Do biotic interactions shape both sides of the humped‐back model of species richness in plant communities?

A humped-back relationship between species richness and community biomass has frequently been observed in plant communities, at both local and regional scales, although often improperly called a productivity-diversity relationship. Explanations for this relationship have emphasized the role of competitive exclusion, probably because at the time when the relationship was first examined, competition was considered to be the significant biotic filter structuring plant communities. However, over the last 15 years there has been a renewed interest in facilitation and this research has shown a clear link between the role of facilitation in structuring communities and both community biomass and the severity of the environment. Although facilitation may enlarge the realized niche of species and increase community richness in stressful environments, there has only been one previous attempt to revisit the humped-back model of species richness and to include facilitative processes. However, to date, no model has explored whether biotic interactions can potentially shape both sides of the humped-back model for species richness commonly detected in plant communities. Here, we propose a revision of Grime's original model that incorporates a new understanding of the role of facilitative interactions in plant communities. In this revised model, facilitation promotes diversity at medium to high environmental severity levels, by expanding the realized niche of stress-intolerant competitive species into harsh physical conditions. However, when environmental conditions become extremely severe the positive effects of the benefactors wane (as supported by recent research on facilitative interactions in extremely severe environments) and diversity is reduced. Conversely, with decreasing stress along the biomass gradient, facilitation decreases because stress-intolerant species become able to exist away from the canopy of the stress-tolerant species (as proposed by facilitation theory). At the same time competition increases for stress-tolerant species, reducing diversity in the most benign conditions (as proposed by models of competition theory). In this way our inclusion of facilitation into the classic model of plant species diversity and community biomass generates a more powerful and richer predictive framework for understanding the role of plant interactions in changing diversity. We then use our revised model to explain both the observed discrepancies between natural patterns of species richness and community biomass and the results of experimental studies of the impact of biodiversity on the productivity of herbaceous communities. It is clear that explicit consideration of concurrent changes in stress-tolerant and competitive species enhances our capacity to explain and interpret patterns in plant community diversity with respect to environmental severity.     

Burial disturbance leads to facilitation among coastal dune plants

There is growing evidence that interactions among plants can be facilitative as well as competitive, but knowledge of how disturbances influence these interactions and how they vary with species diversity is lacking. We manipulated plant density, species diversity (richness), and a burial disturbance in a controlled, complete factorial experiment to test theories about the relationships among species interactions, disturbance, and richness. The hypotheses tested were 1) burial disturbance reduces plant performance at all levels of density and richness, 2) burial disturbance can cause net plant interactions to become more facilitative, and 3) facilitation increases with species richness. Burial decreased plant survival by 60% and biomass by 50%, supporting the hypothesis that burial reduces plant performance. In the control (unburied) treatment, there was no difference in proportion survival or per plant biomass between low and high density plots, meaning that neither competition nor facilitation was detected. In the buried treatment, however, high density plots had significantly greater survival and greater per plant biomass than the low density plots, indicating net facilitative interactions. Thus facilitation occurred in the buried treatment and not in the unburied control plots, supporting the hypothesis that facilitation increases with increasing disturbance severity. The hypothesis that facilitation increases with increasing species richness was not supported. Richness did not affect survival or biomass, and there was no richness by burial treatment interaction, indicating that richness did not influence the response of the community to burial. The influence of the disturbance on plant interactions was thus consistent across levels of richness, increasing the generality of the relationship between disturbance and facilitation.     

Comparing Neutral and Trade-off Community Models in Shaping the Community Biomass-Diversity Relationship Under Different Disturbance Levels

Among numerous mechanisms shaping the unimodal relationship between diversity and community biomass, the trade-off model of “CRS” theory is the most famous one. However, recent researches indicate that this relationship may also emerge under the neutral model where all species are identical with each other. By using an individual-based spatially-explicit model, we evaluated the underlying mechanisms shaping this curve for both models under different disturbance levels. We found unimodal relationships emerged for both models at low and medium disturbance levels; the richness for the trade-off community was lower than the neutral community for most of the environment severity levels, especially at the benign environment due to the strong competitive exclusions among species. Whereas under high disturbance level, the positive relationships emerged for both models; both communities had similar richness with their curves nearly overlapped with each other, that is, because the high disturbance intensity strongly decreased the competitive exclusions within the trade-off community. Our results indicate that although the underlying mechanisms are totally different, both models will produce the similar relationship between diversity and community biomass under different disturbance levels.     

