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Interactions among species are a fundamental aspect of biodiversity and drive ecosystem functioning and services. Species interactions include direct (pairwise) interactions among two species and indirect interactions that occur when a third species interacts and changes the pairwise direct interaction. In a three‐species interaction network, these interactions can be transitive (where one species outperforms all others) or intransitive (where each species outperforms another). Here, we investigate how direct and indirect interactions influence ecosystem functions in crop systems and how diversification and evolutionary adaptation can influence those interactions and therefore ecosystem functions.A common garden experiment was conducted with crop communities in monocultures, 2‐ and 3‐species mixtures that had either a common or no coexistence history (i.e. co‐adaptation) for the three previous years. Net, direct and indirect interaction intensities were estimated and compared between the diversity levels and coexistence histories. Furthermore, species interaction networks were inspected for transitive/intransitive interactions.We found evidence for less intense competition in mixtures and for reduced negative direct interaction intensity and enhanced facilitative effects upon co‐adaptation. We could further show that indirect interactions were generally less important for co‐adaptation than direct interactions. Additionally, we showed that co‐adaptation has the potential to shift interactions in the species interaction networks from competitive intransitive into pairwise competitive interactions where interactions occurred mainly between two species.Synthesis. Co‐adapted crop species with reduced negative interactions might have the potential to enhance productivity, especially in more diverse cropping systems. This supports the notion that intercropping is a vital part towards a more sustainable agriculture and one with further yield potential when developing cultivars optimised for growth in mixtures. [ABSTRACT FROM AUTHOR]

Inspired by grassland biodiversity experiments studying the impact of plant diversity on primary productivity, the Crop Diversity Experiment setup in 2018 aimed at testing whether these biodiversity benefits also hold for annual crop systems and whether crop mixtures also achieved transgressive overyielding, i.e. yield in mixture that was higher than the most productive monoculture. The first 3 years of the experiment demonstrated that crop mixtures do not only increase yield compared with an average monoculture but often also compared with the highest yielding monoculture. The crop diversity effects were stronger under more stressful environmental conditions and were often achieved in mixtures with legume crops. However, we observed transgressive overyielding also under favorable conditions and in mixtures without legumes. With our investigation of the underlying mechanisms of the yield benefits we found both direct complementarities between crop species and indirect effects via other organisms. The former included chemical, spatial and temporal complementarity in N uptake, complementary root distribution leading to complementary water uptake, as well as spatial and temporal complementarity in light use. Among the indirect mechanisms we identified complementary suppression of weeds and more abundant plant growth-promoting microbes in crop mixtures, apart from complementarity in pest and disease suppression not yet studied in the Crop Diversity Experiment but demonstrated elsewhere. In consequence, the Crop Diversity Experiment supports not only the assumption that the ecological processes identified in biodiversity experiments also hold in crop systems, but that diversification of arable crop systems provides a valuable tool to sustainably produce food. [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]

Interactions among neighbours can alter demography and traits of commingled species via adaptation or plasticity in phenotypic expression, and understanding these two mechanisms in diverse communities is important for determining the ecological and evolutionary consequences of plant–plant interactions.We reciprocally transplanted perennial species (Arenaria armerina and Festuca indigesta) among patches of two foundation shrub species and open ground to assess whether origin microsite (defined as the spatially distinct abiotic and biotic conditions associated with the two shrubs and open ground) determines germination, recruitment and growth that, in turn, promotes fine‐scale distribution of species among microsites. In addition, we tested the effect of origin microsite on traits, competitive ability, drought tolerance and outlier loci to assess whether origin microsite conditions drove differences in traits, strategies and adaptive loci.Germination was consistently greater for seeds planted back into their origin microsite relative to seeds sourced from foreign microsites, although this effect was weakened for recruitment. Plant growth was best in open sites regardless of origin microsite. In the greenhouse, A. armerina had conserved traits within origin microsite but distinct trait values among origins, specifically plants from the most productive microsite (e.g. sufficient light, high nutrients and improved water availability) had distinct trait values. Festuca indigesta had conserved trait responses among microsites while within microsite, individuals had significant trait plasticity to different environmental conditions. The combined field and greenhouse results suggest that fine‐scale distributions are supported by local adaptation among microsites of A. armerina and phenotypic plasticity of F. indigesta.Synthesis. Adaptation or plasticity in phenotypic expression has different implications for demographic rate and persistence of species in changing environments. Local adaptation to neighbours suggests that reductions in foundation species diversity could concomitantly lead to reduced genetic diversity of commingled species while a plastic response indicates a more robust and broad response to changing climatic and biotic conditions. [ABSTRACT FROM AUTHOR]

