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Plants demonstrate a broad range of responses to environmental shifts. One of the most remarkable responses is plasticity, which is the ability of a single plant genotype to produce different phenotypes in response to environmental stimuli. As with all traits, the ability of plasticity to evolve depends on the presence of underlying genetic diversity within a population. A common approach for evaluating the role of genetic variation in driving differences in plasticity has been to study genotype-by-environment interactions (G × E). G × E occurs when genotypes produce different phenotypic trait values in response to different environments. In this review, we highlight progress and promising methods for identifying the key environmental and genetic drivers of G × E. Specifically, methodological advances in using algorithmic and multivariate approaches to understand key environmental drivers combined with new genomic innovations can greatly increase our understanding about molecular responses to environmental stimuli. These developing approaches can be applied to proliferating common garden networks that capture broad natural environmental gradients to unravel the underlying mechanisms of G × E. An increased understanding of G × E can be used to enhance the resilience and productivity of agronomic systems.

Seed mass is an ecologically important trait that often differs considerably among ecotypes. Yet, because few studies examine the impacts of seed mass on adult life-history traits, its role in local adaptation is unclear. In this study, using accessions of Panicum hallii that spanned the two major ecotypes, we examined whether covariation between seed mass, seedling and reproductive traits impacts ecotypic divergence and local adaptation. The perennial grass P. hallii has two distinct ecotypes—a large-seeded upland ecotype adapted to xeric environments and a small-seeded lowland ecotype adapted to mesic environments. In the greenhouse, seed mass varied greatly across P. hallii genotypes in a manner consistent with ecotypic divergence. Seed mass covaried significantly with several seedling and reproductive traits. At field sites representing the habitats of the two ecotypes, seed mass had different impacts on seedling and adult recruitment: selection favoured large seeds in upland habitat and small seeds in lowland habitat, which was consistent with local adaptation. By demonstrating the central role of seed mass in ecotypic differences in P. hallii and its importance to seedling and adult recruitment under field conditions, these studies show that early life-history traits can promote local adaptation and potentially explain ecotype formation.

We present unresolved questions in plant abiotic stress biology as posed by 15 research groups with expertise spanning eco-physiology to cell and molecular biology. We present unresolved questions in plant abiotic stress biology as posed by 15 research groups with expertise spanning eco-physiology to cell and molecular biology. Common themes of these questions include the need to better understand how plants detect water availability, temperature, salinity, and rising carbon dioxide (CO2) levels; how environmental signals interface with endogenous signaling and development (e.g. circadian clock and flowering time); and how this integrated signaling controls downstream responses (e.g. stomatal regulation, proline metabolism, and growth versus defense balance). The plasma membrane comes up frequently as a site of key signaling and transport events (e.g. mechanosensing and lipid-derived signaling, aquaporins). Adaptation to water extremes and rising CO2 affects hydraulic architecture and transpiration, as well as root and shoot growth and morphology, in ways not fully understood. Environmental adaptation involves tradeoffs that limit ecological distribution and crop resilience in the face of changing and increasingly unpredictable environments. Exploration of plant diversity within and among species can help us know which of these tradeoffs represent fundamental limits and which ones can be circumvented by bringing new trait combinations together. Better defining what constitutes beneficial stress resistance in different contexts and making connections between genes and phenotypes, and between laboratory and field observations, are overarching challenges.

