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This work was supported by National Basic Research Program of China (973) (2013CB430402), the National Natural Science Foundation for Innovative Research Group (No. 51721093), the National Natural Science Foundation of China (No. 51279007) and the Fundamental Research Funds for the Central Universities.

Biodiversity is considered to mitigate detrimental impacts of climate change on the functioning of forest ecosystems, such as drought‐induced decline in forest productivity. However, previous studies produced controversial results and experimental evidence is rare. Specifically, the biological mechanisms underlying mitigation effects remain unclear, as existing work focuses on biodiversity effects related to the community scale. Using trait‐based neighbourhood models, we quantified changes in above‐ground wood productivity of 3,397 trees that were planted in a large‐scale tree diversity experiment in subtropical China across gradients of neighbourhood diversity and climatic conditions over a 6‐year period. This approach allowed us to simultaneously assess to what extent functional traits of a focal tree and biodiversity at the local neighbourhood scale mediate the growth response of individual trees to drought events. We found that neighbourhood tree species richness can mitigate for drought‐induced growth decline of young trees. Overall, positive net biodiversity effects were strongest during drought and increased with increasing taxonomic diversity of neighbours. In particular, drought‐sensitive species (i.e. those with a low cavitation resistance) benefitted the most from growing in diverse neighbourhoods, suggesting that soil water partitioning among local neighbours during drought particularly facilitated most vulnerable individuals. Thus, diverse neighbourhoods may enhance ecosystem resistance to drought by locally supporting drought‐sensitive species in the community. Synthesis. Our findings demonstrate that mechanisms operating at the local neighbourhood scale are a key component for regulating forests responses to drought and improve insights into how local species interactions vary along stress gradients in highly diverse tree communities. We found that neighbourhood diversity improves resistance of forests to drought by providing greatest support for most vulnerable individuals in the community. Our finding of increasing positive net biodiversity effects during drought suggests that mechanisms operating at the local neighbourhood scale are a key component for regulating the response of forest ecosystems to climate change. (Photo credit: Werner Härdtle)

1. Environmental gradients may influence a plant's physiological status and morphology, which in turn may affect plant-plant interactions. However, little is known about the relationship between environmental variation, physiological and morphological variability of plants and variation in the balance between competition and facilitation. 2. Mountain ranges in dry environments have opposing altitudinal environmental gradients of temperature and aridity, which limit plant growth at high and low elevations. This makes them particularly suitable for exploring the relationships between environmental conditions, plant phenotype and plant-plant interactions. We hypothesized that different environmental Stressors will differently affect the physiological status of a nurse plant. This, then, manifests itself as variation in nurse plant morphological traits, which in turn mediates plant-plant interactions by altering microhabitat conditions for the nurse and associated species. 3. In an observational study, we measured a series of functional traits of Arenaria tetraquetra cushions as indicators of its physiological status (e.g. specific leaf area, relative water content) and morphology (e.g. cushion compactness, size). Measurements were taken along the entire elevation range where A. tetraquetra occurs. Furthermore, we analysed how these functional traits related to soil properties beneath cushions and the number of associated species and individuals compared with open areas. 4. Cushions at high elevation showed good physiological status; they were compact and large, had higher soil water and organic matter content compared with open areas and showed the strongest facilitation effect of the whole elevation gradient — that is, the highest increase in species richness and abundance of beneficiaries compared with open areas. Physiological data at low elevation indicated stressful abiotic conditions for A. tetraquetra, which formed loose and small cushions. These cushions showed less improved soil conditions and had reduced facilitative effects compared with those at high elevation. 5. Synthesis. Functional traits of the nurse species varied distinctively along the two opposing stress gradients, in parallel to the magnitude of differences in microenvironmental conditions between cushions and the surrounding open area, and also to the facilitation effect of cushions. Our data, therefore, provides a strong demonstration of the generally overlooked importance of a nurse plant's vigour and morphology for its facilitative effects.

1. The stress-gradient hypothesis (SGH) predicts that the frequency of facilitative and competitive interactions will vary inversely across abiotic stress gradients, with facilitation being more common in conditions of high abiotic stress relative to more benign abiotic conditions. With notable exceptions, most tests of the SGH have studied the interaction between a single pair or a few pairs of species, and thus have evaluated shifts in the magnitude and direction of pair-wise interactions along stress gradients, rather than shifts in the general frequency of interactions. 2. The SGH has been supported by numerous studies in many ecosystems, has provided a crucial foundation for studying the interplay between facilitation and competition in plant communities, and has a high heuristic value. However, recent empirical research indicates that factors like the variation among species and the nature of the stress gradient studied add complexity not considered in the SGH, creating an opportunity to extend the SGH's general conceptual framework. 3. We suggest that one approach for extending the SGH framework is to differentiate between the original idea of how 'common' interactions might be along stress gradients and the ubiquitous empirical approach of studying shifts in the strength of pair-wise interactions. Furthermore, by explicitly considering the life history of the interacting species (relative tolerance to stress vs. competitive ability) and the characteristics of the stress factor (resource vs. non-resource) we may be able to greatly refine specific predictions relevant to the SGH. 4. We propose that the general pattern predicted by the SGH would hold more frequently for some combinations of life histories and stress factor, particularly when the benefactor and beneficiary species are mostly competitive and stress-tolerant, respectively. However, we also predict that other combinations are likely to yield different results. For example, the effect of neighbours can be negative at both ends of the stress gradient when both interacting species have similar 'competitive' or 'stress-tolerant' life histories and the abiotic stress gradient is driven by a resource (e.g. water). 5. Synthesis. The extension of the SGH presented here provides specific and testable hypotheses to foster research and helps to reconcile potential discrepancies among previous studies. It represents an important step in incorporating the complexity and species-specificity of potential outcomes into models and theories addressing how plant-plant interactions change along stress gradients.

