Herbivore–vegetation feedbacks can expand the range of savanna persistence: insights from a simple theoretical model

Theoretical models of tree–grass coexistence in savannas have focused primarily on the role of resource availability and fire. It is clear that herbivores heavily impact vegetation structure in many savannas, but their role in driving tree–grass coexistence and the stability of the savanna state has received less attention. Theoretical models of tree–grass dynamics tend to treat herbivory as a constant rather than a dynamic variable, yet herbivores respond dynamically to changes in vegetation structure in addition to modifying it. In particular, many savannas host two distinct herbivore guilds, grazers and browsers, both of which have the potential to exert profound effects on tree/grass balance. For example, grazers may indirectly favor tree recruitment by suppressing the destructive effects of fire, and browsers may facilitate the expansion of grassland by reducing the competitive dominance of trees. We use a simple theoretical model to explore the role of grazer and browser dynamics on savanna vegetation structure and stability across fire and resource availability gradients. Our model suggests that herbivores may expand the range of conditions under which trees and grasses are able to stably coexist, as well as having positive reciprocal effects on their own niche spaces. In addition, we suggest that given reasonable assumptions, indirect mutualisms can arise in savannas between functional groups of herbivores because of the interplay of consumption and ecosystem feedbacks.     

Scale matters: fire–vegetation feedbacks are needed to explain tropical tree cover at the local scale

At a broad (regional to global) spatial scale, tropical vegetation is controlled by climate; at the local scale, it is believed to be determined by interactions between disturbance, vegetation and local conditions (soil and topography) through feedback processes. It has recently been suggested that strong fire–vegetation feedback processes may not be needed to explain tree‐cover patterns in tropical ecosystems and that climate–fire determinism is an alternative possibility. This conclusion was based on the fact that it is possible to reproduce observed patterns in tropical regions (e.g. a trimodal frequency distribution of tree cover) using a simple model that does not explicitly incorporate fire–vegetation feedback processes. We argue that these two mechanisms (feedbacks versus fire–climate control) operate at different spatial and temporal scales; it is not possible to evaluate the role of a process acting at fine scales (e.g. fire–vegetation feedbacks) using a model designed to reproduce regional‐scale pattern (scale mismatch). While the distributions of forest and savannas are partially determined by climate, many studies are providing evidence that the most parsimonious explanation for their environmental overlaps is the existence of feedback processes. Climate is unlikely to be an alternative to feedback processes; rather, climate and fire–vegetation feedbacks are complementary processes at different spatial and temporal scales.     

Coupling microscale vegetation–soil water and macroscale vegetation–precipitation feedbacks in semiarid ecosystems

At macroscale, land–atmosphere exchange of energy and water in semiarid zones such as the Sahel constitutes a strong positive feedback between vegetation density and precipitation. At microscale, however, additional positive feedbacks between hydrology and vegetation such as increase of infiltration due to increase of vegetation, have been reported and have a large impact on vegetation distribution and spatial pattern formation. If both macroscale and microscale positive feedbacks are present in the same region, it is reasonable to assume that these feedback mechanisms are connected. In this study, we develop and analyse a soil‐vegetation‐atmosphere model coupling large‐scale evapotranspiration–precipitation feedback with a model of microscale vegetation–hydrology feedback to study the integration of these nonlinearities at disparate scales. From our results, two important conclusions can be drawn: (1) it is important to account for spatially explicit vegetation dynamics at the microscale in climate models (the strength of the precipitation feedback increased up to 35% by accounting for these microscale dynamics); (2) studies on resilience of ecosystems to climate change should always be cast within a framework of possible large‐scale atmospheric feedback mechanism (substantial changes in vegetation resilience resulted from incorporating macroscale precipitation feedback). Analysis of full‐coupled modelling shows that both type of feedbacks markedly influence each other and that they should both be accounted for in climate change models.     