Plant community succession is structured by initial richness, plant consumer pressure and soil resource supply. These drivers influence species' trait trade‐offs that underlie temporal changes in plant community diversity. Importantly, how these drivers interact with each other and through time and whether they act on different facets of plant community diversity by promoting different plant trade‐off strategies remains poorly understood. We experimentally manipulated initial plant richness, consumer pressure via pesticide spraying and soil resource supply via fertilization across 4 years of old field succession. We then allowed natural colonization and extinction to occur and examined how the three drivers influenced succession. Specifically, we examined changes in temporal dynamics by conducting yearly taxonomic cover surveys and measuring light penetration to the ground. In the third year, we measured vegetative height and specific leaf area (SLA), and investigated seed mass using a trait database. Higher initial richness, lower consumer pressure and increased soil resource supply all decreased colonizing species richness and light availability and variably altered species evenness. These effects generally acted additively rather than interactively in driving community diversity during succession. However, soil resource supply suppressed consumer pressure effects on species richness and light availability, while consumer pressure and soil resource supply modified initial richness effects on light availability. Treatments acted on different trait identities, revealing different mechanisms underlying taxonomic responses. Initial richness effects on seed mass of colonizing species were modified by soil resource supply. Decreased consumer pressure increased intraspecific community height and decreased interspecific SLA. Increased soil resource supply increased community height, SLA and seed mass. Synthesis. Our results suggest species' resource strategies underlie plant diversity responses to consumer pressure and soil resource supply. Resource addition promoted resource‐acquisitive species; consumer pressure disadvantaged resource‐conservative species. Meanwhile, initial richness altered subsequent community composition primarily through persistence of early residents. We show that community responses to drivers of succession depend on underlying trait trade‐offs of resident species, and these trade‐offs influence community diversity across succession.

Plant responses to major environmental drivers like precipitation can influence important aspects of carbon (C) cycling like soil CO2 efflux ( urn:x-wiley:00220477:media:jec13382:jec13382-math-0001 ). These responses may be predicted by two independent classes of drivers: plant size—larger plants respire more and produce a larger quantity of labile C, and plant economics—plants possessing more acquisitive plant economics strategies (i.e. high metabolic rate and tissue nutrient content) produce higher‐quality tissue that respires rapidly and decomposes quickly. At two sites in central Texas, USA with similar climates and differing soil characteristics, we examined the response of eight Panicum virgatum genotypes to three annual precipitation levels defined by the driest, average and wettest years from each site's precipitation history. We evaluated the individual and joint influence of plant genotypes and precipitation on urn:x-wiley:00220477:media:jec13382:jec13382-math-0002 and traits related to plant economics and plant size. We then used confirmatory path analysis to evaluate whether effects of precipitation on urn:x-wiley:00220477:media:jec13382:jec13382-math-0003 were in part related to effects of precipitation on plant economics traits or size (‘mediated’ effects). These genotypes exhibited variation in plant economics traits and above‐ground net primary productivity (ANPP), an above‐ground measure of plant size. Increasing precipitation increased urn:x-wiley:00220477:media:jec13382:jec13382-math-0004 and ANPP more than plant economics traits. At both sites, ANPP was the best predictor of urn:x-wiley:00220477:media:jec13382:jec13382-math-0005 . Moreover, the sites differed in the ways that plant size and plant economics traits combined with precipitation to influence urn:x-wiley:00220477:media:jec13382:jec13382-math-0006 . At the Austin site, the positive effect of precipitation on urn:x-wiley:00220477:media:jec13382:jec13382-math-0007 was mediated primarily by ANPP, offset by a smaller effect of leaf nitrogen content; no direct precipitation effect was detected. At the Temple site, increasing precipitation had positive direct and ANPP‐mediated effects on urn:x-wiley:00220477:media:jec13382:jec13382-math-0008 . This suggests that greater water limitation at Austin may strengthen the links between plant size and urn:x-wiley:00220477:media:jec13382:jec13382-math-0009 . Synthesis. Estimates of C cycling can be improved by accounting for mediation of precipitation effects on urn:x-wiley:00220477:media:jec13382:jec13382-math-0010 by plant economics traits and plant size in resource‐limited environments.

Disease may modulate variation in host community structure by modifying the interplay of deterministic and stochastic processes. For instance, deterministic processes like ecological selection can benefit species less impacted by disease. When disease consistently selects for certain host species, this can reduce variation in host community composition. On the other hand, when host communities are less impacted by disease and selection is weaker, stochastic processes (e.g., drift, dispersal) may play a bigger role in host community structure, which can increase variation in structure among communities. While effects of disease on host community structure have been quantified in field experiments, few have addressed the role of disease in modulating variation in structure among host communities. To address this, we conducted a field experiment spanning three years, using a tractable system: foliar fungal pathogens in an old-field grassland community dominated by the grass Lolium arundinaceum, tall fescue. We reduced foliar fungal disease burden in replicate host communities (experimental plots in intact vegetation) in three fungicide regimens that varied in the duration of fungicide exposure and included a fungicide-free control. We measured host diversity, biomass, and variation in community structure among replicate communities. Disease reduction generally decreased plant richness and increased aboveground biomass relative to communities experiencing ambient levels of disease. Despite changes in structure of the plant communities over the experiment’s three years, the effects of disease reduction on plant richness and biomass were consistent across years. However, disease reduction did not reduce variation in host community structure, providing little evidence for ecological selection by competition or other deterministic processes. Instead, disease reduction tended to amplify variation in host community structure among replicate communities (i.e., within fungicide treatment groups), suggesting that disease diminished the degree to which host communities were structured by stochastic processes. These results of experimental disease reduction both highlight the potential importance of stochastic processes in plant communities and reveal the potential for disease to regulate variation in host community structure.

