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Summary: There is a drive to improve the sustainability of agricultural systems including the biodiversity component. Cultivar mixtures offer yield benefits from the same land area, but the mechanisms behind this overyielding have not been completely worked out. One potential mechanism is improved competition with weeds. We use an experimental approach of varying barley (Hordeum vulgare L.) genotypic and phenotypic diversity to test the hypothesis that increases in diversity have an impact on weed growth strategies and community assembly, or if responses are driven by barley performance. There was no effect of increasing barley mixture diversity on weed traits, either in terms of species means or the community‐weighted mean. However, Functional Richness of the weed community decreased with increasing barley mixture diversity mainly as a result of reduced specific leaf area Functional Richness. This pattern was driven by a reduction in species richness of the weed community rather than by reduced variation within species. Whilst barley phenotype had different impacts on weed traits, there were no specific mixtures of phenotypes or genotypes that had consistent effects on community assembly or weed species responses. The competitive exclusion of weed species could have agronomic and environmental benefits, through better targeting or less frequent use of herbicides. Growing crop mixtures is one of many strategies available to improve agricultural sustainability and resilience, and one that has clear benefits. [ABSTRACT FROM AUTHOR]

Growing crops in more diverse crop systems (i.e., intercropping) is one way to produce food more sustainably. Even though intercropping, compared to average monocultures, is generally more productive, the full yield potential of intercropping might not yet have been achieved as modern crop cultivars are bred to be grown in monoculture. Breeding plants for more familiarity in mixtures, i.e., plants that are adapted to more diverse communities (i.e., adaptation) or even to coexist with each other (i.e., coadaptation) might have the potential to sustainably enhance productivity. In this study, the productivity benefits of familiarity through evolutionary adaptation and coevolutionary coadaptation were disentangled in a crop system through an extensive common garden experiment. Furthermore, evolutionary and coevolutionary effects on species-level and community-level productivity were linked to corresponding changes in functional traits. We found evidence for higher productivity and trait convergence with increasing familiarity with the plant communities. Furthermore, our results provide evidence for the coevolution of plants in mixtures leading to higher productivity of coadapted species. However, with the functional traits measured in our study, we could not fully explain the productivity benefits found upon coevolution. Our study investigated coevolution among randomly interacting plants and was able to demonstrate that coadaptation through coevolution of coexisting species in mixtures occurs and promotes ecosystem functioning (i.e., higher productivity). This result is particularly relevant for the diversification of agricultural and forest ecosystems, demonstrating the added value of artificially selecting plants for the communities they are familiar with., Competing Interests: Competing interests statement:The authors declare no competing interest.

Foundational cushion plants can re-organize community structures and sustain a prominent proportion of alpine biodiversity, but they are sensitive to climate change. The loss of cushion species can have broad consequences for associated biota. The potential plant community changes with the population dynamics of cushion plants remain, however, unclear. Using eight plant communities along a climatic and community successional gradient, we assessed cushion population dynamics, the underlying ecological constraints and hence associated plant community changes in alpine communities dominated by the foundational cushion plant Arenaria polytrichoides. The population dynamics of Arenaria are attributed to ecological constraints at a series of life history stages. Reproductive functions are constrained by increasing associated beneficiary plants; subsequent seedling establishment is constrained by temperature, water and light availability, extreme climate events, and interspecific competition; strong competitive exclusion may accelerate mortality and degeneration of cushion populations. Along with cushion dynamics, species composition, abundance and community structure gradually change. Once cushion plants completely degenerate, previously cushion-dominated communities shift to relatively stable communities that are overwhelmingly dominated by sedges. Climate warming may accelerate the degeneration process of A. polytrichoides. Degeneration of this foundational cushion plant will possibly induce massive changes in alpine plant communities and hence ecosystem functions in alpine ecosystems. The assessment of the population dynamics of foundation species is critical for an effective conservation of alpine biodiversity., (© 2023. Science China Press.)

