Asymmetric coexistence: bidirectional abiotic and biotic effects between goose barnacles and mussels

1. Species coexistence depends on the net effect of interacting species, representing the sum of multiple interaction components that may act simultaneously and vary independently depending on ambient environmental conditions. Consequently, for a comprehensive understanding of the compound nature of species interactions and coexistence, a mechanistic approach that allows a separate evaluation of each interaction component is required. 2. Two sessile filter-feeders, the goose barnacle Capitulum mitella and the mussel Septifer virgatus, coexist on moderately wave-exposed rocky shores in south-western Japan. In the upper intertidal, Capitulum positively influenced Septifer survivorship and growth through amelioration of thermal stress and of physical disturbance. On the other hand, these species are potential competitors as they have similar body sizes and modes of resource utilization. These opposite processes, facilitation and competition, are based on abiotic characteristics and biotic functions of the two species, respectively. 3. In order to quantify the bidirectional abiotic, biotic and net effects, a series of experimental manipulations was conducted involving the use of living neighbours with both abiotic and biotic effects, and artificial mimics to simulate abiotic effects without biotic effects. 4. Capitulum had strong positive abiotic effects on the mussel survivorship in most experimental periods, while the biotic effect was negligible or weakly negative, suggesting that the net effect of Capitulum on mussel survival was largely attributable to the abiotic effect. In contrast, a significantly negative biotic effect on the mussel growth rate was always present, though this was cancelled out by the larger, positive abiotic effect. In the case of Septifer, its abiotic and biotic effects on the survivorship of goose barnacles were negligible, while those on the growth rate showed temporal variation. 5. With respect to the relationship between species interaction and environmental conditions, the strength of abiotic facilitative effect of Capitulum on mussel survival increased with increasing abiotic stress, while the strength of biotic effect was negligible or weakly negative. As regards the effects of mussels on goose barnacles, our study indicated no obvious relationship. These results point to the importance of decomposing interaction for an accurate, mechanistic understanding of species relations and coexistence.     

Temperature‐dependent shifts in herbivore performance and interactions drive nonlinear changes in crop damages

Understanding how and to what extent the influence of temperature on physiological performance scales up to interspecific interactions and process rate patterns remains a major scientific challenge faced by ecologists. Here, we combined approaches developed by two conceptual frameworks in ecology, the stress‐gradient hypothesis (SGH), and the biodiversity–ecosystem functioning relationship (B‐EF), to test the hypothesis that interspecific difference in thermal performance modulates multiple species interactions along a thermal stress (SGH) and the subsequent richness effects on process rates (B‐EF). We designed an experiment using three species of herbivorous agricultural pests with different thermal optima for which we determined how temperature influences the direction and the strength of interaction and subsequent richness effects on crop damage (7 species interaction treatments × 6 temperature treatments × 10 replicates). We showed that both biotic interactions and species richness effects drive variations in crop damages along a thermal stress gradient, and thus have the potential to drive agro‐system responses to climate change. To help explain and generalize underlying mechanisms of richness effects on process rates, we further proposed a conceptual model that views interaction outcomes as shifting between positive and negative along a thermal stress depending on species thermal optima. Overall, our study demonstrates that nonlinear effects of temperature on process rates must be a major concern in terms of prediction and management of the consequences of global warming.     

The interplay of stress and mowing disturbance for the intensity and importance of plant interactions in dry calcareous grasslands

Background and Aims There is still debate regarding the direction and strength of plant interactions under intermediate to high levels of stress. Furthermore, little is known on how disturbance may interact with physical stress in unproductive environments, although recent theory and models have shown that this interplay may induce a collapse of plant interactions and diversity. The few studies assessing such questions have considered the intensity of biotic interactions but not their importance, although this latter concept has been shown to be very useful for understanding the role of interactions in plant communities. The objective of this study was to assess the interplay between stress and disturbance for plant interactions in dry calcareous grasslands. Methods A field experiment was set up in the Dordogne, southern France, where the importance and intensity of biotic interactions undergone by four species were measured along a water stress gradient, and with and without mowing disturbance. Key Results The importance and intensity of interactions varied in a very similar way along treatments. Under undisturbed conditions, plant interactions switched from competition to neutral with increasing water stress for three of the four species, whereas the fourth species was not subject to any significant biotic interaction along the gradient. Responses to disturbance were more species-specific; for two species, competition disappeared with mowing in the wettest conditions, whereas for the two other species, competition switched to facilitation with mowing. Finally, there were no significant interactions for any species in the disturbed and driest conditions. Conclusions At very high levels of stress, plant performances become too weak to allow either competition or facilitation and disturbance may accelerate the collapse of interactions in dry conditions. The results suggest that the importance and direction of interactions are more likely to be positively related in stressful environments.     

