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Several studies have found that increased nitrogen (N) deposition leads to a decline in species richness in semi-natural grasslands, mainly due to the loss of species typical of nutrient-poor soils. However, after reaching a peak around 1990, N deposition has decreased in Europe over the last 30 years. In this study, we investigated the changes in species number and composition of semi-natural grasslands during this period of declining N deposition. To this end, we compared the data from the first survey (2001-2005) of 147 grassland sites in Switzerland with those from the third survey (2011-2015). We further analysed the vegetation development of a specific hay meadow from 1992 to 2013. In this grassland, total vegetation cover and the cover of graminoid species decreased, while the cover of oligotrophic species increased. At the 147 grassland sites, total species number decreased at sites with still high levels of N deposition and it tended to increase at sites with low N deposition, i. e. below the critical load for N deposition. The number of oligotrophic grassland species increased at sites with a large decrease in N deposition and strong inclination. Thus, the results of this study indicate that the reduction of N emissions had a measurable positive effect on species diversity in these semi-natural grasslands. Most of the grasslands surveyed appear to be quite resilient against N deposition, i. e. they do not shift to an alternative low diversity state dominated by a few competitive species, and recovery of the species composition as a result of the decrease in N deposition seems possible, especially on steep slopes. Furthermore, the study underlines the importance of regular management of semi-natural, unfertilised, low-productivity grassland to maintain the diversity of oligotrophic grassland species. [ABSTRACT FROM AUTHOR]

Intercropping, by capitalizing on positive biodiversity–productivity relationships, represents a promising option to increase agricultural sustainability. However, the complexity and context-dependency of plant–plant interactions can make it challenging for farmers to find suitable crop combinations. Furthermore, intercropping is usually implemented with standard inter-row spacing and plant densities based on monoculture practices, which might not be the ideal configuration to maximize yield. Here we present a spatially-explicit yield analysis method based on plant ecological interaction models that allowed to optimize crop species combinations and spatial configurations for maximal yield in intercropped systems. We tested this method with three crop species, namely oat, lupine, and camelina. In a first step, field experiments in which crop density and adjacent crop type were varied provided us with indications on which species would compete more with each other. The results showed us that oat and camelina strongly competed with each other. In addition, the distance experiments allowed us to understand how the changes in yield associated with the presence of neighbors vary with distance. This allowed us to find the sets of parameters (identity of neighbors, sowing density, distances between individuals) that optimise intercrop yield (measured as Land Equivalent Ratio [LER]) for the three considered species. Specifically, we show that alternating rows of species led to higher LERs than a homogeneous species mixing, and that 3-species combinations are not necessarily more performant than the best 2-species combinations. In addition, we show that increasing the density of oat is generally beneficial for LER, while increasing the density of lupine is not. By modelling crop yield from simple and reproducible density and distance experiments, our results allow to optimize crop mixtures in terms of species combinations and spatial configurations, for maximal crop yield. [ABSTRACT FROM AUTHOR]

Enhancing diversity within crop systems can have benefits including increased resource use efficiency and productivity, and increased control of weeds, pests and diseases. Some benefits are expected to operate through biodiversity-driven insurance effects, whereby enhanced diversity increases the chance that a system component can compensate for the impacts of adverse environmental conditions. Studies of insurance effects in natural and agricultural systems have provided equivocal results. As insurance effects are expected to play a key role in helping to maintain crop production in more variable future climates (for example under periodic drought), it is essential to know when and how they operate and interact with other potentially beneficial biodiversity-function effects. Using barley as a model crop, and pot-based plant communities, we studied the interactive effects of barley cultivar diversity and drought stress on plant productivity and the response of agricultural weeds, fungal disease, and aphids. Drought reduced barley and weed biomass, but there were no interactive effects of drought and cultivar diversity on plant productivity. Increased cultivar diversity enhanced weed suppression, potentially as a result of reduced functional space availability, and reduced disease severity on a susceptible cultivar; these effects were consistent irrespective of drought. Aphid responses were more complex, with idiosyncratic response patterns on individual cultivars. Overall, we found no evidence of an insurance effect of enhanced cultivar diversity for the negative impact of drought on crop productivity, but our results indicate that other positive biodiversity effects (weed and disease suppression) are maintained under drought. However, it is clear that not all potentially-beneficial biodiversity effects respond in the same manner. Field trials are now needed to explore whether a range of responses also occur in crop field settings, whether these can be expected to occur predictably under a range of environmental conditions, and how these then impact on crop production. [ABSTRACT FROM AUTHOR]

