Mechanisms of forest resilience
Ecosystems are dynamic systems with complex responses to environmental variation. In response to pervasive stressors of changing climate and disturbance regimes, many ecosystems are realigning rapidly across spatial scales, in many cases moving outside of their observed historical range of variation into alternative ecological states. In some cases, these new states are transitory and represent successional stages that may ultimately revert to the pre-disturbance condition; in other cases, alternative states are persistent and potentially self-reinforcing, especially under conditions of altered climate, disturbance regimes, and influences of non-native species. These reorganized states may appear novel, but reorganization is a characteristic ecosystem response to environmental variation that has been expressed and documented throughout the paleoecological record. Resilience, the ability of an ecosystem to recover or adapt following disturbance, is an emergent property that results from the expression of multiple mechanisms operating across levels of organism, population, and community. We outline a unifying framework of ecological resilience based on ecological mechanisms that lead to outcomes of persistence, recovery, and reorganization. Persistence is the ability of individuals to tolerate exposure to environmental stress, disturbance, or competitive interactions. As a direct expression of life history evolution and adaptation to environmental variation and stress, persistence is manifested most directly in survivorship and continued growth and reproduction of established individuals. When persistence has been overcome (e.g., following mortality from stress, disturbance, or both), populations must recover by reproduction. Recovery requires the establishment of new individuals from seed or other propagules following dispersal from the parent plant. When recovery fails to re-establish the pre-disturbance community, the ecosystem will assemble into a new state. Reorganization occurs along a gradient of magnitude, from changes in the relative dominance of species present in a community, to individual species replacements within an essentially intact community, to complete species turnover and shift to dominance by plants of different functional types, e.g. transition from forest to shrub or grass dominance. When this latter outcome is persistent and involves reinforcing mechanisms, the resulting state represents a vegetation type conversion (VTC), which in this framework represents an end member of reorganization processes. We explore reorganization in greater detail as this phase is increasingly observed but the least understood of the resilience responses. This resilience framework provides a direct and actionable basis for ecosystem management in a rapidly changing world, by targeting specific components of ecological response and managing for sustainable change.