Questions: Positive and negative associations among species influence the structure of plant communities. Yet how these plant associations are assembled at the community level is poorly understood. We propose a new approach that combines spatial ecology, network theory and trait‐based ecology to examine the assembly of plant–plant associations at the community level. Location: Gemmipass, Swiss Alps. Methods: We fully mapped an alpine plant community at the individual plant level, recording both plant coordinates and functional traits for each individual. We identified non‐random species associations using spatial point‐pattern analysis and partialled out the effect of abiotic heterogeneity. We then analysed the plant network structure and used plant traits to predict species associations. Results: We identified 36 significant spatial associations between plant species, 34 positive and two negatives. Dominant, stress‐tolerant species such as Dryas octopetala, Linaria alpina and Leontodon montanus were highly connected in the network, whereas rare, water‐ and nutrient‐demanding species such as Saxifraga aizoides, Galium anisophyllon and Thymus praecox were less connected compared to random expectation. The plant network was clustered, meaning that species were overall more connected among each other than expected by chance. Conclusions: Positive associations among species characterized the studied plant community. Besides the primary effect of associations of the "foundation" species D. octopetala with other species, these "subordinate" plants were also associated with each other. Our study reveals the assembly of plant communities as driven by positive associations among stress‐tolerant pioneer species, highlighting their role in supporting the cohesiveness of alpine plant communities. We unveiled the spatial organization of an alpine plant community by combining spatial pattern analysis and network analysis. Plant species were highly connected through many positive interactions. Dominant, stress‐tolerant species were the most important plants for supporting the network structure of plant community. [ABSTRACT FROM AUTHOR]

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Question: In plant communities, the presence of a species has consequences for other species, with some being competitively excluded while others benefit from the close vicinity of neighbours. Even though such specificity in plant interactions is common and known, there is no empirical assessment of the mechanisms that would help us understand its importance for plant diversity. Here we asked whether analysing spatial associations between plant traits known to affect the environment (i.e. effect traits) and those known to respond to the environment (i.e. response traits) might explain plant-plant interactions and their role in community assembly. Location: Sierra Nevada Mountains, Spain. Methods: In a field study, we addressed the specificity of plant-plant interactions by quantifying effect traits of three co-occurring cushion-forming species and response traits of their associated plant assemblages. Traits were measured at the individual level and then aggregated to trait metrics (mean, range, dispersion) at the plot level. Finally, plot-level metrics of effect traits were related to response traits and the species composition of plant communities. Results: Each cushion-forming species had a distinctive combination of effect traits and harboured a unique plant community with an exclusive composition of response traits. With multivariate statistics we showed that differences in effect traits (branch density and canopy height) among and within cushion species significantly explained response traits (specific leaf area, leaf dry matter content) of associated species and the local-scale species composition. Conclusions: Using effect and response traits measured at the individual level, we provide a mechanistic understanding of the species specificity of plant interactions and demonstrate how important such specificity is for species diversity in an ecosystem. [ABSTRACT FROM AUTHOR]

