Interacting water, nutrients, and shrub age control steppe grass-on-shrub competition: Implications for restoration

Abstract

Understanding the factors that control grass–shrub interactions and coexistence is critical to the design of dryland management and restoration strategies. Using Stipa tenacissima tussocks as adult neighbor and Olea europaea var. sylvestris as target woody species, we performed a 5-year-long, large-scale manipulative experiment to investigate the independent and combined effects of water and nutrient availability, and the modulating effects of O. europaea ontogeny on the net outcome of grass–shrub interaction. At two contrasting dryland sites, we conducted experimental plantings of Olea seedlings on two microsites: contiguous to a Stipa tussock (Stipa microsite) and on the intertussock bare-soil areas (Open microsite), and manipulated resource availability by combining nutrient and water addition. The experiment followed a full factorial design, with all treatment combinations applied at each site. We monitored survival, height, and stem basal diameter of Olea individuals over a study period of 5 years. All across the wide range of conditions considered, negative effects of Stipa on Olea largely prevailed during the first 2 years after the plantation. We found that competition was stronger in the drier steppe, where extra inputs of water favored neutral interactions. Conversely, nutrient addition increased competition strength, particularly when combined with water inputs, pointing to the contrasting role of different resources and the importance of interactions between them in the control of plant–plant interaction outcome. The competition effects of Stipa faded with time and gradually shifted to neutral interaction as Olea individuals aged. Our results are particularly relevant to guide restoration efforts in formerly wooded xeric grasslands and rangelands.