Response of the herbaceous layer to natural disturbance in North American forests
Most work on the disturbance ecology of North American forests has focused on the tree canopy and woody understory, with little consideration of the herbaceous layer. Understanding how this species-rich and ecologically important layer responds to natural disturbances is needed as a guide for conservation. As a framework for this review, selected natural disturbances common to North American forests are evaluated in terms of a three-axis model of disturbance severity: percentage of tree canopy removed, percentage of understory vegetation removed, and percentage of forest floor and soil removed or disrupted. Eight factors, which control herbaceous-layer response, are linked to the three axes; these factors vary in dominance, depending on disturbance type and severity. The relative importances of different species' regeneration mechanisms also vary along severity gradients. The disturbance-severity model, along with the linked response factors and regeneration mechanisms, provide a framework for evaluating impacts of both natural and anthropogenic disturbances. Long-term studies of herbaceous-layer response to disturbance are needed, particularly mechanistic studies that focus on the relative importance of the eight factors controlling response. The conceptual model proposed herein needs to be tested across the full spectrum of severity levels. Finally, comparisons between natural and anthropogenic disturbances are needed.Key words: disturbance, forests, herbaceous layer, life-history characteristics, succession, understory.