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.     

Context‐dependent interactions between adult shrubs and seedlings in a semi‐arid shrubland

Abstract. Question: In semi‐arid systems, rainfall gradients can cause plant‐plant interactions to shift from negative to positive or vice versa. However, the importance of a second major abiotic factor, soil nutrients, has rarely been considered. We consider different combinations of both factors and ask: do net adult‐seedling interactions become less competitive and more facilitative with increasing overall abiotic harshness? Location: Succulent Karoo, Western Cape, South Africa. Methods: We examined the interactions between seedlings and adult shrubs at two sites. Sites differ in rainfall, and each contain two habitats: Nutrient‐rich mounds associated with underground termitaria and a relatively nutrient‐poor matrix. We carried out a spatial pattern analysis of community‐wide seedling‐adult associations. We then conducted field and greenhouse experiments to test the effects of soil and the presence of neighbouring shrubs on the growth and survival of six seedling species. Results: At the higher rainfall site, both competitive and facilitative effects of adults on seedlings were found but did not differ by habitat, despite the more benign conditions in the mound habitat. At the lower rainfall site, adult shrubs generally had neutral effects on seedlings in the matrix habitat. In the nutrient‐rich mound habitat, however, adult shrubs had strong and consistently competitive effects on seedlings. Conclusion: Seedling‐adult interactions could not be predicted by a simple overall gradient of abiotic harshness, demonstrating the need for more complex, mechanistic models to predict plant‐plant interactions. We suggest that rainfall and soil nutrients affect seedling‐adult relations through their interactive effects on the life‐history attributes of the species involved.     

Disentangling direct and indirect effects of a legume shrub on its understorey community

Direct and indirect interactions among plants contribute to shape community composition through above‐ and belowground processes. However, we have not disentangled yet the direct and indirect soil and canopy effects of dominants on understorey species. We addressed this issue in a semi‐arid system from southeast Spain dominated by the legume shrub Retama sphaerocarpa. During a year with an exceptionally dry spring, we removed the shrub canopy to quantify aboveground effects and compared removed‐canopy plots to open plots between shrubs to quantify soil effects, both with and without watering. We added a grass removal treatment in order to separate direct from indirect shrub effects and quantified biomass, abundance, richness and composition of the forb functional group. With watering, changes in forb biomass were primarily driven by indirect shrub effects, with contrasting negative soil and positive aboveground indirect effects; changes in forb abundance and composition were more influenced by direct shrub soil effects with contrasting species composition between open and Retama patches. As community composition was different between open and Retama patches the indirect effects of Retama on forb species did not concern forbs from the open community but forbs from Retama patches. Indirect effects are, thus, important at the functional group level rather than at the species level. Without watering, there were no significant interactions. Changes in species richness between treatments were weak and seldom significant. We conclude that shrub effects on understorey forbs are primarily due to their influence on soil properties, directly affecting forb species composition but indirectly affecting the biomass of the forbs of the Retama patches, and only with sufficient water.     

Shrub encroachment in the Alps gives rise to the convergence of sub‐alpine communities on a regional scale

Question: How does Alnus alnobetula colonization affect plant communities in sub‐alpine grasslands undergoing land abandonment on a regional scale? Location: Savoie, French Alps. Methods: Variations within and between communities were monitored in 243 plots within three types of stands representing an A. alnobetula colonization chronosequence from grassland to dense vegetation. They were tested for species richness, species strategies and species distribution through multivariate analysis. Regional variations were assessed at three sites along a rain shadow gradient. Results: We detected a convergence of the understorey communities along an A. alnobetula colonization gradient at all sites, associated with a reduction in species richness and the replacement of stress related strategists by competitor strategists. This convergence was unaffected by the rain shadow gradient although it was a key factor of plant distribution. Conclusions: The dense cover and the soil properties provided by A. alnobetula stands buffered the heterogeneity of sub‐alpine communities induced by a rain shadow gradient throughout the study region. It facilitated the development a homogeneous understorey community dominated by competitor related strategists while excluding grasslands stress adapted species through competition processes. The resulting community, although species‐poor, has an interesting conservation value. Accordingly, as a ‘native invader’, A. alnobetula does not display typical traits of biological invaders. Future experimental research on the interactions between A. alnobetula and components of the understorey would be particularly interesting in a severe environment that promotes facilitation among plants.     