A field test of the stress‐gradient hypothesis along an aridity gradient

Aims: The stress‐gradient hypothesis (SGH) predicts how plant interactions change along environmental stress gradients. We tested the SGH in an aridity gradient, where support for the hypothesis and the specific shape of its response curve is controversial. Location: Almería, Cáceres and Coimbra, three sites in the Iberian Peninsula that encompass the most arid and wet habitats in the distribution range of a nurse shrub species –Retama sphaerocarpa L. (Boiss) – in Europe. Methods: We analysed the effect of Retama on its understorey plant community and measured the biomass and species richness beneath Retama and in gaps. We estimated the frequency (changes in species richness), importance and intensity of the Retama effects, and derived the severity–interaction relationship pattern, analysing how these interaction indices changed along this aridity gradient. Results and conclusions: The intensity and frequency of facilitation by Retama increased monotonically with increasing environmental severity, and the importance tended to have a similar pattern, overall supporting the SGH. Our data did not support predictions from recent revisions of the SGH, which may not apply to whole plant communities like those studied here or when interactions are highly asymmetrical. Facilitation by Retama influenced community composition and species richness to the point that a significant fraction of species found at the most arid end of the gradient were only able to survive beneath the nurse shrub, whereas some of these species were able to thrive in gaps at more mesic sites, highlighting the ecological relevance of facilitation by nurse species in mediterranean environments, especially in the driest sites.     

Is the productivity of vegetation plots higher or lower when there are more species? Variable predictions from interaction of the ‘sampling effect’ and ‘competitive dominance effect’ on the habitat templet

Using a habitat templet model, we predict that the productivity (total biomass) of plots within a plant community may be positively, negatively or not at all related to variation in the number of species per plot, depending on successional stage (time since major disturbance) and habitat carrying capacity (reflecting the total resource supplying power of the habitat). For plots of a given size, a positive relationship between productivity and species richness is predicted in recently disturbed habitats because local neighbourhoods here will have been assembled largely stochastically, usually from a pool of available species with a right‐skewed size frequency distribution. Hence, in the earliest stages of succession, plots will have relatively high total biomass only if they contain at least some of the relatively uncommon larger species which will, in turn, be more likely in those neighbourhoods that contain more species (the sampling effect). Among these will also be some of the more common smaller species; hence, these high biomass, species‐rich plots should have relatively low species evenness, in contrast to what is predicted under effects involving species complementarity. In late succession, the plots with high total biomass will still be those that contain relatively large species but these plots will now contain relatively few species owing to increased competitive exclusion over time (the competitive dominance effect). In intermediate stages of succession, no relationship between plot productivity and species richness is predicted because the opposing sampling and competitive dominance effects cancel each other out. We predict that the intensity of both the sampling and competitive dominance effects on the productivity/species richness relationship will decrease with decreasing habitat carrying capacity (e.g. decreasing substrate fertility) owing to the inherently lower variance in between‐plot productivity that is predicted for more resource‐impoverished habitats.     

