• competition coefficients;
  • competitive interactions;
  • complex forests;
  • complex stands;
  • crowding;
  • interspecific competition;
  • intraspecific competition;
  • neighbourhood models;
  • shading;
  • size-dependency of growth


  • 1
    The neutral theory debate has highlighted the scarcity of robust empirical estimates of the magnitude of competitive effects and responses within guilds of co-occurring tree species. Our analysis quantifies the relative magnitude of all possible pairwise competitive interactions within a guild of nine co-occurring tree species in temperate forests of northern, interior British Columbia, and explicitly partitions the competitive effects of neighbours into the effects of shading versus the residual effects of ‘crowding’, assumed to reflect below-ground competition.
  • 2
    Models that treated neighbours as equivalent in their competitive effects were the most parsimonious for the five species with the smallest sample sizes. For the remaining species (samples sizes of > 150 individuals), the best models estimated separate competition coefficients for all nine species of neighbours. We take this as evidence that species do indeed differ in their competitive effects, but that there can be a minimum sample size required to discriminate between them.
  • 3
    There was a strong size-dependency in potential growth. Six species showed an optimal growth at a size between 5 and 20 cm diameter. Potential growth declined moderately to strongly as diameter increased. Sensitivity to crowding varied as a function of tree size for five of the nine species; however, this response was not consistent by tree species.
  • 4
    The magnitude of reduction in growth due to crowding was greater on average than the reduction in growth due to shading, except for the two least shade tolerant conifers. Sensitivity to shading among the conifer species was correlated with their shade tolerance.
  • 5
    The per capita effects of crowding by different species of neighbours varied widely. A large number of the estimated pairwise per capita competition coefficients were very low. The relative magnitude of the strength of intra- versus interspecific competition also varied widely among the tree species.
  • 6
    Synthesis. Model selection techniques effectively separated above- and below-ground competition in complex forests, and allowed us to assess differences among species in competitive effects and responses. While below-ground effects were strong, they were due to proximity of neighbours from a very specific (and small) subset of strong competitors within the guild. Response to crowding varied with tree size but the nature of the relationship varied widely among the species.