Mechanisms of succession along the emersion gradient in intertidal rocky shore assemblages

Theoretical models predict that the relative importance of competition and facilitation vary inversely along gradients of abiotic stress, with facilitation dominating under harsh conditions (the so called "stress-gradient hypothesis"). To date, very few studies have tested this hypothesis in the framework of succession. Moreover, the generality of the hypothesis is currently under debate and the necessity to examine responses at the community level and using different stress levels has been emphasized. We evaluate the mechanisms of succession across the emersion gradient in a rocky shore system. After complete clearing of experimental plots at four shore heights, we removed two separate components of the assemblage: ephemeral green algae, early colonists and canopy-forming species, main space holders in undisturbed assemblages. We aimed to test two hypotheses: (a) the net effect of ephemeral algae on canopy-forming species shift along the emersion gradient according to the predictions of the stress-gradient hypothesis and (b) the net effect of the canopy-forming group on associate species change, from neutral/negative to positive, as succession progresses and plants grow, especially at the highest shore heights. Our results did not support the predictions of the stress gradient hypothesis in the framework of succession. We did detect changes through time in the effect of canopy-forming species, from negative to positive, but this was not dependent on the shore height. The outcome of interactions depended on the identity and life-stage of the species. Moreover, indirect interactions among species could also create a less predictable relationship between successional mechanisms and the environmental gradient.     

Canopy interactions and physical stress gradients in subtidal communities

Species interactions are integral drivers of community structure and can change from competitive to facilitative with increasing environmental stress. In subtidal marine ecosystems, however, interactions along physical stress gradients have seldom been tested. We observed seaweed canopy interactions across depth and latitudinal gradients to test whether light and temperature stress structured interaction patterns. We also quantified interspecific and intraspecific interactions among nine subtidal canopy seaweed species across three continents to examine the general nature of interactions in subtidal systems under low consumer pressure. We reveal that positive and neutral interactions are widespread throughout global seaweed communities and the nature of interactions can change from competitive to facilitative with increasing light stress in shallow marine systems. These findings provide support for the stress gradient hypothesis within subtidal seaweed communities and highlight the importance of canopy interactions for the maintenance of subtidal marine habitats experiencing environmental stress.     

What does the stress‐gradient hypothesis predict? Resolving the discrepancies

In recent years the importance of facilitative interactions in ecological communities is increasingly recognized. This phenomenon has been observed repeatedly, particularly in vegetation communities, in a wide range of environmental conditions. The current hypothesis predicts that the role of facilitation becomes increasingly important in conjunction with increasing stress. Several empirical studies, however, failed to detect such patterns, particularly at the extreme ends of the stress gradients. Herein, we present a conceptual model that may resolve discrepancies between expected and observed and provides a more precise framework of the existing hypotheses. By relaxing two common assumptions commonly used by the stress‐gradient hypothesis (SGH) we are able to demonstrate that under some circumstances the importance of facilitation may be less at the extreme ends of these gradients. Namely, we first re‐emphasize the notion that physiological response is not linear with respect to environmental changes along stress gradients. Second, it is argued that the net outcome of facilitative and competitive interactions is reflected in the fitness of individuals as a product of these two processes, in contrast to the commonly applied assumption of additivity. Accordingly, a synthesis of the concepts of population biology (measures of fitness) and plant physiology (nonlinear responses) with the stress gradient hypothesis while retaining the original simplicity of the SGH model contributes to a better specification of the predictions of the stress‐gradient hypothesis and the resolution of observed contradictions.     

