Relative contribution of canopy and soil effects between plants with different metal tolerance along a metal pollution gradient.

Multiple effects, operating either on the long-term (soil-engineering effects) or on the short-term during plant life (microclimate modification or resources pre-emption), can act simultaneously and determine the outcome of plant-plant interactions. These diverse effects have not been disentangled along a gradient of metal/metalloid pollution, although this is crucial for understanding the dominant species turnover along the gradient, and thus the driving processes of facilitation recurrently found in metalliferous ecosystems, which could help improving ecological restoration of these degraded ecosystems. Here, we experimentally assessed different short-term effects of two dominant forbs of highly polluted habitats (Hutchinsia alpina and Arenaria multicaulis, tolerant to metal stress) and two grasses of less polluted habitats (Agrostis capillaris and Festuca rubra, less tolerant to metal stress) on target plant species (the same as the dominant species mentioned above) transplanted along a large metal pollution gradient. Additionally, in highly polluted environments, we differentiated short- from long-term effects of the two metallicolous forbs, which had different abilities to concentrate metals in their leaves. In line with other studies along metal gradients, variation of short-term interactions appeared to follow the Stress Gradient Hypothesis for plants less adapted to metal pollution (p = 0.030), with positive interactions dominating in most severe areas. Regarding long-term effects, the species with highest leaf metal-accumulation showed no negative effect contrary to the Elemental allelopathy Hypothesis. Long-term effects of the species with lower leaf-metal accumulation could not be determined because of the occurrence of an unexpected difference in micro-habitat conditions (soil depth and humidity) for this species along the metal pollution gradient. Increasing short-term facilitation along metal pollution gradients, which confirmed previous studies, is promising for improving conditions and restoring the most polluted environments. However, long-term results stressed the difficulty to quantify these effects given that these areas are highly fragmented and heterogeneous.

The role of ecotypic variation for plant facilitation in a metal-polluted system: Stress-intolerant target ecotypes are the best beneficiaries and stress-tolerant nurse ecotypes the best benefactors.

Disentangling competitive-response and -effect abilities has strongly improved our understanding of the role of competition for the diversity and composition of plant communities. Much less is known about the relative importance of facilitative-effect and -response abilities in harsh ecosystems. Here, we aim to fill this gap by simultaneously assessing the facilitative-response and -effect abilities of different species and ecotypes in former mining sites in the French Pyrenees, both in naturally occurring communities and in a common-garden designed on a slag heap. The response of two ecotypes of the target species Festuca rubra with contrasting metal-stress tolerances and the facilitative effects of two ecotypes with contrasting metal-stress tolerances of four different metallicolous nurse species were assessed. The results revealed that the response of the Festuca ecotype with lower metal-stress tolerance shifted from competitive (RII = -0.24) to facilitative (RII = 0.29) as pollution increased, consistently with the stress-gradient-hypothesis. The Festuca ecotype with high metal-stress tolerance did not show any facilitative response. Regarding facilitative effect ability assessed in the common-garden, nurse ecotypes from highly polluted habitats (RII = 0.04) had a significantly higher facilitative effects than ecotypes from less polluted habitats (RII = -0.05). Metal-intolerant target ecotypes of Festuca rubra were the most sensitive to the positive effects of neighbours, while metal-tolerant nurse ecotypes were the best benefactors. Facilitative-response ability appeared to be driven by a trade-off between stress-tolerance and facilitative response of target ecotypes. In contrast, facilitative-effect ability was positively correlated to the stress-tolerance of nurse plants. The results of this study show that the highest restoration success of highly metal-stressed systems should be found when highly stress-tolerant nurse ecotypes are associated with less stress-tolerant target ecotypes.

Contrasting effects of two phenotypes of an alpine cushion plant on understory species drive community assembly.

