The relative importance of relative nonlinearity and the storage effect in the lottery model

Although it is likely that many coexistence mechanisms contribute to maintenance of species diversity, most approaches to understanding species coexistence proceed as if only one mechanism would be present. In studies of species coexistence in a temporally fluctuating environment, the storage effect, believed to be the most important coexistence mechanism, has been the focus. Although a different coexistence mechanism--relative nonlinearity--is also predicted to arise frequently with environmental variation, its effect has been overshadowed by the storage effect. The relatively nonlinear growth rates on which the mechanism depends arise simply from differences in life history traits. Many kinds of temporal variation can then interact with these nonlinearity differences to create the relative nonlinearity coexistence mechanism. Much is unknown about when this mechanism is important and its total neglect is not justified. Here, we use the lottery model to provide a much needed quantitative assessment of the relative and combined effects of relative nonlinearity and the storage effect. Our analysis takes advantage of recently developed techniques for quantifying coexistence mechanisms when multiple mechanisms operate in concert. We find that relative nonlinearity is able to contribute substantially to species coexistence in the lottery model when two conditions are satisfied: (1) species must differ greatly in their adult death rates, (2) sensitivity of recruitment to environmental variation must be greater for species with larger adult death rates. In addition, relative nonlinearity has a critical role in compensating for a weakened storage effect when there is high correlation between species in their responses to the varying environment. In some circumstances relative nonlinearity is stronger than the storage effect or is even the sole mechanism of coexistence.