Cover crop species alter tallgrass prairie community assembly.

Cover crops are increasingly being used in ecological restoration projects, and are hypothesized to facilitate establishment of sown species by reducing weed abundances without competing with the target mix. We tested these predictions and examined the role of cover crop species on later species composition and diversity using cover crop seed treatments. Treatments included a fall seeding of one annual (Raphanus sativus or Avena sativa), one biennial (Oenothera biennis), one perennial species (Elymus canadensis), two grass-forb species combinations, or nothing as a control. All plots received the same diverse tallgrass prairie seed mix in March of the following year. Plant communities were sampled through five growing seasons. We found that cover crop treatments influenced community assembly, and that cover crop species varied in their effectiveness at reducing weed abundances, with the perennial grass E. canadensis being especially effective at reducing weeds. After 5 years, treatments were neutral in their effects on species diversity. However, composition of establishing plants (not including the cover crop) differed significantly among cover crop treatments on all sampling dates, indicating that treatments had long-term effects. Plots containing the C3 grass E. canadensis had greater abundance of C3 grasses and forbs, and reduced C4 grass abundance compared to other treatments. Our results indicate that cover crop species differ in their effects on assembling tallgrass prairie communities and that niche modification (i.e., species altering the abiotic environment in a way that favors species that benefit from the alterations) occurs when the cover crop is a perennial grass.

Long-term, amplified responses of soil organic carbon to nitrogen addition worldwide.

Soil organic carbon (SOC) is the largest carbon sink in terrestrial ecosystems and plays a critical role in mitigating climate change. Increasing reactive nitrogen (N) in ecosystems caused by anthropogenic N input substantially affects SOC dynamics. However, uncertainties remain concerning the effects of N addition on SOC in both organic and mineral soil layers over time at the global scale. Here, we analysed a large empirical data set spanning 60 years across 369 sites worldwide to explore the temporal dynamics of SOC to N addition. We found that N addition significantly increased SOC across the globe by 4.2% (2.7%-5.8%). SOC increases were amplified from short- to long-term N addition durations in both organic and mineral soil layers. The positive effects of N addition on SOC were independent of ecosystem types, mean annual temperature and precipitation. Our findings suggest that SOC increases largely resulted from the enhanced plant C input to soils coupled with reduced C loss from decomposition and amplification was associated with reduced microbial biomass and respiration under long-term N addition. Our study suggests that N addition will enhance SOC sequestration over time and contribute to future climate change mitigation.

Exotic species drive patterns of plant species diversity in 93 restored tallgrass prairies.

A primary goal of restoration ecology is to understand the factors that generate variability in species diversity and composition among restorations. Plant communities may assemble deterministically toward a common community type, or they may assemble stochastically, ending differently because of weather conditions during establishment, soil legacy effects, or exotic species propagule pressure. To test these alternative hypotheses, we sampled plant communities and soil at 93 randomly selected restored prairies distributed throughout Iowa, USA. Five remnant sites were sampled as a reference. We tested our hypotheses using multiple regressions and investigated the strength of direct and indirect effects on species diversity and richness using structural equation models. The prairie restorations were highly variable in their age, size, diversity, soil characteristics, and how they were managed post-seeding. The strongest predictor of plant species richness and diversity was the degree of invasion, as measured by the abundance of exotic species. Restorations planted with species-rich seed mixes had reduced exotic species abundance, which led indirectly to higher species richness of restorations. Sites with higher organic matter and a more linear shape had a direct positive effect on exotic abundance, which in turn decreased diversity. We found little support for deterministic assembly, and diversity did not increase with the age of planting. Our results indicate that restored prairie communities tend to assemble into states of high or low diversity, driven by invasion from exotic plant species. Management of exotic species is essential for maximizing species diversity in temperate grassland restorations.

Reversal of nitrogen-induced species diversity declines mediated by change in dominant grass and litter.

Atmospheric nitrogen (N) deposition reduces plant diversity. However, it often remains unclear how dominant species and litter accumulation feedbacks mediate N-induced plant diversity declines. We tested mechanisms of N-induced diversity change through dominant grasses and litter in a 7-year field experiment. Nitrogen addition reduced species richness, Shannon-Wiener diversity (H') and evenness from the second to the fourth year, however, surprisingly, increased them in the sixth and seventh year. The reversal in the response of diversity to N addition was explained by changes in grass dominance and standing litter accumulation. The diversity recovery during later years in fertilized plots was attributed to a decrease in the dominant grass and an increase in standing litter: standing litter reduced bud numbers of the dominant grass by decreasing light availability. The decreased light availability by standing litter reduced completion from the dominant species, which resulted in diversity increase. The negative feedback between dominant grasses and standing litter led to transient N-induced diversity loss in the short-term, but recovery of plant diversity in the long-term. Grassland management that affects litter accumulation, such as firing, grazing and mowing, can therefore, have substantial effects on the long-term response of plant diversity to N deposition.

