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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]

Abstract Plants with crassulacean acid metabolism ( CAM) are increasing their abundance in drylands worldwide. The drivers and mechanisms underlying the increased dominance of CAM plants and CAM expression (i.e., nocturnal carboxylation) in facultative CAM plants, however, remain poorly understood. We investigated how nutrient and water availability affected competition between Mesembryanthemum crystallinum (a model facultative CAM species) and the invasive C3 grass Bromus mollis that co-occur in California's coastal grasslands. Specifically we investigated the extent to which water stress, nutrients, and competition affect nocturnal carboxylation in M. crystallinum. High nutrient and low water conditions favored M. crystallinum over B. mollis, in contrast to high water conditions. While low water conditions induced nocturnal carboxylation in 9-week-old individuals of M. crystallinum, in these low water treatments, a 66% reduction in nutrient applied over the entire experiment did not further enhance nocturnal carboxylation. In high water conditions M. crystallinum both alone and in association with B. mollis did not perform nocturnal carboxylation, regardless of the nutrient levels. Thus, nocturnal carboxylation in M. crystallinum was restricted by strong competition with B. mollis in high water conditions. This study provides empirical evidence of the competitive advantage of facultative CAM plants over grasses in drought conditions and of the restricted ability of M. crystallinum to use their photosynthetic plasticity (i.e., ability to switch to CAM behavior) to compete with grasses in well-watered conditions. We suggest that a high drought tolerance could explain the increased dominance of facultative CAM plants in a future environment with increased drought and nitrogen deposition, while the potential of facultative CAM plants such as M. crystallinum to expand to wet environments is expected to be limited. [ABSTRACT FROM AUTHOR]

The coexistence of woody plants and grasses in savannas is determined by a complex set of interacting factors that determine access to resources and demographic dynamics, under the control of external drivers and vegetation feedbacks with the physical environment. Existing theories explain coexistence mainly as an effect of competitive relations and/or disturbances. However, theoretical studies on the way facilitative interactions resulting from hydraulic lift affect tree-grass coexistence and the range of environmental conditions in which savannas are stable are still lacking., We investigated the role of hydraulic lift in the stability of tree-grass coexistence in savannas. To that end, we developed a new mechanistic model that accounts for both competition for soil water in the shallow soil and fire-induced disturbance., We found that hydraulic lift favors grasses, which scavenge the water lifted by woody plants. Thus, hydraulic lift expands (at the expenses of woodlands) the range of environmental conditions in which savannas are stable., These results indicate that hydraulic lift can be an important mechanism responsible for the coexistence of woody plants and grasses in savannas. Grass facilitation by trees through the process of hydraulic lift could allow savannas to persist stably in mesic regions that would otherwise exhibit a forest cover. [ABSTRACT FROM AUTHOR]