• High-elevation environments in the Himalayas are stressful for plant life. • Himalayan endemic Berberis angulosa is a nurse shrub that facilitates beneficiary species. • Facilitation by Berberis shrubs promotes plant diversity, and protects high-value and threatened medicinal species. • The nurse shrub plays a critical role in structuring plant communities in high-elevation Himalayas. Facilitation is a global phenomenon that occurs when one species promotes the growth, survival, or reproduction of another species, mostly in stressful environments. However, the importance of facilitation by shrubs in maintaining plant community diversity is not well evaluated in the Himalayas, especially for the richness and conservation of medicinal and human-valued species. Therefore, we aimed to explore the facilitative role of a dwarf shrub species, Berberis angulosa , in maintaining plant composition and richness of human-valued species in the Langtang valley of Nepal's Himalayas. We censused plant species in open patches and beneath Berberis during monsoon and post-monsoon (dry) seasons at three elevations. Total species richness and richness of human-valued species were significantly higher inside the Berberis canopy than in gaps; the former being 39% and the latter 46% greater under shrubs than in open sites. Facilitation by Berberis shrubs promoted plant community diversity irrespective of season and elevation; however, higher differences in mean species richness for both total plant species and human-valued species during the dry season and at high elevation indicated increased facilitation intensity under more stressful conditions. The facilitative effect of Berberis shrubs increased, combining both seasons, overall plant diversity by 19% (total=105), and human-valued species by 16% (total=56). Our results show the importance of facilitation by nurse shrubs in structuring plant communities and protecting medicinal and socio-ecologically important plants, thus enriching ecosystem services in the Himalayas. These results suggest nurse plant species should be incorporated into conservation policies and management strategies for effective biodiversity conservation and sustainability, especially in the face of climate change. [ABSTRACT FROM AUTHOR]

In alpine environments facilitator species buffer environmental extremes while building up soil resources above that of open areas. These modulating effects are critical for the persistence of species out of their optimal range and contribute to increase community-level plant species richness and phylogenetic diversity. We analyzed the effects of seven potential facilitator species with contrasting morphologies on subordinate plant species along a crossed environmental gradient, linking such effects to canopy effects. We also used these patches consisting of multiple shrub species to evaluate the effects of the whole shrub community on species richness and phylogenetic diversity, and whether such shrub community effects differed along the gradient. We used ecological and phylogenetic data of alpine plant communities along two altitudinal gradients on opposing aspects of the Sierra Nevada Mountains (Spain). As expected, shrubs buffered harsh abiotic conditions by decreasing mean temperatures and increasing relative humidity with regard to open areas. Composition of subordinate plant communities differed among shrubs and among sites, and correlated with relative humidity along the gradient pointing to the dependence of subordinate species on micro-environments created by shrubs. There were a variety of shrub effects on overall plant abundance and richness depending on shrub identity. In the most extreme sites we recorded generally positive effects of the shrub community, which promoted whole-community species richness and phylogenetic diversity despite species-specific effects ranging from clearly negative to positive. Our data therefore show that the effect of different shrub species on plant community richness and phylogenetic diversity is not redundant, as every shrub species may host unique communities, thereby affecting the structure and composition of the whole community. [ABSTRACT FROM AUTHOR]

