Myrmecophytic Cecropia: antiherbivore defenses under different nutrient treatments.

In greenhouse experiments, we compared putative biotic, chemical, physical and phenological defenses of six myrmecophytic Cecropia species cultivated under high and low nutrient regimes. We tested the intraspecific predictions of the C:N balance hypothesis for a broader range of defenses than included in other studies to date. Treatment effects on defenses appear to depend on the nutrient constituents of those defenses. Only strictly carbon-based defenses such as tannins and phenolics reached higher concentrations at the lower nutrient level. The production of glycogen-rich and membrane-bound Müllerian body ant rewards (MBs) increased with greater levels of both nutrients (this study) and light (Folgarait and Davidson 1994). In contrast, lipid- and amino acid-rich pearl body food rewards (PBs) were produced in greater numbers under conditions of high nutrient levels (this study) and low light (Folgarait and Davidson 1994), both of which should have contributed to a relative excess of nitrogen. Nutrient effects on toughness and leaf expansion rates (perhaps serving as phenological defenses) were inconsistent with the predictions of the C:N balance hypothesis. Mature leaves are protected principally by chemical and physical defenses, and new leaves, by biotic defenses. As in a previous study, interspecific comparisons agreed with the resource availability theory of plant defense. Plant investment in immobile defenses (tannins and phenolics, and leaf toughness), and in a defense with high initial construction costs (trichilia differentiated to produce MBs) were greater in each of three comparatively slow-growing "gap" Cecropia typical of small openings in primary forest, than in closely related and fast-growing "pioneer" species of large-scale disturbances (riparian edge and land slips). In contrast, both production of PBs (with negligible initial construction costs) and leaf expansion rates were greater in pioneers than in gap species. Rapid onset of biotic defenses during new leaf development (earlier in pioneers) may reduce new leaf herbivory in all species.