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To evaluate how increased anthropogenic nutrient inputs alter carbon cycling in grasslands, we conducted a litter decomposition study across 20 temperate grasslands on three continents within the Nutrient Network, a globally distributed nutrient enrichment experiment We determined the effects of addition of experimental nitrogen (N), phosphorus (P) and potassium plus micronutrient (Kμ) on decomposition of a common tree leaf litter in a long-term study (maximum of 7 years; exact deployment period varied across sites). The use of higher order decomposition models allowed us to distinguish between the effects of nutrients on early- versus late-stage decomposition. Across continents, the addition of N (but not other nutrients) accelerated early-stage decomposition and slowed late-stage decomposition, increasing the slowly decomposing fraction by 28% and the overall litter mean residence time by 58%. Synthesis. Using a novel, long-term cross-site experiment, we found widespread evidence that N enhances the early stages of above-ground plant litter decomposition across diverse and widespread temperate grassland sites but slows late-stage decomposition. These findings were corroborated by fitting the data to multiple decomposition models and have implications for N effects on soil organic matter formation. For example, following N enrichment, increased microbial processing of litter substrates early in decomposition could promote the production and transfer of low molecular weight compounds to soils and potentially enhance the stabilization of mineral-associated organic matter. By contrast, by slowing late-stage decomposition, N enrichment could promote particulate organic matter (POM) accumulation. Such hypotheses deserve further testing.

Grasslands are subject to considerable alteration due to human activities globally, including widespread changes in populations and composition of large mammalian herbivores and elevated supply of nutrients. Grassland soils remain important reservoirs of carbon (C) and nitrogen (N). Herbivores may affect both C and N pools and these changes likely interact with increases in soil nutrient availability. Given the scale of grassland soil fluxes, such changes can have striking consequences for atmospheric C concentrations and the climate. Here, we use the Nutrient Network experiment to examine the responses of soil C and N pools to mammalian herbivore exclusion across 22 grasslands, under ambient and elevated nutrient availabilities (fertilized with NPK + micronutrients). We show that the impact of herbivore exclusion on soil C and N pools depends on fertilization. Under ambient nutrient conditions, we observed no effect of herbivore exclusion, but under elevated nutrient supply, pools are smaller upon herbivore exclusion. The highest mean soil C and N pools were found in grazed and fertilized plots. The decrease in soil C and N upon herbivore exclusion in combination with fertilization correlated with a decrease in aboveground plant biomass and microbial activity, indicating a reduced storage of organic matter and microbial residues as soil C and N. The response of soil C and N pools to herbivore exclusion was contingent on temperature - herbivores likely cause losses of C and N in colder sites and increases in warmer sites. Additionally, grasslands that contain mammalian herbivores have the potential to sequester more N under increased temperature variability and nutrient enrichment than ungrazed grasslands. Our study highlights the importance of conserving mammalian herbivore populations in grasslands worldwide. We need to incorporate local-scale herbivory, and its interaction with nutrient enrichment and climate, within global-scale models to better predict land-atmosphere interactions under future climate change.

Environmental change can result in substantial shifts in community composition. The associated immigration and extinction events are likely constrained by the spatial distribution of species. Still, studies on environmental change typically quantify biotic responses at single spatial (time series within a single plot) or temporal (spatial beta diversity at single time points) scales, ignoring their potential interdependence. Here, we use data from a global network of grassland experiments to determine how turnover responses to two major forms of environmental change - fertilisation and herbivore loss - are affected by species pool size and spatial compositional heterogeneity. Fertilisation led to higher rates of local extinction, whereas turnover in herbivore exclusion plots was driven by species replacement. Overall, sites with more spatially heterogeneous composition showed significantly higher rates of annual turnover, independent of species pool size and treatment. Taking into account spatial biodiversity aspects will therefore improve our understanding of consequences of global and anthropogenic change on community dynamics.