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Better understanding of the mechanistic basis of plant plasticity will enhance efforts to breed crops resilient to predicted climate change. However, complexity in plasticity's conceptualisation and measurement may hinder fruitful crossover of concepts between disciplines that would enable such advances. We argue active adaptive plasticity is particularly important in shaping the fitness of wild plants, representing the first line of a plant's defence to environmental change. Here, we define how this concept may be applied to crop breeding, suggest appropriate approaches to measure it in crops, and propose a refocussing on active adaptive plasticity to enhance crop resilience. We also discuss how the same concept may have wider utility, such as in ex situ plant conservation and reintroductions. Increasing environmental uncertainty is focussing research interest on plant plasticity. But despite calls for plasticity concepts to be adopted in crop breeding, this does not appear to have happened. Plasticity is a broad and multifaceted concept, making it potentially difficult to identify those aspects of previous research most relevant to the crop breeding context. Given the challenges posed by climate change and the different evolutionary contexts in natural and crop systems, we identify active adaptive plasticity as a key issue for further investigation by crop breeders. We outline and illustrate the experimental and statistical analytical approaches necessary to begin to assess active adaptive plasticity, and we highlight benefits that might arise in other fields from a fuller understanding of the role and regulation of this aspect of plasticity. [ABSTRACT FROM AUTHOR]

Climate change and associated environmental alterations affect plant communities, potentially decoupling links between plants and their associated soil microbial communities. This may in turn affect processes like litter decomposition, an important function that controls nutrient and carbon cycling as well as many other ecosystem processes. Microbial decomposers have been proposed to specialize, being able to easier decompose litter from its 'home' community than litter from other communities, in what is termed the home field advantage (HFA) hypothesis. We report a litter decomposition experiment including four alpine meadow communities spread along a geographical gradient that ranged from the Atlas in Morocco to the Iberian Peninsula to the Swiss Alps. We tested whether litter decomposition rate would differ depending on litter and soil origin, expecting interactions between litter and soil microbial communities to be at the core of responses. We expected to find HFA in all four alpine communities. HFA ranged from very negative to neutral to positive along our geographical gradient, in a variation that could be attributed to the interaction of microbial communities with soil and plant traits, and linked to the relative abundance of microbial groups performing specific functions. Litter decomposition depended on litter quality and on the fungal community, which seemed adapted to deal with poor-quality, recalcitrant litter, leading to HFA. Climate plays a role as well, indirectly through plant community composition. Phyllosphere communities competed with soil communities when in interaction, a fact that may have blurred results of some previous HFA experiments. In summary, there is a clear HFA in systems with low-quality litter that requires a specialised fungal community to maximize decomposition. By contrast, high-quality, easy to decompose litter would be dealt with by any decomposer community, not showing evidence of HFA. • Litter decomposition depends on links between plants and soil microbial communities. • Microbes might decompose easier litter from its own community than from elsewhere. • Patterns of this so-called 'home-field advantage' (HFA) are unclear. • Systems with low–quality litter require specialised fungal communities, showing HFA. • Systems with high-quality litter have no specialised communities and show no HFA. [ABSTRACT FROM AUTHOR]