In alpine systems, cushion plants act as foundation species by ameliorating local environmental conditions. Empirical studies indicate that contrasting phenotypes of alpine cushion species have different effects on understory plant species, either facilitative or competitive. Furthermore, dependent species within each community type might also exhibit different responses to each cushion phenotype, which can be clustered into several "response groups". Additionally, these species-groups specific responses to alpine cushion species phenotypes could alter community assembly. However, very few studies have assessed responses of dependent communities at species-group levels, in particular for both above- and below-ground communities. Here, we selected a loose and a tight phenotype of the alpine cushion species Thylacospermum caespitosum in two sites of northwest China, and use the relative intensity of interactions index to quantify cushion plant effects on subordinate communities of plants and soil fungi and bacteria. We assessed variations in responses of both above- and below-ground organisms to cushion plant effects at species-group level. Species-group level analyses showed that the effects of the phenotype varied among groups of each of the three community types, and different species-groups were composed by unique taxa. Additionally, we found that loose cushions enhanced stochastic processes in community assembly, for plants and soil fungi but not for soil bacteria. These variations of phenotypic effects on different species-group induced contrasting taxonomic composition between groups, and alter community assembly thereby. Our study highlights the occurrence of contrasting effects of two phenotypes of a foundation cushion plant on understory plants, soil fungi and bacteria community composition, but not necessarily on their richness. We also showed that assessing responses of understory species at the species-group level allows a more realistic and mechanistic understanding of biotic interactions both for above- and below-ground communities.

Disentangling the effects of biomass and productivity in plant competition.

The relationship between competition and productivity in plant communities is unclear, and this is likely to be due to (1) a confusion in the literature between productivity and biomass, (2) the lack of studies assessing variation in competition in all combinations of biomass and productivity. We assessed the outcome of plant-plant interactions by removing the neighbors around five focal species in 14 herbaceous communities with contrasting biomasses and productivities: meadows with high biomass and productivity, heathlands with high biomass and low productivity, understory communities of deciduous forests with low biomass and high productivity and calcareous grasslands with low biomass and low productivity. Competition intensity was quantified with the relative interaction index (RII) calculated for both survival and growth of the transplanted targets assessed with the increase in leaf number. To examine which traits better explain variation in competition and what drives variation in diversity, we also quantified litter decomposition rate, species composition and diversity and six morphological traits related to plant size and growth rate for eight dominant species of each community. Our main questions were: (1) Is competition mostly related to biomass or productivity? (2) Which traits of the community dominants better explain variation in competition? (3) Is variation in competition and related traits correlated with variation in diversity? Competition for survival significantly increased with increasing community biomass (but not productivity). In addition, competition for survival increased with the size traits and competitive effects of the dominant species of the communities, whereas diversity decreased. Competition for growth also increased with increasing productivity, but only for high-biomass communities. Additionally, the increase in competition for growth with increasing soil fertility, as measured with litter decomposition rate, was only due to an increase in target growth in plots without neighbors and was unrelated to community competitive effects and species diversity. The results of our study illustrate how the confusion between productivity and biomass could have contributed to the long-standing debate on variation in competition along productivity gradients and its consequence for diversity.

Within-species variation of seed traits of dune engineering species across a European climatic gradient.

Within-species variation is a key component of biodiversity and linking it to climatic gradients may significantly improve our understanding of ecological processes. High variability can be expected in plant traits, but it is unclear to which extent it varies across populations under different climatic conditions. Here, we investigated seed trait variability and its environmental dependency across a latitudinal gradient of two widely distributed dune-engineering species (Thinopyrum junceum and Calamagrostis arenaria). Seed germination responses against temperature and seed mass were compared within and among six populations exposed to a gradient of temperature and precipitation regimes (Spiekeroog, DE; Bordeaux, FR; Valencia, ES; Cagliari, IT, Rome, IT; Venice, IT). Seed germination showed opposite trends in response to temperature experienced during emergence in both species: with some expectation, in populations exposed to severe winters, seed germination was warm-cued, whereas in populations from warm sites with dry summer, seed germination was cold-cued. In C. arenaria, variability in seed germination responses disappeared once the seed coat was incised. Seed mass from sites with low precipitation was smaller than that from sites with higher precipitation and was better explained by rainfall continentality than by aridity in summer. Within-population variability in seed germination accounted for 5 to 54%, while for seed mass it was lower than 40%. Seed trait variability can be considerable both within- and among-populations even at broad spatial scale. The variability may be hardly predictable since it only partially correlated with the analyzed climatic variables, and with expectation based on the climatic features of the seed site of origin. Considering seed traits variability in the analysis of ecological processes at both within- and among-population levels may help elucidate unclear patterns of species dynamics, thereby contributing to plan adequate measures to counteract biodiversity loss.

Tree genotypes affect rock lichens and understory plants: examples of trophic-independent interactions.