Background: Legumes can fix atmospheric nitrogen (N) and facilitate N availability to their companion plants in crop mixtures. However, biological nitrogen fixation (BNF) of legumes in intercrops varies largely with the identity of the legume species. The aim of our study was to understand whether BNF and concentration of plant nutrients by common bean is influenced by the identity of the companion plant species in crop mixtures. In this greenhouse pot study, common beans were cultivated with another legume (chickpea) and a cereal (Sorghum). We compared BNF, crop biomass and nutrient assimilation of all plant species grown in monocultures with plants grown in crop mixtures., Results: We found beans to exhibit low levels of BNF, and to potentially compete with other species for available soil N in crop mixtures. The BNF of chickpeas however, was enhanced when grown in mixtures. Furthermore, biomass, phosphorous and potassium values of chickpea and Sorghum plants were higher in monocultures, compared to in mixtures with beans; suggesting competitive effects of beans on these plants. Concentration of calcium, magnesium and zinc in beans was higher when grown with chickpeas than with Sorghum., Conclusions: It is generally assumed that legumes benefit their companion plant species. Our study highlights the contrary and shows that the specific benefits of cereal-legume mixtures are dependent on the growth rate of the species concerned. We further highlight that the potential of legume-legume mixtures is currently undervalued and may play a strong role in increasing N use efficiency of intercrop-based systems., (© 2023. The Author(s).)

Plant recruitment interactions (i.e., what recruits under what) shape the composition, diversity, and structure of plant communities. Despite the huge body of knowledge on the mechanisms underlying recruitment interactions among species, we still know little about the structure of the recruitment networks emerging in ecological communities. Modeling and analyzing the community-level structure of plant recruitment interactions as a complex network can provide relevant information on ecological and evolutionary processes acting both at the species and ecosystem levels. We report a data set containing 143 plant recruitment networks in 23 countries across five continents, including temperate and tropical ecosystems. Each network identifies the species under which another species recruits. All networks report the number of recruits (i.e., individuals) per species. The data set includes >850,000 recruiting individuals involved in 118,411 paired interactions among 3318 vascular plant species across the globe. The cover of canopy species and open ground is also provided. Three sampling protocols were used: (1) The Recruitment Network (RN) protocol (106 networks) focuses on interactions among established plants ("canopy species") and plants in their early stages of recruitment ("recruit species"). A series of plots was delimited within a locality, and all the individuals recruiting and their canopy species were identified; (2) The paired Canopy-Open (pCO) protocol (26 networks) consists in locating a potential canopy plant and identifying recruiting individuals under the canopy and in a nearby open space of the same area; (3) The Georeferenced plot (GP) protocol (11 networks) consists in using information from georeferenced individual plants in large plots to infer canopy-recruit interactions. Some networks incorporate data for both herbs and woody species, whereas others focus exclusively on woody species. The location of each study site, geographical coordinates, country, locality, responsible author, sampling dates, sampling method, and life habits of both canopy and recruit species are provided. This database will allow researchers to test ecological, biogeographical, and evolutionary hypotheses related to plant recruitment interactions. There are no copyright restrictions on the data set; please cite this data paper when using these data in publications., (© 2022 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

By capitalising on positive biodiversity - productivity relationships, intercropping provides opportunities to improve agricultural sustainability. Intercropping is generally implemented using commercial seeds that were bred for maximal productivity in monocultures, thereby ignoring the ability of plants to adapt over generations to the surrounding neighbourhood, notably through increased complementarity, that is reduced competition or increased facilitation. This is why using monoculture-adapted seeds for intercropping might limit the benefits of crop diversity on yield. However, the adaptation potential of crops and the corresponding changes in complementarity have not been explored in annual crop systems. Here we show that plant - plant interactions among annual crops shifted towards reduced competition and/or increased facilitation when the plants were growing in the same community type as their parents did in the previous two generations. Total yield did not respond to this common coexistence history, but in fertilized conditions, we observed increased overyielding in mixtures with a common coexistence history. Surprisingly, we observed character convergence between species sharing the same coexistence history for two generations, in monocultures but also in mixtures: the six crop species tested converged towards taller phenotypes with lower leaf dry matter content. This study provides the first empirical evidence for the potential of parental diversity affecting plant - plant interactions, species complementarity and therefore potentially ecosystem functioning of the following generations in annual cropping systems. Although further studies are required to assess the context - dependence of these results, our findings may still have important implications for diversified agriculture as they illustrate the potential of targeted cultivars to increase complementarity of species in intercropping, which could be achieved through specific breeding for mixtures., Competing Interests: LS, NE, CS No competing interests declared, (© 2022, Stefan et al.)

