Summary 1.Once neglected, the role of facilitative interactions in plant communities has received considerable attention in the last two decades, and is now widely recognized. It is timely to consider the progress made by research in this field. 2. We review the development of plant facilitation research, focusing on the history of the field, the relationship between plant-plant interactions and environmental severity gradients, and attempts to integrate facilitation into mainstream ecological theory. We then consider future directions for facilitation research. 3. With respect to our fundamental understanding of plant facilitation, clarification of the relationship between interactions and environmental gradients is central for further progress, and necessitates the design and implementation of experiments that move beyond the clear limitations of previous studies. 4. There is substantial scope for exploring indirect facilitative effects in plant communities, including their impacts on diversity and evolution, and future studies should connect the degree of non-transitivity in plant competitive networks to community diversity and facilitative promotion of species coexistence, and explore how the role of indirect facilitation varies with environmental severity. 5. Certain ecological modelling approaches (e.g. individual-based modelling), although thus far largely neglected, provide highly useful tools for exploring these fundamental processes. 6. Evolutionary responses might result from facilitative interactions, and consideration of facilitation might lead to re-assessment of the evolution of plant growth forms. 7. Improved understanding of facilitation processes has direct relevance for the development of tools for ecosystem restoration, and for improving our understanding of the response of plant species and communities to environmental change drivers. 8. Attempts to apply our developing ecological knowledge would benefit from explicit recognition of the potential role of facilitative plant-plant interactions in the design and interpretation of studies from the fields of restoration and global change ecology. 9. Synthesis: Plant facilitation research provides new insights into classic ecological theory and pressing environmental issues. Awareness and understanding of facilitation should be part of the basic ecological knowledge of all plant ecologists.

Interactions among species determine local-scale diversity, but local interactions are thought to have minor effects at larger scales. However, quantitative comparisons of the importance of biotic interactions relative to other drivers are rarely made at larger scales. Using a data set spanning 78 sites and five continents, we assessed the relative importance of biotic interactions and climate in determining plant diversity in alpine ecosystems dominated by nurse-plant cushion species. Climate variables related with water balance showed the highest correlation with richness at the global scale. Strikingly, although the effect of cushion species on diversity was lower than that of climate, its contribution was still substantial. In particular, cushion species enhanced species richness more in systems with inherently impoverished local diversity. Nurse species appear to act as a 'safety net' sustaining diversity under harsh conditions, demonstrating that climate and species interactions should be integrated when predicting future biodiversity effects of climate change.

Biotic interactions can shape phylogenetic community structure (PCS). However, we do not know how the asymmetric effects of foundation species on communities extend to effects on PCS. We assessed PCS of alpine plant communities around the world, both within cushion plant foundation species and adjacent open ground, and compared the effects of foundation species and climate on alpha (within-microsite), beta (between open and cushion) and gamma (open and cushion combined) PCS. In the open, alpha PCS shifted from highly related to distantly related with increasing potential productivity. However, we found no relationship between gamma PCS and climate, due to divergence in phylogenetic composition between cushion and open sub-communities in severe environments, as demonstrated by increasing phylo-beta diversity. Thus, foundation species functioned as micro-refugia by facilitating less stress-tolerant lineages in severe environments, erasing a global productivity - phylogenetic diversity relationship that would go undetected without accounting for this important biotic interaction.

SummaryFacilitative interactions are defined as positive effects of one species on another, but bidirectional feedbacks may be positive, neutral, or negative. Understanding the bidirectional nature of these interactions is a fundamental prerequisite for the assessment of the potential evolutionary consequences of facilitation.In a global study combining observational and experimental approaches, we quantified the impact of the cover and richness of species associated with alpine cushion plants on reproductive traits of the benefactor cushions.We found a decline in cushion seed production with increasing cover of cushion-associated species, indicating that being a benefactor came at an overall cost. The effect of cushion-associated species was negative for flower density and seed set of cushions, but not for fruit set and seed quality. Richness of cushion-associated species had positive effects on seed density and modulated the effects of their abundance on flower density and fruit set, indicating that the costs and benefits of harboring associated species depend on the composition of the plant assemblage.Our study demonstrates 'parasitic' interactions among plants over a wide range of species and environments in alpine systems, and we consider their implications for the possible selective effects of interactions between benefactor and beneficiary species.