Patch dynamics integrate mechanisms for savanna tree–grass coexistence

Many mechanisms have been suggested to explain the coexistence of woody species and grasses in savannas. However, evidence from field studies and simulation models has been mixed. Patch dynamics is a potentially unifying mechanism explaining tree–grass coexistence and the natural occurrence of shrub encroachment in arid and semi-arid savannas. A patch-dynamic savanna consists of a spatial mosaic of patches. Each patch maintains a cyclical succession between dominance of woody species and grasses, and the succession of neighbouring patches is temporally asynchronous. Evidence from empirical field studies supports the patch dynamics view of savannas. As a basis for future tests of patch dynamics in savannas, several hypotheses are presented and one is exemplarily examined: at the patch scale, realistically parameterized simulation models have generated cyclical succession between woody and grass dominance. In semi-arid savannas, cyclical successions are driven by precipitation conditions that lead to mass recruitment of shrubs in favourable years and to simultaneous collapse of shrub cohorts in drought years. The spatiotemporal pattern of precipitation events determines the scale of the savanna vegetation mosaic in space and time. In a patch-dynamic savanna, shrub encroachment is a natural, transient phase corresponding to the shrub-dominated phase during the successional cycle. Hence, the most promising management strategy for encroached areas is a large-scale rotation system of rangelands. In conclusion, patch dynamics is a possible scale-explicit mechanism for the explanation of tree–grass coexistence in savannas that integrates most of the coexistence mechanisms proposed thus far for savannas.     

Spatial heterogeneity of plant–soil feedbacks increases per capita reproductive biomass of species at an establishment disadvantage

Plant–soil feedbacks have been widely implicated as a driver of plant community diversity, and the coexistence prediction generated by a negative plant–soil feedback can be tested using the mutual invasibility criterion: if two populations are able to invade one another, this result is consistent with stable coexistence. We previously showed that two co-occurring Rumex species exhibit negative pairwise plant–soil feedbacks, predicting that plant–soil feedbacks could lead to their coexistence. However, whether plants are able to reproduce when at an establishment disadvantage (“invasibility”), or what drivers in the soil might correlate with this pattern, are unknown. To address these questions, we created experimental plots with heterogeneous and homogeneous soils using field-collected conditioned soils from each of these Rumex species. We then allowed resident plants of each species to establish and added invader seeds of the congener to evaluate invasibility. Rumex congeners were mutually invasible, in that both species were able to establish and reproduce in the other’s resident population. Invaders of both species had twice as much reproduction in heterogeneous compared to homogeneous soils; thus the spatial arrangement of plant–soil feedbacks may influence coexistence. Soil mixing had a non-additive effect on the soil bacterial and fungal communities, soil moisture, and phosphorous availability, suggesting that disturbance could dramatically alter soil legacy effects. Because the spatial arrangement of soil patches has coexistence implications, plant–soil feedback studies should move beyond studies of mean effects of single patch types, to consider how the spatial arrangement of patches in the field influences plant communities.     

Abrupt shifts in African savanna tree cover along a climatic gradient

Aim To describe patterns of tree cover in savannas over a climatic gradient and a range of spatial scales and test if there are identifiable climate‐related mean structures, if tree cover always increases with water availability and if there is a continuous trend or a stepwise trend in tree cover. Location Central Tropical Africa. Methods We compared a new analysis of satellite tree cover data with botanical, phytogeographical and environmental data. Results Along the climatic transect, six vegetation structures were distinguished according to their average tree cover, which can co‐occur as mosaics. The resulting abrupt shifts in tree cover were not correlated to any shifts in either environmental variables or in tree species distributions. Main conclusions A strong contrast appears between fine‐scale variability in tree cover and coarse‐scale structural states that are stable over several degrees of latitude. While climate parameters and species pools display a continuous evolution along the climatic gradient, these stable structural states have discontinuous transitions, resulting in regions containing mosaics of alternative stable states. Soils appear to have little effect inside the climatic stable state domains but a strong action on the location of the transitions. This indicates that savannas are patch dynamics systems, prone to feedbacks stabilizing their coarse‐scale structure over wide ranges of environmental conditions.     

Tree–grass coexistence in savannas revisited – insights from an examination of assumptions and mechanisms invoked in existing models

Several explanations for the persistence of tree–grass mixtures in savannas have been advanced thus far. In general, these either concentrate on competition‐based mechanisms, where niche separation with respect to limiting resources such as water lead to tree–grass coexistence, or demographic mechanisms, where factors such as fire, herbivory and rainfall variability promote tree–grass persistence through their dissimilar effects on different life‐history stages of trees. Tests of these models have been largely site‐specific, and although different models find support in empirical data from some savanna sites, enough dissenting evidence exists from others to question their validity as general mechanisms of tree–grass coexistence. This lack of consensus on determinants of savanna structure and function arises because different models: (i) focus on different demographic stages of trees, (ii) focus on different limiting factors of tree establishment, and (iii) emphasize different subsets of the potential interactions between trees and grasses. Furthermore, models differ in terms of the most basic assumptions as to whether trees or grasses are the better competitors. We believe an integration of competition‐based and demographic approaches is required if a comprehensive model that explains both coexistence and the relative productivity of the tree and grass components across the diverse savannas of the world is to emerge. As a first step towards this end, we outline a conceptual framework that integrates existing approaches and applies them explicitly to different life‐history stage of trees.     