From selection to complementarity: the shift along the abiotic stress gradient in a controlled biodiversity experiment

The effects of principal mechanisms (selection and complementarity) of biodiversity on ecosystem functionality have been well studied. However, it remains unknown how environmental conditions affect the relative strength of these two mechanisms. To answer this question, a controlled pot experiment was conducted in which species diversity was manipulated in low (natural soil) and high stress (mine tailing) plots, respectively. Our results demonstrate that the principal mechanism underlying the increasing biomass shifts from the selection to complementarity with increasing abiotic stress. The shift occurs because species interactions varied with increasing abiotic stress. Competition prevails in low stress plots, while facilitation dominates in high stress plots. In low stress plots, the monoculture biomass of a specific species is a good indicator of the competitive ability of that species in the mixture, and the dominant species significantly affects the plot biomass. In high stress plots, the tolerance indexes of all individual species increase with the manipulated species richness, providing clear evidence for the increasing role of facilitation.     

Establishment, competition and the distribution of native grasses among Michigan old-fields

1 In this study the potential role of competition in influencing the distribution of three displaced native perennial grasses across complex gradients of plant productivity and species composition was investigated in Michigan old-fields. To do this plant removal and propagule addition experiments were conducted at nine old-field sites to examine the effects of living plant neighbours and litter on seedling establishment and growth of target species in relation to community biomass.
2 For two target species, Andropogon gerardi and Schizachyrium scoparium , living plant neighbours suppressed establishment from seed at most sites, and suppressed the growth of transplants at all sites.
3 Plant litter strongly inhibited the seedling establishment of both Andropogon and Schizachyrium at sites of high community biomass and litter accumulation, but had little impact on the growth rate of transplants at any of the sites.
4 The total suppressive effect of the plant community on seedling establishment and transplant growth of both Andropogon and Schizachyrium increased in magnitude in a non-linear fashion with community biomass. These effects increased in magnitude more rapidly across sites of low to medium biomass than sites of medium to high biomass.
5 The results suggest that these native grasses may be restricted to low productivity habitats within this landscape because of strong competitive interference with establishment by the existing vegetation in the most productive sites.     

Microhabitat amelioration and reduced competition among understorey plants as drivers of facilitation across environmental gradients: Towards a unifying framework

Studies of facilitative interactions as drivers of plant richness along environmental gradients often assume the existence of an overarching stress gradient that equally affects the performance of all the species in a given community. However, co-existing species differ in their ecophysiological adaptations, and do not experience the same stress level under particular environmental conditions. Moreover, these studies assume a unimodal relationship between richness and biomass, which is not as general as previously thought. We ignored these assumptions to assess changes in plant–plant interactions and their effect on local species richness across environmental gradients in semi-arid areas of Spain and Australia. We aimed to understand the relative importance of direct (microhabitat amelioration) and indirect (changes in the competitive relationships among the understorey species: niche segregation, competitive exclusion or intransitivity) mechanisms that might underlie the effects of nurse plants on local species richness. By jointly studying these direct and indirect mechanisms using a unifying framework, we found that nurse plants (trees, shrubs and tussock grasses) increased local richness not only by expanding the niche of neighbouring species but also by increasing niche segregation among them, though the latter was not important in all cases. The outcome of the competition-facilitation continuum varied depending on the study area, likely because the different types of stress gradient considered. When driven by both rainfall and temperature, or rainfall alone, the community-wide importance of nurse plants remained constant (Spanish sites), or showed a unimodal relationship along the gradient (Australian sites). This study expands our understanding of the relative roles of plant–plant interactions and environmental conditions as drivers of local species richness in semi-arid environments. The results can also be used to refine predictions about the response of plant communities to environmental change, and to clarify the relative importance of biotic interactions as drivers of such responses.     

Density may alter diversity–productivity relationships in experimental plant communities