While there is a large consensus that plant–plant interactions are a crucial component of the response of plant communities to the effects of climate change, available data remain scarce, particularly in alpine systems. This represents an important obstacle to making consistent predictions about the future of plant communities. Here, we review current knowledge on the effects of climate change on facilitation among alpine plant communities and propose directions for future research. In established alpine communities, while warming seemingly generates a net facilitation release, earlier snowmelt may increase facilitation. Some nurse plants are able to buffer microenvironmental changes in the long term and may ensure the persistence of other alpine plants through local migration events. For communities migrating to higher elevations, facilitation should play an important role in their reorganization because of the harsher environmental conditions. In particular, the absence of efficient nurse plants might slow down upward migration, possibly generating chains of extinction. Facilitation–climate change relationships are expected to shift along latitudinal gradients because (1) the magnitude of warming is predicted to vary along these gradients, and (2) alpine environments are significantly different at low vs. high latitudes. Data on these expected patterns are preliminary and thus need to be tested with further studies on facilitation among plants in alpine environments that have thus far not been considered. From a methodological standpoint, future studies will benefit from the spatial representation of the microclimatic environment of plants to predict their response to climate change. Moreover, the acquisition of long-term data on the dynamics of plant–plant interactions, either through permanent plots or chronosequences of glacial recession, may represent powerful approaches to clarify the relationship between plant interactions and climate change.

International audienceQuestions : Does facilitation among plants increase with elevation in a humid tropical alpine system in which climatic and ecological conditions differ from other alpine environments? What mechanisms are involved in the interactions? Location : Volcano Antisana, Eastern Cordillera of the Ecuadorian Andes (00°28′S, 78°09′W). Methods : We selected the cushion-forming Azorella aretioides as a potential nurse plant along an altitudinal gradient (4400, 4550 and 4700 m) in the high tropical Andes. We quantified its effects on other plants at species and community levels by comparing the product vegetation cover × number of individuals of every vascular species found inside and outside 265 cushions, using the relative interaction index. We inferred potential mechanisms behind the interactions through analysis of microclimate, soil moisture and soil nutrient measurements inside and outside cushions. Results : Predictions of the stress gradient hypothesis (SGH) were corroborated at community level, with transition from competitive or neutral effects of A. aretioides at 4400 and 4550 m to facilitative effects at 4700 m. Strong species-specific effects were observed along the altitudinal gradient, with a substantial effect of local habitat disturbance on the outcome of plant-plant interactions. Surprisingly, cushions lowered air and soil temperatures and air humidity, which reduced at higher elevations. Facilitation appeared to be caused by higher soil moisture and nutrient content beneath cushions. Conclusions : Our data extend the framework of the SGH by corroborating it for the first time in a tropical alpine system. However, the mechanisms underlying plant-plant interactions differed from those generally reported from alpine environments, with facilitation varying according to resource-mediated stress (nutrients). It remains to be tested whether this is specific to tropical alpine systems

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Summary1. The stress-gradient hypothesis (SGH) states that environmental stress modulates species interactions, causing a shift from negative interactions to net positive interactions with increasing stress. 2. Potentially, this modulation of species interactions could in turn influence biodiversity-ecosystem function (B-EF) relationships along stress gradients. Although the SGH has been extensively discussed in plant community ecology in the past two decades, it has received little attention from animal ecologists. 3. To explore whether the SGH could be applied to animal communities, we conducted a litter decomposition experiment with aquatic detritivorous invertebrates in which we manipulated litter quality and measured species interactions along this resource quality gradient. Litter quality was manipulated by presenting detritivores with leaves of plant species varying in specific leaf area and decomposition rate in streams. 4. We found a switch from negative to neutral interactions with increasing resource quality stress, in line with the SGH. However, by re-examining other published results with aquatic detritivores from the perspective of the SGH, we found that a diversity of patterns seem to characterize detritivore interactions along stress gradients. 5. Although the basic pattern proposed by the SGH may not apply to animal systems in general, we show that aquatic detritivore interactions do change along stress gradients, which underlines the importance of incorporating environmental stressors more explicitly in B-EF research.