The effect of natural disturbances on forest biodiversity: an ecological synthesis.

Disturbances alter biodiversity via their specific characteristics, including severity and extent in the landscape, which act at different temporal and spatial scales. Biodiversity response to disturbance also depends on the community characteristics and habitat requirements of species. Untangling the mechanistic interplay of these factors has guided disturbance ecology for decades, generating mixed scientific evidence of biodiversity responses to disturbance. Understanding the impact of natural disturbances on biodiversity is increasingly important due to human-induced changes in natural disturbance regimes. In many areas, major natural forest disturbances, such as wildfires, windstorms, and insect outbreaks, are becoming more frequent, intense, severe, and widespread due to climate change and land-use change. Conversely, the suppression of natural disturbances threatens disturbance-dependent biota. Using a meta-analytic approach, we analysed a global data set (with most sampling concentrated in temperate and boreal secondary forests) of species assemblages of 26 taxonomic groups, including plants, animals, and fungi collected from forests affected by wildfires, windstorms, and insect outbreaks. The overall effect of natural disturbances on α-diversity did not differ significantly from zero, but some taxonomic groups responded positively to disturbance, while others tended to respond negatively. Disturbance was beneficial for taxonomic groups preferring conditions associated with open canopies (e.g. hymenopterans and hoverflies), whereas ground-dwelling groups and/or groups typically associated with shady conditions (e.g. epigeic lichens and mycorrhizal fungi) were more likely to be negatively impacted by disturbance. Across all taxonomic groups, the highest α-diversity in disturbed forest patches occurred under moderate disturbance severity, i.e. with approximately 55% of trees killed by disturbance. We further extended our meta-analysis by applying a unified diversity concept based on Hill numbers to estimate α-diversity changes in different taxonomic groups across a gradient of disturbance severity measured at the stand scale and incorporating other disturbance features. We found that disturbance severity negatively affected diversity for Hill number q = 0 but not for q = 1 and q = 2, indicating that diversity-disturbance relationships are shaped by species relative abundances. Our synthesis of α-diversity was extended by a synthesis of disturbance-induced change in species assemblages, and revealed that disturbance changes the ß-diversity of multiple taxonomic groups, including some groups that were not affected at the α-diversity level (birds and woody plants). Finally, we used mixed rarefaction/extrapolation to estimate biodiversity change as a function of the proportion of forests that were disturbed, i.e. the disturbance extent measured at the landscape scale. The comparison of intact and naturally disturbed forests revealed that both types of forests provide habitat for unique species assemblages, whereas species diversity in the mixture of disturbed and undisturbed forests peaked at intermediate values of disturbance extent in the simulated landscape. Hence, the relationship between α-diversity and disturbance severity in disturbed forest stands was strikingly similar to the relationship between species richness and disturbance extent in a landscape consisting of both disturbed and undisturbed forest habitats. This result suggests that both moderate disturbance severity and moderate disturbance extent support the highest levels of biodiversity in contemporary forest landscapes.

Assessing the ecological impacts of biomass harvesting along a disturbance severity gradient.

Disturbance is a central driver of forest development and ecosystem processes with variable effects within and across ecosystems. Despite the high levels of variation in disturbance severity often observed in forests following natural and anthropogenic disturbance, studies quantifying disturbance impacts often rely on categorical classifications, thus limiting opportunities to examine potential gradients in ecosystem response to a given disturbance or management regime. Given the potential increases in disturbance severity associated with global change, as well as shifts in management regimes related to procurement of biofuel feedstocks, there is an increasing need to quantitatively describe disturbance severity and associated responses of forest development, soil processes, and structural conditions. This study took advantage of two replicated large-scale studies of forest biomass harvesting in Populus tremuloides and Pinus bansksiana forests, respectively, to develop and test the utility of a continuous, quantitative, disturbance severity index (DSI) for describing postharvest response of plant communities and nutrient pools to different levels of biomass removal and legacy retention (i.e., live trees and coarse woody material). There was a high degree of variability in DSI within categorical treatments associated with different levels of legacy retention and regression models using DSI as a predictor explained a portion of the variation (>50%) for many of the ecosystem- and community-level responses to biomass harvesting examined. Nutrient losses associated with biomass harvesting were positively related to disturbance severity, particularly in P. tremuloides forests, with postharvest nutrient availability generally declining along the gradient of impacts. Consistent with expectations from ecological theory, species richness and diversity of woody plant communities were greatest at intermediate disturbance severities and regeneration densities of dominant trees species were most abundant at highest levels of disturbance. Although categorical benchmarks will continue to be the primary way through which management guidelines are conveyed to practitioners, evaluation of their effectiveness at sustaining ecosystem functioning should be through continuous analyses, such as the DSI approach used in this study, to allow for the more precise identification of thresholds that ensure a range of desirable outcomes exist across managed landscapes.

Stand-replacing wildfires increase nitrification for decades in southwestern ponderosa pine forests.

Stand-replacing wildfires are a novel disturbance within ponderosa pine (Pinus ponderosa) forests of the southwestern United States, and they can convert forests to grasslands or shrublands for decades. While most research shows that soil inorganic N pools and fluxes return to pre-fire levels within a few years, we wondered if vegetation conversion (ponderosa pine to bunchgrass) following stand-replacing fires might be accompanied by a long-term shift in N cycling processes. Using a 34-year stand-replacing wildfire chronosequence with paired, adjacent unburned patches, we examined the long-term dynamics of net and gross nitrogen (N) transformations. We hypothesized that N availability in burned patches would become more similar to those in unburned patches over time after fire as these areas become re-vegetated. Burned patches had higher net and gross nitrification rates than unburned patches (P < 0.01 for both), and nitrification accounted for a greater proportion of N mineralization in burned patches for both net (P < 0.01) and gross (P < 0.04) N transformation measurements. However, trends with time-after-fire were not observed for any other variables. Our findings contrast with previous work, which suggested that high nitrification rates are a short-term response to disturbance. Furthermore, high nitrification rates at our site were not simply correlated with the presence of herbaceous vegetation. Instead, we suggest that stand-replacing wildfire triggers a shift in N cycling that is maintained for at least three decades by various factors, including a shift from a woody to an herbaceous ecosystem and the presence of fire-deposited charcoal.