Temporal environmental variation tips the balance between facilitation and interference in desert plants

Recently, numerous studies have pointed to the importance of positive interactions in natural communities. There is now a broad consensus that the balance between negative and positive interactions should shift along environmental gradients, with competition prevailing under environmentally benign conditions and positive interactions dominating under harsh conditions. A commonly cited example of the importance of facilitation in harsh environments is the preference of desert annual plants for the areas under the canopy of shrubs. The recognition of apparently positive effects of desert shrubs on annuals, however, has been mostly based on density measurements, while fitness parameters of the understory plants have been ignored. Also, the temporal consistency of such effects has not been previously tested. Based on conceptual ideas about the balance between interference and facilitation, we predicted that positive effects of the shrubs on the understory should dominate in dry years, while in favorable years, negative effects would be stronger. We tested our hypothesis by measuring the direction and magnitude of the shrub effect on demographic responses of four desert annual plant species during four consecutive seasons of differing rainfall. The results contradicted our initial hypothesis. Depending on the species, the effect of the shrubs shifted from either negative to neutral or from neutral to positive with increasing annual rainfall. However, this trend was stronger for the effect of shrubs on plant reproductive success than on their densities. Our data highlight the importance of measuring fitness parameters in studies of plant-plant interactions. We suggest that the negative effects of shrubs on plant fitness were due to rainfall interception, while positive effects were related to increased nutrient availability beneath shrubs. However, the mechanisms by which the shrubs and annuals interact can only be resolved using an experimental approach. Our results contradict previous hypotheses about the relative importance of positive and negative interactions along environmental gradients. A simple conceptual model summarizing the proposed role of rainfall in determining the direction of shrub effects on their understory annuals is presented.