A mathematical model of plants as ecosystem engineers

Understanding the structure and dynamics of plant communities in water-limited systems often calls for the identification of ecosystem engineers--key species that modify the landscape, redistribute resources and facilitate the growth of other species. Shrubs are excellent examples; they self-organize to form patterns of mesic patches which provide habitats for herbaceous species. In this paper we present a mathematical model for studying ecosystem engineering by woody plant species in drylands. The model captures various feedbacks between biomass and water including water uptake by plants' roots and increased water infiltration at vegetation patches. Both the uptake and the infiltration feedbacks act as mechanisms for vegetation pattern formation, but have opposite effects on the water resource; the former depletes the soil-water content under a vegetation patch, whereas the latter acts to increase it. Varying the relative strength of the two feedbacks we find a trade-off between the engineering capacity of a plant species and its resilience to disturbances. We further identify two basic soil-water distributions associated with engineering at the single patch level, hump-shaped and ring-shaped, and discuss the niches they form for herbaceous species. Finally, we study how pattern transitions at the landscape level feedback to the single patch level by affecting engineering strength.     

Self‐thinning and Tree Competition in Savannas

This paper examines the feasibility of applying self‐thinning concepts to savannas and how competition with herbaceous vegetation may modify self‐thinning patterns among woody plants in these ecosystems. Competition among woody plants has seldom been invoked as a major explanation for the persistence of herbaceous vegetation in mixed tree–grass ecosystems. On the contrary, the primary resource‐based explanations for tree–grass coexistence are based on tree–grass competition (niche‐separation) that assumes that trees are inferior competitors unless deeper rooting depths provide them exclusive access to water. Alternative nonresource‐based hypotheses postulate that trees are the better competitors, but that tree populations are suppressed by mortality related to fire, herbivores, and other disturbances. If self‐thinning of woody plants can be detected in savannas, stronger evidence for resource‐limitation and competitive interactions among woody plants would suggest that the primary models of savannas need to be adjusted. We present data from savanna sites in South Africa to suggest that self‐thinning among woody plants can be detected in low‐disturbance situations, while also showing signs that juvenile trees, more so than adults, are suppressed when growing with herbaceous vegetation in these ecosystems. This finding we suggest is evidence for size‐asymmetric competition in savannas.     

Sixteen hundred years of increasing tree cover prior to modern deforestation in Southern Amazon and Central Brazilian savannas

Tropical ecosystems are under increasing pressure from land‐use change and deforestation. Changes in tropical forest cover are expected to affect carbon and water cycling with important implications for climatic stability at global scales. A major roadblock for predicting how tropical deforestation affects climate is the lack of baseline conditions (i.e., prior to human disturbance) of forest–savanna dynamics. To address this limitation, we developed a long‐term analysis of forest and savanna distribution across the Amazon–Cerrado transition of central Brazil. We used soil organic carbon isotope ratios as a proxy for changes in woody vegetation cover over time in response to fluctuations in precipitation inferred from speleothem oxygen and strontium stable isotope records. Based on stable isotope signatures and radiocarbon activity of organic matter in soil profiles, we quantified the magnitude and direction of changes in forest and savanna ecosystem cover. Using changes in tree cover measured in 83 different locations for forests and savannas, we developed interpolation maps to assess the coherence of regional changes in vegetation. Our analysis reveals a broad pattern of woody vegetation expansion into savannas and densification within forests and savannas for at least the past ~1,600 years. The rates of vegetation change varied significantly among sampling locations possibly due to variation in local environmental factors that constrain primary productivity. The few instances in which tree cover declined (7.7% of all sampled profiles) were associated with savannas under dry conditions. Our results suggest a regional increase in moisture and expansion of woody vegetation prior to modern deforestation, which could help inform conservation and management efforts for climate change mitigation. We discuss the possible mechanisms driving forest expansion and densification of savannas directly (i.e., increasing precipitation) and indirectly (e.g., decreasing disturbance) and suggest future research directions that have the potential to improve climate and ecosystem models.     

