Understanding change in benthic marine systems.

Background: The unprecedented influence of human activities on natural ecosystems in the 21st century has resulted in increasingly frequent large-scale changes in ecological communities. This has heightened interest in understanding such changes and effective means to manage them. Accurate interpretation of state changes is challenging because of difficulties translating theory to empirical study, and most theory emphasizes systems near equilibrium, which may not be relevant in rapidly changing environments.

Scope: We review concepts of long-transient stages and phase shifts between stable community states, both smooth, continuous and discontinuous shifts, and the relationships among them. Three principal challenges emerge when applying these concepts. The first is how to interpret observed change in communities - distinguishing multiple stable states from long transients, or reversible shifts in the phase portrait of single attractor systems. The second is how to quantify the magnitudes of three sources of variability that cause switches between community states: (1) 'noise' in species' abundances, (2) 'wiggle' in system parameters and (3) trends in parameters that affect the topography of the basin of attraction. The third challenge is how variability of the system shapes evidence used to interpret community changes. We outline a novel approach using critical length scales to potentially address these challenges. These concepts are highlighted by a review of recent examples involving macroalgae as key players in marine benthic ecosystems.

Conclusions: Real-world examples show three or more stable configurations of ecological communities may exist for a given set of parameters, and transient stages may persist for long periods necessitating their respective consideration. The characteristic length scale (CLS) is a useful metric that uniquely identifies a community 'basin of attraction', enabling phase shifts to be distinguished from long transients. Variabilities of CLSs and time series data may likewise provide proactive management measures to mitigate phase shifts and loss of ecosystem services. Continued challenges remain in distinguishing continuous from discontinuous phase shifts because their respective dynamics lack unique signatures.