Interaction intensity and importance along two stress gradients: adding shape to the stress-gradient hypothesis

The stress-gradient hypothesis (SGH) predicts that the community-wide prevalence of positive interactions, relative to negative interactions, is greater under more severe environmental conditions. Because the frequency of positive and negative interactions within a community is the aggregate of multiple pair-wise interactions, one approach to testing the SGH is to examine how pair-wise interactions vary along severity gradients. While the SGH suggests that the net outcome of an interaction should monotonically become more positive with increasing environmental severity, recent studies have suggested that the severity-interaction relationship (SIR) may rather be unimodal. We tested which of the proposed shapes of the SIR best fits the variation in the interaction between two species along two types of severity gradients on sub-Antarctic Marion Island. This was done by comparing the performance of the grass Agrostis magellanica in the presence and absence of the cushion plant Azorella selago, along both species’ entire altitudinal ranges (transects spanning 4–8 km), and along a shorter (transect = 0.4 km) wind exposure gradient. Along the altitudinal transects the relative intensity, but not the absolute intensity or the importance, of the Azorella selago–Agrostis magellanica interaction increased with altitude, consistently forming a plateau-shaped SIR with a positive asymptote. Thus, while the performance of Agrostis magellanica was negatively affected by Azorella selago at low altitudes, the grass benefited from growing on the cushion plant under greater environmental severity. Along the wind exposure gradient the intensity of the interaction also became more positive with increasing environmental severity for most performance measures. This suggests that the switch from a net negative to a net positive interaction can occur across both short and long distances. Therefore, this study provides strong evidence for a plateau-shaped SIR, and confirms that the SIR is unimodal along the particular non-resource severity gradients of this study.     

The combined effects of energy and disturbance on species richness in protist microcosms

The supply of energy, and the frequency of disturbance can both affect species diversity, often, although not always, producing unimodal diversity patterns along an energy or disturbance gradient. However, it has been suggested that diversity is a combined function of both factors, and the dynamic equilibrium model proposed by Huston (1979) is one formalization of this suggestion. This idea has received little direct testing. Here we carry out such a test using protist microcosms, in a factorial manipulation of six levels of energy (quantity of organic resource) and five levels of disturbance (frequency of temperature shock). Species richness responded to both energy supply and disturbance frequency, although not always unimodally. Patterns of response changed over time, and there was some evidence of an interaction between energy and disturbance at two of six time intervals. However, the specific form of this interaction differs from that predicted by the dynamic equilibrium model, and overall, the results provide little support for the model.     

Variation in the effects of vegetation and litter on recruitment across productivity gradients

1 We tested predictions about how the effect of vegetation and litter on seedling establishment varies among sites and herbaceous community types (sand barrens, prairies, fens). For both vegetation and litter, we also separated direct interactions from indirect interactions and interaction modifications along the gradient.
2 Although the intensity of the effects varied across sites, the direct effects of vegetation or litter alone were consistently facilitative along the productivity gradient. Predominance of facilitative effects may be due to the focus on the seedling establishment phase.
3 However, inclusion of indirect interactions and interaction modifications caused the net effects of both vegetation and litter to become largely negative. While one layer of biomass may be advantageous to ameliorate some moisture stress, the addition of another layer may be disadvantageous if this layer limits light proportionally more than it relieves moisture stress.
4 One exception to this pattern occurred at high productivity when the net effect of vegetation, even in the presence of litter, remained facilitative. The net effect of vegetation was competitive at low productivity and grew increasingly facilitative with productivity. Thus, indirect effects of litter may alter interaction patterns across this gradient.     

The relative importance of disturbance and environmental stress at local and regional scales in French coastal sand dunes

Questions: 1. Is there a primary role of disturbance at local scale and of environmental stress at regional scale? 2. Does disturbance increase or decrease environmental stress at local scale? Location: The Atlantic coastal dune system of the Aquitaine Region (France). Methods: Species biomass and 16 environmental variables were sampled in 128 quadrats along a local beach‐inland gradient and a regional North‐South gradient. Environmental data were analysed with ANOVAs and vegetation‐environment relationships with Canonical Correspondence Analysis. Results: At the local scale community composition was primarily driven by disturbance due to sand burial, whereas water and nutrient stress better explained regional differences. However, random biogeographical events are very likely to also affect community composition at the largest scale. The main interaction between environmental stress and disturbance was the mitigation of nutrient stress induced by disturbance at a local scale. This was due to a positive direct effect of sand burial and a positive indirect effect of wind (decrease in VPD by ocean spray). Although wind had also a significant effect on soil conductivity and pH, there was no evidence that these factors had any role in community composition. Conclusions: Our results support the hypothesis that disturbance had a primary role at local scale and environmental stress at regional scale but further research is needed to separate the effect of stress from that of dispersal at regional scale. We also demonstrated that environmental stress in primary succession may not always decline with decreasing disturbance.     

