The path back: oaks (Quercus spp.) facilitate longleaf pine (Pinus palustris) seedling establishment in xeric sites

Understanding plant–plant facilitation is critical for predicting how plant community function will respond to changing disturbance and climate. In longleaf pine (Pinus palustris Mill.) ecosystems of the southeastern United States, understanding processes that affect pine reproduction is imperative for conservation efforts that aim to maintain ecosystem resilience across its wide geographic range and edaphic gradients. Variation in wildland fire and plant–plant interactions may be overlooked in “coarse filter” restoration management, where actions are often prescribed over a variety of ecological conditions with an assumed outcome. For example, hardwood reduction techniques are commonly deemed necessary for ecological restoration of longleaf pine ecosystems, as hardwoods are presumed competitors with longleaf pine seedlings. Natural regeneration dynamics are difficult to test experimentally given the infrequent and irregular mast seed events of the longleaf pine. Using a longterm, largescale restoration experiment and a longterm monitoring data site at Eglin Air Force Base, Florida (USA), this study explores the influence of native fireintolerant oaks on longleaf regeneration. We test for historical observations of hardwood facilitation against the null hypothesis of competitive exclusion. Our results provide evidence of hardwood facilitation on newly germinated longleaf pine seedlings (<2 yr old) after two mast seeding events (1996, 2011). Using regressiontree and Kaplan–Meier survival analyses, we found that deciduous oak midstory density was the most significant variable associated with longleaf pine seedling survival rates in the first 2 yr after germination. We found that as few as 43 oak midstory stems ha−1 were sufficient to facilitate seedling survival, but as many as 1400 stems ha−1 maintained facilitation without competitive exclusion of seedlings. We found that 1.5yrold pine seedlings were more moisture stressed under more open canopy conditions when compared to those immediately adjacent to a midstory oak canopy. Recognition that deciduous oaks are important facilitators of longleaf seedling establishment on xeric sites represents a significant departure from conventional wisdom and current management practices that has largely focused on competitive exclusion. This points to a critical role of a deciduous oak midstory of moderate densities for longterm ecosystem resilience in xeric longleaf pine ecosystems in light of climate