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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. [ABSTRACT FROM AUTHOR]

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]

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. [ABSTRACT FROM AUTHOR]

More diverse plant communities are generally more productive than monocultures. This benefit of species diversity is supposed to stem from resource partitioning of species in mixtures where different species use the resources spatially, temporally, or chemically in distinct ways. With respect to water, the simultaneous cultivation of crops with distinct water uptake patterns might reduce niche overlaps and thus result in higher productivity. However, little is known about whether and how spatial water uptake patterns of crop species differ among different planting arrangements and whether these changes result in increased niche partitioning and explain overyielding in mixtures.Stable isotopes of water and a Bayesian model were used to investigate the spatial water uptake patterns of six different crop species and how these patterns change depending on the planting arrangement (monocultures vs mixtures). Niche overlaps and niche widths in spatial water uptake were compared among the different crop diversity levels and linked to productivity. Furthermore, spatial water uptake was related to competition intensity and overyielding in mixtures.We found evidence for increased niche partitioning in spatial water uptake, and therefore complementary spatial root distributions of crop species, and higher expected productivity in mixtures compared to expected productivity in monocultures both due to inherent species‐level differences in water uptake and plasticity in the water uptake pattern of species. We also found a significant relationship of competition and overyielding with observed patterns in spatial water uptake. These results suggest that competition was most intense in shallow soil layers and enhanced overyielding was related to a gradual increase of water uptake in deeper soil layers. Thus, overyielding might be related to a more complete spatial exploitation of available water sources.Synthesis. Differences in spatial water uptake and niche partitioning of intercropped species, driven most likely by a complementary spatial root distribution, might explain why mixtures outperform monocultures. These findings underpin the potential of intercropping systems for a more sustainable agriculture with a more efficient use of soil resources and hence reduced input demands. [ABSTRACT FROM AUTHOR]

Facilitation is an interaction where one species (the benefactor) positively impacts another (the beneficiary). However, the reciprocal effects of beneficiaries on their benefactors are typically only documented using short‐term datasets. We use Azorella selago, a cushion plant species and benefactor, and a co‐occurring grass species, Agrostis magellanica, on sub‐Antarctic Marion Island, comparing cushion plants and the grasses growing on them over a 13‐year period using a correlative approach. We additionally compare the feedback effect of A. magellanica on A. selago identified using our long‐term dataset with data collected from a single time period. We hypothesized that A. selago size and vitality would be negatively affected by A. magellanica cover and that the effect of A. magellanica on A. selago would become more negative with increasing beneficiary cover and abiotic‐severity, due to, e.g. more intense competition for resources. We additionally hypothesized that A. magellanica cover would increase more on cushion plants with greater dead stem cover, since dead stems do not inhibit grass colonization or growth. The relationship between A. magellanica cover and A. selago size and vitality was not significant in the long‐term dataset, and the feedback effect of A. magellanica on A. selago did not vary significantly with altitude or aspect; however, data from a single time period did not consistently identify this same lack of correlation. Moreover, A. selago dead stem cover was not significantly related to an increase in A. magellanica cover over the long term; however, we observed contrasting results from short‐term datasets. Long‐term datasets may, therefore, be more robust (and practical) for assessing beneficiary feedback effects than conventional approaches, particularly when benefactors are slow‐growing. For the first time using a long‐term dataset, we show a lack of physical cost to a benefactor species in a facilitative interaction, in contrast to the majority of short‐term studies. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Abstract: Questions: The exceptional occurrence of tall rain forest patches on foggy coastal mountaintops, surrounded by extensive xerophytic shrublands, suggests an important role of plant–plant interactions in the origin and persistence of these patches in semi‐arid Chile. We asked whether facilitation by shrubs can explain the growth and survival of rain forest tree species, and whether shrub effects depend on the identity of the shrub species itself, the drought tolerance of the tree species and the position of shrubs in regard to wind direction. Location: Open area–shrubland–forest matrix, Fray Jorge Forest National Park, Chile. Methods: We recorded survival after 12 years of a ~3600 tree saplings plantation (originally ~30‐cm tall individuals) of Aextoxicon punctatum, Myrceugenia correifolia and Drimys winteri placed outside forests, beneath the shrub Baccharis vernalis, and in open (shrub‐free) areas. We assessed the effects of neighbouring shrubs and soil humidity on survival and growth along a gradient related to the direction of fog movement. Results: B. vernalis had a clear facilitative effect on tree establishment and survival since, after ~12 years, saplings only survived beneath the shrub canopy. Long‐term survival strongly depended on tree species identity, drought tolerance and position along the soil moisture gradient, with higher survival of A. punctatum (>35%) and M. correifolia (>14%) at sites on wind‐ and fog‐exposed shrubland areas. Sites occupied by the shrub Aristeguietia salvia were unsuitable for trees, presumably due to drier conditions than under B. vernalis. Conclusions: Interactions between shrubs and fog‐dependent tree species in dry areas revealed a strong, long‐lasting facilitation effect on planted tree's survival and growth. Shrubs acted as benefactors, providing sites suitable for tree growth. Sapling mortality in the shrubland interior was caused by lower soil moisture, the consequence of lower fog loads in the air and thus insufficient facilitation. While B. vernalis was a key ecosystem engineer (nurse) and intercepted fog water that dripped to trees planted underneath, drier sites with A. salvia were unsuitable for trees. Consequently, nurse effects related to water input are strongly site and species specific, with facilitation by shrubs providing a plausible explanation for the initiation of forest patches in this semi‐arid landscape. [ABSTRACT FROM AUTHOR]

