An experimental analysis of the effects of light and zooplankton on coral zonation.

In Panamá, vertical zonation of coral species is well marked; branching corals (Pocillopora spp.) predominate in shallow (1-6 m) water while massive forms (Pavona spp.) occupy the deeper (6-10 m) areas of the reef. To test the hypothesis that this zonation represents differential resource utilization (i.e., niche partitioning of food resources), one year manipulative field experiments were conducted at two depths (1 m and 7 m below Mean Lower Sea Level) assessing the relative contribution of light and zooplankton to the nutrition of three Pacific corals: Pocillopora damicornis, Pavona clavus and Pavona gigantea. Also tested were the related hypotheses that (i) energy for the maintenance and growth of corals comes mainly from light, independent of zooplankton supply and (ii) Porter's model, in which coral morphology is a predictor of the phototrophic-heterotrophic capabilities of a particular species. That is, corals with a branching morphology and a small polyp diameter (and short tentacle length) should be primarily phototrophic while those species with larger polyps and/or a massive form should show a greater degree of heterotrophy.The comparison of caged versus control corals indicated that the branching coral, Pocillopora damicornis (polyp diameter - 1 mm) grew independent of zooplankton supply (>95μ) but was markedly affected by shading. Pavona clavus (massive form with intermediate size polyps - 2 mm) was also negatively affected by shading but this effect was minimized when zooplankton was present. Surprisingly, under ambient light conditions, the growth of this specics was independent of zooplankton (>95μ); perhaps indicating a facultative reliance on zooplankton especially in shallow water. The third species, Pavona gigan-tea (massive form with largest polyps - 3 mm) was highly dependent on both light and zooplankton and was unable to compensate for the effects of shading by zooplankton feeding. This high dependence on light was unexpected since large polyps, according to Porter's model, would predict a more heterotrophic existence.Under all treatments, in both shallow and deep water, Pocillopora was found to grow more rapidly than either massive species. Thus, this study indicates that zonation on eastern Pacific reefs cannot be explained by depth-related differences in nutritional requirements among the species present. These results do provide the first long-term, experimental field evidence that corals are largely phototrophic organisms. The validity of Porter's model attempting to correlate phototrophic-heterotrophic abilities to morphological characteristics is, at least, partially verified: species with larger tentacles do appear to utilize zooplankton to a greater extent than those species with smaller tentacles. However, the significance of corallum morphology in predicting the mode of nutrient acquisition is unclear. Rather, behavioral patterns (tentacle expansion-contraction cycles) and location on the reef may be more important factors to consider.