The leaf economics spectrum (LES) is hypothesized to result from a trade-off between resource acquisition and conservation. Yet few studies have examined the evolutionary mechanisms behind the LES, perhaps because most species exhibit relatively specialized leaf economics strategies. In a genetic mapping population of the phenotypically diverse grass Panicum virgatum, we evaluate two interacting mechanisms that may drive LES evolution: 1) genetic architecture, where multiple traits are coded by the same gene (pleiotropy) or by genes in close physical proximity (linkage), and 2) correlational selection, where selection acts non-additively on combinations of multiple traits. We found evidence suggesting that shared genetic architecture (pleiotropy) controls covariation between two pairs of leaf economics traits. Additionally, at five common gardens spanning 17 degrees of latitude, correlational selection favored particular combinations of leaf economics traits. Together, these results demonstrate how the LES can evolve within species.

Successful colonization and growth of trees within herbaceous communities may result from different interactions with the herbaceous community. First, colonizing trees compete against larger, established herbs, while subsequent growth occurs among similarly sized or smaller herbs. This shift from colonization to growth may lead three drivers of community dynamics-nutrients, consumers, and herbaceous diversity-to differentially affect tree colonization and, later, tree performance. Initially, these drivers should favor larger, established herbs, reducing tree colonization. Later, when established trees can better compete with herbs, these drivers should benefit trees and increase their performance. In a 4-year study in a southeastern US old field, we added nutrients to, excluded aboveground consumers from, and manipulated initial richness of, the herbaceous community, and then allowed trees to naturally colonize these communities (from intact seedbanks or as seed rain) and grow. Nutrients and consumers had opposing effects on tree colonization and performance: adding nutrients and excluding consumers reduced tree colonization rate, but later increased the size of established trees (height, basal diameter). Adding nutrients and excluding consumers also restricted tree colonization to earlier years of study, which partially explained the effect of nutrient addition on plant size. Together, this shows differing impacts of nutrients and consumers: factors that initially limited tree colonization also resulted in larger established trees. This suggests that succession of grasslands that are either eutrophied or have diminished consumer pressure may experience lags and pulses in woody encroachment, leading to an extended period of herbaceous dominance followed by accelerated woody growth.

Correction to: Nature Communications https://doi.org/10.1038/s41467-020-19870-y, published online 27 November 2020.

Terrestrial ecosystems are increasingly enriched with resources such as atmospheric CO(2) that limit ecosystem processes. The consequences for ecosystem carbon cycling depend on the feedbacks from other limiting resources and plant community change, which remain poorly understood for soil CO(2) efflux, J(CO2), a primary carbon flux from the biosphere to the atmosphere. We applied a unique CO(2) enrichment gradient (250 to 500 µL L(−1)) for eight years to grassland plant communities on soils from different landscape positions. We identified the trajectory of J(CO2) responses and feedbacks from other resources, plant diversity [effective species richness, exp(H)], and community change (plant species turnover). We found linear increases in J(CO2) on an alluvial sandy loam and a lowland clay soil, and an asymptotic increase on an upland silty clay soil. Structural equation modeling identified CO(2) as the dominant limitation on J(CO2) on the clay soil. In contrast with theory predicting limitation from a single limiting factor, the linear J(CO2) response on the sandy loam was reinforced by positive feedbacks from aboveground net primary productivity and exp(H), while the asymptotic J(CO2) response on the silty clay arose from a net negative feedback among exp(H), species turnover, and soil water potential. These findings support a multiple resource limitation view of the effects of global change drivers on grassland ecosystem carbon cycling and highlight a crucial role for positive or negative feedbacks between limiting resources and plant community structure. Incorporating these feedbacks will improve models of terrestrial carbon sequestration and ecosystem services.

