SummaryIntercropping is a farming practice involving two or more crop species, or genotypes, growing together and coexisting for a time. On the fringes of modern intensive agriculture, intercropping is important in many subsistence or low-input/resource-limited agricultural systems. By allowing genuine yield gains without increased inputs, or greater stability of yield with decreased inputs, intercropping could be one route to delivering 'sustainable intensification'. We discuss how recent knowledge from agronomy, plant physiology and ecology can be combined with the aim of improving intercropping systems. Recent advances in agronomy and plant physiology include better understanding of the mechanisms of interactions between crop genotypes and speciesfor example, enhanced resource availability through niche complementarity. Ecological advances include better understanding of the context-dependency of interactions, the mechanisms behind disease and pest avoidance, the links between above-and below-ground systems, and the role of microtopographic variation in coexistence. This improved understanding can guide approaches for improving intercropping systems, including breeding crops for intercropping. Although such advances can help to improve intercropping systems, we suggest that other topics also need addressing. These include better assessment of the wider benefits of intercropping in terms of multiple ecosystem services, collaboration with agricultural engineering, and more effective interdisciplinary research.

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.

SummaryHere, we incorporate facilitation into trait-based community assembly theory by testing two mutually compatible facilitative mechanisms: changes in the environmental filter, causing either an increase in the range of trait values (i.e. a range expansion effect) and/or a shift in trait distributions (i.e. a range shift effect); and changes in trait spacing, suggesting an effect on niche differentiation.We analyzed the distribution of three functional traits -leaf dry matter content, specific leaf area and lateral spread -of plant communities dominated by a cushion-forming foundation species at four sites differing in elevation and aspect.We found support for environmental filtering and niche differentiation mechanisms by cushions, with filtering effects (in particular range shifts) increasing with environmental severity at higher elevation. The effect size of cushions on trait distribution was similar to that of environmental gradients caused by elevation and aspect. The consideration of intraspecific trait variability improved the detection of cushion effects on trait distributions.Our results highlight the importance of facilitation in the modification of taxonomic and functional diversity of ecological communities, and indicate that facilitation can occur through combined effects on environmental filtering and niche differentiation, with strong environmental context dependence of each mechanism.

Summary1. 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.