New evidence demonstrates that facilitation plays a crucial role even at the edge of life in Maritime Antarctica. These findings are interpreted as support for the stress‐gradient hypothesis (SGH) – a dominant theory in plant community ecology that predicts that the frequency of facilitation directly increases with stress. A recent development to this theory, however, proposed that facilitation often collapses at the extreme end of stress and physical disturbance gradients. In this paper, we clarify the current debate on the importance of plant interactions at the edge of life by illustrating the necessity of separating the two alternatives to the SGH, namely the collapse of facilitation, and the switch from facilitation to competition occurring in water‐stressed ecosystems. These two different alternatives to the SGH are currently often amalgamated with each other, which has led to confusion in recent literature. We propose that the collapse of facilitation is generally due to a decrease in the effect of the nurse plant species, whilst the switch from facilitation to competition is driven by environmental conditions and strategy of the response species. A clear separation between those two alternatives is particularly crucial for predicting the role of plant–plant interactions in mediating species responses to global change.

A central theme of range‐limit theory (RLT) posits that abiotic factors form high‐latitude/altitude limits, whereas biotic interactions create lower limits. This hypothesis, often credited to Charles Darwin, is a pattern widely assumed to occur in nature. However, abiotic factors can impose constraints on both limits and there is scant evidence to support the latter prediction. Deviations from these predictions may arise from correlations between abiotic factors and biotic interactions, as a lack of data to evaluate the hypothesis, or be an artifact of scale. Combining two tenets of ecology—niche theory and predator–prey theory—provides an opportunity to understand how biotic interactions influence range limits and how this varies by trophic level. We propose an expansion of RLT, interactive RLT (iRLT), to understand the causes of range limits and predict range shifts. Incorporating the main predictions of Darwin's hypothesis, iRLT hypothesizes that abiotic and biotic factors can interact to impact both limits of a species’ range. We summarize current thinking on range limits and perform an integrative review to evaluate support for iRLT and trophic differences along range margins, surveying the mammal community along the boreal‐temperate and forest‐tundra ecotones of North America. Our review suggests that range‐limit dynamics are more nuanced and interactive than classically predicted by RLT. Many (57 of 70) studies indicate that biotic factors can ameliorate harsh climatic conditions along high‐latitude/altitude limits. Conversely, abiotic factors can also mediate biotic interactions along low‐latitude/altitude limits (44 of 68 studies). Both scenarios facilitate range expansion, contraction or stability depending on the strength and the direction of the abiotic or biotic factors. As predicted, biotic interactions most often occurred along lower limits, yet there were trophic differences. Carnivores were only limited by competitive interactions (n = 25), whereas herbivores were more influenced by predation and parasitism (77%; 55 of 71 studies). We highlight how these differences may create divergent range patterns along lower limits. We conclude by (a) summarizing iRLT; (b) contrasting how our model system and others fit this hypothesis and (c) suggesting future directions for evaluating iRLT. Understanding why range limits form and shift is an important pursuit. The authors introduce a novel extension to a classic theory and provide evidence that range dynamics are the result of the interaction between abiotic and biotic factors. The approach is especially useful for predicting climate change impacts on species distributions.

1. The stress-gradient hypothesis (SGH) predicts an increasing importance of facilitative mechanisms relative to competition along gradients of increasing environmental stress. Although developed across a variety of ecosystems, the SGH's relevance to the dynamic tree—grass systems of global savannas remains unclear. Here, we present a meta-analysis of empirical studies to explore emergent patterns of tree—grass relationships in global savannas in the context of the SGH. 2. We quantified the net effect of trees on understorey grass production relative to production away from tree canopies along a rainfall gradient in tropical and temperate savannas and compared these findings to the predictions of the SGH. We also analysed soil and plant nutrient concentrations in subcanopy and open-grassland areas to investigate the potential role of nutrients in determining grass production in the presence and absence of trees. 3. Our meta-analysis revealed a shift from net competitive to net facilitative effects of trees on subcanopy grass production with decreasing annual precipitation, consistent with the SGH. We also found a significant difference between sites from Africa and North America, suggesting differences in tree—grass interactions in the savannas of tropical and temperate regions. 4. Nutrient analyses indicate no change in nutrient ratios along the rainfall gradient, but consistent nutrient enrichment under tree canopies. 5. Synthesis. Our results help to resolve questions about the SGH in semi-arid systems, demonstrating that in mixed tree—grass systems, trees facilitate grass growth in drier regions and suppress grass growth in wetter regions. Relationships differ, however, between African and North American sites representing tropical and temperate bioclimates, respectively. The results of this meta-analysis advance our understanding of tree—grass interactions in savannas and contribute a valuable data set to the developing theory behind the SGH.

