The importance of small-scale heterogeneity in boreal forests: variation in diversity in forest-floor invertebrates across the succession gradient

Heterogeneity in species assemblages of forest-floor arthropods — carabid beetles, ants and spiders — within and between different forest age classes was studied in the southern Finnish taiga. The importance of processes operating on the local scale (within the movement radius of the species) vs on the regional scale (among forest stands) in determining the observed variation was assessed. Four data sets with different spatial resolutions in mesic forests in the same general study area were used. The material consists of 18 283 carabids of 51 species, 48 769 spiders of 212 species, and 126 718 worker ants of 23 species. Analyses of abundance variation and species complementarity among successional stages revealed that in all the three taxa species occurring in the mature forest were prevalent in the younger successional stages as well, constituting more than half of the catch in any age class. A great majority of carabid and spider species were widely distributed across the forest age classes, whereas ants include a higher proportion of species with a narrower amplitude across the succession gradient. Comparisons of similarity between samples at increasing distance from one another on the local scale within forest stands (a few tens of meters to a few hundreds of meters) showed a quite consistent pattern in carabids and spiders: there was more variation between sampling sites in young successional forests than in the mature forest. Furthermore, only in the mature forest a slight, albeit statistically not significant, negative relationship between similarity of samples and distance between sampling sites was detected. In carabids and spiders, comparisons between samples located at a distance of 10–15 m from each other showed considerable heterogeneity, the mean percentage similarity being c. 0.6 (in ants c. 0.8). On the regional scale, systematic variation between young and mature forest stands is a major element increasing the total diversity (species turnover c. 50% in carabids and spiders; compositional similarity c. 0.3–0.4 in carabids, 0.2–0.3 in spiders), but variation within forest stands on a spatial scale of 10–15 meters is another important component in the total heterogeneity. The results suggest that regional abundance variation is a primary factor influencing the composition of local assemblages; a set of hypotheses elaborating this conclusion is formulated. The result implies that maintenance of habitat heterogeneity on a small scale (10–15 m) is needed to preserve biodiversity in managed forests.