Persistent effects of fire severity on ponderosa pine regeneration niches and seedling growth

Several recent studies have documented how fire severity affects the density and spatial patterns of tree regeneration in western North American ponderosa pine forests. However, less is known about the effects of fire severity on fine-scale tree regeneration niche attributes such as understory plant composition and cover, surface fuel abundance, and soil properties, or how these attributes in turn affect regenerating ponderosa pine growth. Using 1-m(2) plots centered on 360 ponderosa pine seedlings that regenerated naturally after the Pumpkin Fire in 2000 in Arizona, we quantified regeneration niche attributes 13 years post-fire and measured their associations with seedling growth 11-16 years post-fire. Plots were established in a) unburned, b) moderate-severity, and two types of high-severity (100% tree mortality) burns, either c) adjacent to residual live forest edges (high-severity edge plots), or d) > 200 m from any residual live trees (high-severity interior plots). We found that all burned plots had greater understory plant species richness, percent cover of forbs, exotic plants and coarse wood, as well as higher soil pH, sand and gravel content, and lower soil clay content compared to unburned plots. High-severity burn plots had the greatest total understory plant and shrub cover, the most herbaceous fine fuel biomass, and tended to have the highest soil nitrogen content compared to other burn severity classes. Ponderosa pine growth (i.e., stem diameter at root collar (DRC) and length of terminal leader) was lowest in the unburned compared to burned plots, and ponderosa pine terminal leader growth was consistently greater in the high-severity edge plots compared to other seventies. Finally, niche characteristics such as overstory tree canopy cover ( - ), soil phosphate (+), and cover of coarse wood ( +), forbs (+), and the native grasses, mountain muhly (+) and squirreltail (-), were important explanatory variables of ponderosa pine growth. Exotic plant cover did not have a negative association with ponderosa pine growth. These results suggest that if ponderosa pine seeds can disperse and germinate, and if seedlings can survive the first few critical years after germination, then low overstory canopy cover and abundant forbs or coarse wood may be associated with increased growth rates. Alternatively, forbs may be responding to the same site benefits as the seedling; and abundant forbs, coarse wood, and fine fuels might also put seedlings at increased risk of mortality from subsequent fire, at least until they are taller and more fire resistant.