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]

Plant interactions are fundamental processes for structuring plant communities and are an important mechanism governing the response of plant species and communities to environmental changes. Thus, understanding the role played by the interaction network in modulating the impact of environmental changes on plant community composition and diversity is crucial. Here, we aimed to develop a new analytical and conceptual framework to evaluate the responses of plant communities to environmental changes., This framework uses functional traits as sensitivity measures for simulated environmental changes and assesses the consequences of microhabitat loss. We show here its application to an alpine plant community where we recorded functional traits [specific leaf area ( SLA) and leaf dry matter content ( LDMC)] of all plants associated with three foundation species or the surrounding open areas. We then simulated primary species loss based on different scenarios of environmental change and explored community persistence to the loss of foundation species., Generally, plant community responses differed among environmental change scenarios. In a scenario of increasing drought alone (i.e. species with lower LDMC were lost first) or increasing drought with increasing temperature (i.e. species with lower LDMC and higher SLA were lost first), the plant community resisted because drought-tolerant foundation species tolerated those deteriorating conditions. However, in scenarios with increasing nitrogen input (i.e. species having lower SLA were lost earlier), foundation species accelerated species loss due to their early primary extinctions and the corresponding secondary extinctions of species associated to their microhabitat., The resistance of a plant community depends on the driver of environmental change, meaning that the prediction of the fate of this system is depending on the knowledge of the main driver of environmental change. Our framework provides a mechanistic understanding of an ecosystem response to such environmental changes thanks to the integration of biology-informed criteria of species sensitivities to environmental factors into a network of interacting species., A is available for this article. [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]

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]

Questions What is the community-level consequence of biotic processes, in particular the importance of facilitation, in determining patterns of diversity in alpine plant communities of the Himalayan Hengduan Mountains? Does facilitation intensity change with environmental severity, and are these changes due to environmental severity or nurse trait effects? Location Eleven alpine plant communities dominated by cushion plants in the Himalayan Hengduan Mountains, within the mountain system of south-central China. Methods We determined plant species richness and abundance in habitats created by cushions and cushion-free areas, and assessed the cushion effects on species richness and abundance with rarefaction curves and the relative interaction index, respectively. We examined the relationship between cushion effects on diversity and habitat severity and tested if changes in the net cushion effects along the severity gradient were due to a change in the performance of species without cushions (environmental severity effect) and/or with cushions (nurse trait-mediated effect). Results The presence of cushion plants increased species richness and abundance in most studied plant communities. The net facilitation effect of cushions increased with increasing habitat severity, an effect that could be attributed mainly to a reduction in species richness in cushion-free areas with increasing severity. However, the changes in magnitude of facilitation of particular cushion species growing in high and low severity habitats could be attributed to both environmental severity and neighbour trait effects. Despite the overall positive effect of cushion plants on plant species richness, not all cushion species contributed to higher species richness. The deviation of individual cushion species from the general pattern of increasing facilitation with increasing severity indicated the species specificity of facilitation depends on the traits of nurse species. Conclusion The presence of cushion plants generally increased species richness in alpine plant communities of the Himalayan Hengduan Mountains, with the importance of the cushion effect increasing with habitat severity due to a buffering effect by cushions of the negative effect of habitat severity on species richness observed in cushion-free areas. This indicates the pivotal role of facilitative interactions among plant species in supporting high diversity in these severe environments. [ABSTRACT FROM AUTHOR]