A dynamic model of facilitation on environmental stress gradients

Theories based on competition for resources in animals and other non‐sessile organisms rarely consider the role of facilitative interactions. Yet these interactions are important for community assembly, especially under stressful environments (e.g. the stress‐gradient hypothesis, SGH). To make an explicit link between species interaction theory and SGH patterns, I used a classic resource competition model promoting coexistence between a beneficiary and its facilitator sharing a common resource along a stress gradient. I compared model outcomes for two fundamentally different mechanisms of facilitation (alleviation of resource versus non‐resource stress), and also tested the effect of a reciprocal cost of facilitation from the beneficiary. I then tested model's biological relevance using experimental data from two tuber moth species (Lepidoptera, Gelechiidae) for which facilitation in resource access was previously established. Simulation outcomes revealed that both the mode of facilitation and the incorporation of facilitation costs affected the shape of the facilitation–stress relationship. These predictions are in line with current SGH observations and experiments on both plants and animals and reconcile the frequently reported variability of this relationship in nature. Moreover, a sensitivity analysis of model's parameters confirmed the robustness of the modelling framework to uncover the mechanisms responsible for observed species interaction–stress patterns. Finally, when parameterized with tuber moth demographic data, model's results corresponded to observed interaction outcomes along resource stress gradients. Overall, having a common model for plants and animals may simplify assumptions in SGH studies, allow contrasting the shapes of different consumer–resource relationships and specifying the conditions that favour one type of interaction outcome over another.     

Thermal Carrying Capacity for a Thermally-Sensitive Species at the Warmest Edge of Its Range

Anthropogenic environmental change is causing unprecedented rates of population extirpation and altering the setting of range limits for many species. Significant population declines may occur however before any reduction in range is observed. Determining and modelling the factors driving population size and trends is consequently critical to predict trajectories of change and future extinction risk. We tracked during 12 years 51 populations of a cold-water fish species (brown trout Salmo trutta) living along a temperature gradient at the warmest thermal edge of its range. We developed a carrying capacity model in which maximum population size is limited by physical habitat conditions and regulated through territoriality. We first tested whether population numbers were driven by carrying capacity dynamics and then targeted on establishing (1) the temperature thresholds beyond which population numbers switch from being physical habitat- to temperature-limited; and (2) the rate at which carrying capacity declines with temperature within limiting thermal ranges. Carrying capacity along with emergent density-dependent responses explained up to 76% of spatio-temporal density variability of juveniles and adults but only 50% of young-of-the-year''s. By contrast, young-of-the-year trout were highly sensitive to thermal conditions, their performance declining with temperature at a higher rate than older life stages, and disruptions being triggered at lower temperature thresholds. Results suggest that limiting temperature effects were progressively stronger with increasing anthropogenic disturbance. There was however a critical threshold, matching the incipient thermal limit for survival, beyond which realized density was always below potential numbers irrespective of disturbance intensity. We additionally found a lower threshold, matching the thermal limit for feeding, beyond which even unaltered populations declined. We predict that most of our study populations may become extinct by 2100, depicting the gloomy fate of thermally-sensitive species occurring at thermal range margins under limited potential for adaptation and dispersal.     

Spatial variation in plant interactions across a severity gradient in the sub-Antarctic

The stress–gradient hypothesis predicts that the intensity of interspecific positive interactions increases along environmental severity (i.e. stress and disturbance) gradients faster than the intensity of negative interactions. This study is the first to test if the stress–gradient hypothesis is supported for a location in the climatically extreme and species-poor sub-Antarctic. To do so, we investigate the fine-scale spatial distribution of plant species across altitude- and aspect-related abiotic severity gradients on a scoria cone on Marion Island. A clear altitudinal severity gradient was observed across the scoria cone, with lower temperatures, stronger winds and greater soil movement at higher altitudes. The altitudinal severity gradient was matched by stronger interspecific spatial association between the four dominant species at higher altitudes and in areas of lower vegetation cover. This suggests that, relative to the intensity of competition, the intensity of facilitation is greater under more severe conditions, supporting the stress–gradient hypothesis at the community level (i.e. for multiple pairs of species) and corroborating its usefulness for predicting variation in plant interactions at high latitudes and altitudes. Furthermore, the directional intraspecific aggregation and interspecific association plant cover patterns found within the gradient suggest that protection from the prevailing wind and from burial by loose substrate are the dominant facilitative mechanisms. Thus, plants benefit from the presence of neighbours when they provide shelter and substrate stability, and the relative intensity of this positive interaction is greatest at higher altitudes, and varies between species pairs. This study, therefore, not only provides support for the stress–gradient hypothesis in the sub-Antarctic, but also demonstrates fine-scale directional spatial patterns between multiple species nested within the severity gradient. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Does disturbance drive the collapse of biotic interactions at the severe end of a diversity–biomass gradient?

