Kailiang Yu

Hydraulic lift as a determinant of tree–grass coexistence on savannas

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.

An ecohydrological framework for grass displacement by woody plants in savannas

During the past several decades, woody plants have been encroaching into grasslands around the world. This transition in plant dominance is often explained as a state shift in bistable ecosystem dynamics induced by fire-vegetation feedbacks. These feedbacks occur when woody plants are able to displace grasses because of their better access to soil water and light. On the other hand, grasses can displace woody plants because of their ability to increase fire frequency and of the higher susceptibility of woody plants to fire-induced mortality. In this study, we present an ecohydrological framework to investigate the displacement of grasses by woody plants. Considering the effect of lateral root spread and of soil water and light limitations, we found that woody plant encroachment can substantially suppress grass production even without the presence of grazers. Bistable dynamics emerge as a result of the grass-fire feedback for a wide range of rainfall conditions, fire susceptibility, and woody plant growth rates.

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.

Dust-rainfall feedback in West African Sahel

Drought persistence in West African Sahel has often been explained as an effect of positive vegetation-atmosphere feedback associated with surface albedo or the partitioning of solar radiation into sensible and latent heat fluxes. An often overlooked aspect of land-atmosphere coupling results from vegetation controls on dust emissions and the ability of mineral aerosols to suppress precipitation. Here we first consider the case of local (endogenous) dynamics within the Sahel, whereby enhanced dust emissions resulting from a decrease in vegetation partly suppress precipitation, thereby further reducing vegetation cover. We then account for teleconnections between Sahel precipitation and exogenous (i.e., Saharan) dust emissions due to an increase in Saharan wind speed in years of above average Sahel precipitation. We find that in both cases vegetation-climate dynamics may have two stable states, one with low precipitation and high concentration of atmospheric dust and the other with high precipitation and lower levels of atmospheric dust.

Effects of competition on induction of crassulacean acid metabolism in a facultative CAM plant

Abiotic drivers of environmental stress have been found to induce CAM expression (nocturnal carboxylation) in facultative CAM species such as Mesembryanthemum crystallinum. The role played by biotic factors such as competition with non-CAM species in affecting CAM expression, however, remains largely understudied. This research investigated the effects of salt and water conditions on the competition between M. crystallinum and the C3 grass Bromus mollis with which it is found to coexist in California’s coastal grasslands. We also investigated the extent to which CAM expression in M. crystallinum was affected by the intensity of the competition with B. mollis. We found that M. crystallinum had a competitive advantage over B. mollis in drought and saline conditions, while B. mollis exerted strong competitive effects on M. crystallinum in access to light and soil nutrients in high water conditions. This strong competitive effect even outweighed the favorable effects of salt or water additions in increasing the biomass and productivity of M. crystallinum in mixture. Regardless of salt conditions, M. crystallinum did not switch to CAM photosynthesis in response to this strong competitive effect from B. mollis. Disturbance (i.e., grass cutting) reduced the competitive pressure by B. mollis and allowed for CAM expression in M. crystallinum when it was grown mixed with B. mollis. We suggest that moderate competition with other functional groups can enhance CAM expression in M. crystallinum, thereby affecting its plasticity and ability to cope with biological stress.

The effect of nitrogen availability and water conditions on competition between a facultative CAM plant and an invasive grass

Abstract 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 Californias 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.

Effects of fertilization, burning, and grazing on plant community in the long-term fenced grasslands

Fencing is the common management practice to restore degraded grasslands. However, long-term fencing decreases grassland productivity and species diversity. The study was therefore conducted as a three-year (2011-2013) experiment with a randomized complete block in a grassland fenced for 20 years in the Loess Plateau of China, and the effects of fertilization, burning and grazing on aboveground biomass, species and functional group composition, species and some functional group diversity were analysed. Our results showed that the functional group of perennial bunchgrasses dominated the grassland regardless of management practices. However, burning altered species composition (i.e. the unpalatable species, Artemisia sacrorum) more significantly than fertilization or grazing, and surprisingly, nearly quadrupled the functional group of shrubs and semi-shrubs. Fertilization had a positive effect on the aboveground biomass (44.0%), while clearly reducing species diversity (21.9%). Grazing decreased aboveground biomass, but increased species diversity by 15.9%. This study indicated that fertilization influenced plant community through its impact on aboveground biomass, while burning changed plant community by altering dominant species. Thus, it was concluded that fertilizer could further improve community biomass while burning reduced the edibility of grass. Grazing could be carried out to enhance the biodiversity in the long-term fenced grasslands.

The Enemy of My Enemy Hypothesis: Why Coexisting with Grasses May Be an Adaptive Strategy for Savanna Trees

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.

Response of a facultative CAM plant and its competitive relationship with a grass to changes in rainfall regime

Background and aimsWe investigated the response of a model facultative CAM plant (Mesembryanthemum crystallinum) and its competition with a C3 grass (Bromus mollis) to changes in rainfall regime.MethodsSeedlings of M. crystallinum and B. mollis in both monoculture and mixtures growing in shallow and deep pots were subjected to three levels of intra-seasonal rainfall variability and rainfall seasonality in both high water and low water conditions. Response of plants were evaluated by nocturnal carboxylation and biomass.ResultsA high rate of water drainage beneath root zones in coarse soil led to a negative response of M. crystallinum and B. mollis in monoculture under increased intra-seasonal rainfall variability. Seasonal rainfall shifts to later dates during the growing season generally favored the growth of M. crystallinum and B. mollis in monoculture, with the exception of high water stress conditions whereby drought-intolerant species B. mollis was disfavored. Rainfall seasonality but not intra-seasonal rainfall variability affected nocturnal carboxylation by M. crystallinum in monoculture.ConclusionsWe suggest that soil texture, root depth, and rainfall gradient are important mediators of plant growth under increased intra-seasonal rainfall variability. Drought severity and the ability of a plant to tolerate drought and can greatly affect its response to the seasonal timing of rainfall. Nocturnal carboxylation by M. crystallinum in response to rainfall variability depends on the timescale.