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. [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]

Facilitation is known as the positive effect of one species (benefactor) on associated neighbouring species (beneficiaries). Although the beneficial part of this interaction has received considerable research interest, there is a gap of knowledge on the bidirectional nature of these interactions; in particular, the physiological and fitness consequences for both beneficiaries and benefactors., Alpine cushion plants are generally strong benefactors, increasing species richness and abundance on a global scale, and provide a suitable system to study the physiological effects of bidirectional interactions and its consequences for reproduction. Current knowledge suggests that species improve their fitness when associated to a benefactor cushion species, whereas cushions may receive predominantly negative feedbacks., We measured physiological and reproductive traits of the cushion species Arenaria tetraquetra ssp. amabilis and three other forbs ( Eryngium glaciale, Lotus corniculatus ssp. glacialis and Plantago nivalis) in the dry Sierra Nevada Mountains, southern Spain. All four species were studied either when growing alone or when the three forbs were associated with the cushion plant., The three forb species improved their water status when associated with the cushion, and Lotus and Plantago significantly increased their seed set. In contrast, Arenaria showed poorer water status and reduced flower density and seed set with increasing cover of beneficiary species. There was a clear relationship between physiological and reproductive traits in Arenaria growing without beneficiaries but not in Arenaria with beneficiaries. Control cushions (without beneficiary species) showed increased seed set and seed mass with increasing photosynthetic and water use efficiencies, respectively, the latter being positively related to leaf nitrogen content. In contrast, cushions with a large cover of beneficiary species did not show such relations. The missing relationship between physiological and reproductive traits for these facilitating cushions indicates that reproductive output in heavily colonized cushions may not be directly related to the plant physiological status and the availability of resources., Our results revealed the antagonistic behaviour underlying the interaction between beneficiary species and facilitating cushion plants, similar to parasitic interactions. They also provide indications for changes in cushions' resource allocation pattern in response to colonization by beneficiaries. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Altitudinal gradients provide a useful space-for-time substitution to examine the capacity for plant competition and facilitation to mediate responses to climate change. Decomposing net interactions into their facilitative and competitive components, and quantifying the performance of plants with and without neighbours along altitudinal gradients, may prove particularly informative in understanding the mechanisms behind plant responses to environmental change., To decouple the inherent responses of species to climate from the responses of plant-plant interactions to climate, we conducted a meta-analysis. Using data from 16 alpine experiments, we tested if changes in net interactions along altitudinal gradients were due to a change in the performance of target species without neighbours (i.e. environmental severity effects only) or with neighbours (neighbour trait mediated effects)., There was a global shift from competition to facilitation with increasing altitude driven by both environmental severity and neighbour trait effects. However, this global pattern was strongly influenced by the high number of studies in mesic climates and driven by competition at low altitude in temperate climates (neighbour trait effect), and facilitation at high altitude in arctic and temperate climates (environmental severity effect)., In Mediterranean systems, there was no significant effect of competition, and facilitation increased with decreasing altitude. Changes in facilitation with altitude could not unambiguously be attributed to either neighbour trait effects or environmental severity effects, probably because of the opposing stress gradients of cold and aridity in dry environments., Partitioning net interactions along altitudinal gradients led to the prediction that climate change should decrease the importance of facilitation in mesic alpine communities, which might in turn exacerbate the negative effects of climate change in these regions. In xeric climates, the importance of facilitation by drought-tolerant species should increase at low altitudes which should mitigate the negative effect of climate change. However, the importance of facilitation by cold-tolerant species at high altitudes may decrease and exacerbate the effects of climate change. [ABSTRACT FROM AUTHOR]

Background:In those alpine regions where growing season precipitation is decreasing due to climate change, the capacity of soils to retain water may become an important factor for the persistence of plant species. However, the importance of soil water storage capacity (WSC) for plant species diversity has not been studied so far. Aims:We aim to evaluate the relevance of WSC for species diversity of alpine plant communities in relation to temperature and length of growing season. Methods:Species diversity was determined in 150 plots from a broad range of alpine vegetation types in the calcareous western part of the central Swiss Alps. WSC of soil cores sampled in every plot was determined, as well as rooting zone temperature and snowmelt date. Linear mixed models were used to assess the relationship between environmental data and species diversity. Results:Species diversity was most strongly and positively related to WSC, followed by mean daily minimum temperature (Tmin) of the growing season. Species diversity was significantly related to date of snowmelt only in sites with high WSC and/or Tmin. Conclusions:WSC represents an integrative measure for habitat quality and accounts for differences in species diversity within the study region. In order to understand and predict responses of plant species to climate change in high mountain regions, it may be crucial to also take changes in plant water supply into account. [ABSTRACT FROM PUBLISHER]