The influence of Prosopis canopies on understorey vegetation: Effects of landscape position

Abstract. The influence of canopy trees and shrubs on under‐storey plants is complex and context‐dependent. Canopy plants can exert positive, negative or neutral effects on production, composition and diversity of understorey plant communities, depending on local environmental conditions and position in the landscape. We studied the influence of Prosopis velutina (mesquite) on soil moisture and nitrogen availability, and understorey vegetation along a topographic gradient in the Sonoran Desert. We found significant increases in both soil moisture and N along the gradient from desert to riparian zone. In addition, P. velutina canopies had positive effects, relative to open areas, on soil moisture in the desert, and soil N in both desert and intermediate terrace. Biomass of understorey vegetation was highest and species richness was lowest in the riparian zone. Canopies had a positive effect on biomass in both desert and terrace, and a negative effect on species richness in the terrace. The effect of the canopy depended on landscape position, with desert canopies more strongly influencing soil moisture and biomass and terrace canopies more strongly influencing soil N and species richness. Individual species distributions suggested interspecific variation in response to water‐ vs. N‐availability; they strongly influence species composition at both patch and landscape position levels.     

Interactive effects of grazing and shrubs on the annual plant community in semi‐arid Mediterranean shrublands

Question: We studied the interactive effects of grazing and dwarf shrub cover on the structure of a highly diverse annual plant community. Location: Mediterranean, semi‐arid shrubland in the Northern Negev desert, Israel. Methods: Variation in the biomass and plant density of annual species in the shrub and open patches was monitored during four years, inside and outside exclosures protected from sheep grazing, in two contrasting topographic sites: north and south‐facing slopes that differed in their dominant dwarf shrubs species: Sarcopoterium spinosus and Corydothymus capitatus, respectively. Results: Above‐ground biomass, density and richness of annual species were lower under the canopy of both shrub species compared to the adjacent open patches in the absence of grazing. Grazing reduced the biomass of annuals in open patches of both topographic sites, but not in the shrub patches. On the north‐facing slope, grazing also reduced plant density and richness in the open patches, but increased plant density in the shrub patches. At the species level, various response patterns to the combined effects of grazing and patch type were exhibited by different annuals. Protection against the direct impacts of grazing by shrub cover as well as species‐specific interactions between shrubs and annuals were observed. A conceptual mechanistic model explaining these interactions is proposed. Conclusion: In semi‐arid Mediterranean shrublands grazing and dwarf shrub cover interact in shaping the structure of the annual plant community through (1) direct impacts of grazing restricted to the open patches, (2) species‐specific facilitation/ interference occurring in the shrub patches and (3) subsequent further processes occurring among the interconnected shrub and open patches mediated through variation in seed flows between patches.     

Seed Bank and Understorey Species Composition in a Semi-arid Environment: The Effect of Shrub Age and Rainfall

Understorey vegetation in patches of Retama sphaerocarpa shrubsin semi-arid environments is dependent on the overstorey shrublife history. Community structure changes with shrub age asa result of physical amelioration of environmental conditionsby the canopy and organic matter accumulation in the soil. Weinvestigated the effect of the canopy on understorey speciesdiversity in the field and its relationships with the soil seedbank under 50 shrubs from 5 to 25+ years old, and compared speciescomposition in the field in a wet and a dry year. Species compositionof the soil seed bank under R. sphaerocarpa shrubs did not differsignificantly with shrub age, but seed density increased asthe shrubs aged. In the field, community composition changedwith shrub age, increasing species richness in a process thatdepended on the amount of spring rainfall. Our results suggestthat the soil seed bank is rather uniform and that the shrubcanopy strongly selects which species appear in the understorey.There were seeds of many species present under both young andold shrubs but which only established under old shrubs. Thisshowed dispersal was not limiting species abundance and suggestedthat the canopy was an important sorting factor for speciespresent in the understorey. Less frequent species contributedthe most to patch diversity, and rainfall effectively controlledspecies emergence. Understorey community composition dependedon multiple interspecific interactions, such as facilitationby the shrub and competition from neighbours, as well as ondispersal processes. Facilitation in this environment is a keyfeature in the structuring of plant communities and in governingecosystem functioning. Copyright 2000 Annals of Botany Company Community structure, competition, dispersal, facilitation, species composition, rainfall variability, Retama sphaerocarpa, seed bank, semi-arid environments     