Genetic variation in foundation tree species can strongly influence communities of trophic-dependent organisms, such as herbivorous insects, pollinators, and mycorrhizal fungi. However, the extent and manner in which this variation results in unexpected interactions that reach trophic-independent organisms remains poorly understood, even though these interactions are essential to understanding complex ecosystems. In pinyon-juniper woodland at Sunset Crater (Arizona, USA), we studied pinyon (Pinus edulis) that were either resistant or susceptible to stem-boring moths (Dioryctria albovittella). Moth herbivory alters the architecture of susceptible trees, thereby modifying the microhabitat beneath their crowns. We tested the hypothesis that this interaction between herbivore and tree genotype extends to affect trophic-independent communities of saxicolous (i.e., growing on rocks) lichens and bryophytes and vascular plants beneath their crowns. Under 30 pairs of moth-resistant and moth-susceptible trees, we estimated percent cover of lichens, bryophytes, and vascular plants. We also quantified the cover of leaf litter and rocks as well as light availability. Four major findings emerged. (1) Compared to moth-resistant trees, which exhibited monopodial architecture, the microhabitat under the shrub-like susceptible trees was 60% darker and had 21% more litter resulting in 68% less rock exposure. (2) Susceptible trees had 56% and 87% less cover, 42% and 80% less richness, and 38% and 92% less diversity of saxicolous and plant communities, respectively, compared to resistant trees. (3) Both saxicolous and plant species accumulated at a slower rate beneath susceptible trees, suggesting an environment that might inhibit colonization and/or growth. (4) Both saxicolous and plant communities were negatively affected by the habitat provided by susceptible trees. The results suggest that herbivory of moth-susceptible trees generated litter at high enough rates to reduce rock substrate availability, thereby suppressing the saxicolous communities. However, our results did not provide a causal pathway explaining the suppression of vascular plants. Nonetheless, the cascading effects of genetic variation in pinyon appear to extend beyond trophic-dependent moths to include trophic-independent saxicolous and vascular plant communities that are affected by specific tree-herbivore interactions that modify the local environment. We suggest that such genetically based interactions are common in nature and contribute to the evolution of complex communities.

Morphological and ecological responses of a managed coastal sand dune to experimental notches.

In northern Europe, coastal dune remobilization by restoring natural processes is considered by some to maintain the coastal dune in chronically eroding sectors by migrating landward and to restore dune ecology. In wet climatic contexts, this nature-based solution has been shown to induce an increase in both sand bare areas and vegetation diversity. However, it has never been tested in the coastal dunes of southern Europe with a drier climate and, thus, more stressful conditions, where disturbance may inversely decrease vegetation diversity. An original experiment was set up in 2018 on a 4-km stretch of coastal dune in southwest France where Experimental Notches (EN) were excavated in the incipient foredune, referred to as West Experimental Notch (WEN), and in the established foredune, referred as to East Experimental Notch (EEN). Morphological and ecological responses were monitored using UAV photogrammetry and vegetation sampling along transects during two years with contrasted winter storm conditions. During the first winter characterized by calm wind conditions, a rapid filling of the WENs and the initiation of deposition lobes landward of the EENs were observed. Stronger winds during the second winter led to the development of deposition lobes of the EENs, increasing both their volume, up to 6 times, and their cross-shore elongation. The increase in disturbance induced by the notches had a significant impact on vegetation. New sandy bares were colonized by pioneer species leading to an increase in species richness and rejuvenation, in particular landward of the EENs. Although longer-term monitoring is required to draw conclusions, these results suggest that the excavation of foredune notches are able to re-establish an ecomorphological dynamic in the dunes of southwest France on the time scales of years, promoting landward sand transport and, thus, the foredune landward translation, while not threatening diversity. Such approach may become a relevant adaptation strategy to sea level rise and increased erosion in this region of the world.

Network motifs involving both competition and facilitation predict biodiversity in alpine plant communities.

Biological diversity depends on multiple, cooccurring ecological interactions. However, most studies focus on one interaction type at a time, leaving community ecologists unsure of how positive and negative associations among species combine to influence biodiversity patterns. Using surveys of plant populations in alpine communities worldwide, we explore patterns of positive and negative associations among triads of species (modules) and their relationship to local biodiversity. Three modules, each incorporating both positive and negative associations, were overrepresented, thus acting as "network motifs." Furthermore, the overrepresentation of these network motifs is positively linked to species diversity globally. A theoretical model illustrates that these network motifs, based on competition between facilitated species or facilitation between inferior competitors, increase local persistence. Our findings suggest that the interplay of competition and facilitation is crucial for maintaining biodiversity.

Direct and indirect facilitation affect community productivity through changes in functional diversity in an alpine system.