Terrestrial ecosystems are increasingly enriched with resources such as atmospheric CO2 that limit ecosystem processes. The consequences for ecosystem carbon cycling depend on the feedbacks from other limiting resources and plant community change, which remain poorly understood for soil CO2 efflux, JCO2, a primary carbon flux from the biosphere to the atmosphere. We applied a unique CO2 enrichment gradient (250 to 500 µL L−1) for eight years to grassland plant communities on soils from different landscape positions. We identified the trajectory of JCO2 responses and feedbacks from other resources, plant diversity [effective species richness, exp(H)], and community change (plant species turnover). We found linear increases in JCO2 on an alluvial sandy loam and a lowland clay soil, and an asymptotic increase on an upland silty clay soil. Structural equation modeling identified CO2 as the dominant limitation on JCO2 on the clay soil. In contrast with theory predicting limitation from a single limiting factor, the linear JCO2 response on the sandy loam was reinforced by positive feedbacks from aboveground net primary productivity and exp(H), while the asymptotic JCO2 response on the silty clay arose from a net negative feedback among exp(H), species turnover, and soil water potential. These findings support a multiple resource limitation view of the effects of global change drivers on grassland ecosystem carbon cycling and highlight a crucial role for positive or negative feedbacks between limiting resources and plant community structure. Incorporating these feedbacks will improve models of terrestrial carbon sequestration and ecosystem services.

For succession to proceed from herbaceous to woody dominance, trees must colonize herbaceous communities and grow. Success across these two phases of succession might result from different interactions with the herbaceous community. First, colonizing trees must compete against larger, established herbs, while subsequent growth occurs among similarly sized or smaller herbs. This shift from colonization to growth may cause three drivers of secondary succession— nutrients, consumers, and herbaceous diversity—to differentially affect tree colonization and growth. Initially, these drivers should favor larger, established herbs, reducing colonization. Later, when established trees can better compete with herbs, these drivers should benefit trees and increase their growth. In a four-year study, we added nutrients to, excluded aboveground consumers from, and manipulated initial richness of, the herbaceous community, then allowed trees to naturally colonize these communities (from intact seedbanks or as seed-rain) and grow. Nutrients and consumers had opposing effects on tree colonization and growth: adding nutrients and excluding consumers reduced tree colonization, but later increased established tree growth (height, basal diameter). Together, this shows stage-specific impacts of nutrients and consumers that may improve predictions of the rate and trajectory of succession: factors that initially limited tree colonization later helped established trees to grow.

Human activities are transforming grassland biomass via changing climate, elemental nutrients, and herbivory. Theory predicts that food-limited herbivores will consume any additional biomass stimulated by nutrient inputs (‘consumer-controlled’). Alternatively, nutrient supply is predicted to increase biomass where herbivores alter community composition or are limited by factors other than food (‘resource-controlled’). Using an experiment replicated in 58 grasslands spanning six continents, we show that nutrient addition and vertebrate herbivore exclusion each caused sustained increases in aboveground live biomass over a decade, but consumer control was weak. However, at sites with high vertebrate grazing intensity or domestic livestock, herbivores consumed the additional fertilization-induced biomass, supporting the consumer-controlled prediction. Herbivores most effectively reduced the additional live biomass at sites with low precipitation or high ambient soil nitrogen. Overall, these experimental results suggest that grassland biomass will outstrip wild herbivore control as human activities increase elemental nutrient supply, with widespread consequences for grazing and fire risk., It is unclear whether terrestrial herbivores are able to consume the extra plant biomass produced under nutrient enrichment. Here the authors test this in grasslands using a globally distributed network of coordinated field experiments, finding that wild herbivore control on grassland production declines under eutrophication.