Facilitation by nurse plants plays an important role in determining community composition in severe environments. Although the unidirectional effect of nurses on beneficiary species has received considerable research interest, nurse-mediated interactions among beneficiary species (so-called indirect interactions) are less known. Consequently, community composition in nurse plant systems is generally considered as a simple consequence of the facilitative effect of the nurse even though beneficiary species may significantly contribute to community assembly and modulate the direct nurse effects on the community. In an observational study we assessed nurse effects and nurse-mediated beneficiary interactions in two contrasting nurse plant systems in dry environments using a newly developed framework. We quantified plant-plant interaction intensity using the relative interaction index (RII) at the community and species level for three Retama sphaerocarpa shrub size-classes in a semiarid shrubland and four Arenaria tetraquetra cushion plant communities differing in aspect and elevation in dry alpine gravel habitats. The observed RII was split into nurse and beneficiary effects, and related to individual mass, species frequency and abundance using generalized linear mixed models. Results showed predominantly positive nurse effects and negative beneficiary interactions. The effect size of nurse plants, however, was significantly higher than the effect size of beneficiary species in both systems. Individual plant mass and abundance of species was dependent on the combined effects of nurse and beneficiary species whereas species occurrence was related to nurse effects only. Despite evident differences, the semiarid and alpine nurse plant systems showed strong functional parallelisms. We found interdependence between the effects of nurse and beneficiary species on beneficiary plant assemblages emphasizing their combined role on community assembly in both systems. Our results highlight the need to consider indirect interactions to understand fully plant community dynamics. [ABSTRACT FROM AUTHOR]

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

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. [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]

Alpine snowbeds are habitats where the major limiting factors for plant growth are herbivory and a small time window for growth due to late snowmelt. Despite these limitations, snowbed vegetation usually forms a dense carpet of palatable plants due to favourable abiotic conditions for plant growth within the short growing season. These environmental characteristics make snowbeds particularly interesting to study the interplay of facilitation and competition. We hypothesised an interplay between resource competition and facilitation against herbivory. Further, we investigated whether these predicted neighbour effects were species-specific and/or dependent on ontogeny, and whether the balance of positive and negative plant-plant interactions shifted along a snowmelt gradient. We determined the neighbour effects by means of neighbour removal experiments along the snowmelt gradient, and linear mixed model analyses. The results showed that the effects of neighbour removal were weak but generally consistent among species and snowmelt dates, and depended on whether biomass production or survival was considered. Higher total biomass and increased fruiting in removal plots indicated that plants competed for nutrients, water, and light, thereby supporting the hypothesis of prevailing competition for resources in snowbeds. However, the presence of neighbours reduced herbivory and thereby also facilitated survival. For plant growth the facilitative effects against herbivores in snowbeds counterbalanced competition for resources, leading to a weak negative net effect. Overall the neighbour effects were not species-specific and did not change with snowmelt date. Our finding of counterbalancing effects of competition and facilitation within a plant community is of special theoretical value for species distribution models and can explain the success of models that give primary importance to abiotic factors and tend to overlook interrelations between biotic and abiotic effects on plants. [ABSTRACT FROM AUTHOR]

Alpine snowbeds are characterised by a very short growing season. However, the length of the snow-free period is increasingly prolonged due to climate change, so that snowbeds become susceptible to invasions from neighbouring alpine meadow communities. We hypothesised that spatial distribution of species generated by plant interactions may indicate whether snowbed species will coexist with or will be out-competed by invading alpine species -- spatial aggregation or segregation will point to coexistence or competitive exclusion, respectively. We tested this hypothesis in snowbeds of the Swiss Alps using the variance ratio statistics. We focused on the relationships between dominant snowbed species, subordinate snowbed species, and potentially invading alpine grassland species. Subordinate snowbed species were generally spatially aggregated with each other, but were segregated from alpine grassland species. Competition between alpine grassland and subordinate snowbed species may have caused this segregation. Segregation between these species groups increased with earlier snowmelt, suggesting an increasing importance of competition with climate change. Further, a dominant snowbed species (Alchemilla pentaphyllea) was spatially aggregated with subordinate snowbed species, while two other dominants (Gnaphalium supinum and Salix herbacea) showed aggregated patterns with alpine grassland species. These dominant species are known to show distinct microhabitat preferences suggesting the existence of hidden microhabitats with different susceptibility to invaders. These results allow us to suggest that alpine snowbed areas are likely to be reduced as a consequence of climate change and that invading species from nearby alpine grasslands could outcompete subordinate snowbed species. On the other hand, microhabitats dominated by Gnaphalium or Salix seem to be particularly prone to invasions by non-snowbed species. [ABSTRACT FROM AUTHOR]