High plant diversity is needed to maintain ecosystem services.

Biodiversity is rapidly declining worldwide, and there is consensus that this can decrease ecosystem functioning and services. It remains unclear, though, whether few or many of the species in an ecosystem are needed to sustain the provisioning of ecosystem services. It has been hypothesized that most species would promote ecosystem services if many times, places, functions and environmental changes were considered; however, no previous study has considered all of these factors together. Here we show that 84% of the 147 grassland plant species studied in 17 biodiversity experiments promoted ecosystem functioning at least once. Different species promoted ecosystem functioning during different years, at different places, for different functions and under different environmental change scenarios. Furthermore, the species needed to provide one function during multiple years were not the same as those needed to provide multiple functions within one year. Our results indicate that even more species will be needed to maintain ecosystem functioning and services than previously suggested by studies that have either (1) considered only the number of species needed to promote one function under one set of environmental conditions, or (2) separately considered the importance of biodiversity for providing ecosystem functioning across multiple years, places, functions or environmental change scenarios. Therefore, although species may appear functionally redundant when one function is considered under one set of environmental conditions, many species are needed to maintain multiple functions at multiple times and places in a changing world.

Exotic grassland species have stronger priority effects than natives regardless of whether they are cultivated or wild genotypes.

During community assembly, early arriving exotic species might suppress other species to a greater extent than do native species. Because most exotics were intentionally introduced, we hypothesize there was human selection on regeneration traits during introduction. This could have occurred at the across- or within-species level (e.g. during cultivar development). We tested these predictions by seeding a single species that was either native, exotic 'wild-type' (from their native range), or exotic 'cultivated' using 28 grassland species in a glasshouse experiment. Priority effects were assessed by measuring species' effect on establishment of species from a seed mix added 21 d later. Exotic species had higher germination and earlier emergence dates than native species, and differences were found in both 'wild' and 'cultivated' exotics. Exotic species reduced biomass and species diversity of later arriving species much more than native species, regardless of seed source. Results indicate that in situations in which priority effects are likely to be strong, effects will be greater when an exotic species arrives first than when a native species arrives first; and this difference is not merely a result of exotic species cultivation, but might be a general native-exotic difference that deserves further study.

Differences in beta diversity between exotic and native grasslands vary with scale along a latitudinal gradient.

Biodiversity can be partitioned into alpha, beta, and gamma components, and beta diversity is not as clearly understood. Biotic homogenization predicts that exotic species should lower beta diversity at global and continental scales, but it is still unclear how exotic species impact beta diversity at smaller scales. Exotic species could theoretically increase or decrease beta diversity relative to natives depending on many factors, including abiotic conditions, community assembly history, management, dispersal rates of species, and connectivity among patches. We sampled plant species abundances in 42 novel, exotic- and native-dominated (remnant) grasslands across a latitudinal gradient in the tallgrass prairie region, and tested whether exotic and native grasslands differed in beta diversity at three scales: across sites within the entire biome, across sites within regions, and across locations within sites. Exotic-dominated grasslands differed from native-dominated grasslands in beta diversity at all scales, but the direction of the difference changed from positive to negative as scales went from large to small. Contrary to expectations, exotic-dominated grasslands had higher beta diversity than native-dominated grasslands at the largest scale considered. This occurred because the identity of dominant exotic species varied across the latitudinal gradient, with many exotic grassland pairs exhibiting zero similarity, whereas native-dominated grasslands differed more gradually with distance. Beta diversity among sites within a region was variable, with exotic-dominated grasslands having 29% higher beta diversity than native grasslands in the south and 33% lower beta diversity in the north. Within sites, beta diversity was 26% lower in exotic-dominated than native grasslands. Our results provide evidence that different regional identities and abundances of exotics, and lack of connectivity in fragmented landscapes can alter beta diversity in unexpected ways across spatial scales.

Invaded grassland communities have altered stability-maintenance mechanisms but equal stability compared to native communities.