Contemporary biodiversity experiments, in which plant species richness is manipulated and aboveground productivity of the system measured, generally demonstrate that lowering plant species richness reduces productivity. However, we propose that community density may in part compensate for this reduction of productivity at low diversity. We conducted a factorial experiment in which plant functional group richness was held constant at three, while plant species richness increased from three to six to 12 species and community density from 440 to 1050 to 2525 seedlings m⁻². Response variables included density, evenness and above- and belowground biomass at harvest. The density gradient converged slightly during the course of the experiment due to about 10% mortality at the highest sowing density. Evenness measured in terms of aboveground biomass at harvest significantly declined with density, but the effect was weak. Overall, aboveground, belowground and total biomass increased significantly with species richness and community density. However, a significant interaction between species richness and community density occurred for both total and aboveground biomass, indicating that the diversity–productivity relationship was flatter at higher than at lower density. Thus, high species richness enabled low-density communities to reach productivity levels otherwise seen only at high density. The relative contributions of the three functional groups C₃, C₄ and nitrogen-fixers to aboveground biomass were less influenced by community density at high than at low species richness. We interpret the interaction effects between community density and species richness on community biomass by expanding findings about constant yield and size variation from monocultures to plant mixtures.     

An unfortunate alliance: Native shrubs increase the abundance,performance, and apparent impacts of Bromus tectorum across a regional aridity gradient

Interspecific facilitation contributes to the assembly of desert plant communities. However, we know little of how desert communities invaded by exotic species respond to facilitation along regional-scale aridity gradients. These measures are essential for predicting how desert plant communities might respond to concomitant plant invasion and environmental change. Here, we evaluated the potential for Bromus tectorum (a dominant invasive plant species) and the broader herbaceous plant community to form positive associations with native shrubs along a substantial aridity gradient across the Great Basin, Mojave, and San Joaquin Deserts in North America. Along this gradient, we sampled metrics of abundance and performance for B. tectorum, all native herbaceous species combined, all exotic herbaceous species combined, and the total herbaceous community using 180 pairs of shrub and open microsites. Across the gradient, B. tectorum formed strong positive associations with native shrubs, achieving 1.6–2.2 times greater abundance, biomass, and reproductive output under native shrubs than away from shrubs, regardless of relative aridity. In contrast, the broader herbaceous community was not positively associated with native shrubs. Interestingly, increasing B. tectorum abundance corresponded to decreasing native abundance, native species richness, exotic species richness, and total species richness under but not away from shrubs. Taken together, these findings suggest that native shrubs have considerable potential to directly (by increasing abundance and performance) and indirectly (by increasing competitive effects on neighbors) facilitate B. tectorum invasion across a large portion of the non-native range.     

Seed limitation interacts with biotic and abiotic factors to constrain novel species' impact on community biomass and richness

Seed limitation can narrow down the number of coexisting plant species, limit plant community productivity, and also constrain community responses to changing environmental and biotic conditions. In a 10-year full-factorial experiment of seed addition, fertilisation, warming and herbivore exclusion, we tested how seed addition alters community richness and biomass, and how its effects depend on seed origin and biotic and abiotic context. We found that seed addition increased species richness in all treatments, and increased plant community biomass depending on nutrient addition and warming. Novel species, originally absent from the communities, increased biomass the most, especially in fertilised plots and in the absence of herbivores, while adding seeds of local species did not affect biomass. Our results show that seed limitation constrains both community richness and biomass, and highlight the importance of considering trophic interactions and soil nutrients when assessing novel species immigrations and their effects on community biomass.     

Environmental stress alters native-nonnative relationships at the community scale

The invasion of natural habitats by nonnative species is affected by both native biodiversity and environmental conditions; however few tests of facilitation between native community members and nonnative species have been conducted along disturbance and stress gradients. There is strong evidence for an increase in facilitation between native plant species with increasing levels of natural environmental stress, however it is unknown whether these same positive interactions occur between nonnative invaders and native communities. I investigated the effects of natural stress on community interactions between native heathland species and nonnative species with two field studies conducted at the landscape and community scale. At the landscape scale of investigation, nonnative species richness was positively related to native species richness. At the community level, nonnative invaders experienced facilitation with natives in the most stressful zones, whereas they experienced competition with native plants in the less stressful zones of the heathlands. Due to the observational nature of the landscape scale data, it is unclear whether nonnative diversity levels are responding positively to extrinsic factors or to native biodiversity. The experimental component of this research suggests that native community members may ameliorate stressful environmental conditions and facilitate invasion into high stress areas. I present a conceptual model which is a modification of the Shea and Chesson diversity-invasibility model and includes both facilitation as well as competition between the native community and nonnative invaders at the community level, summing to an overall positive relationship at the landscape scale.     