Over the last 20 years, several studies comparing recent survey data with historical data from the early 20th century documented an increase in species numbers on high mountain summits of the European Alps. This increase has more or less explicitly been attributed to an upward migration of plant species due to anthropogenic climate warming. However, a reconsideration of the historical and recent data has revealed that more than 90% of the recent species occurrences on mountain summits concern species that were already present at the same or even at higher altitudes within the study region at the time of the historical surveys. This finding suggests that suitable habitats already occurred on these summits under the mesoclimatic conditions prevailing at the beginning of the 20th century and that these habitats were, at least in part, occupied by these plant species. Consequently, the observed increase in species number during the last century does not require the additional temperature increase due to anthropogenic climate change. We therefore consider the phenomenon of increasing species number on high mountain summits to be primarily the result of a natural dispersal process that was triggered by the temperature increase at the end of the Little Ice Age and that is still in progress mostly due to the dispersal limitation of the species involved. Since both the natural dispersal process and a potential upward migration due to anthropogenic climate warming would take place at the same time, we suggest seeding and transplanting experiments in order to assess their respective roles in the increase in species number on mountain summits. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The authors discuss the impact of ontogeny on the plant-plant interactions along the environmental stress gradients. They relate the study by P. C. le Roux and colleagues which examines the interactions between Azorella selago and Agrostis magellanica in a subantarctic island ecosystem. They note that the study shows that the outcome of plant-plant interactions along stress gradients can be affected by ontogeny.

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. [ABSTRACT FROM AUTHOR]

Abstract: The impact of plant–plant interactions on species diversity patterns has been broadly addressed in stressful environments, such as alpine ecosystems, where foundation species promote species richness by creating habitat for other species. However, foundation species with contrasting phenotypes might modify the microhabitat differently, which would alter the subordinate community composition, and coincide with distinct feedback effects of those subordinate species on the foundation species. However, the precise interaction mechanisms that facilitate species are not fully understood, especially the relative contribution of above‐ and below‐ground compartments of foundation species to subordinate species and the potential feedbacks they receive. We explored whether two contrasted canopy phenotypes (tight and loose) of the shrub Cytisus galianoi differed in their effects on the microhabitat and on subordinate plant community composition in a dry subalpine system. We also experimentally distinguished the relative contribution of above‐ (canopy) and below‐ground (soil) effects of C. galianoi on the most frequent subordinate species, Festuca indigesta, as well as the reciprocal effects of F. indigesta on C. galianoi. We performed observational and manipulative experiments to assess the influence of phenotypic differences of the shrub on understorey microhabitat and subordinate plant community composition. Reciprocal effects were assessed by removing either F. indigesta from the understorey of the two shrub phenotypes or the C. galianoi canopy from the immediate vicinity of F. indigesta. The two C. galianoi phenotypes differed in mean values of functional traits (like stem density or plant height), modified their understorey microhabitats differently and hosted distinct subordinate communities. Loose phenotypes had more positive effects on community composition and diversity than tight phenotypes. In addition, tight phenotypes simultaneously showed both more positive above‐ground and more negative below‐ground effects on F. indigesta than loose phenotypes. There were no significant feedback effects of F. indigesta on C. galianoi. The two phenotypes of the foundation species C. galianoi showed contrasting effects on the subordinate plant community: Compared to the tight phenotype, the loose phenotype had higher associated species diversity and reduced reciprocal interaction intensities above‐ and below‐ground with the subordinate species F. indigesta. This highlights the impact of phenotypic variation for plant interactions and community‐level diversity. A plain language summary is available for this article. [ABSTRACT FROM AUTHOR]

