Spatial patterns in different forest development stages of an intact old-growth Oriental beech forest in the Caspian region of Iran

Spatial patterns of trees are important structural characteristics that can provide insights into forest dynamics and may be related to the development stages of forests. The spatial patterns of three development stages (i.e., initial, optimal, and decay) of an intact old-growth beech (Fagus orientalis Lipsky) forest in the Caspian region of Iran were quantified within one-hectare permanent plots. All trees with a diameter at breast height of more than 7.5 cm were measured, stem-mapped, and then assigned to one of four diameter size classes (small, medium, large, and extra-large timbers). Spatial patterns of all trees and spatial associations among tree size classes were analyzed using Ripley's K-function, and the spatial dependence of tree diameters was analyzed using variograms. Results showed that trees in the small size class exhibited an aggregated distribution in every development stage, which matched the overall spatial pattern of all trees in each stage. However, the degree of aggregation of all trees as well as association patterns among the size classes differed among the three development stages. Further, the average spatial dependence of tree diameters was 24, 14, and 19 m in the initial, optimal, and decay stages, respectively. Differences in spatial patterns among the development stages in this beech old-growth forest are consistent with the gap-dynamics paradigm and likely reflect different canopy disturbance events coupled with associated regeneration, release, and competition processes (e.g., shade-tolerance characteristics, seed dispersal limitation, and intraspecific competition).