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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]

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]

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]

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]

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]

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.

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]