Bottleneck stages determine resilience in successional communities

Successional communities often recover slowly because progression stalls at persistent stages that resist replacement. Here, we show that resilience in such systems is frequently governed by a single bottleneck stage with the lowest effective exit rate. Using empirical transition ma-trices from intertidal and plant communities, we demonstrate that altering the bottleneck has a much larger effect on resilience than modifying any other stage. We then show that this bottle-neck principle emerges naturally from both Markov and continuous-time models of succession. Specifically that the slowest return to equilibrium is controlled primarily by the stage with the smallest effective exit rate, which also dominates the mean first-passage time to late succession. These results provide a simple biological interpretation of resilience in successional communities and suggest that management efforts are most effective when they target the stage that limits the pace of succession.