It has been recently proposed that the decrease in diversity towards the severe end of the humped-back diversity–biomass model of Grime was driven by a collapse of facilitation due to extreme conditions of either stress or physical disturbance. In order to test the hypothesis that disturbance is the primary direct factor driving the collapse of interactions occurring along environmental severity gradients, we conducted a removal experiment in the highly stressed French coastal dunes along a gradient of disturbance due to sand burial. Four dune species were used as targets and transplanted with and without neighbours in four communities along the gradient. The experiment was conducted twice, a dry and an average year. Results of the experiment showed that during the dry year the effect of the environment was prominent and only one species was facilitated for survival in the least disturbed community. During the average year, interactions for growth were important only in the same community, with positive or negative responses depending on the natural position of the target species within the coastal dune gradient. In accordance with our hypothesis, most interactions for both survival and growth were observed in the least disturbed community exhibiting the highest diversity. There were no interactions in the most disturbed community with the lowest diversity.     

Response of wetland herbaceous communities to gradients of light and substrate following disturbance by thermal pollution

The influence of thermal disturbance and site characteristics on distribution of herbs was studied in portions of a 3020 ha wetland in the southeastern USA. Presence-absence of 52 species in 130 0.25 m² plots was determined from four sites with different disturbance histories and from an undisturbed site. Data from the four disturbed sites were ordinated by detrended correspondence analysis. Differences in species composition among sites (coarse scale) were associated with water depth, light, and substrate type. Within a site (at a fine scale), correlation of environmental variables with ordination scores at a chronically disturbed site was weakly correlated with light (r=0.50). At two sites with episodic disturbance, species composition correlated significantly and positively with substrate and water depth. At a recovering site, vegetation patterns were moderately correlated with water depth (r=?0.52). Species richness was correlated with substrate type along the disturbance gradient. Our results are consistent the intermediate disturbance hypothesis and the subsidy-stress gradient concept.     

Plant species responses along a grazing disturbance gradient in Australian grassland

Abstract. Floristic data from paired roadside-paddock analyses from grassland in central Queensland, Australia, were ordinated. The mean direction of the vectors between these pairs was almost perfectly aligned with the indirect gradient represented by the first axis of Non-metric Multi-dimensional Scaling. It confirms anecdotal evidence of a trend from infrequently grazed roadsides to constantly grazed paddocks. The increasing abundance of annual herbs and grasses along this putative gradient is consistent with documented trends from elsewhere in the world. The response patterns of individual species along the disturbance gradient is consistent with ecological theory predicting unimodal peaks in abundance along physical environmental gradients. The ancestral perennial dominants of the grasslands, Dichanthium sericeum and D. queenslandicum, exhibited a declining response to grazing disturbance. Even the generally unpalatable perennial grass Aristida leptopoda declined considerably in the upper segments of the grazing disturbance gradient. A suite of herbaceous trailing legumes had peaks in their abundance near the middle of the grazing disturbance gradient, trends that can be readily explained by the combination of their palatability and intolerance to competition from tall perennial grasses. Several species including the noxious exotic herb Parthenium hysterophorus showed increasing abundance along the grazing disturbance gradient. The methodology may have application as a rapid method of assessing disturbance impacts elsewhere, and is most suited where a management differential between paired plots can be reliably generalized and where the physical environment is relatively monotonous.     