Higher plant species diversity decreases variability of plant community productivity. The stabilizing effect of plant diversity can result from species‐specific responses to environmental fluctuations and from shifts in competitive hierarchies. Evolutionary adaptation of species to surrounding plant diversity could further decrease productivity variability.We used a three‐year dataset from a crop diversity experiment with seven species to assess the effect of crop diversity and selection history on temporal variability of yield.We found contrasting patterns of temporal variability: Yield of species varied more in mixtures than in monocultures over years. However, at community‐level, we found lower yield variability in crop mixtures compared to monocultures, although only in combination with fertilizer application under Mediterranean climate. Furthermore, we found that a mixture selection history can increase yield productivity and decrease its variability, although only in monocultures. This suggests that the interspecific interactions among crops in mixtures act as an evolutionary selective force, promoting niche complementarity.Synthesis. Our results highlight the ecological and evolutionary role of plant interactions in crop mixtures, which can affect yield stability, while also reflecting on the importance of climate and resource availability in modifying the diversity‐stability relationship. [ABSTRACT FROM AUTHOR]

Determining which drivers lead to a specific species assemblage is a central issue in community ecology. Although many processes are involved, plant-plant interactions are among the most important. The phylogenetic limiting similarity hypothesis states that closely related species tend to compete stronger than distantly related species, although evidence is inconclusive. We used ecological and phylogenetic data on alpine plant communities along an environmental severity gradient to assess the importance of phylogenetic relatedness in affecting the interaction between cushion plants and the whole community, and how these interactions may affect community assemblage and diversity. We first measured species richness and individual biomass of species growing within and outside the nurse cushion species, Arenaria tetraquetra. We then assembled the phylogenetic tree of species present in both communities and calculated the phylogenetic distance between the cushion species and its beneficiary species, as well as the phylogenetic community structure. We also estimated changes in species richness at the local level due to the presence of cushions. The effects of cushions on closely related species changed from negative to positive as environmental conditions became more severe, while the interaction with distantly related species did not change along the environmental gradient. Overall, we found an environmental context-dependence in patterns of phylogenetic similarity, as the interaction outcome between nurses and their close and distantly-related species showed an opposite pattern with environmental severity. [ABSTRACT FROM AUTHOR]

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]

We review the need for increasing agricultural sustainability, how this can in part be delivered by positive biodiversity–ecosystem function (BEF) effects, the role within these of plant–plant facilitation, and how a better understanding of this role may help to deliver sustainable crop (particularly arable) production systems.Major challenges facing intensive arable production include overall declines in biodiversity, poor soil structure and health, nutrient and soil particle run‐off, high greenhouse gas emissions, and increasing costs of synthetic inputs including herbicides, pesticides and fertilisers.Biodiversity–ecosystem function effects have the potential to deliver win–wins for arable food production, whereby enhanced biodiversity is associated with 'good outcomes' for farming sustainability, albeit sometimes through negative BEF effects for some components of the system. Although it can be difficult to separate explicitly from niche differentiation, evidence indicates facilitation can be a key component of these BEF effects.Explicit recognition of facilitation's role brings benefits to developing sustainable crop systems. First, it allows us to link fundamental ecological studies on the evolution of facilitation to the selection of traits that can enhance functioning in crop mixtures. Second, it provides us with analytical frameworks which can be used to bring structure and testable hypotheses to data derived from multiple (often independent) crop trials.Before concrete guidance can be provided to the agricultural sector as to how facilitation might be enhanced in crop systems, challenges exist with respect to quantifying facilitation, understanding the traits that maximise facilitation and integrating these traits into breeding programmes, components of an approach we suggest could be termed 'Functional Ecological Selection'.Synthesis. Ultimately, better integration between ecologists and crop scientists will be essential in harnessing the benefits of ecological knowledge for developing more sustainable agriculture. We need to focus on understanding the mechanistic basis of strong facilitative interactions in crop systems and using this information to select and breed for improved combinations of genotypes and species as part of the Functional Ecological Selection approach. [ABSTRACT FROM AUTHOR]