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., (Copyright © 2022 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Increasing plant diversity in agricultural systems provides promising solutions for sustainably increasing crop yield. It remains unclear; however, how plant-plant interactions in diverse systems are mediated by plant genetic variation. We conducted a greenhouse experiment in which we grew three varieties of common beans with three companion plant species (chickpeas, sorghum, and sunflower) in different combinations (crop mixtures, bean cultivar mixtures, and monocultures), with and without drought stress. We hypothesized that under drought stress, the effect of companion plant species on bean yield would be mediated by the drought tolerance potential of the species. We further hypothesized that this effect would vary across different bean cultivars. Overall, we show that the effect of companion plant species on bean yield was not influenced by drought stress; instead, it was dependent on the identity of the bean variety. This could partially be explained by variation in growth rate between bean varieties, where the fastest growing variety recorded the highest yield increase in plant mixtures. The effect of companion plant species on chickpea biomass, however, was potentially influenced by chickpea drought tolerance potential; chickpea biomass was recorded to be higher in plant mixtures than in its monoculture under drought conditions. Our study highlights that to develop plant mixtures, it is not only important to consider the functional traits of the interacting plant species, but also those of the different plant varieties. We further suggest that stress tolerance can be a useful trait for initial selection of plant varieties when developing crop mixtures., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Singh, Lehner and Schöb.)

Increasing biodiversity generally enhances productivity through selection and complementarity effects not only in natural, but also in agricultural, systems. However, the quest to explain why diverse cropping systems are more productive than monocultures remains a central goal in agricultural science. In a mesocosm experiment, we constructed monocultures, two- and four-species mixtures from eight crop species with or without fertilizer and both in temperate Switzerland and dry, Mediterranean Spain. We measured physical factors and plant traits and related these in structural equation models to selection and complementarity effects to explain seed yield differences between monocultures and mixtures. Increased crop diversity increased seed yield in Switzerland. This positive biodiversity effect was driven to almost the same extent by selection and complementarity effects, which increased with plant height and specific leaf area (SLA), respectively. Also, ecological processes driving seed yield increases from monocultures to mixtures differed from those responsible for seed yield increases through the diversification of mixtures from two to four species. Whereas selection effects were mainly driven by one species, complementarity effects were linked to larger leaf area per unit leaf weight. Seed yield increases due to mixture diversification were driven only by complementarity effects and were not mediated through the measured traits, suggesting that ecological processes beyond those measured in this study were responsible for positive diversity effects on yield beyond two-species mixtures. By understanding the drivers of positive biodiversity-productivity relationships, we can improve our ability to predict species combinations that enhance ecosystem functioning and can promote sustainable agricultural production., (© 2021 The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Resource allocation to reproduction is a critical trait for plant fitness 1,2 . This trait, called harvest index in the agricultural context 3-5 , determines how plant biomass is converted to seed yield and consequently financial revenue from numerous major staple crops. While plant diversity has been demonstrated to increase plant biomass 6-8 , plant diversity effects on seed yield of crops are ambiguous 9 and dependent on the production syndrome 10 . This discrepancy might be explained through changes in the proportion of resources invested in reproduction in response to changes in plant diversity, namely through changes in species interactions and microenvironmental conditions 11-14 . Here, we show that increasing crop plant diversity from monocultures over two- to four-species mixtures increased annual primary productivity, resulting in overall higher plant biomass and, to a lesser extent, higher seed yield in mixtures compared with monocultures. The difference between the two responses to diversity was due to a reduced harvest index of the eight tested crop species in mixtures, possibly because their common cultivars have been bred for maximum performance in monoculture. While crop diversification provides a sustainable measure of agricultural intensification 15 , the use of currently available cultivars may compromise larger gains in seed yield. We therefore advocate regional breeding programmes for crop varieties to be used in mixtures that should exploit complementarity 16 among crop species., (© 2021. The Author(s), under exclusive licence to Springer Nature Limited.)