Recent meta-analyses demonstrate that intercropping can increase the land use efficiency of crop production by 20–30 % on average, indicating a strong potential contribution to sustainable intensification. However, there is substantial variability around this average: individual studies range from half to double the land productivity of monocrops. Legume-cereal intercrops and intercrops with high temporal niche separation tend to be more productive than the average, but these two combination types are not always suitable. There is a need to explore other possibilities to achieve productive intercrops. We explored whether two simple functional traits involved in radiation use, plant vegetative height and specific leaf area (SLA), could be used to predict intercrop productivity. Height and SLA together are associated with key plant life-history and resource economy strategies determining competitiveness and tolerance of competition, especially with regard to light, and could therefore be expected to underpin overyielding in intercrops. In the first year of our study, we grew crops as monocrops at one site in Kenya and measured their height and SLA. In the second year, we grew crops in monocrop, intercrop, and single plant treatments at two sites in Kenya and one site in Nigeria. Together, these treatments allowed us to identify whether each intercrop combination overyielded or underyielded, and whether any overyielding was driven by facilitation and/or differences in inter- vs intraspecific competition. We then related the strength of these interactions to the two traits. We found that intercrop grain yields varied in relation to the height and SLA of each species in the intercrop, but together these traits explained less than a third of variation in intercrop land equivalence ratios (LER). More variation could be explained by allowing for the effect of site, suggesting that the two traits interact with site conditions to determine yield. Biomass LERs responded differently to grain LERs, suggesting that plasticity in resource allocation in response to intercropping conditions may further influence yields. Our study found some evidence that combining species with traits indicating contrasting responses to competition (an avoidant species with a tolerant species) could increase resource use complementarity and thus intercrop overyielding. However, it was clear that other factors (such as additional traits, or the trait by site interaction) are needed to refine our understanding of intercrop productivity. A trait-based framework has potential to predict intercrop productivity, but simple measures of height and SLA alone are insufficient. • Intercrop yields vary from much lower than sole crops to much higher across different studies. • Plant traits may mediate competition and facilitation between intercrops, and predict yields. • Two simple traits, vegetative height and specific leaf area (SLA), explained some variance in yields. • Yields were higher when tall species with a low SLA were paired with short species with a high SLA. • Height and SLA alone cannot reliably predict intercrop yields: other traits should be explored. [ABSTRACT FROM AUTHOR]

Understanding the mechanisms of community assembly may provide evidence to improve crop management, and in particular how weeds impact on crop yields. Focussing on plant functional traits and their diversity, we analysed a crop–weed interaction study with different levels of weed species and barley cultivar diversity to assess how weed species and barley cultivars respond to competition. Pre-emption of light resources by the taller barley did not impact on the weeds, with both weeds and barley showing similar order of magnitude shifts in height, specific leaf area and leaf dry matter content in response to crop–weed competition. These shifts were to a more conservative growth pattern, and suggest in this study a greater importance of below- than above-ground interactions in driving trait responses. The mixture of barley cultivars shifted the weeds to a more conservative growth pattern compared to the cultivar monocultures. The results indicate that cultivar mixtures could result in less need for weed control in arable fields, and possibly that the development of complementary cultivar mixtures could reinforce this effect. This confirms the results of other studies which show that mixtures either improve yields or make yields less variable in response to weed competition. [ABSTRACT FROM AUTHOR]

Cushion plants' significant role for alpine biodiversity has been demonstrated in particular through their positive effects on plant diversity. However, their effects on higher trophic levels (e.g. insects) remain largely unclear. In this study, by field sampling in the Hengduan Mountains (HDM) in southwestern China, we evaluated the effects of an alpine gynodioecious cushion species, Arenaria polytrichoides (Carophyllaceae), on insect richness, abundance and population dynamics at two different elevations (4427 m vs. 4732 m) separately at two time periods (day vs. night) and in two growing seasons (early vs. late). The results showed that the total insect diversity decreases from low to high elevation sites. Some insect species were exclusively detected within A. polytrichoides cushions, leading to an increase in local insect richness from 7% to 35%. The positive effects of cushions on insect diversity could be attributed to unique biotic properties provided by cushions. Firstly, the effects were determined by the sexual dimorphism of the cushion with hermaphroditic cushions supporting higher insect diversity than female cushions. This could be because hermaphroditic cushions provide more resources, such as nectar and pollen grains, for insects than female cushions. Secondly, the cushions' associated beneficiary plants can also provide extra resources for attracting more insects, but this effect was mediated by the micro-environmental conditions. Finally, the magnitude of cushions' positive effects on insect dynamics were stronger under higher than under lower environmental stress. This study confirmed that facilitation by A. polytrichoides cushions in HDM plays an important role in constructing the alpine insect community and further regulating its dynamics. Moreover, the positive effects of cushions on insect dynamics increase with increasing environmental stress. Therefore, the distribution range of insects would quite possibly be expanded into higher elevation under future climate changes, which will induce new challenges for the local alpine ecosystems. Unlabelled Image • Insect diversity decreases with increasing elevation in the Hengduan Mountains. • The cushion plant Arenaria hosts exclusive insects, thereby increasing local insect diversity. • The positive effects of Arenaria were attributed to biotic properties provided by the cushion. • Cushions' positive effects on insects were determined by environmental severity. [ABSTRACT FROM AUTHOR]