Host and parasite richness are generally positively correlated, but the stability of this relationship in response to global change remains poorly understood. Rapidly changing biotic and abiotic conditions can alter host community assembly, which in turn, can alter parasite transmission. Consequently, if the relationship between host and parasite richness is sensitive to parasite transmission, then changes in host composition under various global change scenarios could strengthen or weaken the relationship between host and parasite richness. To test the hypothesis that host community assembly can alter the relationship between host and parasite richness in response to global change, we experimentally crossed host diversity (biodiversity loss) and resource supply to hosts (eutrophication), then allowed communities to assemble. As previously shown, initial host diversity and resource supply determined the trajectory of host community assembly, altering post-assembly host species richness, richness-independent host phylogenetic diversity, and colonization by exotic host species. Overall, host richness predicted parasite richness, and as predicted, this effect was moderated by exotic abundance-communities dominated by exotic species exhibited a stronger positive relationship between post-assembly host and parasite richness. Ultimately, these results suggest that, by modulating parasite transmission, community assembly can modify the relationship between host and parasite richness. These results thus provide a novel mechanism to explain how global environmental change can generate contingencies in a fundamental ecological relationship-the positive relationship between host and parasite richness.

Host and parasite richness are generally positively correlated, but the stability of this relationship during community assembly remains untested. The composition of host communities can alter parasite transmission, and the relationship between host and parasite richness is sensitive to parasite transmission. Thus, changes in composition during host community assembly could strengthen or weaken the relationship between host and parasite richness. Host community assembly, in turn, can be driven by many processes, including resource enrichment. To test the hypothesis that host community assembly can alter the relationship between host and parasite richness, we experimentally crossed host diversity and resource supply to hosts, then allowed communities to assemble. As previously shown, initial host diversity and resource supply determined the trajectory of host community assembly, altering post-assembly host species richness, richness-independent host phylogenetic diversity, and colonization by exotic host species. Throughout community assembly, host richness predicted parasite richness. As predicted, this effect was moderated by exotic abundance: communities dominated by exotic species exhibited a stronger positive relationship between post-assembly host and parasite richness. Ultimately, these results suggest that, by modulating parasite transmission, community assembly can modify the relationship between host and parasite richness, providing a novel mechanism to explain contingencies in this relationship.

High-resource environments typically favor quick-growing, poorly defended plants, while resource-poor environments typically favor slow-growing, well-defended plants. The prevailing hypothesis explaining this pattern states that, as resource availability increases, well-defended, slow-growing species are replaced by poorly defended, fast-growing species. A second hypothesis states that greater resource availability increases allocation to growth at the expense of defense, within species. Regardless of mechanism, if exotic species are released from enemies relative to natives, shifts in allocation to growth and defense both within and among species could differ by geographic provenance. To test whether resource availability alters growth or defense, within and among species, and whether any such effects differ between natives and exotics, we manipulated soil nutrient supply and access of aboveground insect herbivores and fungal pathogens under field conditions to individuals of six native and six exotic grass species that co-occurred in a North Carolina old field. The prevailing hypothesis’ prediction—that species-level enemy impact increases with species’ nutrient responsiveness—was confirmed. Moreover, this relationship did not differ between native and exotic species. The second hypothesis’ prediction—that individual-level enemy impact increases with nutrient supply, after accounting for species-level variation in performance—was not supported. Together, these results support the idea, across native and exotic species, that plant species turnover is the primary mechanism underlying effects of nutrient enrichment on allocation to growth and defense in plant communities.

Invasive plants can seriously threaten biodiversity and ecosystem function. Thus, controlling the abundance and mitigating the deleterious effects of invasive plants is imperative. The effectiveness of removal methods could vary based on potential damage to non‐target species and on the ecological context of invasion. For example, species‐poor communities might benefit disproportionately from invasive removal because they tend to be more heavily invaded than species‐rich communities. In this study, we tested the efficacy of two common methods of invasive plant control, hand removal and 2,4‐D herbicide application, against the invasive forb, Galium verum. We further tested whether the community response to G. verum removal varied across communities that differed in native grass richness (one, three or six species) and composition. Grass diversity did not interact with G. verum removal to influence any response. Instead, hand removal and herbicide application each reduced G. verum stem density by >50% relative to controls. Furthermore, hand removal of G. verum increased species richness by 25% relative to controls and herbicide‐sprayed plots. Despite this benefit, hand removal is labour‐intensive and impractical for large areas. These results also suggest that applying herbicide may control G. verum spread without substantially reducing forb diversity in invaded tallgrass prairie. Thus, herbicide application may be most suitable for grasslands with low native forb diversity, especially in managed grasslands and pastures where maintaining native forb diversity is a lower priority than maintaining high grass productivity.

