Mechanisms of succession along the emersion gradient in intertidal rocky shore assemblages

Theoretical models predict that the relative importance of competition and facilitation vary inversely along gradients of abiotic stress, with facilitation dominating under harsh conditions (the so called ‘‘stress-gradient hypothesis’’). To date, very few studies have tested this hypothesis in the framework of succession. Moreover, the generality of the hypothesis is currently under debate and the necessity to examine responses at the community level and using different stress levels has been emphasized. We evaluate the mechanisms of succession across the emersion gradient in a rocky shore system. After complete clearing of experimental plots at four shore heights, we removed two separate components of the assemblage: ephemeral green algae, early colonists and canopy-forming species, main space holders in undisturbed assemblages. We aimed to test two hypotheses: (a) the net effect of ephemeral algae on canopy-forming species shift along the emersion gradient according to the predictions of the stress-gradient hypothesis and (b) the net effect of the canopy-forming group on associate species change, from neutral/negative to positive, as succession progresses and plants grow, especially at the highest shore heights. Our results did not support the predictions of the stress gradient hypothesis in the framework of succession. We did detect changes through time in the effect of canopy-forming species, from negative to positive, but this was not dependent on the shore height. The outcome of interactions depended on the identity and life-stage of the species. Moreover, indirect interactions among species could also create a less predictable relationship between successional mechanisms and the environmental gradient.