Optimizing the complexity of phytoplankton functional group modeling: An allometric approach

Elucidating patterns and mechanisms that shape phytoplankton assemblages is a popular area of research for empirical and theoretical ecologists. Despite the daunting complexity of phytoplankton dynamics, much of our current understanding has been based on simple models describing food-web interactions with few differential equations. Skeptical views in the literature raise concerns about the increasing model complexity and advice to seek parsimony rather than simplicity. To address this controversy (simple versus complex models), we propose the introduction of an extra layer of causality into plankton models by connecting algal processes (maximum growth rates, nutrient kinetics, settling velocities, metabolic rates) with species-specific morphological features (cell volume, surface-to-volume ratio, shape). In this study, we demonstrate the capacity of a size-based plankton model to reproduce observed water quality patterns (phosphate, total phosphorus, nitrate, total ammonia, total nitrogen, chlorophyll a, and total zooplankton biomass) in the Hamilton Harbour, Ontario. Consistent with empirical evidence, our modeling analysis showed that small algal species have a distinct competitive advantage in summer epilimnetic environments across the range of cell volume and nutrient loading conditions examined; especially, when they are characterized by higher optimal temperature for growth. Strong top-down pressure mediated by high zooplankton abundance effectively controls the standing biomass of phytoplankton species that can otherwise realize high growth rates under the conditions typically prevailing in the end-of-summer epilimnetic environments (e.g., higher temperature optima, higher tolerance in low water clarity). Under high zooplankton control, the secondary variations of phytoplankton are modulated by the ambient phosphorus levels and the size-based strategies for resources procurement, such as the regulation of nutrient transport kinetics. By contrast, when the summer algal assemblage is released by the zooplankton grazing, the exceedance of critical phytoplankton biomass levels and the likelihood of harmful algal blooms are determined by the multitude of factors that shape inter-specific competition patterns (e.g., relative abundance of competing species, nutrient uptake kinetics). Our study evaluates the strengths and weaknesses of this approach and identifies future directions that would provide operational models founded upon concepts of allometry.