Xerophytic hardwood retention promotes competition over facilitation in longleaf pine woodlands in the absence of fire
Recent reports of xerophytic hardwood facilitation of longleaf pine seedlings challenge the traditional paradigm that co-occurring hardwood tree species constrain longleaf pine (Pinus palustris Mill.) regeneration. To better understand this dynamic, we established an experiment examining the effects of hardwood retention duration (no retention, one-year retention, four-year retention), overstory basal area (2-to-41 m2 ha-1), and understory vegetation cover on microclimatic conditions and longleaf pine seedling survival, development, physiological performance, and brownspot needle blight (Mycosphaerella dearnessii) infection in the Sandhills Ecoregion of North Carolina, USA. Light availability (μmol m-2 s-1) at the forest floor (~20 cm) was reduced by 27% in the four-year retention treatment which significantly exceeded reductions found in the one-year (<1%) and no retention (<1%) treatments, respectively, post midstory removal. However, midstory treatment had little effect on soil moisture and temperature at 15 and 30 cm depth. Longleaf pine seedling survival (85%), aboveground biomass (7.51 g dry weight), and root collar size (1.53 cm) were significantly highest in the no retention treatment compared to the four-year retention treatment. Between treatments with midstory retention (one-year vs. four-year retention), longleaf pine seedlings were statistically larger in the one-year retention treatment (5.37 g dry weight) compared to the four-year retention treatment (4.84 g dry weight). Seedling brownspot infection did not statistically vary among midstory retention treatments. Across all midstory treatments, overstory basal area had a stronger negative effect on longleaf pine seedling aboveground biomass development, root collar development and brownspot infection compared to midstory hardwood stem density or understory vegetation cover. Moreover, for most measured metrics, overstory basal area had a stronger negative effect on longleaf pine physiological performance compared to midstory hardwood stem density. Collectively, these results indicate that hardwood retention did not facilitate longleaf pine survival and development in the grass stage and that asymmetric competition from the overstory represents a stronger impediment to longleaf pine sapling recruitment than midstory hardwoods.