Aim: Large-scale patterns linking energy availability, biological productivity and diversity form a central focus of ecology. Despite evidence that the activity and abundance of animals may be limited by climatic variables associated with regional biological productivity (e.g. mean annual precipitation and annual actual evapotranspiration), it is unclear whether plant-granivore interactions are themselves influenced by these climatic factors across broad spatial extents. We evaluated whether climatic conditions that are known to alter the abundance and activity of granivorous animals also affect rates of seed removal. Location: Eleven sites across temperate North America. Methods: We used a common protocol to assess the removal of the same seed species (Avena sativa) over a 2-day period. Model selection via the Akaike information criterion was used to determine a set of candidate binomial generalized linear mixed models that evaluated the relationship between local climatic data and post-dispersal seed predation. Results: Annual actual evapotranspiration was the single best predictor of the proportion of seeds removed. Annual actual evapotranspiration and mean annual precipitation were both positively related to mean seed removal and were included in four and three of the top five models, respectively. Annual temperature range was also positively related to seed removal and was an explanatory variable in three of the top four models. Main conclusions: Our work provides the first evidence that energy and precipitation, which are known to affect consumer abundance and activity, also translate to strong, predictable patterns of seed predation across a continent. More generally, these findings suggest that future changes in temperature and precipitation could have widespread consequences for plant species composition in grasslands, through impacts on plant recruitment.

Aboveground-belowground interactions exert critical controls on the composition and function of terrestrial ecosystems, yet the fundamental relationships between plant diversity and soil microbial diversity remain elusive. Theory predicts predominantly positive associations but tests within single sites have shown variable relationships, and associations between plant and microbial diversity across broad spatial scales remain largely unexplored. We compared the diversity of plant, bacterial, archaeal and fungal communities in one hundred and forty-five 1 m(2) plots across 25 temperate grassland sites from four continents. Across sites, the plant alpha diversity patterns were poorly related to those observed for any soil microbial group. However, plant beta diversity (compositional dissimilarity between sites) was significantly correlated with the beta diversity of bacterial and fungal communities, even after controlling for environmental factors. Thus, across a global range of temperate grasslands, plant diversity can predict patterns in the composition of soil microbial communities, but not patterns in alpha diversity.

Terrestrial ecosystem productivity is widely accepted to be nutrient limited(1). Although nitrogen (N) is deemed a key determinant of aboveground net primary production (ANPP)(2,3), the prevalence of co-limitation by N and phosphorus (P) is increasingly recognized(4-8). However, the extent to which terrestrial productivity is co-limited by nutrients other than N and P has remained unclear. Here, we report results from a standardized factorial nutrient addition experiment, in which we added N, P and potassium (K) combined with a selection of micronutrients (K+μ), alone or in concert, to 42 grassland sites spanning five continents, and monitored ANPP. Nutrient availability limited productivity at 31 of the 42 grassland sites. And pairwise combinations of N, P, and K+μ co-limited ANPP at 29 of the sites. Nitrogen limitation peaked in cool, high latitude sites. Our findings highlight the importance of less studied nutrients, such as K and micronutrients, for grassland productivity, and point to significant variations in the type and degree of nutrient limitation. We suggest that multiple-nutrient constraints must be considered when assessing the ecosystem-scale consequences of nutrient enrichment.

Exotic species dominate many communities; however the functional significance of species' biogeographic origin remains highly contentious. This debate is fuelled in part by the lack of globally replicated, systematic data assessing the relationship between species provenance, function and response to perturbations. We examined the abundance of native and exotic plant species at 64 grasslands in 13 countries, and at a subset of the sites we experimentally tested native and exotic species responses to two fundamental drivers of invasion, mineral nutrient supplies and vertebrate herbivory. Exotic species are six times more likely to dominate communities than native species. Furthermore, while experimental nutrient addition increases the cover and richness of exotic species, nutrients decrease native diversity and cover. Native and exotic species also differ in their response to vertebrate consumer exclusion. These results suggest that species origin has functional significance, and that eutrophication will lead to increased exotic dominance in grasslands.

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. Nutrient addition consistently reduced local diversity through light limitation, and herbivory rescued diversity at sites where it alleviated light limitation. Thus, species loss from anthropogenic eutrophication can be ameliorated in grasslands where herbivory increases ground-level light.

Abstract: Plant stoichiometry, the relative concentration of elements, is a key regulator of ecosystem functioning and is also being altered by human activities. In this paper we sought to understand the global drivers of plant stoichiometry and compare the relative contribution of climatic vs. anthropogenic effects. We addressed this goal by measuring plant elemental (C, N, P and K) responses to eutrophication and vertebrate herbivore exclusion at eighteen sites on six continents. Across sites, climate and atmospheric N deposition emerged as strong predictors of plot‐level tissue nutrients, mediated by biomass and plant chemistry. Within sites, fertilization increased total plant nutrient pools, but results were contingent on soil fertility and the proportion of grass biomass relative to other functional types. Total plant nutrient pools diverged strongly in response to herbivore exclusion when fertilized; responses were largest in ungrazed plots at low rainfall, whereas herbivore grazing dampened the plant community nutrient responses to fertilization. Our study highlights (1) the importance of climate in determining plant nutrient concentrations mediated through effects on plant biomass, (2) that eutrophication affects grassland nutrient pools via both soil and atmospheric pathways and (3) that interactions among soils, herbivores and eutrophication drive plant nutrient responses at small scales, especially at water‐limited sites. [ABSTRACT FROM AUTHOR]

