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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.

Light limitation is a major driver of succession and an important determinant of the performance of shade-intolerant tree seedlings. Shade intolerance may result from a resource allocation strategy characterized by rapid growth and high metabolic costs, which may make shade-intolerant species particularly sensitive to nutrient limitation and pathogen pressure. In this study, we evaluated the degree to which nitrogen availability and fungal pathogen pressure interact to influence plant performance across different light environments. To test this, we manipulated nitrogen availability (high, low) and access by foliar fungal pathogens (sprayed with fungicide, unsprayed) to seedlings of the shade-intolerant tree, Liquidambar styraciflua, growing at low and high light availability, from forest understory to adjacent old field. Foliar fungal damage varied with light and nitrogen availability; in low light, increasing nitrogen availability tripled foliar damage, suggesting that increased nutrient availability in low light makes plants more susceptible to disease. Despite higher foliar damage under low light, spraying fungicide to exclude pathogens promoted 14% greater plant height only under high light conditions. Thus, although nitrogen availability and pathogen pressure each influenced aspects of plant performance, these effects were context dependent and overwhelmed by light limitation. This suggests that failure of shade-intolerant species to invade closed-canopy forest can be explained by light limitation alone.

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.

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 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.

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.