Direct and Indirect Facilitation of Plants with Crassulacean Acid Metabolism (CAM)

Plants with crassulacean acid metabolism (CAM) are increasing their cover in many dryland regions around the world. Their increased dominance has been related to climate warming and atmospheric CO2 fertilization, while the effects of interspecies interactions and the role of CAM plant facilitation by trees and grasses remain poorly understood. Woody plants are known for their ability to directly facilitate CAM plants through amelioration of the abiotic environment. Mechanisms of indirect facilitation of trees on CAM plants in tree-grass-CAM associations, however, have received less attention. It is also unclear whether grasses might facilitate CAM plants in mixed tree-grass-CAM communities. For instance, the inclusion of grasses in tree-CAM associations could enhance hydraulic lift and facilitate CAM plants in their access to shallow soil moisture at the expenses of deep-rooted trees. If this effect outweighs the competitive effects of grasses on CAM plants, grasses could overall facilitate CAM plants through hydraulic lift. Here we develop a process-based ecohydrological model to investigate the direct and indirect facilitation in tree-CAM-grass associations; the model quantifies transpiration of CAM plants when isolated as well as in associations with trees and/or grasses. It is found that woody plants having a high root overlap with CAM plants indirectly facilitate CAM plants by significantly reducing grass transpiration in shaded conditions. For situations of a low-to-moderate root overlap, facilitation may occur both directly and indirectly. Conversely, grasses are unable to indirectly facilitate CAM plants through the mechanism of hydraulic lift because the competitive effects of grasses on CAM plants outweigh the facilitation induced by hydraulic lift.