Evidence for a Trade-Off Strategy in Stone Oak (Lithocarpus) Seeds between Physical and Chemical Defense Highlights Fiber as an Important Antifeedant

Trees in the beech or oak family (Fagaceae) have a mutualistic relationship with scatter-hoarding rodents. Rodents obtain nutrients and energy by consuming seeds, while providing seed dispersal for the tree by allowing some cached seeds to germinate. Seed predation and caching behavior of rodents is primarily affected by seed size, mechanical protection, macronutrient content, and chemical antifeedants. To enhance seed dispersal, trees must optimize trade-offs in investment between macronutrients and antifeedants. Here, we examine this important chemical balance in the seeds of tropical stone oak species with two substantially different fruit morphologies. These two distinct fruit morphologies in Lithocarpus differ in the degree of mechanical protection of the seed. For 'acorn' fruit, a thin exocarp forms a shell around the seed while for 'enclosed receptacle' (ER) fruit, the seed is embedded in a woody receptacle. We compared the chemical composition of numerous macronutrient and antifeedant in seeds from several Lithocarpus species, focusing on two pairs of sympatric species with different fruit morphologies. We found that macronutrients, particularly total non-structural carbohydrate, was more concentrated in seeds of ER fruits while antifeedants, primarily fibers, were more concentrated in seeds of acorn fruits. The trade-off in these two major chemical components was more evident between the two sympatric lowland species than between two highland species. Surprisingly, no significant difference in overall tannin concentrations in the seeds was observed between the two fruit morphologies. Instead, the major trade-off between macronutrients and antifeedants involved indigestible fibers. Future studies of this complex mutualism should carefully consider the role of indigestible fibers in the foraging behavior of scatter-hoarding rodents.