Shrub facilitation drives tree establishment in a semiarid fog‐dependent ecosystem

Questions

The exceptional occurrence of tall rain forest patches on foggy coastal mountaintops, surrounded by extensive xerophytic shrublands, suggests an important role of plant–plant interactions in the origin and persistence of these patches in semi‐arid Chile. We asked whether facilitation by shrubs can explain the growth and survival of rain forest tree species, and whether shrub effects depend on the identity of the shrub species itself, the drought tolerance of the tree species and the position of shrubs in regard to wind direction.

Location

Open area–shrubland–forest matrix, Fray Jorge Forest National Park, Chile.

Methods

We recorded survival after 12 years of a ~3600 tree saplings plantation (originally ~30‐cm tall individuals) of Aextoxicon punctatum, Myrceugenia correifolia and Drimys winteri placed outside forests, beneath the shrub Baccharis vernalis, and in open (shrub‐free) areas. We assessed the effects of neighbouring shrubs and soil humidity on survival and growth along a gradient related to the direction of fog movement.

Results

B. vernalis had a clear facilitative effect on tree establishment and survival since, after ~12 years, saplings only survived beneath the shrub canopy. Long‐term survival strongly depended on tree species identity, drought tolerance and position along the soil moisture gradient, with higher survival of A. punctatum (>35%) and M. correifolia (>14%) at sites on wind‐ and fog‐exposed shrubland areas. Sites occupied by the shrub Aristeguietia salvia were unsuitable for trees, presumably due to drier conditions than under B. vernalis.

Conclusions

Interactions between shrubs and fog‐dependent tree species in dry areas revealed a strong, long‐lasting facilitation effect on planted tree's survival and growth. Shrubs acted as benefactors, providing sites suitable for tree growth. Sapling mortality in the shrubland interior was caused by lower soil moisture, the consequence of lower fog loads in the air and thus insufficient facilitation. While B. vernalis was a key ecosystem engineer (nurse) and intercepted fog water that dripped to trees planted underneath, drier sites with A. salvia were unsuitable for trees. Consequently, nurse effects related to water input are strongly site and species specific, with facilitation by shrubs providing a plausible explanation for the initiation of forest patches in this semi‐arid landscape.     

Spatial patterns of desert annuals in relation to shrub effects on soil moisture

Questions: What are the effects of a shrub (Haloxylon ammodendron) on spatial patterns of soil moisture in different seasons? How does productivity of understorey annuals respond to these effects? Are such effects always positive for annuals under shrubs? Location: South Gurbantunggut Desert, northwest China. Methods: Using geostatistics, we explored seasonal patterns of topsoil moisture in a 12 × 9‐m plot over the growing season. To determine spatial patterns of understorey annuals in response to H. ammodendron presence, biomass of annuals was recorded in four 0.2 × 5.0‐m transects from the centre of a shrub to the space between shrubs (interspace). We also investigated vertical distribution of root biomass for annuals and soil moisture dynamics across soil profiles in shrub‐canopied areas and interspaces. Results: Topsoil moisture changed from autocorrelation in the wet spring to random structure in the dry season, while soil moisture below 20 cm was higher in shrub‐canopied areas. Across all microhabitats, soil moisture in upper soil layers was higher than in deeper soil layers during the spring wet season, but lower during summer drought. Topsoil was close to air‐dry during the dry season and developed a ‘dry sand layer’ that reduced evaporative loss of soil water from deeper layers recharged by snowmelt in spring. Aboveground biomass of understorey annuals was lowest adjacent to shrub stems and peaked at the shrub margin, forming a ‘ring’ of high herbaceous productivity surrounding individual shrubs. To acclimate to drier conditions, annuals in interspaces invested more root biomass in deeper soil with a root/shoot ratio (R/S) twice that in canopied areas. Conclusions: Positive and negative effects of shrubs on understorey plants in arid ecosystems are commonly related to nature of the environmental stress and tested species. Our results suggest there is also microhabitat‐dependence in the Gurbantunggut Desert. Soil water under H. ammodendron is seasonally enriched in topsoil and deeper layers. Understorey annuals respond to the effect of shrubs on soil water availability with lower R/S and less root biomass in deeper soil layers and develop a ‘ring’ of high productivity at the shrub patch margin where positive and negative effects of shrubs are balanced.     

