Wildfire disturbance and productivity as drivers of plant species diversity across spatial scales

Wildfires influence many temperate terrestrial ecosystems worldwide. Historical environmental heterogeneity created by wildfires has been altered by human activities and will be impacted by future climate change. Our ability to predict the impact of wildfire-created heterogeneity on biodiversity is limited because few studies have investigated variation in community composition (beta-diversity) in response to fire. Wildfires may influence beta-diversity through several ecological mechanisms. First, highseverity fires may decrease beta-diversity by homogenizing species composition when they create landscapes dominated by disturbance-tolerant or rapidly colonizing species. In contrast, mixed-severity fires may increase beta-diversity by creating mosaic landscapes containing habitats that support species with differing environmental tolerances and dispersal traits. Moreover, the effects of fire severity on betadiversity may change depending on site conditions. Disturbance is hypothesized to increase local species richness at higher productivity and decrease local species richness at lower productivity, a process that can have important, but largely unexamined, consequences on beta-diversity in fire-prone ecosystems. We tested these hypotheses by comparing patterns of beta-diversity and species richness across 162 plant communities in three sites that span a large-scale gradient in climate and productivity in the Northern Rockies of Montana. Within each site, we used spatially explicit fire-severity data to stratify sampling across unburned forests and forests burned with mixed-and high-severity wildfires. We found that betadiversity (community dispersion) of forbs was higher in mixed-severity compared to high-severity fire, regardless of productivity. Counter to our predictions, local species richness of forbs was higher in burned landscapes compared to unburned landscapes at the low-productivity site, but lower in burned landscapes at the high-productivity site. This pattern may be explained by rapid regeneration of woody plants after fire in high-productivity forests. Moreover, forbs and woody plants had disproportionately higher overall species richness in mixed-severity fire compared to high-severity fire, but only at the low-productivity site. These patterns suggest that mixed-severity fires promote higher landscape-level biodiversity in lowproductivity sites by increasing species turnover across landscapes with a diverse mosaic of habitats. Our study illustrates the importance of understanding the mechanisms by which patterns of wildfire severity interact with environmental gradients to influence patterns of biodiversity across spatial scales.