Theory predicts that stability should increase with diversity via several mechanisms. We tested predictions in a 5-year experiment that compared low-diversity exotic to high-diversity native plant mixtures under two irrigation treatments. The study included both wet and dry years. Variation in biomass across years (CV) was 50% lower in mixtures than monocultures of both native and exotic species. Growth among species was more asynchronous and overyielding values were greater during and after a drought in native than exotic mixtures. Mean-variance slopes indicated strong portfolio effects in both community types, but the intercept was higher for exotics than for natives, suggesting that exotics were inherently more variable than native species. However, this failed to result in higher CV's in exotic communities because species that heavily dominated plots tended to have lower than expected variance. Results indicate that diversity-stability mechanisms are altered in invaded systems compared to native ones they replaced.

Impacts of climate change drivers on C4 grassland productivity: scaling driver effects through the plant community.

Climate change drivers affect plant community productivity via three pathways: (i) direct effects of drivers on plants; (ii) the response of species abundances to drivers (community response); and (iii) the feedback effect of community change on productivity (community effect). The contribution of each pathway to driver-productivity relationships depends on functional traits of dominant species. We used data from three experiments in Texas, USA, to assess the role of community dynamics in the aboveground net primary productivity (ANPP) response of C4 grasslands to two climate drivers applied singly: atmospheric CO2 enrichment and augmented summer precipitation. The ANPP-driver response differed among experiments because community responses and effects differed. ANPP increased by 80-120g m(-2) per 100 µl l(-1) rise in CO2 in separate experiments with pasture and tallgrass prairie assemblages. Augmenting ambient precipitation by 128mm during one summer month each year increased ANPP more in native than in exotic communities in a third experiment. The community effect accounted for 21-38% of the ANPP CO2 response in the prairie experiment but little of the response in the pasture experiment. The community response to CO2 was linked to species traits associated with greater soil water from reduced transpiration (e.g. greater height). Community effects on the ANPP CO2 response and the greater ANPP response of native than exotic communities to augmented precipitation depended on species differences in transpiration efficiency. These results indicate that feedbacks from community change influenced ANPP-driver responses. However, the species traits that regulated community effects on ANPP differed from the traits that determined how communities responded to drivers.

Biodiversity, photosynthetic mode, and ecosystem services differ between native and novel ecosystems.

Human activities have caused non-native plant species with novel ecological interactions to persist on landscapes, and it remains controversial whether these species alter multiple aspects of communities and ecosystems. We tested whether native and exotic grasslands differ in species diversity, ecosystem services, and an important aspect of functional diversity (C3:C4 proportions) by sampling 42 sites along a latitudinal gradient and conducting a controlled experiment. Exotic-dominated grasslands had drastically lower plant diversity and slightly higher tissue N concentrations and forage quality compared to native-dominated sites. Exotic sites were strongly dominated by C4 species at southern and C3 species at northern latitudes with a sharp transition at 36-38°, whereas native sites contained C3:C4 mixtures. Large differences in C3:C4 proportions and temporal niche partitioning were found between native and exotic mixtures in the experiment, implying that differences in C3:C4 proportions along the latitudinal gradient are caused partially by species themselves. Our results indicate that the replacement of native- by exotic-dominated grasslands has created a management tradeoff (high diversity versus high levels of certain ecosystem services) and that models of global change impacts and C3/C4 distribution should consider effects of exotic species.

Predicting ecosystem stability from community composition and biodiversity.

As biodiversity is declining at an unprecedented rate, an important current scientific challenge is to understand and predict the consequences of biodiversity loss. Here, we develop a theory that predicts the temporal variability of community biomass from the properties of individual component species in monoculture. Our theory shows that biodiversity stabilises ecosystems through three main mechanisms: (1) asynchrony in species' responses to environmental fluctuations, (2) reduced demographic stochasticity due to overyielding in species mixtures and (3) reduced observation error (including spatial and sampling variability). Parameterised with empirical data from four long-term grassland biodiversity experiments, our prediction explained 22-75% of the observed variability, and captured much of the effect of species richness. Richness stabilised communities mainly by increasing community biomass and reducing the strength of demographic stochasticity. Our approach calls for a re-evaluation of the mechanisms explaining the effects of biodiversity on ecosystem stability.

Biodiversity simultaneously enhances the production and stability of community biomass, but the effects are independent.