Implementing sustainable weed control strategies is a major challenge in agriculture. Intercropping offers a potential solution to control weed pressure by reducing the resources available for weeds; however, available research on the relationship between crop diversity and weed pressure and its consequences for crop yield is not yet fully conclusive. In this study, we performed an extensive intercropping experiment using eight crop species and 40 different species mixtures to examine how crop diversity affects weed communities and how the subsequent changes in weeds influence crop yield. Mesocosm experiments were carried out under field conditions in Switzerland and in Spain, which differ drastically in terms of climate, soil and weed community, and included monocultures, two- and four-species mixtures, and a control treatment without crops. Weed communities were assessed in terms of biomass, species number and evenness, and community composition. Results indicate that intercropping reduces weed biomass and diversity in Spain but not in Switzerland. In Switzerland, despite the lack of a crop diversity effect on weeds, crop yield increased with crop species number. Moreover, in Switzerland, where soil resources were abundant, increasing crop yield correlated with reduced weed biomass. In Spain, where water and nutrients were limited, crop yield was not related to weed biomass or diversity. The presented research applies plant community ecology in the context of agricultural crop production systems. We demonstrate that, in our study, increased crop yield in mixtures was not due to increased weed suppression in diverse crop communities, and so must be the result of other ecological processes. We further show that crop-weed relationships vary across environmental conditions; more specifically, our study shows that weeds are less detrimental to crop yield in harsher environments compared to benign abiotic conditions, where alternative strategies are needed to control weed pressure and ensure the yield benefits provided by intercropping., (© 2021 by the Ecological Society of America.)

Intercropping, i.e., the simultaneous cultivation of different crops on the same field, has demonstrated yield advantages compared to monoculture cropping. These yield advantages have often been attributed to complementary resource use, but few studies quantified the temporal complementarity of nutrient acquisition and biomass production. Our understanding of how nutrient uptake rates of nitrogen (N) and phosphorous (P) and biomass accumulation change throughout the growing season and between different neighbors is limited. We conducted weekly destructive harvests to measure temporal trajectories of N and P uptake and biomass production in three crop species (oat, lupin, and camelina) growing either as isolated single plants, in monocultures or as intercrops. Additionally, we quantified organic acid exudation in the rhizosphere and biological N 2 -fixation of lupin throughout the growing season. Logistic models were fitted to characterize nutrient acquisition and biomass accumulation trajectories. Nutrient uptake and biomass accumulation trajectories were curtailed by competitive interactions, resulting in earlier peak rates and lower total accumulated nutrients and biomass compared to cultivation as isolated single plants. Different pathways led to overyielding in the two mixtures. The oat-camelina mixture was characterized by a shift from belowground temporal niche partitioning of resource uptake to aboveground competition for light during the growing season. The oat-lupin mixture showed strong competitive interactions, where lupin eventually overyielded due to reliance on atmospheric N and stronger competitiveness for soil P compared to oat. Synthesis: This study demonstrates temporal shifts to earlier peak rates of plants growing with neighbors compared to those growing alone, with changes in uptake patterns suggesting that observed temporal shifts in our experiment were driven by competitive interactions rather than active plant behavior to reduce competition. The two differing pathways to overyielding in the two mixtures highlight the importance of examining temporal dynamics in intercropping systems to understand the underlying mechanisms of overyielding., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Engbersen, Brooker, Stefan, Studer and Schöb.)

