Structural heterogeneity and spatial autocorrelation in a natural mature Pinus sylvestris dominated forest

Abstract

The tree species composition, vertical stratification and patterns of spatial autocorrelation at the tree and quadrate (25 × 25 m) scales were studied in a natural mature PinuS sylvestris dominated forest in eastern Finland. For the analyses we mapped the locations and dimensions of trees taller than 10 m in a 9 ha (300 × 300 m) area, and within this area we mapped all trees taller than 0.3 m on a core plot of 4 ha (200 × 200 m). The overall tree size distribution was bimodal. the dominant layer and the understory forming the peak frequencies. Pinus sylvestris dominated the main canopy, together with scattered Betula pendula and Picea abies. Alnus incana, Populus tremula, Salix caprea, Sorbus aucuparia and Juniperus communis occurred only in the under‐ and middlestories. Autocorrelation analysis (semivarianee) of tree size variation revealed spatial patterns, which were strongly dependent on the size of trees included in the analysis. When all living trees, including the understory regeneration, were taken into account, the autocorrelation pattern ranged up to 35 m inter‐tree distances, reflecting the spatial scale of understory regeneration patches. Competitive interaction among middle‐ and upperstory trees (height>10 m) had contrasting effects on autocorrelation pattern depending on spatial scale. At the fine scale, dominant trees suppressed their smaller close neighbors (asymmetric competition), which was shown as increased tree size variation at small inter‐tree distances (<2 m). At slightly larger inter‐tree distances, specifically among large trees of similar size, competition was more symmetrical, which resulted in decreased tree size variation at these inter‐tree distances (3–4 m). This effect was seen most clearly in the dominant trees, there being a clear autocorrelation pattern in tree size up to inter‐tree distances of ∼4 m. At the quadrate scale (25 × 25 m) the analysis revealed high local variation in structural characteristics such as tree height diversity (THD), tree species diversity (H) and autocorrelation of tree height. The analysis suggests that naturally developed P. sylvestris forests exhibit complex small‐scale patterns of structural heterogeneity and spatial autocorrelation in tree size. These patterns may be important for stand‐scale habitat diversity and can have aggregated effects on ecosystem dynamics at larger spatial scales though their influence on the spread of disturbance and regeneration after disturbance.