To predict the ecological consequences of biodiversity loss, researchers have spent much time and effort quantifying how biological variation affects the magnitude and stability of ecological processes that underlie the functioning of ecosystems. Here we add to this work by looking at how biodiversity jointly impacts two aspects of ecosystem functioning at once: (1) the production of biomass at any single point in time (biomass/area or biomass/ volume), and (2) the stability of biomass production through time (the CV of changes in total community biomass through time). While it is often assumed that biodiversity simultaneously enhances both of these aspects of ecosystem functioning, the joint distribution of data describing how species richness regulates productivity and stability has yet to be quantified. Furthermore, analyses have yet to examine how diversity effects on production covary with diversity effects on stability. To overcome these two gaps, we reanalyzed the data from 34 experiments that have manipulated the richness of terrestrial plants or aquatic algae and measured how this aspect of biodiversity affects community biomass at multiple time points. Our reanalysis confirms that biodiversity does indeed simultaneously enhance both the production and stability of biomass in experimental systems, and this is broadly true for terrestrial and aquatic primary producers. However, the strength of diversity effects on biomass production is independent of diversity effects on temporal stability. The independence of effect sizes leads to two important conclusions. First, while it may be generally true that biodiversity enhances both productivity and stability, it is also true that the highest levels of productivity in a diverse community are not associated with the highest levels of stability. Thus, on average, diversity does not maximize the various aspects of ecosystem functioning we might wish to achieve in conservation and management. Second, knowing how biodiversity affects productivity gives no information about how diversity affects stability (or vice versa). Therefore, to predict the ecological changes that occur in ecosystems after extinction, we will need to develop separate mechanistic models for each independent aspect of ecosystem functioning.

Increasing native, but not exotic, biodiversity increases aboveground productivity in ungrazed and intensely grazed grasslands.

Species-rich native grasslands are frequently converted to species-poor exotic grasslands or pastures; however, the consequences of these changes for ecosystem functioning remain unclear. Cattle grazing (ungrazed or intensely grazed once), plant species origin (native or exotic), and species richness (4-species mixture or monoculture) treatments were fully crossed and randomly assigned to plots of grassland plants. We tested whether (1) native and exotic plots exhibited different responses to grazing for six ecosystem functions (i.e., aboveground productivity, light interception, fine root biomass, tracer nitrogen uptake, biomass consumption, and aboveground biomass recovery), and (2) biodiversity-ecosystem functioning relationships depended on grazing or species origin. We found that native and exotic species exhibited different responses to grazing for three of the ecosystem functions we considered. Intense grazing decreased fine root biomass by 53% in exotic plots, but had no effect on fine root biomass in native plots. The proportion of standing biomass consumed by cattle was 16% less in exotic than in native grazed plots. Aboveground biomass recovery was 30% less in native than in exotic plots. Intense grazing decreased aboveground productivity by 25%, light interception by 14%, and tracer nitrogen uptake by 54%, and these effects were similar in native and exotic plots. Increasing species richness from one to four species increased aboveground productivity by 42%, and light interception by 44%, in both ungrazed and intensely grazed native plots. In contrast, increasing species richness did not influence biomass production or resource uptake in ungrazed or intensely grazed exotic plots. These results suggest that converting native grasslands to exotic grasslands or pastures changes ecosystem structure and processes, and the relationship between biodiversity and ecosystem functioning.

Biodiversity, productivity and the temporal stability of productivity: patterns and processes.

Theory predicts that the temporal stability of productivity, measured as the ratio of the mean to the standard deviation of community biomass, increases with species richness and evenness. We used experimental species mixtures of grassland plants to test this hypothesis and identified the mechanisms involved. Additionally, we tested whether biodiversity, productivity and temporal stability were similarly influenced by particular types of species interactions. We found that productivity was less variable among years in plots planted with more species. Temporal stability did not depend on whether the species were planted equally abundant (high evenness) or not (realistically low evenness). Greater richness increased temporal stability by increasing overyielding, asynchrony of species fluctuations and statistical averaging. Species interactions that favoured unproductive species increased both biodiversity and temporal stability. Species interactions that resulted in niche partitioning or facilitation increased both productivity and temporal stability. Thus, species interactions can promote biodiversity and ecosystem services.

Biodiversity maintenance mechanisms differ between native and novel exotic-dominated communities.

In many systems, native communities are being replaced by novel exotic-dominated ones. We experimentally compared species diversity decline between nine-species grassland communities under field conditions to test whether diversity maintenance mechanisms differed between communities containing all exotic or all native species using a pool of 40 species. Aboveground biomass was greater in exotic than native plots, and this difference was larger in mixtures than in monocultures. Species diversity declined more in exotic than native communities and declines were explained by different mechanisms. In exotic communities, overyielding species had high biomass in monoculture and diversity declined linearly as this selection effect increased. In native communities, however, overyielding species had low biomass in monoculture and there was no relationship between the selection effect and diversity decline. This suggests that, for this system, yielding behaviour is fundamentally different between presumably co-evolved natives and coevolutionarily naive exotic species, and that native-exotic status is important to consider.

