Recruitment facilitation can drive alternative states on temperate reefs

How the combination of positive and negative species interactions acts to drive community dynamics is a fundamental question in ecology. Here we explore one aspect of this question by expanding the theory of predator-mediated coexistence to include the potential role of facilitation between the predator and inferior competitor. To motivate and illustrate our simple model, we focus on sea-urchin-algae interactions in temperate rocky reef systems and incorporate recruitment facilitation, a common characteristic of marine systems. Specifically, the model represents sea urchin grazing on macroalgae, macroalgal competition with crustose coralline algae (CCA), and facilitation of sea urchin recruitment to CCA. These interactions generate alternative stable states, one dominated by macroalgae and the other by urchins, which do not occur when recruitment facilitation of urchins to CCA is ignored. Therefore, recruitment facilitation provides a possible mechanism for alternative kelp forest and urchin barren states in temperate marine systems, where storm events or harvesting of urchins or their predators can drive switches between states that are difficult to reverse. In systems with such dynamics, spatial management such as no-take marine reserves may play a crucial role in protecting community structure by increasing the resilience to shifts between states.