• Nurse plants in alpine systems may enhance species survival under climate change. • We simulated climate change effects on an alpine cushion species. • We also assessed consequences of new climate on its facilitation effect. • Warming enhanced cushion performance but did not alter its facilitation effect. • Global change may alter carbon fixation and respiration, threatening plant survival. Climate change is more pronounced in high-elevation habitats than elsewhere, potentially causing disruptions in plant community structure and dynamics through changes in plant interactions. We tested the effect of warming and rainfall manipulations on growth and gas exchange of a cushion plant species, Arenaria tetraquetra , in the Sierra Nevada Mountains in SE Spain and assessed its consequences for plant interactions. Arenaria tetraquetra is a facilitator that often hosts a variety of beneficiary species within its canopy. Warming and drought could alter cushion growth patterns and canopy compaction, which in turn affect facilitation. We increased temperature using acrylic open-top chambers (OTC) and increased or decreased water availability with rainfall collectors. Our rainfall treatments did not have significant effects on cushion growth or facilitation. Three years of warming enhanced cushion photosynthesis and respiration, leading to larger cushions with thinner leaves, but did not change other cushion traits nor alter its facilitation effects. Some traits, however, readily responded to inter-annual climate variability. Global warming will change cushion plant physiology, as higher temperatures at times increase plant respiration above the level of carbon fixation. However, we did not find changes in interaction between benefactor and beneficiary species, perhaps due to the short-term nature of our experiment. Recorded physiological and morphological changes might be, however, signs of more pronounced, long-term consequences of warming on plant survival and community composition. [ABSTRACT FROM AUTHOR]

• Rhododendron rupicola shrubs in the Hengduan mountains can modify abiotic microenvironmental conditions. • R. rupicola can increase diversity of soil associated fungi and modify their community composition. • R. rupicola facilitates growth and survival of treeline seedlings in a species-specific manner. • R. rupicola facilitates upward migration of the treeline and modifies its species composition. The alpine treeline is shifting upward due to climate warming. However, the treeline species composition and the pace of its upward migration can be mediated by ecological interactions. In particular, so-called ecosystem engineers, i.e. species that modulate the microscale environmental conditions, at the treeline may play a crucial role. We conducted a three-year seedling transplant experiment at the alpine treeline ecotone in southwest China to study how the shrub Rhododendron rupicola modifies the microscale physical and biotic environments and thus influences the establishment and performance of the two treeline species Larix potaninii and Picea likiangensis. Seedlings were transplanted to the current timberline and treeline, as well as above the current treeline in order to determine the responses of the two tree species to the shrub with respect to the current tree distribution. R. rupicola modified the microenvironment by increasing soil moisture and nutrient contents, buffering soil temperature fluctuations, and by increasing richness and changing the composition of root-associated fungi. As a result, tree seedlings planted under shrubs had significantly higher survival, growth rates and nutrient accumulations than those planted in open ground. Furthermore, seedlings planted at lower elevations performed better than those planted at higher elevations. Beyond the treeline, seedling survival was very low on open ground but strongly facilitated by the shrub. Finally, facilitation effects were species-specific, with Larix benefitting more from the shrub than Picea , while Picea had less mortality than Larix in the absence of the shrub. This study demonstrates that shrubs, through the amelioration of physical and biotic microenvironmental conditions, can act as stepping stones for the establishment of selective tree species beyond the current treeline. This suggests that biotic interactions can strongly modify the treeline species composition and push the treeline beyond its current climatic limits, thereby facilitating the upward shift with ongoing climate warming. [ABSTRACT FROM AUTHOR]