• alpine meadow;
  • density-dependent mortality;
  • Elymus nutans;
  • facilitation;
  • individual-based model;
  • plant–plant interactions;
  • size inequality


1. Large variation in the size of individuals is a ubiquitous feature of natural plant populations. While the role of competition in generating this variation has been studied extensively, the potential effects of positive interactions among plants, which are common in high-stress environments, have not been investigated.

2. Using an individual-based ‘zone-of-influence’ model, we investigate the effects of competition, abiotic stress and facilitation on size inequality in plant monocultures. In the model, stress reduces the growth rate of plants, and facilitation ameliorates the effects of stress. Both facilitation and competition occur in overlapping zones of influence. We tested some of the model’s predictions with a field experiment using the clonal grass Elymus nutans in an alpine meadow.

3. Facilitation increased the size inequality of model populations when there was no density-dependent mortality. This effect decreased with density as competition overwhelmed facilitation. The lowest size inequality was found at intermediate densities both with the model and in the field.

4. When density-dependent mortality was included in the model, stress delayed its onset and reduced its rate by reducing growth rates, so the number of survivors at any point in time was higher under harsh than under more benign conditions. Facilitation increased size inequality during self-thinning.

5. Synthesis. Our results demonstrate that facilitation interacts with abiotic stress and competition to influence the degree of size inequality in plant populations. Facilitation increased size inequality at low to intermediate densities and during self-thinning.