Facilitation by nurse plants increases understorey diversity and supports ecological communities. In turn, biodiversity shapes ecological networks and enhances ecosystem functioning. However, whether and how facilitation and increased biodiversity jointly influence community structure and ecosystem functioning remains unclear.We performed a field experiment disentangling the relative contribution of nurse plants and increasing understorey plant diversity in driving pollination interactions. Both the presence of nurse shrubs and increased understorey plant diversity increased pollinator diversity and visitation rates. While nurse and understorey diversity effects on pollinator visitation rates did not interact, the effects of increasing understorey plant diversity on pollinator diversity were stronger in the absence than in the presence of shrubs, meaning that nurse shrubs attenuated the effects of high understorey diversity and buffered the effects of low understorey diversity.We also found positive complementarity effects among understorey species as well as complementarity between nurse plants and understorey species at high diversity. Results also indicate negative selection effects, suggesting that species with generally few pollinators benefit the most in the polyculture while a species (possibly the nurse plant) with generally lots of pollinators does not. The corresponding changes in pollination networks with the experimental treatments were due to both changes in the frequency of visits and turnover in pollinator community composition.Synthesis. Plant–plant facilitative systems, where a nurse plant increases understorey plant diversity, are common in stressful environments. Here, we show that these facilitative systems positively influence mutualistic interactions with pollinators via both direct nurse effects and indirect positive effects of increasing plant diversity. Conserving and supporting nurse plant systems is crucial not only for maintaining plant diversity but also for supporting ecosystem functions and services. [ABSTRACT FROM AUTHOR]

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. [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]

Plant diversity fosters productivity in natural ecosystems. Biodiversity effects might increase agricultural yields at no cost in additional inputs. However, the effects of diversity on crop assemblages are inconsistent, probably because crops and wild plants differ in a range of traits relevant to plant–plant interactions. We tested whether domestication has changed the potential of crop mixtures to over‐yield by comparing the performance and traits of major crop species and those of their wild progenitors under varying levels of diversity. We found stronger biodiversity effects in mixtures of wild progenitors, due to larger selection effects. Variation in selection effects was partly explained by within‐mixture differences in leaf size. Our results indicate that domestication might disrupt the ability of crops to benefit from diverse neighbourhoods via reduced trait variance. These results highlight potential limitations of current crop mixtures to over‐yield and the potential of breeding to re‐establish variance and increase mixture performance. [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]

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]

Understanding how ecological networks are organised over the course of an organism’s lifetime is crucial for predicting the dynamics of interacting populations and communities across temporal scales. However, most studies so far considered only one life history stage at a time, such as adult, when studying networks of interacting species. Therefore, knowledge about how multiple life history stages affect the development and stability of plant–plant association networks is lacking. We measured the understory adult plant community and the soil seed bank across a plant age gradient of the nurse shrub Retama sphaerocarpa in an arid ecosystem in Spain. Using a multilayer network approach, we built adult understory–nurse and seed bank–nurse networks and analysed how network nestedness, species’ role, and species specificity varied between them and with nurse plant age. We found that seed bank and adult understory networks changed depending on nurse plant age in two different ways. With increasing nurse plant age, adult understory networks became significantly more nested than seed bank networks. The nested architecture of seed bank networks was therefore a poor predictor of adult understory network nestedness. The contribution and specificity of species to network nestedness increased with increasing nurse plant age more in the adult understory than in seed bank networks, despite high species turnover. Our data show that life history and ontogeny affect the development of plant–plant association networks. Niche construction and environmental filtering along nurse ontogeny seem pivotal mechanisms structuring adult understory networks while the assembly of seed bank networks seems rather stochastic. We highlight the importance of mature plant communities for maintaining rare species populations and supporting the stability of ecological communities through time. [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]

Abstract: How the patterns of niche occupancy vary from species‐poor to species‐rich communities is a fundamental question in ecology that has a central bearing on the processes that drive patterns of biodiversity. As species richness increases, habitat filtering should constrain the expansion of total niche volume, while limiting similarity should restrict the degree of niche overlap between species. Here, by explicitly incorporating intraspecific trait variability, we investigate the relationship between functional niche occupancy and species richness at the global scale. We assembled 21 datasets worldwide, spanning tropical to temperate biomes and consisting of 313 plant communities representing different growth forms. We quantified three key niche occupancy components (the total functional volume, the functional overlap between species and the average functional volume per species) for each community, related each component to species richness, and compared each component to the null expectations. As species richness increased, communities were more functionally diverse (an increase in total functional volume), and species overlapped more within the community (an increase in functional overlap) but did not more finely divide the functional space (no decline in average functional volume). Null model analyses provided evidence for habitat filtering (smaller total functional volume than expectation), but not for limiting similarity (larger functional overlap and larger average functional volume than expectation) as a process driving the pattern of functional niche occupancy. Synthesis. Habitat filtering is a widespread process driving the pattern of functional niche occupancy across plant communities and coexisting species tend to be more functionally similar rather than more functionally specialized. Our results indicate that including intraspecific trait variability will contribute to a better understanding of the processes driving patterns of functional niche occupancy. [ABSTRACT FROM AUTHOR]

