Species-specific spatial structure, species coexistence and mortality pattern in natural, uneven-aged Scots pine ( Pinus sylvestris L.)-dominated forest

Abstract

Complexity of uneven-aged forests results from the heterogeneity of their structure reflected among others by the spatial pattern of their components. Forest structure is usually modified by various processes operating at different scales and time. Structure and processes are not independent, and both are important drivers of forest dynamics. The impact of natural processes on forest structure manifested in the specific spatial pattern of trees can be quantified by point pattern analysis applied to long-term repeatedly measured stem-mapped plots. Such studies are relatively scarce in the literature although they provide better insight into the mechanisms affecting forest dynamics. Our study is focused on the spatiotemporal analysis of the structure of mixed uneven-aged Scots pine-dominated forest located at the Kampinoski National Park (Poland). Univariate analysis showed that the initial pattern of all live trees was initially random and it shifted toward more uniform with forest aging. Spatial patterns of individual tree species varied from that stated for all forest community. We observed changes in spatial pattern of Scots pine and common oak from random toward more clumped (pine) or uniform (oak) pattern. In case of black alder and common birch, the initial aggregated pattern was maintained over the examined 14-year period of the forest succession. Bivariate analysis showed that the most common interspecific association between pairs of tree species was spatial segregation (pine vs. alder, alder vs. birch and oak vs. birch) followed by spatial independence (pine vs. oak and oak vs. alder). The positive association was stated only for pine and birch and only for certain spatial scales (> 5 m). Simultaneously, at small distances they showed reciprocal repulsion. Changes in spatial relationships between tree species were negligible over 14-year period of forest succession. Our results confirmed the density-dependent mortality process in the uneven-aged Scots pine-dominated forest over 14-year period of forest development. Our study showed that spatial interactions between individuals along with species-specific ecological requirements should be incorporated into realistic models of forest development, helping to manage the forest ecosystems toward their greater structural complexity.

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