Highlights: We studied the tree community spatial structure in three 1.2-ha plots representing naturally developed northern boreal forests of varying ages; Spatial structure showed little differences between the mid-successional, late-successional and old-growth stands; The occurrence of Picea abies relative to Betula spp. indicated a mosaic-like spatial assembly; Mosaics are likely maintained by species-specific replacement, not reciprocal replacement as thought earlier.
Development of species composition during succession is well studied in natural boreal forests, but empirical assessments of how within-stand spatial structure develops in late-successional stages are few. Here, we quantified spatial patterns in three unmanaged stands consisting of Picea abies (L.) Karst. and Betula pubescens Ehrh. and Betula pendula Roth (hereafter Betula spp.) in northern boreal Fennoscandia. We conducted a comprehensive analysis of small-scale spatial point patterns in three fully mapped 1.2-ha sample plots, representing different forest developmental stages: mid-successional, late-successional and old-growth forest. We used several variants of Ripley’s K-function to analyze the spatial point patterns along the successional gradient. Univariate analyses showed that mature trees of both species were either randomly distributed or clumped. P. abies saplings were clumped, and Betula spp. saplings occurred in a random or clumped manner. In the bivariate analyses, saplings were more likely to be found in the surroundings of mature trees of the same species, but occurred independent of the individuals of other tree species. Mature trees showed interspecific repulsion. Only modest differences occurred in the univariate patterns between the three successional stages, but in the bivariate analyses the most evident patterns, i.e. intraspecific attraction and interspecific repulsion, were stronger in the older successional stages. Overall, the studied stands appear structured as species-specific mosaics. These mosaics, along with mixed species composition, seem to be maintained by species self-replacement, which contrasts with findings from earlier studies.