Exotic plants are often most successful in high resource environments. By drawing down available resources, species-rich communities may be able to reduce exotic success when resource supply is elevated. We tested the prediction that exotic success would be greatest in species-poor communities when nitrogen availability is high. We also tested two underlying assumptions of this prediction: species-rich communities draw down soil nitrogen availability more than species-poor communities following fertilization and exotic success increases when soil nitrogen availability is high. In a restored grassland where native grass diversity was manipulated (one, three, or five-species) seven years earlier to form a gradient in species richness, we manipulated nitrogen availability directly via fertilization, and indirectly via burning. We then examined the success of the exotic forb Galium verum L. Contrary to our prediction, diversity and nutrient treatments did not jointly influence exotic success. Instead, one-time fertilization increased exotic biomass in the first year of the study. This likely occurred because the effect of nutrient treatments on nitrogen availability was independent of diversity treatment. Thus, we found no evidence that species-rich communities are better able to reduce exotic biomass when nitrogen is added than are species-poor communities. This suggests that in some systems, the effects of increasing species richness can be overwhelmed by the effects of nutrient addition that promote exotic success.

Infectious disease risk is often influenced by host diversity, but the causes are unresolved. Changes in diversity are associated with changes in community structure, particularly during community assembly; therefore, by incorporating change over time, host community assembly may provide a framework to resolve causation. In turn, community assembly can be driven by many processes, including resource enrichment. To test the hypothesis that community assembly causally links host diversity to future disease, we experimentally manipulated host diversity and resource supply to hosts, then allowed communities to assemble for 2 years (surveyed 2012-2014). Initially, host diversity increased disease. Subsequently, host diversity did not directly alter disease. However, host diversity determined the trajectory of host community assembly, altering colonisation by exotic host species and richness-independent host phylogenetic diversity, which together reversed the initial increase in disease. Ultimately, incorporating the temporal dimension of community assembly revealed novel mechanisms linking host diversity to future disease.

Many factors can promote exotic plant success. Three of these factors-greater pressure from natural enemies on natives, increased soil nutrient supply, and low native species richness-may interact during invasions. To test for independent and interactive effects of these drivers, we planted herbaceous perennial communities at two levels of native richness (monocultures and five-species polycultures). We then factorially manipulated soil nutrient supply and access to these communities by aboveground foliar enemies (fungal pathogens and insect herbivores), and allowed natural colonization to proceed for four years. We predicted that nutrient addition would increase exotic success, while enemy exclusion and increasing native richness would reduce exotic success. Additionally, we expected that enemy exclusion would reduce the benefits of nutrient addition to exotic species most in species-poor communities, and that this effect would be weaker in species-rich communities. In total, we found no evidence that nutrient supply, enemy access, and native richness interacted to influence exotic success. Furthermore, native richness had no effect on exotic success. Instead, nutrient addition increased, and enemy exclusion decreased, exotic success independently. As predicted, enemy exclusion reduced exotic success, primarily by slowing the decline in abundance of planted native species. Together, these results demonstrate that multiple drivers of exotic success can act independently within a single system.

Worldwide, ecosystems are increasingly dominated by exotic plant species, a shift hypothesized to result from numerous ecological factors. Two of these, increased resource availability and enemy release, may act in concert to increase exotic success in plant communities (Resource-Enemy Release Hypothesis, R-ERH). To test this, we manipulated the availability of soil nutrients and access of vertebrate herbivores, insect herbivores, and fungal pathogens to intact grassland communities containing both native and exotic species. Our results supported both conditions necessary for R-ERH. First, exotics were less damaged than natives, experiencing less foliar damage (insect herbivory and fungal disease) than native species, particularly in communities where soil nutrients were added. Second, fertilization increased foliar damage on native species, but not exotic species. As well as fulfilling both conditions for R-ERH, these results demonstrate the importance of considering the effects of resource availability when testing for enemy release. When both conditions are fulfilled, R-ERH predicts that increasing resource availability will increase exotic abundance only in the presence of enemies. Our results fully supported this prediction for vertebrate herbivores—fertilization increased exotic cover only in communities exposed to vertebrate herbivores. Additionally, the prediction was partially supported for insect herbivores and fungal pathogens—excluding these enemies reduced exotic cover as predicted, but inconsistent with R-ERH, this effect occurred only in unfertilized communities. These results highlight the need to consider the influence of multiple enemy guilds on community processes like exotic plants invasions. Moreover, this study experimentally demonstrates that resource availability and natural enemies can jointly influence exotic success in plant communities. This article is protected by copyright. All rights reserved.

