Thresholds in woody and herbaceous component co-existence inform the restoration of a fire-dependent community

Questions A paradoxical co-existence challenges woodland and savanna restoration worldwide: How are shade-intolerant, flammable herbaceous layers promoted while maintaining the shade-casting, more fire-sensitive woody regeneration that sustains overstorey structure? Where restoration success consisted of robust, diverse herbaceous layers and vigorous, well-stocked Pinus echinata regeneration (hereafter, shortleaf-bluestem response), we asked: are there targetable conditions of overstorey, understorey, topography, and proximity to mature Pinus echinata that simultaneously maximize desired woody and herbaceous understorey components? Do these conditions and dependent responses differ across canopy disturbance level and fire season? Location Cumberland Plateau, Tennessee, USA (36 degrees 04 ' 8.11 " N, 84 degrees 50 ' 38.36 " W). Methods We measured 12 shortleaf-bluestem response and 17 explanatory condition variables at 345 plots spanning an experimental restoration gradient (canopy disturbance level and fire season combinations). We ordinated variation and identified response thresholds using a multivariate regression tree. Differences across tree groupings and splits associated response thresholds with specific explanatory conditions. Results Pockets of substantial Pinus echinata regeneration (>3,000 stems/ha), C-4 grass density (>40,000 ramets/ha), and herbaceous diversity (increase from 22 to 205 species) occurred 7-14 years after canopy disturbance and 3-8 fires. Such shortleaf-bluestem response was maximized at 3 m(2)/ha residual tree basal area, 11% canopy closure, reduced midstorey density (5,000 small-sapling stems/ha), and southwesterly aspects within 70 m of mature Pinus echinata. In contrast, shortleaf-bluestem response was negligible at 11.3 m(2)/ha basal area and 68% canopy closure. Fire season, snag basal area, slope, and slope position effects were minimal. Conclusions We identified specific conditions fostering the co-existence of desired herbaceous and woody understorey components, addressing a major woodland and savanna restoration challenge and expanding on previous threshold concept applications by simultaneously considering multiple desired responses. Results can direct the restoration of imperiled shortleaf-bluestem communities east of the Mississippi River, USA, where work has been scarce or ineffective, and similar approaches could inform fire-dependent woodland and savanna restoration worldwide.