A neighborhood analysis of canopy tree competition: effects of shading versus crowding

We have developed extensions of traditional distance-dependent, spatial competition analyses that estimate the magnitude of the competitive effects of neighboring trees on target tree growth as a function of the species, size, and distance to neighboring trees. Our analyses also estimate inter- and intra-specific competition coefficients and explicitly partition the competitive effects of neighbors into the effects of shading versus crowding. We tested the method using data from forests of northern, interior British Columbia dominated by western hemlock (Tsuga heterophylla (Raf.) Sarg.) and western redcedar (Thuja plicata Donn ex D. Don). For both species, the most parsimonious regression models included terms for the effects of tree size, crowding, and shading and separate competitive effects of four different groups of competing species. The models explained 33%–59% of the variation in radial growth of the two species. For both species, growth declined more steeply as a function of crowding than shading. There was striking asymmetry in the strength of interspecific competition between hemlock and redcedar, with crowding by hemlock having a strong per capita effect on redcedar, while crowding by redcedar had relatively little effect on the radial growth of hemlock.