Terrestrial ecosystem productivity is widely accepted to be nutrient limited. Although nitrogen is deemed a key determinant of aboveground net primary production, the prevalence of co-limitation by nitrogen and phosphorus is increasingly recognized. However, the extent to which terrestrial productivity is co-limited by nutrients other than nitrogen and phosphorus, such as potassium and micronutrients, has remained unclear. Here, we report results from a standardized factorial nutrient addition experiment, in which we added nitrogen (N), phosphorus (P), and potassium combined with a selection of micronutrients (K+μ), alone or in concert, to 42 grasslands spanning five continents, and monitored aboveground net primary production. Nutrient availability limited productivity at 31 of the 42 grassland sites studied. Pairwise combinations of N, P, and K+μ co-limited aboveground net primary production at 24 of the 42 sites, and nitrogen limitation peaked in cool, high latitude sites. Our findings indicate widespread variation in the combination of nutrients that limit aboveground grassland productivity, and highlight the importance of potassium and micronutrients, and an apparent absence of limitation by the nutrients we considered at some sites. This variation in nutrient limitation must be considered when assessing the ecosystem-scale consequences of nutrient enrichment.

By altering the strength of intra- and interspecific competition, droughts may reshape plant communities. Furthermore, species may respond differently to drought when other influences, such as herbivory, are considered. To explore this relationship, we conducted a greenhouse experiment measuring responses to inter- and intraspecific competition for two grasses, Schedonorus arundinaceus and Paspalum dilatatum, while varying water availability and simulating herbivory via clipping. We then parameterized population growth models to examine the long-term outcome of competition under these conditions. Under drought, S. arundinaceus was less water stressed than P. dilatatum, which exhibited severe water stress; clipping alleviated this stress, increasing the competitive ability of P. dilatatum relative to S. arundinaceus. Although P. dilatatum competed weakly under drought, clipping reduced water stress in P. dilatatum, thereby enhancing its ability to compete with S. arundinaceus under drought. Supporting these observations, population growth models predicted that P. dilatatum would exclude S. arundinaceus when clipped under drought, while S. arundinaceus would exclude P. dilatatum when unclipped under drought. When the modeled environment varied temporally, environmental variation promoted niche differences that, though insufficient to maintain stable coexistence, prevented unconditional competitive exclusion by promoting priority effects. Our results suggest that it is important to consider how species respond not just to stable, but also to variable, environments. When species differ in their responses to drought, competition, and simulated herbivory, stable environments may promote competitive exclusion, while fluctuating environments may promote coexistence. These interactions are critical to understanding how species will respond to global change.

Theory predicts that increasing biodiversity will dilute the risk of infectious diseases under certain conditions and will amplify disease risk under others. Yet, few empirical studies demonstrate amplification. This contrast may occur because few studies have considered the multivariate nature of disease risk, which includes richness and abundance of parasites with different transmission modes. By combining a multivariate statistical model developed for biodiversity–ecosystem–multifunctionality with an extensive field manipulation of host (plant) richness, composition and resource supply to hosts, we reveal that (i) host richness alone could not explain most changes in disease risk, and (ii) shifting host composition allowed disease amplification, depending on parasite transmission mode. Specifically, as predicted from theory, the effect of host diversity on parasite abundance differed for microbes (more density-dependent transmission) and insects (more frequency-dependent transmission). Host diversity did not influence microbial parasite abundance, but nearly doubled insect parasite abundance, and this amplification effect was attributable to variation in host composition. Parasite richness was reduced by resource addition, but only in species-rich host communities. Overall, this study demonstrates that multiple drivers, related to both host community and parasite characteristics, can influence disease risk. Furthermore, it provides a framework for evaluating multivariate disease risk in other systems.