Ecological theory predicts that positive interactions among organisms will increase across gradients of increasing abiotic stress or consumer pressure. This theory has been supported by empirical studies examining the magnitude of ecosystem engineering across environmental gradients and between habitat settings at local scale. Predictions that habitat setting, by modifying both biotic and abiotic factors, will determine large‐scale gradients in ecosystem engineering have not been tested, however. A combination of manipulative experiments and field surveys assessed whether along the east Australian coastline: (1) facilitation of invertebrates by the oyster Saccostrea glomerata increased across a latitudinal gradient in temperature; and (2) the magnitude of this effect varied between intertidal rocky shores and mangrove forests. It was expected that on rocky shores, where oysters are the primary ecosystem engineer, they would play a greater role in ameliorating latitudinal gradients in temperature than in mangroves, where they are a secondary ecosystem engineer living under the mangrove canopy. On rocky shores, the enhancement of invertebrate abundance in oysters as compared to bare microhabitat decreased with latitude, as the maximum temperatures experienced by intertidal organisms diminished. By contrast, in mangrove forests, where the mangrove canopy resulted in maximum temperatures that were cooler and of greater humidity than on rocky shores, we found no evidence of latitudinal gradients of oyster effects on invertebrate abundance. Contrary to predictions, the magnitude by which oysters enhanced biodiversity was in many instances similar between mangroves and rocky shores. Whether habitat‐context modifies patterns of spatial variation in the effects of ecosystem engineers on community structure will depend, in part, on the extent to which the environmental amelioration provided by an ecosystem engineer replicates that of other co‐occurring ecosystem engineers.

The stress-gradient hypothesis predicts a switch from competition to facilitation, under increasing environmental stress. However, it is unclear how important is the change in competition–facilitation balance (i.e., the net outcome of plant–plant interactions) along the stress gradient in the regulation of community temporal stability (i.e., the inverse of temporal variability in total biomass). Increasing environmental stress may enhance community temporal stability by reduced competition or eventually by leading to facilitative interactions between the dominant and subordinate species. Here, we present the results of a 5-yr mesocosm experiment that demonstrates the effects of interspecific interactions on the temporal stability of a riparian community across different drought-stress scenarios. We constructed artificial communities of dominant species (Carex elata) and three subordinate species to simulate the independent effects of environmental stress and interspecific interactions. Using removal of the dominant species, we evaluated the interplay of various mechanisms regulating the temporal stability of the subordinate species (competition–facilitation balance, species asynchrony, and dominant species stability). By simultaneous testing of these stabilizing mechanisms, we show their importance differs depending on environmental variability and harshness. The predominant role is taken by species asynchrony in a seasonally dry environment, whereas in a permanently dry environment, the importance of reduced competition increases. Reduced competition was stabilizing, in particular through increased total community biomass, whereas species asynchrony increased total community biomass and decreased biomass variation. These results suggest experiments and simulations that exclude interspecific interactions may not offer realistic predictions of the effects of changing hydrological regimes on ecosystem functioning.

Summary The stress‐gradient hypothesis (SGH) predicts that the balance of plant–plant interactions shifts along abiotic environmental gradients, with facilitation becoming more frequent under stressful conditions. However, recent studies have challenged this perspective, reporting that positive interactions are, in some cases, more common at the intermediate level of environmental severity gradients. Here, we test whether and how neighbour effects by Silene acaulis cushions vary along a 700 m wide altitudinal transect, in relation to cushion morphological traits and environmental severity. Field measurements along the gradient, within and outside cushions, included (i) species richness and cover of coexisting vascular plants; (ii) cushion morphology; (iii) above‐ and below‐ground microclimate; and (iv) soil quality. We used the relative interaction index to decouple neighbour trait effects and environmental severity effects on plant diversity at different elevations. The ability of the cushion plant to facilitate heterospecifics shifts considerably along the elevation gradient, being greatest at the intermediate level. On the other hand, Silene morphological traits steadily change along the gradient, from lax, soft and flat‐shaped cushion habits at low elevation to tightly knit and dome‐shaped habits at high elevation. Cushion morphological changes are associated with mitigating effects on microclimate, indicating that cushions effectively act as a heat‐trap at medium and high elevations, while at low elevations the soft and flat cushions avoid excessive heat accumulation by tight coupling with the surrounding atmosphere. At the upper end of the gradient, cushion cespitose–pulvinate compactness and high stem density appear to be critical traits in modulating the net effect of plant–plant interaction, since the space available for hosting other vascular species is considerably reduced. In conclusion, this work provides a mechanistic link between plant morphological traits, associated biogenic microclimate changes and variation in net plant–plant interactions along the explored severity gradient. Our findings support an alternative conceptual model to SGH, with plant facilitation collapsing at the upper extreme of the abiotic stress gradient. Lay Summary