Abstract
Increasingly, overstory retention is being used in forests traditionally managed for single-cohort structure. One rationale for retention is that residual stand structure better resembles the complex structure of forests after natural disturbance, helping to perpetuate ecosystem functions dependent on that structure. The benefits of retention come at the cost of reduced survival and growth of regeneration because of competition with residual trees. We argue that inhibition of regeneration depends not only on the number and size of residual trees, but also on their spatial arrangement, which ranges from dispersed to aggregated. We use a model of competition at the scale of seedlings to hypothesize that maximum stand-level resource availability, seedling growth, and seedling survival occur with aggregate retention, rather than dispersed retention, even with constant residual basal area. We test our hypotheses with a silvicultural experiment in longleaf pine (Pinus palustris) in Georgia, USA. Replicated treatments included an uncut control, dispersed retention, small-aggregate retention, and large-aggregate retention. We measured light, soil nitrogen, soil moisture, and growth of longleaf pine seedlings across the full range of overstory conditions in each treatment. Postharvest basal areas in the cut treatments were similar. Gap light index increased from the control to large-aggregate retention, as did nitrogen availability, measured on exchange resins. Nitrogen mineralization did not differ among treatments, nor did soil moisture or temperature. Seedling biomass increment increased significantly from the control to large-aggregate retention. Survival did not differ among treatments. We argue that these results are a consequence of exponential relationships between overstory competition intensity, resource availability, and seedling growth. Given this relationship, resources and seedling growth are low across a wide range of decreasing overstory competitor abundance but increase exponentially only at very low competitor abundance. This seedling-scale model translates into maximum stand scale resource availability and seedling growth with large-aggregate retention, compared to dispersed retention, because the probability of a seedling occupying a site free of overstory competition is greater with the former. Our research shows that one can improve competitive environments for regeneration by manipulating spatial distribution of residual trees without sacrificing the ecological benefits of overstory retention. Corresponding Editor: D. L. Peterson
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