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Precipitation variability has been predicted to increase in a global warmer climate, and is expected to greatly affect plant growth, interspecies interactions, plant community composition, and other ecosystem processes. Although previous studies have investigated the effect of intra-annual rainfall variability on plant growth and ecosystem dynamics, the impacts of interannual rainfall variability remain understudied. This paper uses satellite data and develops a new mechanistic model to investigate the response of tree-grass composition to increasing interannual rainfall variability in arid to sub-humid ecosystems along the Kalahari Transect in Southern Africa. Both satellite data and model results show that increasing interannual rainfall fluctuations favor deep-rooted trees over shallow-rooted grasses in drier environments (that is, mean annual rainfall, MAP < 900-1000 mm) but favor grasses over trees in wetter environments (that is, MAP > 900-1000 mm). Trees have a competitive advantage over grasses in dry environments because their generally deeper root systems allow them to have exclusive access to the increased deep soil water resources expected to occur in wet years as a result of the stronger interannual rainfall fluctuations. In relatively wet environments, grasses are favored because of their high growth rate that allows them to take advantage of the window of opportunity existing in years with above average precipitation and thus increase fire-induced tree mortality. Thus, under increasing interannual rainfall fluctuations both direct effects on soil water availability and indirect effects mediated by tree-grass interactions and fire dynamics are expected to play an important role in determining changes in plant community composition. [ABSTRACT FROM AUTHOR]

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 CO 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. [ABSTRACT FROM AUTHOR]

Savannas are characterized by the coexistence of trees and flammable grasses. Yet, tree-grass coexistence has been labeled as paradoxical-how do these two functional groups coexist over such an extensive area, despite being generally predisposed to excluding each other? For instance, many trees develop dense canopies that limit grass growth, and many grasses facilitate frequent/intense fires, increasing tree mortality. This study revisits tree-grass coexistence with a model of hierarchical competition between pyrogenic grasses, 'forest trees' adapted to closed-canopy competition, and 'savanna trees' that are inferior competitors in closed-canopy communities, but more resistant to fire. The assumptions of this model are supported by empirical observations, including a systematic review of savanna and forest tree community composition reported here. In general, the model simulations show that when savanna trees exert weaker competitive effects on grasses, a self-reinforcing grass community is maintained, which limits forest tree expansion while still allowing savanna trees to persist (albeit as a subdominant to grasses). When savanna trees exert strong competitive effects on grasses, savanna trees cover increases initially, but as grasses decline their inhibitory effect on forest trees weakens, allowing forest trees to expand and exclude grasses and savanna trees. Rather than paradoxical, these results suggest that having weaker competitive effects on grasses may be advantageous for savanna trees, leading to greater long-term abundance and stability. We label this the 'enemy of my enemy hypothesis,' which might apply to species coexistence in communities defined by hierarchical competition or with species capable of generating strong ecological feedbacks. [ABSTRACT FROM AUTHOR]