The Roles of Competition and Facilitation in Producing Zonation Along an Experimental Flooding Gradient: A Tale of Two Tails with Ten Freshwater Marsh Plants

Plant zonation is conspicuous in wetlands. The cause is frequently assumed to be the direct physiological effects of physical factors (termed ‘stress’), however many experiments show that competition and facilitation also cause zonation patterns. We conducted a field experiment with freshwater marsh emergent plants to test the causes of zonation along a single stress gradient: flooding duration. We constructed an experimental wetland with ten flooding levels to ensure that the environmental conditions represented the full range of potential flooding levels, from never flooded to continually flooded. We planted ten common marsh plants with varied ecology along the flooding duration gradient. We grew them alone and in mixture for three years and measured changes in the minimum and maximum limits, the mode and the range of distribution, and interaction importance. The mode of distribution did not shift, whether species were grown alone or with neighbours. We found strong effects of competition under low flooding stress. We found no effects from facilitation under high flooding stress. Flooding duration alone controlled the lower limits of plants. The effects of competition were intense enough to eliminate half of the species within three growing seasons. Our experiment showed that competition and physical stresses, but not facilitation, controls the zonation of emergent macrophytes along a flooding duration gradient, at least in freshwater wetlands. Models guiding wetland restoration need to include competition as well as flood duration as causal factors, but not facilitation.