Spatial variability and temporal trends in water‐use efficiency of European forests

The increasing carbon dioxide (CO₂) concentration in the atmosphere in combination with climatic changes throughout the last century are likely to have had a profound effect on the physiology of trees: altering the carbon and water fluxes passing through the stomatal pores. However, the magnitude and spatial patterns of such changes in natural forests remain highly uncertain. Here, stable carbon isotope ratios from a network of 35 tree‐ring sites located across Europe are investigated to determine the intrinsic water‐use efficiency (iWUE), the ratio of photosynthesis to stomatal conductance from 1901 to 2000. The results were compared with simulations of a dynamic vegetation model (LPX‐Bern 1.0) that integrates numerous ecosystem and land–atmosphere exchange processes in a theoretical framework. The spatial pattern of tree‐ring derived iWUE of the investigated coniferous and deciduous species and the model results agreed significantly with a clear south‐to‐north gradient, as well as a general increase in iWUE over the 20th century. The magnitude of the iWUE increase was not spatially uniform, with the strongest increase observed and modelled for temperate forests in Central Europe, a region where summer soil‐water availability decreased over the last century. We were able to demonstrate that the combined effects of increasing CO₂ and climate change leading to soil drying have resulted in an accelerated increase in iWUE. These findings will help to reduce uncertainties in the land surface schemes of global climate models, where vegetation–climate feedbacks are currently still poorly constrained by observational data.     

Tree island pattern formation in the Florida Everglades

The Florida Everglades freshwater landscape exhibits a distribution of islands covered by woody vegetation and bordered by marshes and wet prairies. Known as “tree islands”, these ecogeomorphic features can be found in few other low gradient, nutrient limited freshwater wetlands. In the last few decades, however, a large percentage of tree islands have either shrank or disappeared in apparent response to altered water depths and other stressors associated with human impacts on the Everglades. Because the processes determining the formation and spatial organization of tree islands remain poorly understood, it is still unclear what controls the sensitivity of these landscapes to altered conditions. We hypothesize that positive feedbacks between woody plants and soil accretion are crucial to emergence and decline of tree islands. Likewise, positive feedbacks between phosphorus (P) accumulation and trees explain the P enrichment commonly observed in tree island soils. Here, we develop a spatially-explicit model of tree island formation and evolution, which accounts for these positive feedbacks (facilitation) as well as for long range competition and fire dynamics. It is found that tree island patterns form within a range of parameter values consistent with field data. Simulated impacts of reduced water levels, increased intensity of drought, and increased frequency of dry season/soil consuming fires on these feedback mechanisms result in the decline and disappearance of tree islands on the landscape.     

A probabilistic analysis of fire-induced tree-grass coexistence in savannas

Fires play an important role in determining the composition and structure of vegetation in semiarid ecosystems. The study of the interactions between fire and vegetation requires a stochastic approach because of the random and unpredictable nature of fire occurrences. To this end, this article develops a minimalist probabilistic framework to investigate the impact of intermittent fire occurrences on the temporal dynamics of vegetation. This framework is used to analyze the emergence of statistically stable conditions favorable to tree-grass coexistence in savannas. It is found that these conditions can be induced and stabilized by the stochastic fire regime. A decrease in fire frequency leads to bush encroachment, while more frequent and intense fires favor savanna-to-grassland conversions. The positive feedback between fires and vegetation can convert states of tree-grass coexistence in semiarid savannas into bistable conditions, with both woodland and grassland as possible, though mutually exclusive, stable states of the system.     

Bistability and regular spatial patterns in arid ecosystems

A variety of patterns observed in ecosystems can be explained by resource–concentration mechanisms. A resource–concentration mechanism occurs when organisms increase the lateral flow of a resource toward them, leading to a local concentration of this resource and to its depletion from areas farther away. In resource–concentration systems, it has been proposed that certain spatial patterns could indicate proximity to discontinuous transitions where an ecosystem abruptly shifts from one stable state to another. Here, we test this hypothesis using a model of vegetation dynamics in arid ecosystems. In this model, a resource–concentration mechanism drives a positive feedback between vegetation and soil water availability. We derived the conditions leading to bistability and pattern formation. Our analysis revealed that bistability and regular pattern formation are linked in our model. This means that, when regular vegetation patterns occur, they indicate that the system is along a discontinuous transition to desertification. Yet, in real systems, only observing regular vegetation patterns without identifying the pattern-driving mechanism might not be enough to conclude that an ecosystem is along a discontinuous transition because similar patterns can emerge from different ecological mechanisms.     