Community diversity outweighs effect of warming on plant colonization

Abiotic environmental change, local species extinctions and colonization of new species often co‐occur. Whether species colonization is driven by changes in abiotic conditions or reduced biotic resistance will affect community functional composition and ecosystem management. We use a grassland experiment to disentangle effects of climate warming and community diversity on plant species colonization. Community diversity had dramatic impacts on the biomass, richness and traits of plant colonists. Three times as many species colonized the monocultures than the high diversity 17 species communities (~30 vs. 10 species), and colonists collectively produced 10 times as much biomass in the monocultures than the high diversity communities (~30 vs. 3 g/m²). Colonists with resource‐acquisitive strategies (high specific leaf area, light seeds, short heights) accrued more biomass in low diversity communities, whereas species with conservative strategies accrued most biomass in high diversity communities. Communities with higher biomass of resident C4 grasses were more resistant to colonization by legume, nonlegume forb and C3 grass colonists, but not by C4 grass colonists. Compared with effects of diversity, 6 years of 3°C‐above‐ambient temperatures had little impact on plant colonization. Warmed subplots had ~3 fewer colonist species than ambient subplots and selected for heavier seeded colonists. They also showed diversity‐dependent changes in biomass of C3 grass colonists, which decreased under low diversity and increased under high diversity. Our findings suggest that species colonization is more strongly affected by biotic resistance from residents than 3°C of climate warming. If these results were extended to invasive species management, preserving community diversity should help limit plant invasion, even under climate warming.     

Does macrophyte fractal complexity drive invertebrate diversity,biomass and body size distributions?

Habitat structure is one of the fundamental factors determining the distribution of organisms at all spatial scales, and vegetation is of primary importance in shaping the structural environment for invertebrates in many systems. In the majority of biotopes, invertebrates live within vegetation stands of mixed species composition, making estimates of structural complexity difficult to obtain. Here we use fractal indices to describe the structural complexity of mixed stands of aquatic macrophytes, and these are employed to examine the effects of habitat complexity on the composition of free-living invertebrate assemblages that utilise the habitat in three dimensions. Macrophytes and associated invertebrates were sampled from shallow ponds in southwest England, and rapid digital image analysis was used to quantify the fractal complexity of all plant species recorded, allowing the complexity of vegetation stands to be reconstructed based on their species composition. Fractal indices were found to be significantly related to both invertebrate biomass–body size scaling and overall invertebrate biomass; more complex stands of macrophytes contained a greater number of small animals. Habitat complexity was unrelated to invertebrate taxon richness and macrophyte surface area and species richness were not correlated with any of the invertebrate community parameters. The biomass–body size scaling relationship of lentic macroinvertebrates matched those predicted by models incorporating both allometric scaling of resource use and the fractal dimension of a habitat, suggesting that both habitat fractal complexity and allometry may control density–body size scaling in lentic macroinvertebrate communities.     

The species richness–biomass relationship in herbaceous plant communities: what difference does the incorporation of root biomass data make?