BACKGROUND AND AIMS: Facilitation is an important ecological process for plant community structure and functional composition. Although direct facilitation has accrued most of the evidence so far, indirect facilitation is ubiquitous in nature and it has an enormous potential to explain community structuring. In this study, we assess the effect of direct and indirect facilitation on community productivity via taxonomic and functional diversity. METHODS: In an alpine community on the Tibetan Plateau, we manipulated the presence of the shrub Dasiphora fruticosa and graminoids in a fenced meadow and a grazed meadow to quantify the effects of direct and indirect facilitation. We measured four plant traits: height, lateral spread, specific leaf area (SLA) and leaf dry matter content (LDMC) of forbs; calculated two metrics of functional diversity [range of trait and community-weighted mean (CWM) of trait]; and assessed the responses of functional diversity to shrub facilitation. We used structural equation modelling to explore how shrubs directly and indirectly drove community productivity via taxonomic diversity and functional diversity. KEY RESULTS: We found stronger effects from herbivore-mediated indirect facilitation than direct facilitation on productivity and taxonomic diversity, regardless of the presence of graminoids. For functional diversity, the range and CWM of height and SLA, rather than lateral spread and LDMC, generally increased due to direct and indirect facilitation. Moreover, we found that the range of traits played a primary role over taxonomic diversity and CWM of traits in terms of shrub effects on community productivity. CONCLUSIONS: Our study reveals that the mechanism of shrub direct and indirect facilitation of community productivity in this alpine community is expanding the realized niche (i.e. expanding range of traits). Our findings indicate that facilitators might increase trait dispersion in the local community, which could alleviate the effect of environmental filters on trait values in harsh environments, thereby contributing to ecosystem functioning.

An experimental approach to assessing the impact of ecosystem engineers on biodiversity and ecosystem functions.

Plants acting as ecosystem engineers create habitats and facilitate biodiversity maintenance within plant communities. Furthermore, biodiversity research has demonstrated that plant diversity enhances the productivity and functioning of ecosystems. However, these two fields of research developed in parallel and independent from one another, with the consequence that little is known about the role of ecosystem engineers in the relationship between biodiversity and ecosystem functioning across trophic levels. Here, we present an experimental framework to study this relationship. We combine facilitation by plants acting as ecosystem engineers with plant-insect interaction analysis and variance partitioning of biodiversity effects. We present a case-study experiment in which facilitation by a cushion-plant species and a dwarf-shrub species as ecosystem engineers increases positive effects of plant functional diversity (ecosystem engineers and associated plants) on ecosystem functioning (flower visitation rate). The experiment, conducted in the field during a single alpine flowering season, included the following treatments: (1) removal of plant species associated with ecosystem engineers, (2) exclusion (covering) of ecosystem engineer flowers, and (3) control, i.e., natural patches of ecosystem engineers and associated plant species. We found both positive and negative associational effects between plants depending on ecosystem engineer identity, indicating both pollination facilitation and interference. In both cases, patches supported by ecosystem engineers increased phylogenetic and functional diversity of flower visitors. Furthermore, complementarity effects between engineers and associated plants were positive for flower visitation rates. Our study reveals that plant facilitation can enhance the strength of biodiversity-ecosystem functioning relationships, with complementarity between plants for attracting more and diverse flower visitors being the likely driver. A potential mechanism is that synergy and complementarity between engineers and associated plants increase attractiveness for shared visitors and widen pollination niches. In synthesis, facilitation among plants can scale up to a full network, supporting ecosystem functioning both directly via microhabitat amelioration and indirectly via diversity effects.

Disentangling Large- and Small-Scale Abiotic and Biotic Factors Shaping Soil Microbial Communities in an Alpine Cushion Plant System.

Microorganisms play a crucial role in biogeochemical cycles and ecosystem processes, but the key factors driving microbial community structure are poorly understood, particularly in alpine environments. In this study, we aim to disentangle the relative contribution of abiotic and biotic factors shaping bacterial and fungal community structure at large and small spatial and integration scales in an alpine system dominated by a stress-tolerant cushion species Thylacospermum ceaspitosum. These effects were assessed in two mountain ranges of northwest China and for two contrasting phenotypes of the cushion species inhabiting two different microtopographic positions. The large- and small-scale abiotic effects include the site and microhabitat effects, respectively, while the large- and small-scale biotic effects include the effects of cushion presence and cushion phenotype, respectively. Soil microbial communities were characterized by Illumina Miseq sequencing. Uni- and multivariate statistics were used to test the effects of abiotic and biotic factors at both scales. Results indicated that the site effect representing the soil pH and abiotic hydrothermal conditions mainly affected bacterial community structure, whereas fungal community structure was mainly affected by biotic factors with an equal contribution of cushion presence and cushion phenotype effects. Future studies should analyze the direct factors contributing to shaping microbial community structure in particular of the cushion phenotypes.