Positive effects of shrubs on plant species diversity do not change along a gradient in grazing pressure in an arid shrubland

Facilitative or positive interactions among species are driven mainly by the environmental amelioration or protection from grazing provided by nurse plants. Some studies have suggested that protection from grazing is inconsequential in water-limited environments because of low herbivore densities and their grazing effects. Others, however, argue that herbivores have a major effect on semi-arid plant communities, and that protection from grazing is a significant factor driving positive plant–plant interactions in such environments. We identified a gradient in grazing pressure in a semi-arid shrubland in south-eastern Australia along which we compared soil condition, incident radiation and plant composition beneath two nurse shrub species with open (shrub-free) interspaces. Our aim was to assess the degree of microclimatic amelioration provided by both shrubs, and changes in the interactions (intensity, importance and frequency) between both nurse shrubs and their understorey species, and their effects on species richness at the community level. Both the relative interaction intensity (RII) and interaction importance (I_imp) indices of plant–plant interactions were generally positive and independent of grazing pressure. Soil beneath both nurse plants had significantly greater indices of nutrient cycling and infiltration, and contained more C and N than soil in the open. Almost twice as many species occurred under the canopies of both shrubs (44 species) than in the open (23 species), and the composition of species differed significantly among microsites. Fifty-four percent of all perennial plant species occurred exclusively under shrubs. Our results suggest that environmental amelioration is a stronger driver of the facilitatory effect of shrubs on their understorey species than protection from grazing. Our conclusions are based on the fact that the substantial effect of plant–plant interactions on plant species richness was largely independent of grazing pressure. Irrespective of the underlying mechanism for this effect, our study illustrates the ecological role of shrubs as refugia for understorey plants in semi-arid environments and cautions against management practices aimed at reducing shrub populations.     

Individual and combined effects of disturbance and N addition on understorey vegetation in a subarctic mountain birch forest

Questions: What are the effects of repeated disturbance and N‐fertilization on plant community structure in a mountain birch forest? What is the role of enhanced nutrient availability in recovery of understorey vegetation after repeated disturbance? How are responses of soil micro‐organisms to disturbance and N‐fertilization reflected in nutrient allocation patterns and recovery of understorey vegetation after disturbance? Location: Subarctic mountain birch forest, Finland. Methods: We conducted a fully factorial experiment with annual treatments of disturbance (two levels) and N‐fertilization (four levels) during 1998–2002. We monitored treatment effects on above‐ground plant biomass, plant community structure and plant and soil nutrient concentrations. Results: Both disturbance and N‐fertilization increased the relative biomass of graminoids. The increase of relative biomass of graminoids in the disturbance treatment was over twice that of the highest N‐fertilization level, and N‐fertilization further increased their relative biomass after disturbance. As repeated disturbance broke the dominance of evergreen dwarf shrubs, it resulted in a situation where deciduous species, graminoids and herbs dominated the plant community. Although relative biomass of deciduous dwarf shrubs declined with N‐fertilization, it did not cause a shift in plant community structure, as evergreen dwarf shrubs remained dominant. Both disturbance and N‐fertilization increased the N concentration in vascular plants, whereas microbial biomass N and C were not affected by the treatments. Concentrations of NH₄⁺, dissolved organic N (DON) and dissolved organic C (DOC) increased in the soil after N‐fertilization, whereas concentrations of NH₄⁺ and DON decreased after disturbance. Conclusions: Disturbances caused by e.g. humans or herbivores contribute more to changes in the understorey vegetation structure than increased levels of N in subarctic vegetation. Fertilization accelerated the recovery potential after repeated disturbance in graminoids. Microbial activities did not limit plant growth.     