Integrating spatial structure and interspecific and intraspecific plant–soil feedback effects and responses into community structure

Plant–soil feedbacks have important effects on plant communities, but most theory has been derived from experiments on intraspecific plant–soil feedbacks. Much less is known about how interspecific plant–soil feedbacks affect coexistence and plant communities, due in part to experimental and analytical challenges. Here, we propose a framework for evaluating plant–soil feedbacks among multiple interacting species that incorporates 1) the average effect each species has on conspecific and heterospecific neighbors via how they modify soil biota, 2) the average response of each species to the soil modifications made by neighboring species, and 3) intraspecific feedback. We refer to this as the ‘effect–response–intraspecific’ (ERI) model. We used individual‐based models to evaluate the relative importance of intraspecific and interspecific soil feedback in determining species abundance ranks in simulated plant communities. To compare the heuristic value of the ERI model to that of an established model in which effects and responses to soil feedback are not explicitly recognized, we evaluated a ‘full‐factorial’ model in which soil feedbacks among five plant species were measured and then explicitly modeled. The ERI model indicated that the response to interspecific plant–soil feedbacks was the key factor for species’ abundance rank without spatial structure. In contrast, interspecific plant–soil feedback had no impact on species abundance with spatial structure, and intraspecific feedback became dominant. Thus, our models predict that the relative importance of intraspecific and interspecific feedbacks changes as a function of the degree of spatial structure in a system. Overall, the ERI model provides a novel and tractable framework for evaluating complex multi‐species plant–soil feedbacks.     

Seasonal,interannual and decadal drivers of tree and grass productivity in an Australian tropical savanna

Tree–grass savannas are a widespread biome and are highly valued for their ecosystem services. There is a need to understand the long‐term dynamics and meteorological drivers of both tree and grass productivity separately in order to successfully manage savannas in the future. This study investigated the interannual variability (IAV) of tree and grass gross primary productivity (GPP) by combining a long‐term (15 year) eddy covariance flux record and model estimates of tree and grass GPP inferred from satellite remote sensing. On a seasonal basis, the primary drivers of tree and grass GPP were solar radiation in the wet season and soil moisture in the dry season. On an interannual basis, soil water availability had a positive effect on tree GPP and a negative effect on grass GPP. No linear trend in the tree–grass GPP ratio was observed over the 15‐year study period. However, the tree–grass GPP ratio was correlated with the modes of climate variability, namely the Southern Oscillation Index. This study has provided insight into the long‐term contributions of trees and grasses to savanna productivity, along with their respective meteorological determinants of IAV.     

Seasonality and facilitation drive tree establishment in a semi-arid floodplain savanna

A popular hypothesis for tree and grass coexistence in savannas is that tree seedlings are limited by competition from grasses. However, competition may be important in favourable climatic conditions when abiotic stress is low, whereas facilitation may be more important under stressful conditions. Seasonal and inter-annual fluctuations in abiotic conditions may alter the outcome of tree–grass interactions in savanna systems and contribute to coexistence. We investigated interactions between coolibah (Eucalyptus coolabah) tree seedlings and perennial C₄ grasses in semi-arid savannas in eastern Australia in contrasting seasonal conditions. In glasshouse and field experiments, we measured survival and growth of tree seedlings with different densities of C₄ grasses across seasons. In warm glasshouse conditions, where water was not limiting, competition from grasses reduced tree seedling growth but did not affect tree survival. In the field, all tree seedlings died in hot dry summer conditions irrespective of grass or shade cover, whereas in winter, facilitation from grasses significantly increased tree seedling survival by ameliorating heat stress and protecting seedlings from herbivory. We demonstrated that interactions between tree seedlings and perennial grasses vary seasonally, and timing of tree germination may determine the importance of facilitation or competition in structuring savanna vegetation because of fluctuations in abiotic stress. Our finding that trees can grow and survive in a dense C₄ grass sward contrasts with the common perception that grass competition limits woody plant recruitment in savannas.     