Human alterations to nutrient cycles and herbivore communities are affecting global biodiversity dramatically. Ecological theory predicts these changes should be strongly counteractive: nutrient addition drives plant species loss through intensified competition for light, whereas herbivores prevent competitive exclusion by increasing ground-level light, particularly in productive systems. Here we use experimental data spanning a globally relevant range of conditions to test the hypothesis that herbaceous plant species losses caused by eutrophication may be offset by increased light availability due to herbivory. This experiment, replicated in 40 grasslands on 6 continents, demonstrates that nutrients and herbivores can serve as counteracting forces to control local plant diversity through light limitation, independent of site productivity, soil nitrogen, herbivore type and climate. 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Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencia Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Damschen, Ellen. University of Wisconsin; Estados Unidos Fil: Davies, Kendi. State University of Colorado - Fort Collins; Estados Unidos Fil: DeCrappeo, Nicole. Forest and Rangeland Ecosystem Science Center; Estados Unidos Fil: Du, Guozhen. Lanzhou University; Estados Unidos Fil: Firn, Jennifer. Queensland University of Technology; Australia Fil: Hautier, Yann. University of Minnesota; Estados Unidos Fil: Heckman, Robert. University of North Carolina at Chapel Hill; Estados Unidos Fil: Hector, Andy. University of Oxford; Reino Unido Fil: Hillerislambers, Janneke. University of Washington; Estados Unidos Fil: Iribarne, Oscar Osvaldo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencia Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Klein, Julia. State University of Colorado - Fort Collins; Estados Unidos Fil: Knops, Johannes M. H.. University of Nebraska; Estados Unidos Fil: La Pierre, Kimberly. University of California; Estados Unidos Fil: Leakey, Andrew D. B.. University of Illinois at Urbana-Champaign; Estados Unidos Fil: Li, Wei. Iowa State University; Estados Unidos Fil: MacDougall, Andrew S.. University of Guelph; Canadá Fil: McCulley, Rebecca. University Of Kentucky; Estados Unidos Fil: Melbourne, Brett. State University of Colorado - Fort Collins; Estados Unidos Fil: Mitchell, Charles. University of North Carolina at Chapel Hill; Estados Unidos Fil: Moore, Joslin. University of Melbourne; Australia Fil: Mortensen, Brent. Iowa State University; Estados Unidos Fil: O'Halloran, Lydia. Oregon State University; Estados Unidos Fil: Orrock, John. University of Wisconsin; Estados Unidos Fil: Pascual, Jesus Maria. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencia Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Prober, Suzanne. CSIRO Ecosystem Sciences; Australia Fil: Pyke, David. Forest and Rangeland Ecosystem Science Center; Canadá Fil: Risch, Anita. Swiss Federal Institute For Forest, Snow And Landscape Research; Suiza Fil: Schuetz, Martin. Swiss Federal Institute For Forest, Snow And Landscape Research; Suiza Fil: Smith, Melinda D.. State University of Colorado - Fort Collins; Estados Unidos Fil: Stevens, Carly. Lancaster University; Reino Unido Fil: Sullivan, Lauren. Iowa State University; Estados Unidos Fil: Williams, Ryan. Iowa State University; Estados Unidos Fil: Wragg, Peter. University of Minnesota; Estados Unidos Fil: Yang, Louie. University of California; Estados Unidos

A fundamental assumption in invasion biology is that most invasive species exhibit enhanced performance in their introduced range relative to their home ranges. This idea has given rise to numerous hypotheses explaining ''invasion success'' by virtue of altered ecological and evolutionary pressures. There are surprisingly few data, however, testing the underlying assumption that the performance of introduced populations, including organism size, reproductive output, and abundance, is enhanced in their introduced compared to their native range. Here, we combined data from published studies to test this hypothesis for 26 plant and 27 animal species that are considered to be invasive. On average, individuals of these 53 species were indeed larger, more fecund, and more abundant in their introduced ranges. The overall mean, however, belied significant variability among species, as roughly half of the investigated species (N ¼ 27) performed similarly when compared to conspecific populations in their native range. Thus, although some invasive species are performing better in their new ranges, the pattern is not universal, and just as many are performing largely the same across ranges.