Neighbourhood-based evidence of tree diversity promotion by beech in an old-growth deciduous-coniferous mixed forest (Eastern Carpathians)

Neighbourhood models are useful tools for understanding the role of positive and negative interactions in maintaining the tree species diversity in mixed forests. Under such a presumption, we aimed at testing several hypotheses concerning the mechanisms of autogenic species coexistence in an old-growth, beech-fir-spruce stand, which is part of the Slatioara forest reserve (Eastern Carpathians). Univariate/bivariate spatial point pattern analyses, the individual tree species-area relationship, the species mingling analysis and generalised linear mixed models of neighbour interference were applied on data concerning the position and allometry of all saplings and trees occurring within a 0.24 ha plot. The monospecific distribution of either beech or spruce saplings did not support the spatial segregation hypothesis. There was no evidence of conspecific negative distance dependence, as no spatial segregation was detected between the saplings and trees of any species. Within 4 m- neighbourhood, the beech saplings appeared as diversity accumulators, which might be indicative of indirect facilitation (e.g., herd protection hypothesis). At tree stage, none of the three species showed either accumulator or repeller patterns in their neighbourhood with respect to sapling species richness. Signals of positive and negative interspecific association were found in tree-sized beech (at scales of 10 to 20 m) and spruce (at scales of 4 to 17 m), respectively. The former, highly interspersed pattern is in accordance with the hypothesis of positive complementary effects, whereas the latter, poorly intermingled pattern is probably linked to the unexpected, positive neighbouring effect of spruce trees on the stem growth of their conspecific saplings. Such self-favouring process might be due to a facilitative below-ground mechanism. Conversely, the beech saplings were suppressed through interference from the neighbouring conspecific trees. The beech appears to be the key promoter of tree species coexistence in the study forest stand, in contrast to the low interspersion of spruce in the overstorey leading to lower local tree diversity.