Plant interactions shape pollination networks via nonadditive effects.

Plants grow in communities where they interact with other plants and with other living organisms such as pollinators. On the one hand, studies of plant-plant interactions rarely consider how plants interact with other trophic levels such as pollinators. On the other, studies of plant-animal interactions rarely deal with interactions within trophic levels such as plant-plant competition and facilitation. Thus, to what degree plant interactions affect biodiversity and ecological networks across trophic levels is poorly understood. We manipulated plant communities driven by foundation species facilitation and sampled plant-pollinator networks at fine spatial scale in a field experiment in Sierra Nevada, Spain. We found that plant-plant facilitation shaped pollinator diversity and structured pollination networks. Nonadditive effects of plant interactions on pollinator diversity and interaction diversity were synergistic in one foundation species networks while they were additive in another foundation species. Nonadditive effects of plant interactions were due to rewiring of pollination interactions. In addition, plant facilitation had negative effects on the structure of pollination networks likely due to increase in plant competition for pollination. Our results empirically demonstrate how different network types are coupled, revealing pervasive consequences of interaction chains in diverse communities.

Harnessing positive species interactions as a tool against climate-driven loss of coastal biodiversity.

Habitat-forming species sustain biodiversity and ecosystem functioning in harsh environments through the amelioration of physical stress. Nonetheless, their role in shaping patterns of species distribution under future climate scenarios is generally overlooked. Focusing on coastal systems, we assess how habitat-forming species can influence the ability of stress-sensitive species to exhibit plastic responses, adapt to novel environmental conditions, or track suitable climates. Here, we argue that habitat-former populations could be managed as a nature-based solution against climate-driven loss of biodiversity. Drawing from different ecological and biological disciplines, we identify a series of actions to sustain the resilience of marine habitat-forming species to climate change, as well as their effectiveness and reliability in rescuing stress-sensitive species from increasingly adverse environmental conditions.

The relative contribution of short-term versus long-term effects in shrub-understory species interactions under arid conditions.

Plant-plant interactions (competition and facilitation) in terrestrial ecosystems include: (1) short-term effects, primarily quantified with experimental removals; and (2) long-term effects, mostly due to soil weathering processes, primarily quantified with observational methods. It has been argued that these effects are likely to vary in contrasting directions with increasing drought stress in arid systems. However, few studies have used appropriate methodology to assess both types of effects and their variation across nurse species and environmental conditions, in particular in arid systems. This knowledge is crucial for predicting variation in the mediating role of facilitation with climate change and assessing the importance of nurse effects in ecological restoration. In the arid climate of central-south Tunisia, understory species' biomass, abundance and composition and soil parameters were compared in shrub-control, shrub-removed and open areas for three shrub species and in two habitats with contrasting soil moisture conditions. Long-term effects were dominant, positive for all three shrub species and associated with increasing nutrient content in shrub patches. Short-term effects, mainly related to water consumption, were weaker, mostly negative and dependent on shrub species. Additionally, long-term effects were less positive and short-term effects more negative in the dry habitat than in the wet habitat. Our study provides evidence of the primary influence of positive (facilitative) long-term effects in this arid system. However, the net effects of shrubs could be less beneficial for other species with increasing aridity under climate change, due to both a decrease in positive long-term effects and an increase in negative short-term effects.

The effects of foundation species on community assembly: a global study on alpine cushion plant communities.

Foundation species can change plant community structure by modulating important ecological processes such as community assembly, yet this topic is poorly understood. In alpine systems, cushion plants commonly act as foundation species by ameliorating local conditions. Here, we analyze diversity patterns of species' assembly within cushions and in adjacent surrounding open substrates (83 sites across five continents) calculating floristic dissimilarity between replicate plots, and using linear models to analyze relationships between microhabitats and species diversity. Floristic dissimilarity did not change across biogeographic regions, but was consistently lower in the cushions than in the open microhabitat. Cushion plants appear to enable recruitment of many relatively stress-intolerant species that otherwise would not establish in these communities, yet the niche space constructed by cushion plants supports a more homogeneous composition of species than the niche space beyond the cushion's influence. As a result, cushion plants support higher α-diversity and a larger species pool, but harbor assemblies with lower ß-diversity than open microhabitats. We conclude that habitats with and without dominant foundation species can strongly differ in the processes that drive species recruitment, and thus the relationship between local and regional species diversity.

