Integrating physiological and biomechanical drivers of population growth over environmental gradients on coral reefs.

Coral reefs exhibit marked spatial and temporal variability, and coral reef organisms exhibit trade-offs in functional traits that influence demographic performance under different combinations of abiotic environmental conditions. In many systems, trait trade-offs are modelled using an energy and/or nutrient allocation framework. However, on coral reefs, differences in biomechanical vulnerability have major demographic implications, and indeed are believed to play an essential role in mediating species coexistence because highly competitive growth forms are vulnerable to physical dislodgment events that occur with high frequency (e.g. annual summer storms). Therefore, an integrated energy allocation and biomechanics framework is required to understand the effect of physical environmental gradients on species' demographic performance. However, on coral reefs, as in most ecosystems, the effects of environmental conditions on organisms are measured in different currencies (e.g. lipid accumulation, survival and number of gametes), and thus the relative contributions of these effects to overall capacity for population growth are not readily apparent. A comprehensive assessment of links between the environment and the organism, including those mediated by biomechanical processes, must convert environmental effects on individual-level performance (e.g. survival, growth and reproduction) into a common currency that is relevant to the capacity to contribute to population growth. We outline such an approach by considering the population-level performance of scleractinian reef corals over a hydrodynamic gradient, with a focus on the integrating the biomechanical determinants of size-dependent coral colony dislodgment as a function of flow, with the effects of flow on photosynthetic energy acquisition and respiration.