Invasive, exotic grasses are increasing in tallgrass prairie and their dominance may be contributing to the decline of grassland butterflies through alterations in forage quality. Tall fescue (Schedonorus arundinaceus (Schreb.) Dumort.), an exotic grass covering millions of acres in the United States, can host a fungal endophyte, Epichloë coenophiala (Morgan-Jones & Gams). Alkaloids produced by the endophyte are known to be toxic to some foliar-feeding pest insects. Endophyte-infected tall fescue is commonly planted in hayfields, pastures, lawns, and is invading natural areas, but effects of the endophyte on nonpest insects such as butterflies are relatively unknown. Our objective was to investigate the role that tall fescue and its endophyte might play in the decline of grass skippers (Hesperiidae). We examined growth and survival parameters of tawny-edged skippers (Polites themistocles (Latreille)) that were reared on endophyte-infected tall fescue (E+), endophyte-free tall fescue (E-), and Kentucky bluegrass (KBG). KBG was included as a comparison because it is a cool season grass known to be palatable to P. themistocles larvae. Interestingly, results showed that the endophyte did not affect growth and survival of larvae compared to uninfected tall fescue, even though significant amounts of loline alkaloids (average 740 ppm) were measured in endophyte-infected plant material. Larvae feeding on KBG grew faster with greater survival rates than larvae on both tall fescue treatments. These results confirm that tall fescue invasion and dominance may be deteriorating the quality of grassland habitats for native pollinators; however, this effect does not appear to be linked to endophyte infection. [ABSTRACT FROM AUTHOR]

Under climate change, alterations to precipitation and temperature regimes are expected to impact ecosystem structure and function. These impacts may in part be determined by feedbacks between plants and associated microbes, including both endophytic fungal symbionts and viral pathogens., To test potential key components of these feedbacks in a grassland system, we conducted a field experiment that factorially manipulated temperature regime, precipitation regime and endophyte infection of a dominant species, Schedonorus arundinaceus. After one growing season, we evaluated the four most frequently occurring grass species for infection by three species of barley and cereal yellow dwarf viruses (B/ CYDVs). For S. arundinaceus, we also tracked changes in viral prevalence over an additional two growing seasons, for a total of three growing seasons., Plant species identity played a strong role in determining virus prevalence; Poa pratensis and Paspalum dilatatum were more frequently infected than either S. arundinaceus (endophyte infected or endophyte free) or Eleusine indica. Both elevated temperature and elevated precipitation impacted virus prevalence, but effects varied among years and interacted with fungal endophyte symbiosis for S. arundinaceus., In the final year of data collection, elevated precipitation increased virus prevalence in endophyte-infected, but not endophyte-free, S. arundinaceus. In the same year, but regardless of endophyte infection, elevated temperature by itself increased virus prevalence for S. arundinaceus. This effect of temperature on virus prevalence may have been driven by vector abundance because the viruses are obligately aphid-transmitted, and aphids were found more frequently in heated plots., Synthesis. Overall, our investigation experimentally demonstrates strong combined impacts of biotic and abiotic factors on disease dynamics in a grassland system. Impacts of climate change on virus prevalence in grasslands may depend on the responses of aphid vectors and the presence of endophytic fungal symbionts. [ABSTRACT FROM AUTHOR]

Invasive species may owe some of their success in competing and co-existing with native species to microbial symbioses they are capable of forming. Tall fescue is a cool-season, non-native, invasive grass capable of co-existing with native warm-season grasses in North American grasslands that frequently experience fire, drought, and cold winters, conditions to which the native species should be better-adapted than tall fescue. We hypothesized that tall fescue’s ability to form a symbiosis with Neotyphodium coenophialum, an aboveground fungal endophyte, may enhance its environmental stress tolerance and persistence in these environments. We used a greenhouse experiment to examine the effects of endophyte infection (E+ vs. E−), prescribed fire (1 burn vs. 2 burn vs. unburned control), and watering regime (dry vs. wet) on tall fescue growth. We assessed treatment effects for growth rates and the following response variables: total tiller length, number of tillers recruited during the experiment, number of reproductive tillers, tiller biomass, root biomass, and total biomass. Water regime significantly affected all response variables, with less growth and lower growth rates observed under the dry water regime compared to the wet. The burn treatments significantly affected total tiller length, number of reproductive tillers, total tiller biomass, and total biomass, but treatment differences were not consistent across parameters. Overall, fire seemed to enhance growth. Endophyte status significantly affected total tiller length and tiller biomass, but the effect was opposite what we predicted (E−>E+). The results from our experiment indicated that tall fescue was relatively tolerant of fire, even when combined with dry conditions, and that the fungal endophyte symbiosis was not important in governing this ecological ability. The persistence of tall fescue in native grassland ecosystems may be linked to other endophyte-conferred abilities not measured here (e.g., herbivory release) or may not be related to this plant-microbial symbiosis. [ABSTRACT FROM AUTHOR]

