Canopy-mediated microclimate refugia do not match narrow regeneration niches in a managed dry conifer forest

Abstract

Structurally heterogeneous forest conditions provide diverse microclimates which can support juvenile tree regeneration, survival, and growth. However, the extent to which juvenile trees depend on moderated microclimates depends on their physical maturity and species-specific tolerances for environmental stresses related to variables like soil moisture, heat, and shade. Linking juvenile responses to microclimates for different life-stages and species facilitates more nuanced understanding of regeneration niches and is especially important in dry conifer forests where structurally heterogeneous conditions are a frequent objective of forest restoration treatments. In this study, we sampled microclimate conditions and planted seeds and greenhouse-grown seedlings across a range of overstory structure and microclimate conditions in a heterogeneous forest restoration treatment in Colorado, USA. We assessed how fine-scale (∼1–20 m) spatial variation in overstory structure and topography related to variability in microclimate conditions and, collectively, how these biophysical conditions influenced survival and growth of ponderosa pine (Pinus ponderosa var. scopulorum) and Douglas-fir (Pseudotsuga menziesii var. glauca) over 3 years. We found evidence of strong canopy buffering of mean daily vapor pressure deficit (VPD), but buffering of daily maximum VPD was moderate, and buffering of soil moisture was weak except in the hottest and driest month of our 3-year study. Although canopy cover provided refuge from hot and dry conditions, survival and growth of juvenile trees across life-stages and species were greatest in above-average canopy openness, with warm and dry microclimate conditions in May. The spatial patterns of seedling establishment relative to canopy cover, driven by early-season microclimate conditions, did not match canopy-mediated, moderate microclimates later in growing seasons when weather conditions were most limiting. Importantly, our results reflect a specific regeneration trajectory initiated by favorable early-growing season conditions in the first year of study, which promoted survival and growth resilience through subsequent limiting conditions, most notably a hot and dry second year of study. The patterns observed in this study are consistent with drivers of landscape-scale regeneration previously identified for dry conifer forests and show that overstory structure and microclimate variation can mediate these patterns at very fine spatial scales. The narrow spatial and temporal ranges of conditions which supported survival and growth in this study highlight opportunities for restoration treatment planning in similar forests, but also underscore the susceptibility of these dominant dry forest species to potential future changes in the timing and intensity of abiotic conditions which limit regeneration success.