So far, in all studies on the much-discussed hump-backed relationship between plant community productivity and species richness, productivity has been assessed through plant shoot biomass, i.e. it has been ignored that frequently most of the biomass is produced below ground. We revisited the 27 grassland and forest field-layer communities, studied earlier by Zobel and Liira, to sample root biomass, plant total biomass and root/shoot allocation, and learn how the incorporation of below-ground biomass data would affect the shape of the hump-backed relationship. In order to avoid scaling artefacts we estimated richness as the average count of species per 500 plant ramets (absolute richness). We also included relative richness measures. Relative richness was defined as richness per 500 ramets/size of the actual species pool (the set of species present in the community), relative pool size was defined as size of the actual species pool/size of the regional species pool (the set of species available in the region and capable of growing in the given community).
The biomass-absolute richness relationship was humped, irrespective of the biomass measure used, the hump being most obvious when plant total biomass was used as the independent variable. Evidently, the unimodal richness–productivity curve is not a sampling artefact, as suspected by Oksanen. However, relative richness was not related to community biomass (above-ground, below-ground or total). The hump-backed curve is shaped by the sizes of actual species pools in communities, implying that processes which are responsible for small-scale diversity pattern mainly operate on the community level.
Neither absolute nor relative richness were significantly related to root/shoot allocation. The presumably stronger (asymmetric) shoot competition at greater allocation to shoots appears not to suppress small-scale richness. However, there is a significant relationship between relative pool size and root/shoot allocation. Relatively more species from regional species pools are able to enter and persist in communities with more biomass allocated into roots.     

Phylogenetic distance among beneficiary species in a cushion plant species explains interaction outcome

Determining which drivers lead to a specific species assemblage is a central issue in community ecology. Although many processes are involved, plant–plant interactions are among the most important. The phylogenetic limiting similarity hypothesis states that closely related species tend to compete stronger than distantly related species, although evidence is inconclusive. We used ecological and phylogenetic data on alpine plant communities along an environmental severity gradient to assess the importance of phylogenetic relatedness in affecting the interaction between cushion plants and the whole community, and how these interactions may affect community assemblage and diversity. We first measured species richness and individual biomass of species growing within and outside the nurse cushion species, Arenaria tetraquetra. We then assembled the phylogenetic tree of species present in both communities and calculated the phylogenetic distance between the cushion species and its beneficiary species, as well as the phylogenetic community structure. We also estimated changes in species richness at the local level due to the presence of cushions. The effects of cushions on closely related species changed from negative to positive as environmental conditions became more severe, while the interaction with distantly related species did not change along the environmental gradient. Overall, we found an environmental context‐dependence in patterns of phylogenetic similarity, as the interaction outcome between nurses and their close and distantly‐related species showed an opposite pattern with environmental severity.     

Nutrient concentrations and fibre contents of plant community biomass reflect species richness patterns along a broad range of land-use intensities among agricultural grasslands

Understanding changes in biodiversity in agricultural landscapes in relation to land-use type and intensity is a major issue in current ecological research. In this context nutrient enrichment has been identified as a key mechanism inducing species loss in Central European grassland ecosystems. At the same time, insights into the linkage between agricultural land use and plant nutrient status are largely missing. So far, studies on the relationship between chemical composition of plant community biomass and biodiversity have mainly been restricted to wetlands and all these studies neglected the effects of land use. Therefore, we analyzed aboveground biomass of 145 grassland plots covering a gradient of land-use intensities in three regions across Germany. In particular, we explored relationships between vascular plant species richness and nutrient concentrations as well as fibre contents (neutral and acid detergent fibre and lignin) in the aboveground community biomass.We found the concentrations of several nutrients in the biomass to be closely linked to plant species richness and land use. Whereas phosphorus concentrations increased with land-use intensity and decreased with plant species richness, nitrogen and potassium concentrations showed less clear patterns. Fibre fractions were negatively related to nutrient concentrations in biomass, but hardly to land-use measures and species richness. Only high lignin contents were positively associated with species richness of grasslands. The N:P ratio was strongly positively related to species richness and even more so to the number of endangered plant species, indicating a higher persistence of endangered species under P (co-)limited conditions. Therefore, we stress the importance of low P supply for species-rich grasslands and suggest the N:P ratio in community biomass to be a useful proxy of the conservation value of agriculturally used grasslands.     