Based on regional-scale studies, aboveground production and litter decomposition are thought to positively covary, because they are driven by shared biotic and climatic factors. Until now we have been unable to test whether production and decomposition are generally coupled across climatically dissimilar regions, because we lacked replicated data collected within a single vegetation type across multiple regions, obfuscating the drivers and generality of the association between production and decomposition. Furthermore, our understanding of the relationships between production and decomposition rests heavily on separate meta-analyses of each response, because no studies have simultaneously measured production and the accumulation or decomposition of litter using consistent methods at globally relevant scales. Here, we use a multi-country grassland dataset collected using a standardized protocol to show that live plant biomass (an estimate of aboveground net primary production) and litter disappearance (represented by mass loss of aboveground litter) do not strongly covary. Live biomass and litter disappearance varied at different spatial scales. There was substantial variation in live biomass among continents, sites and plots whereas among continent differences accounted for most of the variation in litter disappearance rates. Although there were strong associations among aboveground biomass, litter disappearance and climatic factors in some regions (e.g. U.S. Great Plains), these relationships were inconsistent within and among the regions represented by this study. These results highlight the importance of replication among regions and continents when characterizing the correlations between ecosystem processes and interpreting their global-scale implications for carbon flux. We must exercise caution in parameterizing litter decomposition and aboveground production in future regional and global carbon models as their relationship is complex. [ABSTRACT FROM AUTHOR]

The large organic carbon (C) pools found in noncultivated grassland soils suggest that historically these ecosystems have had high rates of C sequestration. Changes in the soil C pool over time are a function of alterations in C input and output rates. Across the Great Plains and at individual sites through time, inputs of C (via aboveground production) are correlated with precipitation; however, regional trends in C outputs and the sensitivity of these C fluxes to annual variability in precipitation are less well known. To address the role of precipitation in controlling grassland C fluxes, and thereby soil C sequestration rates, we measured aboveground and belowground net primary production (ANPP-C and BNPP-C), soil respiration (SR-C), and litter decomposition rates for 2 years, a relatively dry year followed by a year of average precipitation, at five sites spanning a precipitation gradient in the Great Plains. ANPP-C, SR-C, and litter decomposition increased from shortgrass steppe (36, 454, and 24 g C m-2 y-1) to tallgrass prairie (180, 1221, and 208 g C m-2 y-1 for ANPP-C, SR-C, and litter decomposition, respectively). No significant regional trend in BNPP-C was found. Increasing precipitation between years increased rates of ANPP-C, BNPP-C, SR-C, and litter decomposition at most sites. However, regional patterns of the sensitivity of ANPP-C, BNPP-C, SR-C, and litter decomposition to between-year differences in precipitation varied. BNPP-C was more sensitive to between-year differences in precipitation than were the other C fluxes, and shortgrass steppe was more responsive than were mixed grass and tallgrass prairie. [ABSTRACT FROM AUTHOR]

The microbes contained within free‐living organisms can alter host growth, reproduction, and interactions with the environment. In turn, processes occurring at larger scales determine the local biotic and abiotic environment of each host that may affect the diversity and composition of the microbiome community. Here, we examine variation in the diversity and composition of the foliar fungal microbiome in the grass host, Andropogon gerardii, across four mesic prairies in the central United States. Composition of fungal endophyte communities differed among sites and among individuals within a site, but was not consistently affected by experimental manipulation of nutrient supply to hosts (A. gerardii) or herbivore reduction via fencing. In contrast, mean fungal diversity was similar among sites but was limited by total plant biomass at the plot scale. Our work demonstrates that distributed experiments motivated by ecological theory are a powerful tool to unravel the multiscale processes governing microbial community composition and diversity. [ABSTRACT FROM AUTHOR]