Using trait and phylogenetic diversity to evaluate the generality of the stress‐dominance hypothesis in eastern North American tree communities

The stress‐dominance hypothesis (SDH) is a model of community assembly predicting that the relative importance of environmental filtering increases and competition decreases along a gradient of increasing environmental stress. Tests of the SDH at limited spatial scales have thus far demonstrated equivocal support and no prior study has assessed the generality of the SDH at continental scales. We examined over 53 000 tree communities spanning the eastern United States to determine whether functional trait variation and phylogenetic diversity support the SDH for gradients of water and soil nutrient availability. This analysis incorporated two complementary datasets, those of the U.S. Forest Service Forest Inventory and Analysis National program and the Carolina Vegetation Survey, and was based on three ecologically important traits: leaf nitrogen, seed mass, and wood density. We found that mean trait values were weakly correlated with water and soil nutrient availability, but that trait diversity did not vary consistently along either gradient. This did not conform to trait variation expected under the SDH and instead suggested that environmental filters structure tree communities throughout both gradients, without evidence for an increased role of competition in less stressful environments. Phylogenetic diversity of communities was principally driven by the ratio of angiosperms to gymnosperms and therefore did not exhibit the pattern of variation along stress gradients expected under the SDH. We conclude that the SDH is not a general paradigm for all eastern North American tree communities, although it may operate in certain contexts.     

Abundance, diversity and fidelity of macroinvertebrates sheltering beneath rocks during tidal emersion in an intertidal cobble field: Does the intermediate disturbance hypothesis hold for less exposed shores with smaller rocks?

The community structure of macroinvertebrates occurring beneath cobbles during tidal emersion was investigated in an intertidal cobble field on the east coast of the south island of New Zealand. In order to test whether the intermediate disturbance hypothesis (IDH) holds in less exposed shores comprised exclusively of smaller rocks (cobbles) we measured 1) the size frequency distribution of cobbles (mass and basal surface area), 2) the incidence of barnacle encrustation as an indirect measure of cobble disturbance, 3) the abundances of the dominant macroinvertebrates occurring beneath cobbles (the crab Hemigrapsus crenulatus, the chiton Sypharochiton pelliserpentis and the snail Diloma zealandica), and 4) macroinvertebrate fidelity beneath a range of cobble sizes. Only large cobbles were found to be encrusted with barnacles, indicative of their greater resistance to physical disturbance by waves and currents. Mark and relocate experiments indicated fidelity to a given cobble over two tidal cycles was 11-14% for crabs and did not vary with cobble size. In contrast, fidelity to medium and large cobbles was higher for chitons (26-45%) and snails (17-31%) and generally increased with increasing cobble size. In contrast to the predictions of the IDH, there was a positive correlation between increasing cobble size and increasing species diversity of organisms found under the cobbles. This pattern is not simply attributable to greater space available beneath large cobbles as there was a significantly greater density of macroinvertebrates beneath medium and large, compared to the small cobbles. Increasing species diversity with increasing cobble size appears to be independent of patterns of species recolonization associated with small cobbles being subject to high disturbance or interspecific competitive hierarchies associated with low disturbance beneath large cobbles. Instead, factors such as aggregative behaviors, and reducing vulnerability to predators and desiccation stress are likely to explain the relationship between cobbles and their associated macroinvertebrate communities.     

Fine nurse variations explain discrepancies in the stress‐interaction relationship in alpine regions

Despite a large consensus on increasing facilitation among plants with increasing stress in alpine regions, a number of different outcomes of interaction have been observed, which impedes the generalisation of the ‘stress‐gradient hypothesis’ (SGH). With the aim to reconcile the different viewpoints on the stress‐interaction relationship in alpine environments we hypothesized that fine nurse variations within a single life form (cushion) may explain this pattern variability. To test this hypothesis, we compared the magnitude of the stress‐interaction relationship in a single study area with that observed in existing studies involving cushions, worldwide. We characterized the nurse effects of cushions on the whole plant community at inter‐specific, intra‐specific and intra‐individual levels along a stress gradient in the dry, alpine tropics of Bolivia (4400 m, 4700 m and 4900 m a.s.l). Using a relative index of interaction (RII) we included our data in a meta‐analysis on the nurse effects of cushions along alpine gradients, worldwide. At inter‐specific level, the loose cushion Pycnophyllum was a better nurse than the compact Azorella compacta. However, at intra‐individual level facilitation was higher at the periphery than at the centre of cushions, exceeding in magnitude the variation observed at inter‐specific level. This pattern was associated with higher minimum temperature and lower mortality at the periphery of cushions. The net effects of cushions on plant communities became more positive at higher elevation, corroborating the SGH. Within our single site in Bolivia, fine morphological nurse variations captured a similar variability in the stress‐interaction relationship as that observed in a subset of studies on cushions on a worldwide scale. This suggests that fine variations in nurse traits, in general those not considered in protocols dealing with facilitation or in restoration/conservation management plans, explain in part the current discrepancies among SGH studies in alpine regions.