The strength of competition among individual trees and the biomass-density trajectories of the cohort

We simulated the self-thinning of Rhizophora mangle mangrove forests with the spatially explicit simulation model KiWi. This model is an application of the field-of-neighbourhood (FON) approach, which describes an individual tree by a competition function defined on the zone of influence (ZOI) around the stem. The FON causes growth depression of the trees involved. Sustained growth depression results in tree death. We propose a subdivision of the biomass density trajectories (bdt), obtained during the thinning process, into four segments related to characteristic shapes of the stem diameter distribution of the cohort. A positive skewness of the stem diameter distribution, indicating that the majority of the individuals are small and hindered in growth, is necessary for the occurrence of a linear segment within the bdt, the so-called 'self-thinning line'. This segment is the third bdt segment according to our classification. It is reached when the positive skewness of the stem diameter distribution is maximal and ends when the skewness reaches its second zero transition. The thinning line is therefore linked to the homogenisation process, which forces the symmetry of the stem distribution. We show that the ongoing search for a universal slope for the linear segment of the biomass-density trajectory ( bdt) cannot succeed, since it is species-specific and may also be site-dependent. The slope increases with increasing competition strength of the individuals. Nevertheless, the lower limit of the slope is pre-defined by geometrical constraints and modified by the actual strength of the neighbourhood competition. Although the simulations were all carried out with growth parameters of the mangrove species Rhizophora mangle, our results should be qualitatively valid and form a plausible theoretical framework to account for different biomass-density trajectories.