Intensive agriculture has major negative impacts on ecosystem diversity and functioning, including that of soils. The associated reduction of soil biodiversity and essential soil functions, such as nutrient cycling, can restrict plant growth and crop yield. By increasing plant diversity in agricultural systems, intercropping could be a promising way to foster soil microbial diversity and functioning. However, plant-microbe interactions and the extent to which they influence crop yield under field conditions are still poorly understood. In this study, we performed an extensive intercropping experiment using eight crop species and 40 different crop mixtures to investigate how crop diversity affects soil microbial diversity and activity, and whether these changes subsequently affect crop yield. Experiments were carried out in mesocosms under natural conditions in Switzerland and in Spain, two countries with drastically different soils and climate, and our crop communities included either one, two or four species. We sampled and sequenced soil microbial DNA to assess soil microbial diversity, and measured soil basal respiration as a proxy for soil activity. Results indicate that in Switzerland, increasing crop diversity led to shifts in soil microbial community composition, and in particular to an increase of several plant-growth promoting microbes, such as members of the bacterial phylum Actinobacteria . These shifts in community composition subsequently led to a 15 and 35% increase in crop yield in 2 and 4-species mixtures, respectively. This suggests that the positive effects of crop diversity on crop productivity can partially be explained by changes in soil microbial composition. However, the effects of crop diversity on soil microbes were relatively small compared to the effects of abiotic factors such as fertilization (three times larger) or soil moisture (three times larger). Furthermore, these processes were context-dependent: in Spain, where resources were limited, soil microbial communities did not respond to crop diversity, and their effect on crop yield was less strong. This research highlights the potential beneficial role of soil microbial communities in intercropping systems, while also reflecting on the relative importance of crop diversity compared to abiotic drivers of microbiomes and emphasizing the context-dependence of crop-microbe relationships., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Stefan, Hartmann, Engbersen, Six and Schöb.)

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., Competing Interests: Declaration of competing interest The authors declare no competing financial interests or personal relationships in this paper., (Copyright © 2020 Elsevier B.V. All rights reserved.)

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.

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., (© 2020 by the Ecological Society of America.)

Where interspecific facilitation favors the establishment of high densities of a beneficiary species, strong intraspecific competition may subsequently impede beneficiary performance. Consequently, the negative influence of intraspecific competition between beneficiary individuals could potentially outweigh the positive influence of interspecific facilitation when, for example, higher densities of a beneficiary are negated by the negative effect of crowding on beneficiary reproduction. The aim of this study was, therefore, to examine the impact of an interspecific interaction on the outcome of intraspecific interactions within the context of plant-plant facilitation. We used the cushion-forming Azorella selago and a commonly co-occurring dominant perennial grass species, Agrostis magellanica, on sub-Antarctic Marion Island as a model system. We assessed the impact of an interspecific interaction (between A. selago and A. magellanica) on the outcome of intraspecific interactions (between A. magellanica individuals), by testing if the impact of A. magellanica density on A. magellanica performance is mediated by its interaction with A. selago. We observed evidence for competition among A. magellanica conspecifics, with a decreasing proportion of A. magellanica individuals being reproductive under higher conspecific density. This negative intraspecific effect was greater on A. selago than on the adjacent substrate, suggesting that the facilitative effect of A. selago changes the intensity of intraspecific interactions between A. magellanica individuals. However, experimentally reducing A. magellanica density did not affect the species' performance. We also observed that the effect of A. selago on A. magellanica was positive, and despite the negative effect of intraspecific density on the proportion of reproductive A. magellanica individuals, the net reproductive effort of A. magellanica (i.e., the density of reproductive individuals) was significantly greater on A. selago than on the adjacent substrate. These results highlight that, in abiotically severe environments, the positive effects of interspecific facilitation by a benefactor species may outweigh the negative effects of intraspecific competition among beneficiaries. More broadly, these results suggest that both positive inter- and intraspecific biotic interactions may be key to consider when examining spatial and temporal variation in species' performance., (© 2020 by the Ecological Society of America.)

