Chemical defense against diverse coral reef herbivores

Five secondary metabolites from tropical marine algae and one related com- pound from an herbivorous sea-hare (Aplysidae) were coated, at approximately natural concentrations, onto the palatable seagrass Thalassia testudinum and placed on coral reefs where they could be eaten by the diverse group of herbivorous fishes that occur there. Laboratory feeding assays with the herbivorous sea urchin Diadema antillarum were also conducted. When compared to appropriate controls, the following terpenoid compounds significantly reduced the amount of Thalassia eaten by both Diadema and reef fishes: stypotriol, from the brown seaweed Stypopodium zonale; pachydictyol-A, which is pro- duced by several genera of tropical (Dictyota and Dilophus) and warm-temperate (Pachy- dictyon and Glossophora) brown seaweeds; elatol, from the tropical red alga Laurencia obtusa; and isolaurinterol, which is produced by several tropical and warm-temperate species of Laurencia. Under very mild acid conditions, isolaurinterol is converted to a structurally similar compound, aplysin, found in high concentrations in sea-hares that feed on isolaurinterol-containing Laurencia species. Aplysin did not deter feeding by either type of herbivore. Cymopol, a terpenoid bromohydroquinone from the green alga Cymopolia barbata, significantly reduced feeding by reef fishes but significantly stimulated feeding by Diadema. Pharmacological and crude bioactivity tests suggest that several of these compounds function as generalized toxins. However, these generalized laboratory assays are not nec- essarily good predictors of how compounds will affect feeding by herbivores. For example, pachydictyol-A and stypotriol were equally effective at deterring fishes and Diadema, even though pachydictyol-A shows almost no bioactivity in laboratory assays while stypotriol and its oxidation product, stypoldione, are very bioactive. Herbivory on coral reefs is more intense than in any other habitat studied and the diversity of herbivore types is high. It appears that this intense grazing has provided strong selection for seaweeds that synthesize unique secondary metabolites that significantly reduce the consumption of plants exposed to attack by a diverse group of reef herbivores.