The effect of plant litter on ecosystem properties in a Mediterranean semi‐arid shrubland

Abstract. We studied the distribution of litter in a shrubland of the Negev with a semi‐arid Mediterranean climate of less than 200 mm of rainfall per year. Our focus was on the effects of litter on properties of landscape patches relevant to ecosystem processes (water runoff and soil erosion), annual plant community responses (seedling density, biomass production and species richness), and animal activity (soil disturbance by termites). Three 60‐m transects, extending across a pair of opposing north‐ and south‐facing slopes and their drainage channel, showed that litter accumulates not only under shrubs, but to a lesser extent also on the crusted inter‐shrub open areas. We used 35 experimental units (‘cells’, 0.5m × 1 m), each containing a crust and a shrub patch. Because runoff flows from crusted patches and is intercepted by shrub patches, the latter were in the lower third of the cells. Leaf litter was added in single and double amounts providing ca. 0.5 and 1.0 cm litter depth, to either, both, or none of the patches. Litter addition significantly decreased the amount of runoff, regardless of the location and amount of litter applied. Litter on the crust increased species number and seedling density of species with low abundance. Adding a double litter layer increased annual plant biomass production, while a single amount had no effect. Litter addition to the shrub patch affected neither biomass nor species richness. Litter addition to both patches at both quantities caused a large increase in termite activity. Termites caused disturbance by disrupting the crust, which may contribute to the reduction in runoff amounts. In the open, flat crust patches, annual plant communities are limited in their productivity and species richness, as there are few structures stopping the outflow of water, soil and seeds. Litter adds such structures, but affects the plant communities only when added to litter‐free crust. Litter accumulation and its patchy distribution have large impacts on landscape patch properties affecting resource distribution, plant productivity and diversity, and animal activity. Therefore, understanding litter distribution in relation to the patchy structure of the landscape of semi‐arid shrubland should be viewed as an important component of shrubland management.     

Responses of different herb life-history groups to a dominant shrub species along a dune stabilization gradient

Shrubs play a pivotal role in reversing desertification and in promoting ecological rehabilitation in severe environments. However, how the interactions between shrubs and their understorey species change during restoration remains unclear. Here, we report the results of an observational study conducted in the Mu Us Desert of northern China. This study explored how dune stabilization and the size of individual shrubs affect shrub–herb interactions. In particular, we aimed to determine how different life-history groups and performance indicators (e.g., biomass and richness) of subordinate species respond to shrub–herb interactions during dune stabilization. The shrub Artemisia ordosica had positive effects on understorey species in this dune system. The ability of the shrub to promote the growth of understorey species increased with dune stabilization, but decreased from small to large shrubs. This effect was due to an increase in the relative abundance and biomass of perennials and their higher sensitivity to the positive effects of the shrubs. In contrast, the ability of shrubs to improve the richness of understorey species decreased with dune stabilization, but increased from small to large shrubs. This effect occurred because perennials suffered high strain during recruitment in disturbed open patches of the semi-fixed dunes, particularly below small shrubs. Our results support the theory claiming that communities are sets of hidden interaction groups that have contrasting responses (neutral for annuals, but facilitative for perennials) to dominant neighbors in a single community, depending on their functional strategies. Furthermore, our study highlights the high efficiency of A. ordosica in facilitating ecological restoration of dry and disturbed sandy communities.     