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., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Positive biodiversity-ecosystem-functioning (BEF) relationships are commonly found in experimental and observational studies, but how they vary in different environmental contexts and under the influence of coexisting life forms is still controversial. Investigating these variations is important for making predictions regarding the dynamics of plant communities and carbon pools under global change. We conducted this study across 433 shrubland sites in northern China. We fitted structural equation models (SEMs) to analyze the variation in the species-richness-biomass relationships of shrubs and herbs along a wetness gradient and general liner models (GLMs) to analyze how shrub or herb biomass affected the species-richness-biomass relationship of the other life form. We found that the positive species-richness-biomass relationships for both shrubs and herbs became weaker or even negative with higher water availability, likely indicating stronger interspecific competition within life forms under more benign conditions. After accounting for variation in environmental contexts using residual regression, we found that the benign effect of greater facilitation by a larger shrub biomass reduced the positive species-richness-biomass relationships of herbs, causing them to become nonsignificant. Different levels of herb biomass, however, did not change the species-richness-biomass relationship of shrubs, possibly because greater herb biomass did not alter the stress level for shrubs. We conclude that biodiversity in the studied plant communities is particularly important for plant biomass production under arid conditions and that it might be possible to use shrubs as nurse plants to facilitate understory herb establishment in ecological restoration., (© The Authors. Ecology published by Wiley Periodicals, Inc. on behalf of Ecological Society of America.)

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., (© 2019 by the Ecological Society of America.)

Diverse experimental plant communities are more productive than monocultures. The increase of this biodiversity effect over time has been attributed to evolutionary selection for complementarity in mixtures. Here we show that evolutionary selection for enhanced net facilitative plant interactions occurred only in mixtures, while evolutionary selection for reduced net competition occurred in mixtures with mixture coexistence history and monocultures with monoculture coexistence history. Widespread declines in natural and agricultural biodiversity could therefore compromise potential evolution of facilitative interactions, that is, cornerstone processes in nature conservation and the development of sustainable agriculture.

Legumes are characterized as keeping stable nutrient supply under nutrient-limited conditions. However, few studies examined the legumes' stoichiometric advantages over other plants across various taxa in natural ecosystems. We explored differences in nitrogen (N) and phosphorus (P) stoichiometry of different tissue types (leaf, stem, and root) between N 2 -fixing legume shrubs and non-N 2 -fixing shrubs from 299 broadleaved deciduous shrubland sites in northern China. After excluding effects of taxonomy and environmental variables, these two functional groups differed considerably in nutrient regulation. N concentrations and N:P ratios were higher in legume shrubs than in non-N 2 -fixing shrubs. N concentrations were positively correlated between the plants and soil for non-N 2 -fixing shrubs, but not for legume shrubs, indicating a stronger stoichiometric homeostasis in legume shrubs than in non-N 2 -fixing shrubs. N concentrations were positively correlated among three tissue types for non-N 2 -fixing shrubs, but not between leaves and non-leaf tissues for legume shrubs, demonstrating that N concentrations were more dependent among tissues for non-N 2 -fixing shrubs than for legume shrubs. N and P concentrations were correlated within all tissues for both functional groups, but the regression slopes were flatter for legume shrubs than non-N 2 -fixing shrubs, implying that legume shrubs were more P limited than non-N 2 -fixing shrubs. These results address significant differences in stoichiometry between legume shrubs and non-N 2 -fixing shrubs, and indicate the influence of symbiotic nitrogen fixation (SNF) on plant stoichiometry. Overall, N 2 -fixing legume shrubs are higher and more stoichiometrically homeostatic in N concentrations. However, due to excess uptake of N, legumes may suffer from potential P limitation. With their N advantage, legume shrubs could be good nurse plants in restoration sites with degraded soil, but their P supply should be taken care of during management according to our results.

Arenaria polytrichoides (Caryophyllaceae) is a common cushion plant occurring at high elevations in the Himalaya-Hengduan Mountains, SW China. It frequently has other non-cushion species growing within its canopy, forming a contrast with the surrounding areas because it creates patches of higher diversity and greater biomass. In this study, we examined the relationship between the cushions and associated non-cushion species along a gradient of cushion size. A total of 200 A. polytrichoides individuals were selected to fit four size classes. Field measurements were carried out to assess canopy structure, functional traits relevant to growth and reproduction, and soil quality below cushions along the size gradient. Furthermore, the size effect of cushions on the richness and abundance of species and biomass production was also examined. All the morphological variables examined exhibited a positive correlation with cushion size, as did the nutrients under cushions. Large and compact cushions were associated with higher soil nutrient contents compared with small and loose cushions. As a result of these biogenic environmental changes, there was a stronger facilitation effect performed by large cushions. Data pertaining to functional traits revealed that large cushions benefit from the enhanced resources within their compact structure and exhibit greater fitness and a higher reproductive output than small cushions. Our data indicated that interactions occur between cushion species and other plants depending on the size of the cushions, probably because of the greater heterogeneity of conditions beneath larger cushions. These findings provide a clear demonstration of the generally overlooked importance of the traits of nurse plants, such as size and age, in terms of their facilitative effects.

