Leaf size and leaf display of thirty-eight tropical tree species.

Trees forage for light through optimal leaf display. Effective leaf display is determined by metamer traits (i.e., the internode, petiole, and corresponding leaf), and thus these traits strongly co-determine carbon gain and as a result competitive advantage in a light-limited environment. We examined 11 metamer traits of sun and shade trees of 38 coexisting moist forest tree species and determined the relative strengths of intra- and interspecific variation. Species-specific metamer traits were related to two variables that represent important life history variation; the regeneration light requirements and average leaf size of the species. Metamer traits varied strongly across species and, in contrast to our expectation, showed only modest changes in response to light. Intra- and interspecific responses to light were only congruent for a third of the traits evaluated. Four traits, amongst which leaf size, specific leaf area (SLA), and leaf area ratio at the metamer level (LAR) showed even opposite intra- and interspecific responses to light. Strikingly, these are classic traits that are thought to be of paramount importance for plant performance but that have completely different consequences within and across species. Sun trees of a given species had small leaves to reduce the heat load, but light-demanding species had large leaves compared to shade-tolerants, probably to outcompete their neighbors. Shade trees of a given species had a high SLA and LAR to capture more light in a light-limited environment, whereas shade-tolerant species have well-protected leaves with a low SLA compared to light-demanding species, probably to deter herbivores and enhance leaf lifespan. There was a leaf-size-mediated trade-off between biomechanical and hydraulic safety, and the efficiency with which species can space their leaves and forage for light. Unexpectedly, metamer traits were more closely linked to leaf size than to regeneration light requirements, probably because leaf-size-related biomechanical and vascular constraints limit the trait combinations that are physically possible. This suggests that the leaf size spectrum overrules more subtle variation caused by the leaf economics spectrum, and that leaf size represents a more important strategy axis than previously thought.