A multivariate model of plant species richness in forested systems: Old-growth montane forests with a long history of fire

Recently, efforts to develop multivariate models of plant species richness have been extended to include systems where trees play important roles as overstory elements mediating the influences of environment and disturbance on understory richness. We used structural equation modeling to examine the relationship of understory vascular plant species richness to understory abundance, forest structure, topographic slope, and surface fire history in lower montane forests on the North Rim of Grand Canyon National Park, USA based on data from eighty-two 0.1 ha plots. The questions of primary interest in this analysis were: (1) to what degree are influences of trees on understory richness mediated by effects on understory abundance? (2) To what degree are influences of fire history on richness mediated by effects on trees and/or understory abundance? (3) Can the influences of fire history on this system be related simply to time-since-fire or are there unique influences associated with long-term fire frequency? The results we obtained are consistent with the following inferences. First, it appears that pine trees had a strong inhibitory effect on the abundance of understory plants, which in turn led to lower understory species richness. Second, richness declined over time since the last fire. This pattern appears to result from several processes, including (1) a post-fire stimulation of germination, (2) a decline in understory abundance, and (3) an increase over time in pine abundance (which indirectly leads to reduced richness). Finally, once time-since-fire was statistically controlled, it was seen that areas with higher fire frequency have lower richness than expected, which appears to result from negative effects on understory abundance, possibly by depletions of soil nutrients from repeated surface fire. Overall, it appears that at large temporal and spatial scales, surface fire plays an important and complex role in structuring understory plant communities in old-growth montane forests. These results show how multivariate models of herbaceous richness can be expanded to apply to forested systems. Copyright ?? Oikos 2006.