The paradox of the "paradox of the plankton"

One of the central orienting questions in biodiversity theory and ecology is the "paradox of the plankton", which asks how it is possible for many species to coexist on limited resources given the tendency for competition to exclude species. Over the past five decades, ecologists have offered dozens of solutions to the paradox, invoking game theory, chaos, stochastics, and many other concepts. Despite the plentitude of solutions to the paradox, ecologists continue to offer up novel solutions. Ocean modellers are now faced with the opposite paradox: given the overabundance and the diversity of solutions to the paradox, what is the appropriate way to build coexistence into ecosystem models? Ocean ecosystem models have a very standardized form-nutrient-phytoplankton-zooplankton (NPZ)-type systems of differential equations-where competitive exclusion is a common model behaviour. We suggest approaching the problem from the perspective of community-level patterns. We offer a prototype for building coexistence into NPZ models. The model allows for diverse assemblages of phytoplankton or zooplankton groups to persist and produces accurate community-level patterns. The approach is simple, adding only one additional parameter, and allows us to test the effects of trait distributions and environmental variables on diversity.