The role of reproductive plant traits and biotic interactions in the dynamics of semi-arid plant communities

The dynamics of semi-arid plant communities are determined by the interplay between competition and facilitation among plants. The sign and strength of these biotic interactions depend on plant traits. However, the relationships between plant traits and biotic interactions, and the consequences for plant communities are still poorly understood. Our objective here was to investigate, with a modelling approach, the role of plant reproductive traits on biotic interactions, and the consequences for processes such as plant succession and invasion. The dynamics of two plant types were modelled with a spatially-explicit integrodifferential model: (1) a plant with seed dispersal (colonizer of bare soil) and (2) a plant with local vegetative propagation (local competitor). Both plant types were involved in facilitation due to a local positive feedback between vegetation biomass and soil water availability, which promoted establishment and growth. Plants in the system also competed for limited water. The efficiency in water acquisition (dependent on reproductive and growth plant traits) determined which plant type dominated the community at the steady state. Facilitative interactions between plant types also played an important role in the community dynamics, promoting establishment in the driest conditions and recovery from low biomass. Plants with vegetative propagation took advantage of the ability of seed dispersers to establish on bare soil from a low initial biomass. Seed dispersers were good invaders, maintained high biomass at intermediate and high rainfall and showed a high ability in taking profit from the positive feedback originated by plants with vegetative propagation under the driest conditions. However, seed dispersers lost competitiveness with an increasing investment in fecundity. All together, our results showed that reproductive plant traits can affect the balance between facilitative and competitive interactions. Understanding this effect of plant traits on biotic interactions provides insights in processes such as plant succession and shrub encroachment.     

Single introductions of soil biota and plants generate long‐term legacies in soil and plant community assembly

Recent demonstrations of the role of plant–soil biota interactions have challenged the conventional view that vegetation changes are mainly driven by changing abiotic conditions. However, while this concept has been validated under natural conditions, our understanding of the long‐term consequences of plant–soil interactions for above‐belowground community assembly is restricted to mathematical and conceptual model projections. Here, we demonstrate experimentally that one‐time additions of soil biota and plant seeds alter soil‐borne nematode and plant community composition in semi‐natural grassland for 20 years. Over time, aboveground and belowground community composition became increasingly correlated, suggesting an increasing connectedness of soil biota and plants. We conclude that the initial composition of not only plant communities, but also soil communities has a long‐lasting impact on the trajectory of community assembly.     

Soil pathogen communities associated with native and non‐native Phragmites australis populations in freshwater wetlands

Soil pathogens are believed to be major contributors to negative plant–soil feedbacks that regulate plant community dynamics and plant invasions. While the theoretical basis for pathogen regulation of plant communities is well established within the plant–soil feedback framework, direct experimental evidence for pathogen community responses to plants has been limited, often relying largely on indirect evidence based on above‐ground plant responses. As a result, specific soil pathogen responses accompanying above‐ground plant community dynamics are largely unknown. Here, we examine the oomycete pathogens in soils conditioned by established populations of native noninvasive and non‐native invasive haplotypes of Phragmites australis (European common reed). Our aim was to assess whether populations of invasive plants harbor unique communities of pathogens that differ from those associated with noninvasive populations and whether the distribution of taxa within these communities may help to explain invasive success. We compared the composition and abundance of pathogenic and saprobic oomycete species over a 2‐year period. Despite a diversity of oomycete taxa detected in soils from both native and non‐native populations, pathogen communities from both invaded and noninvaded soils were dominated by species of Pythium. Pathogen species that contributed the most to the differences observed between invaded and noninvaded soils were distributed between invaded and noninvaded soils. However, the specific taxa in invaded soils responsible for community differences were distinct from those in noninvaded soils that contributed to community differences. Our results indicate that, despite the phylogenetic relatedness of native and non‐native P. australis haplotypes, pathogen communities associated with the dominant non‐native haplotype are distinct from those of the rare native haplotype. Pathogen taxa that dominate either noninvaded or invaded soils suggest different potential mechanisms of invasion facilitation. These findings are consistent with the hypothesis that non‐native plant species that dominate landscapes may “cultivate” a different soil pathogen community to their rhizosphere than those of rarer native species.     