Species interaction mechanisms maintain grassland plant species diversity.

Development of theory has outpaced experimental tests for most maintenance of diversity mechanisms. Here we demonstrate how data from biodiversity-ecosystem functioning experiments can be used to determine the mechanisms that maintain plant species diversity. We hypothesized that grassland plant diversity is maintained by two classes of mechanisms: (1) equalizing mechanisms, which reduce asymmetric competition by reducing differences in monoculture biomass production among species in mixture, and (2) species interaction mechanisms, which increase overyielding by increasing niche partitioning and facilitation among species in mixture. Specifically, equalizing mechanisms reduce the coefficient of variation in monoculture biomass production among species in mixture. Species interaction mechanisms increase species overyielding in mixture, especially for low-biomass species. We tested these predictions with a seven-year data set from an experiment that varied grassland plant species evenness and richness. We used path analysis to model effects of these mechanisms on annual and multiyear changes in diversity. We found that diversity was frequently maintained by species interaction mechanisms and was infrequently maintained by equalizing mechanisms. Species interaction mechanisms maintained diversity by allowing the species that produced the least biomass in monoculture to benefit the most from species interactions in mixture. Equalizing mechanisms infrequently maintained diversity because asymmetric competition infrequently resulted in competitive exclusion. We propose that this mechanistic framework be used to better understand the specific processes that influence diversity.

Plant invasions differentially affected by diversity and dominant species in native- and exotic-dominated grasslands.

Plant invasions are an increasingly serious global concern, especially as the climate changes. Here, we explored how plant invasions differed between native- and novel exotic-dominated grasslands with experimental addition of summer precipitation in Texas in 2009. Exotic species greened up earlier than natives by an average of 18 days. This was associated with a lower invasion rate early in the growing season compared to native communities. However, invasion rate did not differ significantly between native and exotic communities across all sampling times. The predictors of invasion rate differed between native and exotic communities, with invasion being negatively influenced by species richness in natives and by dominant species in exotics. Interestingly, plant invasions matched the bimodal pattern of precipitation in Temple, Texas, and did not respond to the pulse of precipitation during the summer. Our results suggest that we will need to take different approaches in understanding of invasion between native and exotic grasslands. Moreover, with anticipated increasing variability in precipitation under global climate change, plant invasions may be constrained in their response if the precipitation pulses fall outside the normal growing period of invaders.

Urea additions and defoliation affect plant responses to elevated CO2 in a C3 grass from Yellowstone National Park.

A common grass from Yellowstone National Park, Stipa occidentalis, was grown in a factorial experiment to determine if its response to the direct effects of elevated CO2 would be affected by defoliation, and urea additions simulating the N in a urine hit. Plants were grown in tall pots (to mimic rooting depth in the field) in growth chambers under elevated (700 ppm) and ambient (370 ppm) CO2, were defoliated or left undefoliated, and given N-supply rates based on field mineralization rates (untreated) or with an additional 40 g N/m2. Growth increases in response to elevated CO2 were largest when plants remained unclipped and received urea additions, and were found primarily in crowns and roots (storage organs). Aboveground biomass, which is the part of the plant consumed by grazing mammals, was not affected by elevated CO2. The elevated CO2 treatment caused a reduction in leaf percent N. However, there was a significant interaction between the CO2 and urea treatments, resulting in a larger difference in leaf percent N between urea-treated and control plants under elevated than under ambient CO2. Hence, elevations in atmospheric CO2 may cause an increase in the amount of urine-hit-induced spatial variability in temperate grasslands. Since food quantity remained largely unchanged in response to elevated CO2, and forage N content went down, grazing mammals may be negatively affected by increases in atmospheric CO2.

Will increases in atmospheric CO2 affect regrowth following grazing in C4 grasses from tropical grasslands? A test with Sporobolus kentrophyllus.

We grew a C4 grass from the Serengeti ecosystem under ambient (370 ppm) and elevated (700 ppm) CO2, and under clipped and unclipped conditions to test whether regrowth following grazing would be affected by elevated CO2. Above-ground productivity was slightly decreased under elevated CO2, and was similar between clipped and unclipped plants. Regrowth (clipping offtake) following clipping was similar in the two CO2 treatments, and there was no CO2 by clipping interaction on biomass, productivity, or leaf nutrient concentrations. Based on this evidence, we suggest that C4 grasses from the Serengeti will show little direct response to future increases in atmospheric CO2.