Background: Intensive farming affects farmland biodiversity, and some arable plants in particular. Increasing crop genetic diversity can increase crop productivity or resilience and could also benefit rare arable plants. Aims: We examined whether barley presence, sowing density and genetic diversity impacted the rare plant Valerianella rimosa and explored possible underlying mechanisms. Methods: In a field study near Dundee, Scotland, we sowed plots of five single barley genotypes, and all five genotypes combined, at three densities; we also had barley-free plots. Valerianella seeds were sown into half of all plots. Measured responses included early-season cover and harvest biomass of barley and common weeds, abiotic parameters (soil moisture, light) and establishment, biomass and seed production by V. rimosa. Results: Barley presence promoted V. rimosa establishment early in the growing season, but without barley density or genetic diversity effects. By harvest, the impact of barley presence on V. rimosa abundance was lost; there were no effects on Valerianella seed production. Barley negatively impacted common weeds, but V. rimosa did not benefit from any indirect facilitation by barley, being bigger without barley. Conclusions: Early beneficial effects of barley on V. rimosa abundance appear offset by late-season competition. However, limited impacts of barley on V. rimosa reproductive success, and negative impacts on common weeds, indicate crops might play a role in conservation management of rare arable plants by creating space in the weed community. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Food security is currently considered a major global problem. However, increasing intensity of food production in agricultural systems has driven reductions in farmland biodiversity. A major challenge is to enable biodiversity conservation whilst addressing the problem of food security., Here we describe how facilitative plant-plant interactions in crop systems could be used to help strike this balance. An obvious example is that of intercropping systems, where combinations of crop species can - under some circumstances - deliver reduced inputs of agrochemicals (fertilizers, pesticides) per unit yield, with potential knock-on benefits for biodiversity conservation., Other facilitative processes can also play a role in biodiversity conservation. Increased intraspecific crop genetic diversity can help protect crops from pests and diseases. Although overlooked in facilitation research, we argue that the mechanisms operate in a manner which is directly analogous to associational defence against herbivores, a process well recognized in the facilitation literature. As with intercropping, the benefits to nature conservation arise from reduced pesticide use per unit harvested crop., Crops may have facilitative effects on some arable weed species, particularly those that are currently considered rare in intensive farming systems. Work is in its early stages to understand the underlying mechanisms, but it appears that crops might create niche space to which some weed species are adapted. Increasing plant species diversity through niche space creation may then have cascading benefits for other components of farmland biodiversity., Our new understanding of facilitative processes arising from work on crop systems has lessons for the study of facilitative interactions in natural and semi-natural communities. We argue that, although easier to identify and quantify in crop systems, some of these facilitative processes have to date been overlooked in studies of non-crop systems and certainly deserve further consideration., Finally, we discuss what steps may be needed to move from our understanding of the role of facilitation to the development of new agricultural practice. In some cases the challenge may be one of the encouraging uptake of existing practices, and in others more research is needed to understand how new ecological understanding might deliver more sustainable agricultural practice. [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]

Understanding the relative contribution of local and regional processes to local species richness is an important ecological question and a subject of controversy between macroecologists and community ecologists. We test the hypothesis that the contribution of local and regional processes is dependent on environmental conditions and that the effect of regional processes should be the highest in communities from intermediate positions along environmental severity gradients due to the importance of facilitation. We used the recently developed log-ratio method to analyze the relationship between local species richness (LSR) and regional species richness (RSR) for 13 plant communities from 4 habitat types of France (coastal sand dunes, oceanic heathlands, alpine grasslands, lowland calcareous grasslands). Each habitat type was split in 3-4 communities using multivariate analyses to identify the relative importance of stress, disturbance, competition, and facilitation functioning within the 13 communities. We found that the LSR/RSR relationship was highly dependent on environmental conditions with saturated communities occurring more frequently than unsaturated communities highlighting the relative importance of local drivers on species richness. We argued that competition was most likely the main source of community saturation whilst facilitation likely contributed to enhancing the importance of the regional species pool for all habitat types. However, the effect of facilitation might be stronger in the disturbed than in the stressed systems because unsaturated curves were only observed in the former conditions. In extreme conditions of disturbance LSR was only controlled by the intensity of disturbance. This effect was not observed in extreme stress conditions. Our study provides support for the emerging balance theory that both local and regional processes are important in nature with their relative contribution depending on environmental conditions. Additionally, this synthesis strongly suggests that facilitation contributes to an important process - the influence of regional species pool on local species richness. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]