Weak Evidence of Regeneration Habitat but Strong Evidence of Regeneration Niche for a Leguminous Shrub.

The identification of an ecological niche specific to the regeneration phase has mobilised significant attention. However, the importance of the regeneration niche concept remains unclear. Our main objective was to study the existence of such a regeneration niche for a leguminous shrub, Ulex europaeus. This study was carried out in southwest France in the context of water and nutrient stresses (mainly phosphorus limitation) due to the presence of nutrient-poor sandy soils. We analysed the regeneration of the species from the germination of seeds and emergence of new seedlings until the seedlings reached young shrub size. Our design included a P fertilisation treatment. We also investigated microsite characteristics (micro-topography and vegetation development) as they can interact with meteorological conditions and determine water availability for seeds and seedlings. We found that P availability controlled seedling growth and the time necessary to reach young shrub size. Water availability appeared to impact the species germination and seedlings survival. We also found that P and water availability depended on the interactions between microsite characteristics and climatic variations. Finally we found evidence that P and water availability are important ecological factors shaping the regeneration niche of the species, but we found weak evidence that any microsite would be appropriate for the regeneration of the species in the long term. Future studies regarding regeneration niches need to distinguish more clearly the ecological factors important for regeneration (the regeneration niche per se) and the physical world where the seedlings appear and develop (the regeneration habitat).

Phenotypic differentiation within a foundation grass species correlates with species richness in a subalpine community.

Few studies have examined consequences of ecotypic differentiation within alpine foundation species for community diversity and their feedbacks for the foundation species' fitness. Additionally, no study has quantified ecotypic differences in competitive effects in the field and in controlled conditions to disentangle genetic from plasticity effects in foundation/subordinate species interactions. We focused on a subalpine community of the French Pyrenees including two phenotypes of a cushion-forming species, Festuca gautieri: tight cushions in dry convex outcrops, and loose cushions (exhibiting high subordinate species richness) in wet concave slopes. We assessed, with field and shadehouse experiments, the genetic vs. plasticity basis of differences in: (1) cushion traits and (2) competitive effects on subordinates, and (3) quantified community feedbacks on foundation species' fitness. We found that trait differences across habitats had both genetic and plasticity bases, with stronger contribution of the latter. Field results showed higher competition within loose than tight phenotypes. In contrast, shadehouse results showed higher competitive ability for tight phenotypes. However, as changes in interactions across habitats were due to environmental effects without changes in cushion effects, we argue that heritable and plastic changes in competitive effects maintain high subordinate species diversity through decreasing competition. We showed high reproduction cost for loose cushions when hosting subordinates highlighting the occurrence of community feedbacks. These results suggest that phenotypic differentiation within foundation species may cascade on subordinate species diversity through heritable and plastic changes in the foundation species' competitive effects, and that community feedbacks may affect foundation species' fitness.

The context dependence of beneficiary feedback effects on benefactors in plant facilitation.

Facilitative effects of some species on others are a major driver of biodiversity. These positive effects of a benefactor on its beneficiary can result in negative feedback effects of the beneficiary on the benefactor and reduced fitness of the benefactor. However, in contrast to the wealth of studies on facilitative effects in different environments, we know little about whether the feedback effects show predictable patterns of context dependence. We reanalyzed a global data set on alpine cushion plants, previously used to assess their positive effects on biodiversity and the nature of the beneficiary feedback effects, to specifically assess the context dependence of how small- and large-scale drivers alter the feedback effects of cushion-associated (beneficiary) species on their cushion benefactors using structural equation modelling. The effect of beneficiaries on cushions became negative when beneficiary diversity increased and facilitation was more intense. Local-scale biotic and climatic conditions mediated these community-scale processes, having indirect effects on the feedback effect. High-productivity sites demonstrated weaker negative feedback effects of beneficiaries on the benefactor. Our results indicate a limited impact of the beneficiary feedback effects on benefactor cushions, but strong context dependence. This context dependence may help to explain the ecological and evolutionary persistence of this widespread facilitative system.

A global analysis of bidirectional interactions in alpine plant communities shows facilitators experiencing strong reciprocal fitness costs.

Facilitative 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.