Why plankton communities have no equilibrium: solutions to the paradox

In a classical paper, Hutchinson (1961) argued that the large number of species in most plankton communities is remarkable in view of the competitive exclusion principle, which suggests that in homogeneous, well-mixed environments species that compete for the same resources cannot coexist. Few ideas in aquatic ecology have evoked more research than this `paradox of the plankton'. This review is an effort to put the main solutions to the paradox that have been proposed over the years into perspective. Hutchinson himself already suggested that the explanation could be that plankton communities are not in equilibrium at all due to weather-driven fluctuations. Subsequent research confirmed that such externally imposed variability can allow many species to coexist. Another important point is that in practice the homogeneous well-mixed conditions assumed in the competitive exclusion principle hardly exist. Even the open ocean, for instance, has a spatial complexity resulting from meso-scale vortices and fronts that can facilitate coexistence of species. Perhaps most excitingly, theoretical work on species interactions has given a counter-intuitive new dimension to the understanding of diversity. Various competition and predation models suggest that even in homogeneous and constant environments plankton will never settle to equilibrium. Instead, interactions between multiple species may give rise to oscillations and chaos, with a continuous wax and wane of species within the community. Long-term laboratory experiments support this view. This chaotic behavior implies among other things that plankton dynamics are intrinsically unpredictable in the long run when viewed in detail. Nonetheless, on a higher aggregation level, indicators such as total algal biomass may show quite regular patterns