Understanding species coexistence and regulation of biodiversity are major research challenges, yet there is no consensus on the effects of diversity on diversity, including their mediation through plant-plant interactions., We examined how the diversity of recipient communities impacted on the establishment of colonising species. We ran a greenhouse-based community experiment, creating artificial arable crop communities with varying levels of barley genotype and weed species diversity, analysed with structural equation modelling of responses across trophic levels., Colonising arable weed species establishment was significantly reduced when the recipient communities' weed species richness, but not barley genotype richness, increased. Recipient plant communities with higher species richness occupied more total niche space and showed stronger competition with colonising species. In contrast, recipient plant communities with higher genotype richness, even though they had slightly greater niche space occupancy, had no significant effects on the colonisation of additional species., Responses were species specific: colonising common weed species establishment was less affected by the recipient plant communities than that of colonising rare weed species. Strong responses of colonising rare species were related to the abiotic rather than the biotic environment created by the recipient communities. The stronger response of colonising rare species suggests greater niche overlap with and competitive inferiority to the recipient plant community, whereas the relative independence of colonising common species indicates less niche overlap and/or competitive equality or superiority., We observed a negative species diversity effect on colonising species, with the effect size increasing with increasing rarity of the colonising species. This differential responsiveness of colonising rare and common arable weed species to the presence of and diversity in the recipient arable plant communities may be crucial in understanding rarity and may result from differential evolutionary pathways in response to interactions. [ABSTRACT FROM AUTHOR]

In facilitative interactions, the beneficiary feedback effect ( BFE) has been defined as the effect of beneficiary species (facilitated species) on their benefactor. BFEs have been shown to be dependent on environmental conditions and the composition of the beneficiary community. In alpine cushion systems, BFEs are more negative with more abundant, diverse and phylogenetically aggregated communities of beneficiary species., We tested the hypothesis that the functional composition of the beneficiary communities correlates with the direction and strength of BFE received by alpine cushion benefactors and specifically that a more negative BFE would occur with increasing density of graminoids and a more positive BFE would occur with increasing density of forbs and legumes. Additionally, we predicted that the negative BFE of graminoids would increase with increasing summer aridity., We used a data base of alpine cushion communities from 30 sites throughout the world to assess the overall relationship between the composition of beneficiary communities and the total flower density of cushion benefactors, and its variation with increasing drought. Additionally, in order to assess more precisely the role of the functional composition of the beneficiary communities on BFE in a very dry site with cushion benefactors exhibiting contrasting functional compositions of beneficiary communities, we also designed a field study in the Qilian Shan mountain range (China). At this site with a highly continental climate, we compared the number of flowers and fruits of different phenotypes of the alpine cushion species Thylacospermum caespitosum hosting numerous graminoids, numerous forbs or very few beneficiary species., In the intercontinental study, we found a negative relationship between graminoids and cushion benefactor flower density but no effect of other functional groups. The negative BFE of graminoids increased with increasing summer drought. In the dry Qilian Shan range, we found both a negative effect of graminoids on total flower density and a positive effect of forbs on flower density and fruit set., Our study indicates that the context dependence of BFE may be partially explained by the composition of beneficiary communities and in particular the negative effect of graminoids. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Background: Neighbour-removal experiments (NRE) and spatial pattern analyses (SPA) are commonly used methods to investigate plant–plant interactions. Although they address the same issue, they measure different aspects of plant interactions: experiments indicate contemporary processes, whilst observations of spatial patterns integrate the results of interactions that have prevailed in the past. Aim: The aim of this study was to propose a new conceptual approach that takes into account the chronological order between processes and the arising patterns i.e., the time lag between neighbour effects quantified with NRE and SPA, to detect shifts in the balance of plant interactions due to current environmental change. Methods: This conceptual approach was applied to alpine snowbeds. Data from NRE were used to calculate the importance index of current neighbour interactions. Spatial patterns were quantified using variance ratio statistics and were assumed to reflect historic interactions. Results: The results of the two approaches showed a consistent difference in the prevailing type of plant interactions and suggested a shift towards competition in recent times. Conclusions: The simultaneous application of NRE and SPA allows the detection of a recent shift in the balance of plant interactions and provides a deeper and more accurate insight into the temporal dynamics of plant communities that could not be gained using one method alone. [ABSTRACT FROM PUBLISHER]

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. [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]