, fleshy algae predominate on the roots of the red mangrove, Rhizophora mangle, which do not contact the bottom sediments ("hanging roots"), while calcifying algae dominate on the substratum-penetrating roots and banks (=embedded-root habitat) surrounding the mangrove thickets. Considerable natural-history and floristic information reveals that the fleshy hanging-root species are not specialists, for that type of habitat. Experimental transplants showed that on banks and embedded roots where there typically are abundant macroherbivores (particularly sea urchins), most fleshy algae are eliminated.The dominants of the hanging-root habitat (e.g, Acanthophora spicifera, Spyridia filamentosa, Caulerpa racemosa var. Occidentalis) are 6-20 times more susceptible to herbivores than the dominants of the embedded-root habitat (e.g., Halimeda opuntia f. triloba, H. monile). Consequently, we suggest the former are relegated to the spatial refugia from herbivores (=non-coexistence escapes) provided by the hanging roots. Factors associated with these palatability differences include higher average calorific values (6.5 times) of the fleshy hanging-root dominants, greater proportions of organic content (2.6 times) and the general absence of calcification. The dominants of the embedded-root habitat show reduced edibility as a probable consequence of low calorific values, heavy calcification and potential herbivore-detering secondary metabolites. Correlative evidence and preliminary experimental results tentatively indicate that, in the absence of macroherbivores, the hanging-root dominants, which exhibit production rates 4.7 times greater than the dominants of the embedded-root habitat, are better competitors for space.We suggest that variations in herbivory are responsible, in part, for maintaining greater algal diversity in mangrove systems. At a study site with abundant sea urchins, five algal species were found only in the embedded-root habitat three species were confined to the hanging roots, while three others occurred in both. At an urchin-free site, no macrophytes were found only on embedded-root substrata, while one (in trace amounts) was found only on hanging-root habitat and eight occurred in both. We predict that in the absence of herbivores, the species assemblage characteristic of the hanging-roots would exclude many of the dominants from the embedded-root habitat.