Spatial patterns of living and dead small trees in subalpine Norway spruce forest reserves in Switzerland

Abstract

Spatial patterns can reveal a lot about ecological processes, but our knowledge of the spatial ecology of tree regeneration at a fine scale is quite limited. Therefore, we studied the spatial patterns of living and dead small trees in two subalpine Norway spruce forest reserves in Switzerland (Scatlè and Bödmerenwald) using three types of analyses. First, we investigated the distances of small trees to the nearest large neighboring tree and, by using maximum distances as indicator, inferred the size of forest gaps, detecting mainly forest gaps of small size, although with two exceptions that were driven by large-scale disturbances. Second, we accounted for spatial inhomogeneity in the pattern of small and large trees (i.e., variations in local tree densities) by including environmental covariates in point pattern models. Latitude (within the forest reserve), elevation and aspect contributed significantly to explaining the density of living and dead small trees, and partly of living and dead large trees. Yet, the influence of these environmental covariates varied between the two reserves due to their different topography and peculiar site conditions. Third, we analyzed neighborhood interactions between small and large trees based on the vicinity and size of trees. In both forest reserves, small living trees were randomly dispersed around large dead trees over a broad range of distances and, at certain distances in one reserve, even dispersed away from them. Small living trees further showed clustering around large living trees at short distances and dispersion at large distances. Small dead trees featured mainly a random pattern, although with a tendency to cluster around large neighbors at short distances, irrespective whether these were living or dead. Yet, the weakening of clustering with increasing distances indicates that the influence of large trees on small trees varies with spatial scale and thus that these neighborhood interactions are scale-dependent. Overall, our study contributes to a better understanding of the spatial ecology of mortality in small trees and ultimately of tree regeneration processes and stand dynamics in mountain forests.