Plant‐driven changes in soil microbial communities influence seed germination through negative feedbacks

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Annual plant–shrub interactions along an aridity gradient

Annual plants in semi-arid and arid areas are often closely associated with shrubs. The degree of association largely depends on the balance of negative and positive effects between these contrasting plant life-forms, ranging from interference to facilitation. Since positive interactions are predicted to become less important with increasing rainfall, the interaction balance is expected to shift along aridity gradients. However, this prediction has not been tested on a community level and for different life-history stages across large geographical gradients. Here, we show such changes for annual plant populations and communities in four contrasting sites along a steep climatic gradient, ranging from the arid desert to mesic Mediterranean regions in Israel. Above-ground productivity, richness, seedling density, and seed bank density of the annual plant community, as well as fecundity of annual plant populations, were generally higher under shrubs than in areas between shrubs at the arid end of the gradient, but significantly lower at the humid end. Net effects of shrubs on annuals expressed as relative interaction intensity indicated a steady and consistent shift from net positive or neutral effects in the desert to net negative effects in the mesic part of the gradient. These findings emphasize the usefulness of studies along large-scale gradients encompassing a wide range of environmental conditions for understanding community level interactions among coexisting species.     

Experimental comparison of competition and facilitation in alpine communities varying in productivity

Question. Competitive and facilitative interactions among plant species in different abiotic environments potentially link productivity, vegetation structure, species composition and functional diversity. We investigated these interactions among four alpine communities along an environmental productivity gradient in a generally harsh climate. We hypothesised that the importance of competition would be higher in more productive sites. Location. Mt. M. Khatipara (43°27′N, 41°41′E, altitude 2750 m), NW Caucasus, Russia. Communities ranged from low‐productivity alpine lichen heath (ALH) and snowbed communities (SBC), to intermediate productivity Festuca grassland (FVG), and high‐productivity Geranium‐Hedysarum meadow (GHM). Methods. We quantified the relative influence of competition and facilitation on community structure by expressing biomass of target species within each natural community proportionally to biomass of the species in a “null community” with experimental release from interspecific competition by removing all other species (for 6 years). An overall index of change in community composition due to interspecific interactions was calculated as the sum of absolute or proportional differences of the component species. Results. Species responses to neighbour removal ranged from positive to neutral. There was no evidence of facilitation among the selected dominant species. As expected, competition was generally most important in the most productive alpine community (GHM). The intermediate position for low‐productivity communities of stressful environments (ALH, SBC) and the last position of intermediately productive FVG were unexpected. Conclusions. Our results appear to support the Fretwell‐Oksanen hypothesis in that competition in communities of intermediate productivity was less intense than in low‐ or high‐productive communities. However, the zero net effect of competition and facilitation in FVG might be the result of abiotic stress due to strong sun exposure and high soil temperatures after neighbour removal. Thus, non‐linear relationships between soil fertility, productivity and different abiotic stresses may also determine the balance between competition and facilitation.     

Grazing‐induced changes in plant–soil feedback alter plant biomass allocation

Large vertebrate herbivores, as well as plant–soil feedback interactions are important drivers of plant performance, plant community composition and vegetation dynamics in terrestrial ecosystems. However, it is poorly understood whether and how large vertebrate herbivores and plant–soil feedback effects interact. Here, we study the response of grassland plant species to grazing‐induced legacy effects in the soil and we explore whether these plant responses can help us to understand long‐term vegetation dynamics in the field. In a greenhouse experiment we tested the response of four grassland plant species, Agrostis capillaris, Festuca rubra, Holcus lanatus and Rumex acetosa, to field‐conditioned soils from grazed and ungrazed grassland. We relate these responses to long‐term vegetation data from a grassland exclosure experiment in the field. In the greenhouse experiment, we found that total biomass production and biomass allocation to roots was higher in soils from grazed than from ungrazed plots. There were only few relationships between plant production in the greenhouse and the abundance of conspecifics in the field. Spatiotemporal patterns in plant community composition were more stable in grazed than ungrazed grassland plots, but were not related to plant–soil feedbacks effects and biomass allocation patterns. We conclude that grazing‐induced soil legacy effects mainly influenced plant biomass allocation patterns, but could not explain altered vegetation dynamics in grazed grasslands. Consequently, the direct effects of grazing on plant community composition (e.g. through modifying light competition or differences in grazing tolerance) appear to overrule indirect effects through changes in plant–soil feedback.