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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.

Woody plants may facilitate the establishment of seedlings with crassulacean acid metabolism (CAM) by ameliorating the abiotic environment through an increase in soil water availability. Because of the low transpiration rate in shallow-rooted CAM plants and the consequently high soil water contents in the shallow soil, deep-rooted trees in tree–CAM associations could perform hydraulic descent transporting water from wetter shallow soil to drier deep soil in arid environments. It remains unclear, however, whether a high rate of hydraulic descent can turn the facilitation of CAM plants by woody plants into competition. In this study, we develop a mechanistic model to investigate the facilitation of shallow-rooted CAM plants by deep-rooted woody plants in the access to soil water resources along a rainfall gradient. The model results show that in the case of low-to-moderate root overlap woody plants could facilitate CAM plants in access to soil water; this effect is mainly induced by the reduction in evaporation from the soil surface due to shading. Both shading and hydraulic descent decrease (or hydraulic lift increases) along a rainfall gradient, thereby favoring facilitation. Investment in deep roots by woody plants is usually thought to increase niche differentiation with shallow-rooted plants, thereby reducing the competition and promoting species coexistence. This study indicates that deep root development could also favor competition through the mechanism of hydraulic descent, thereby changing our understanding of the role of root depth in niche differentiation between shallow and deep-rooted plants.

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]

Plants with crassulacean acid metabolism (CAM) are increasing in distribution and abundance in drylands worldwide, but the underlying drivers remain unknown. We investigate the impacts of extreme drought and CO2 enrichment on the competitive relationships between seedlings of Cylindropuntia imbricata (CAM species) and Bouteloua eriopoda (C4 grass), which coexist in semiarid ecosystems across the Southwestern United States. Our experiments under altered water and CO2 water conditions show that C. imbricata positively responded to CO2 enrichment under extreme drought conditions, while B. eriopoda declined from drought stress and did not recover after the drought ended. Conversely, in well‐watered conditions B. eriopoda had a strong competitive advantage on C. imbricata such that the photosynthetic rate and biomass (per individual) of C. imbricata grown with B. eriopoda were lower relative to when growing alone. A meta‐analysis examining multiple plant families across global drylands shows a positive response of CAM photosynthesis and productivity to CO2 enrichment. Collectively, our results suggest that under drought and elevated CO2 concentrations, projected with climate change, the competitive advantage of plant functional groups may shift and the dominance of CAM plants may increase in semiarid ecosystems. [ABSTRACT FROM AUTHOR]