The Effect of Positive Interactions on Temporal Turnover of Community Composition along an Environmental Gradient

It has been demonstrated that the interplay between negative and positive interactions simultaneously shapes community structure and composition. However, few studies have attempted to examine the effect of facilitation on compositional changes in communities through time. Additionally, due to the difficulties in collecting the long-term data, it would be useful to indicate the rate of temporal turnover using a readily obtainable metric. Using an individual-based model incorporating plant strategies, we examined the role of facilitation on the temporal turnover of communities located at different positions along an environmental gradient for three model scenarios: CM without facilitation; CFM-U, a unimodal relationship between facilitation and environmental severity; and CFM-L, a positively linear relationship between facilitation and environmental severity. Our results demonstrated that facilitation could increase, decrease or have no remarkable effect on temporal turnover. The specific outcome depended on the location of the focal community across the environmental gradient and the model employed. Compared with CM, the inclusion of positive interactions (i.e. CFM-U and CFM-L), at intermediate environmental stress levels (such as S = 0.7 and 0.8) resulted in lower Bray-Curtis similarity values; at other severity levels, facilitation slowed down (such as S = 0.3 and 0.4 at low to medium stress levels, and S = 0.9 at high stress levels) or had only a subtle effect (such as at S = 0.1) on temporal turnover. We also found that the coefficient of variation (CV) in species abundances and the rate of temporal variability showed a significant quadratic relationship. Our theoretical analysis contributes to the understanding of factors driving temporal turnover in biotic communities, and presents a potential metric (i.e. CV in species abundances) assessing the consequences of ongoing environmental change on community structure.     

Effect of cattle grazing on soil salinity and vegetation composition along an elevation gradient in a temperate coastal salt marsh of Samborombón Bay (Argentina)

Salt marshes of Samborombón Bay (Argentina) have been grazed sporadically at very low stocking rates, but in the last decade, grazing intensity increased due to agriculture expansion. We investigated the effect of cattle grazing on vegetation and soil salinity on the most extended Spartina densiflora community. This community develops along an elevation gradient where the frequency and duration of tidal flooding and soil salinity increased as elevation decreased. Vegetation and soil data were collected from a national park excluded to cattle grazing for 30 years and from an adjacent commercial livestock farm continuously grazed by cattle. As elevation level decreased, plant cover, richness and diversity of functional groups and species decreased. As we expected, grazing altered soil salinity and vegetation composition in different extent along the elevation gradient. Grazing changed vegetation structure more intensively in the high elevation level because it reduced the competitive exclusion exerted by S. densiflora, allowing the increase in floristic richness. Grazing increased soil salinity and the contribution of salt-tolerant species only in the medium but not in the low elevation level probably because the higher frequency and duration of tidal flooding counterbalanced the increase in evaporation promoted by biomass removal in the low respect to the medium elevation level. While grazing may cause positive impacts for plant conservation in the high elevation level, it may cause negative consequence for livestock production because of the reduction in forage quality along the entire elevation gradient.     

Extreme drought stress shifts net facilitation to neutral interactions between shrubs and sub‐canopy plants in an arid desert

The stress gradient hypothesis (SGH) predicts that the importance or intensity of competition and facilitation will change inversely along abiotic stress gradients. It was originally postulated that increasing environmental stress can induce a monotonic increase in facilitation. However, more recent models predicted that the relationship between severity and interaction exhibits a hump‐shaped pattern, in which positive interactions prevail under moderate stress but decline at the extreme ends of stress gradients. In the present study, we conducted a field experiment along a temporal rainfall gradient for five consecutive years, in order to investigate interactions in a shrub‐herbaceous plant community at the southern edge of the Badain Jaran Desert, and, more specifically, investigated the effects of Calligonum mongolicum, a dominant shrub species, on both abiotic environmental variables and the performance of sub‐canopy plant species. We found that shrubs can improve sub‐canopy water regimes, soil properties, plant biomass, density, cover, and richness and, more importantly, that the positive effect of shrubs on sub‐canopy soil moisture during the summer diminishes as rainfall decreases, a pattern that partly explains the collapse of the positive interaction between shrubs and their understory plants. These results provide empirical evidence that the positive effect of shrubs on understory plant communities in extreme arid environments may decline and become neutral with increasing drought stress.