The stratification theory for plant coexistence promoted by one-sided competition

It is an essential feature of plants that leaves at higher levels have better access to light than those at lower levels. Thus, larger plants generally enjoy greater success in competing for light than smaller ones. We analyse the effect of such size-asymmetry, or one-sided competition, on the successful coexistence of plant species, using an analytically tractable model for stratified populations, in which a plant in the same layer exhibits the same crowding effect as any other, irrespective of species. A two-layer population that is reproductive in upper layer and juvenile in lower layer has a uniquely stable (plant-size-weighted) equilibrium density, as long as its fecundity is sufficient to compensate for its mortality rate. We also calculate a unique threshold lower-layer density of this layered population when there is no upper-layer plant. This threshold lower-layer density is larger than the weighted equilibrium density with upper layer, except for the case of perfect two-sided competition. A two-layer species can stably coexist with a one-layer, understorey species as a result of one-sided, but not two-sided competition. The coexistence condition is that the equilibrium density of the one-layer species lies between the threshold lower-layer density and the equilibrium density of the two-layer species. For an understorey species to coexist successfully with a two-layer species, any advantage in demographic performance, most prominently in a sufficiently high fecundity per plant must offset the disadvantage of living in dark conditions. Results from a model of multi-layer populations suggest that several species differing in terms of maximum layer and fecundity can coexist under conditions of one-sided competition. We demonstrate an example of the stable coexistence of eight species. The inter-specific trade-offs predicted by the model correspond to patterns observed in a rain forest. Synthesis. We propose a stratification theory that explains the generation and maintenance of the successful coexistence of plant species. Under the condition of one-sided competition, a canopy population that takes advantage of escaping from understorey competition shows an ability to invade an understorey with a density higher than its own equilibrium density, and which offers opportunities for an understorey population with high fecundity and/or shade tolerance to coexist. The predicted coexistence of species that share maximum canopy height is most pronounced for trees of tropical rain forests.