A patch-dynamics approach to savanna dynamics and woody plant encroachment – Insights from an arid savanna

The coexistence of woody and grassy plants in savannas has often been attributed to a rooting-niche separation (two-layer hypothesis). Water was assumed to be the limiting resource for both growth forms and grasses were assumed to extract water from the upper soil layer and trees and bushes from the lower layers. Woody plant encroachment (i.e. an increase in density of woody plants often unpalatable to domestic livestock) is a serious problem in many savannas and is believed to be the result of overgrazing in ‘two-layer systems’. Recent research has questioned the universality of both the two-layer hypothesis and the hypothesis that overgrazing is the cause of woody plant encroachment.

We present an alternative hypothesis explaining both tree–grass coexistence and woody plant encroachment in arid savannas. We propose that woody plant encroachment is part of a cyclical succession between open savanna and woody dominance and is driven by two factors: rainfall that is highly variable in space and time, and inter-tree competition. In this case, savanna landscapes are composed of many patches (a few hectares in size) in different states of transition between grassy and woody dominance, i.e. we hypothesize that arid savannas are patch-dynamic systems. We summarize patterns of tree distribution observed in an arid savanna in Namibia and show that these patterns are in agreement with the patch-dynamic savanna hypothesis. We discuss the applicability of this hypothesis to fire-dominated savannas, in which rainfall variability is low and fire drives spatial heterogeneity.

We conclude that field studies are more likely to contribute to a general understanding of tree–grass coexistence and woody plant encroachment if they consider both primary (rain and nutrients) and secondary (fire and grazing) determinants of patch properties across different savannas.     

Numerical investigation of spatial pattern in a vegetation model with feedback function

The vegetative cover in semi-arid lands typically occurs as patches of individual species more or less separated from one another by bare ground. Klausmeier [1999. Regular and irregular patterns in semiarid vegetation. Science 284 (5421), 1826-1828] reported that the vegetation striped patterns can grow lying along the contours of gentle slopes. He has proposed a model of vegetation stripes based on competition for water. In this paper, our main aim is to study the positive feedback effects between the water and biomass on the vegetation spatial pattern formation within a nonsaturated soil, which arises from the suction of water by the roots and processes of water resource redistribution. According to the dispersion relation formula, we discuss the changes of the wavelength, wave speed, as well as the conditions of the spatial pattern formation. Our numerical results show that trees are more sensitive than grasses to the positive feedback function to format the spatial heterogenous pattern, and the stronger positive feedback increases the parameters region where vegetation bands occur, which indicates that the positive feedback raises the possibility of shift from green to desert states in semi-arid areas for the long term. Our numerical results also show that the positive feedback can increase the migration velocity of the vegetation stripes.     

Effects of plant-soil feedback on tree seedling growth under arid conditions

Aims Plants are able to influence their growing environment by changing biotic and abiotic soil conditions. These soil conditions in turn can influence plant growth conditions, which is called plant–soil feedback. Plant–soil feedback is known to be operative in a wide variety of ecosystems ranging from temperate grasslands to tropical rain forests. However, little is known about how it operates in arid environments. We examined the role of plant–soil feedbacks on tree seedling growth in relation to water availability as occurring in arid ecosystems along the west coast of South America.Methods In a two-phased greenhouse experiment, we compared plant–soil feedback effects under three water levels (no water, 10% gravimetric moisture and 15% gravimetric moisture). We used sterilized soil inoculated with soil collected from northwest Peru (Prosopis pallida forests) and from two sites in north-central Chile (Prosopis chilensis forest and scrublands without P. chilensis).Important findings Plant–soil feedbacks differed between plant species and soil origins, but water availability did not influence the feedback effects. Plant–soil feedbacks differed in direction and strength in the three soil origins studied. Plant–soil feedbacks of plants grown in Peruvian forest soil were negative for leaf biomass and positive for root length. In contrast, feedbacks were neutral for plants growing in Chilean scrubland soil and positive for leaf biomass for those growing in Chilean forest soil. Our results show that under arid conditions, effects of plant–soil feedback depend upon context. Moreover, the results suggest that plant–soil feedback can influence trade-offs between root growth and leaf biomass investment and as such that feedback interactions between plants and soil biota can make plants either more tolerant or vulnerable to droughts. Based on dissecting plant–soil feedbacks into aboveground and belowground tissue responses, we conclude that plant–soil feedback can enhance plant colonization in some arid ecosystems by promoting root growth.