BIDIRECTIONAL FACILITATION AND INTERFERENCE BETWEEN SHRUBS AND ANNUALS IN THE MOJAVE DESERT

Abstract

Composition and structure of plant communities can be strongly influenced by plant interactions. Interactions among plants commonly comprise positive and negative effects operating simultaneously and bidirectionally. Thus, a thorough understanding of plant interactions requires experimental separation and quantitative assessment of the bidirectional positive and negative effects that add up to the net effects of plant interactions. Using the close spatial association of annual plants with a desert shrub (Ambrosia dumosa) in a sandy area of the Mojave Desert of California as a test system, we separated and quantified negative and positive effects of annuals on shrubs and of shrubs on annuals. We achieved the separation of negative and positive effects with an experimental design that included reciprocal removals of neighbors and simulations of physical effects of neighbors using artificial structures. All experimental manipulations were conducted on space originally occupied by Ambrosia shrubs to focus on immediate effects of neighbors on water availability rather than on long-term microenvironmental effects (e.g., nutrient accumulation). We quantified positive effects by calculating the difference between performance parameters of neighbors growing with artificial structures (thatch to mimic the physical effects of the presence of annuals and artificial canopies to mimic the physical effects of shrubs) and those of neighbors growing alone (removals). We estimated negative effects by calculating the difference between plant performance on control plots (shrubs and annuals growing together) and performances of plants growing with the artificial structures. Annuals had simultaneously strong negative and weak positive effects on shrub water status, growth, and reproductive output. Annuals also had an impact on the sex expression of shrubs by inducing shifts toward a higher male proportion in inflorescences of monoecious Ambrosia. In contrast, we found strong positive and weak or no negative effects of the shrubs on survival, biomass production, and seed production of the entire annual community and of selected annual species (the abundant native Chaenactis fremontii and the two dominant introduced annual species Bromus madritensis ssp. rubens and Schismus barbatus). Overall, in net effect, the interaction between shrubs and annuals can be described as facilitation or positive net effects of shrubs on annuals, and interference or negative net effects of annuals on shrubs. However, during the growing season, the ratios between positive and negative effects shifted. Annual plants benefited from the presence of shrubs to the greatest extent early in the growing season, and initial negative effects of annuals on shrubs declined as annuals senesced later in the season. Results of this study support the view that an experimental resolution of bidirectional positive and negative effects is necessary to achieve an accurate, mechanistic understanding of species interactions.