The stress-gradient hypothesis predicts a higher frequency of facilitative interactions as resource limitation increases. Under severe resource limitation, it has been suggested that facilitation may revert to competition, and identifying the presence as well as determining the magnitude of this shift is important for predicting the effect of climate change on biodiversity and plant community dynamics. In this study, we perform a meta-analysis to compare temporal differences of species diversity and productivity under a nurse plant ( Retama sphaerocarpa ) with varying annual rainfall quantity to test the effect of water limitation on facilitation. Furthermore, we assess spatial differences in the herbaceous community under nurse plants in situ during a year with below-average rainfall. We found evidence that severe rainfall deficit reduced species diversity and plant productivity under nurse plants relative to open areas. Our results indicate that the switch from facilitation to competition in response to rainfall quantity is nonlinear. The magnitude of this switch depended on the aspect around the nurse plant. Hotter south aspects under nurse plants resulted in negative effects on beneficiary species, while the north aspect still showed facilitation. Combined, these results emphasize the importance of spatial heterogeneity under nurse plants for mediating species loss under reduced precipitation, as predicted by future climate change scenarios. However, the decreased water availability expected under climate change will likely reduce overall facilitation and limit the role of nurse plants as refugia, amplifying biodiversity loss.

Recent studies have shown that accounting for intraspecific trait variation (ITV) may better address major questions in community ecology. However, a general picture of the relative extent of ITV compared to interspecific trait variation in plant communities is still missing. Here, we conducted a meta-analysis of the relative extent of ITV within and among plant communities worldwide, using a data set encompassing 629 communities (plots) and 36 functional traits. Overall, ITV accounted for 25% of the total trait variation within communities and 32% of the total trait variation among communities on average. The relative extent of ITV tended to be greater for whole-plant (e.g. plant height) vs. organ-level traits and for leaf chemical (e.g. leaf N and P concentration) vs. leaf morphological (e.g. leaf area and thickness) traits. The relative amount of ITV decreased with increasing species richness and spatial extent, but did not vary with plant growth form or climate. These results highlight global patterns in the relative importance of ITV in plant communities, providing practical guidelines for when researchers should include ITV in trait-based community and ecosystem studies., (© 2015 John Wiley & Sons Ltd/CNRS.)

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 β-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.

Intercropping 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 species – for 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.

Biodiversity regulates ecosystem functions such as productivity, and experimental studies of species mixtures have revealed selection and complementarity effects driving these responses. However, the impacts of intraspecific genotypic diversity in these studies are unknown, despite it forming a substantial part of the biodiversity. In a glasshouse experiment we constructed plant communities with different levels of barley (Hordeum vulgare) genotype and weed species diversity and assessed their relative biodiversity effects through additive partitioning into selection and complementarity effects. Barley genotype diversity had weak positive effects on aboveground biomass through complementarity effects, whereas weed species diversity increased biomass predominantly through selection effects. When combined, increasing genotype diversity of barley tended to dilute the selection effect of weeds. We interpret these different effects of barley genotype and weed species diversity as the consequence of small vs large trait variation associated with intraspecific barley diversity and interspecific weed diversity, respectively. The different effects of intra- vs interspecific diversity highlight the underestimated and overlooked role of genetic diversity for ecosystem functioning., (© 2014 The Authors New Phytologist © 2014 New Phytologist Trust.)

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., (© 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.)

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., (© 2013 The Authors. New Phytologist © 2013 New Phytologist Trust.)

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., (© 2013 John Wiley & Sons Ltd/CNRS.)

Here, 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., (© 2012 The Authors. New Phytologist © 2012 New Phytologist Trust.)