Many hypotheses dealing with the success of invasive plant species concern plant–herbivore interactions. The invasional meltdown and enemy inversion hypotheses suggest that non-native herbivores may indirectly facilitate the invasion of a non-native plant species by either favorably changing environmental conditions or reducing competition from native plant species. Our objective was to determine the role of herbivory by the non-native snail Otala lactea in structuring California grassland communities. We conducted two experiments to examine the feeding preferences of O. lactea for eight representative grassland species. Overall, O. lactea preferred Brassica nigra, a non-native forb, over all other species tested. Field monocultures of B. nigra supported significantly higher snail densities than monocultures of any of the other species tested. O. lactea also preferred B. nigra over all other species tested in controlled laboratory feeding trials. However, based on trait comparisons of each of the eight grassland species, we cannot pinpoint the preference for B. nigra to a basic nutritional requirement on the part of the herbivore or an allocation to defense on the part of the plants. Our study provides evidence for an antagonistic relationship between a non-native herbivore and a non-native plant species in their invasive range. We term this relationship “invasional antagonism”.

Point 1: The ecological models of Alfred J. Lotka and Vito Volterra have had an enormous impact on ecology over the past century. Some of the earliest-and clearest-experimental tests of these models were famously conducted by Georgy Gause in the 1930s. Although well known, the data from these experiments are not widely available and are often difficult to analyze using standard statistical and computational tools. Point 2: Here, we introduce the gauseR package, a collection of tools for fitting Lotka-Volterra models to time series data of one or more species. The package includes several methods for parameter estimation and optimization, and includes 42 datasets from Gause's species interaction experiments and related work. Additionally, we include with this paper a short blog post discussing the historical importance of these data and models, and an R vignette with a walk-through introducing the package methods. The package is available for download at github.com/adamtclark/gauseR. Point 3: To demonstrate the package, we apply it to several classic experimental studies from Gause, as well as two other well-known datasets on multi-trophic dynamics on Isle Royale, and in spatially structured mite populations. In almost all cases, models fit observations closely and fitted parameter values make ecological sense. Point 4: Taken together, we hope that the methods, data, and analyses that we present here provide a simple and user-friendly way to interact with complex ecological data. We are optimistic that these methods will be especially useful to students and educators who are studying ecological dynamics, as well as researchers who would like a fast tool for basic analyses.

Changes in vegetation structure and composition, particularly due to the invasion of exotic species, are predicted to influence biosphere-atmosphere exchanges of mass and energy. Invasion of Cynara cardunculus (cardoon or artichoke thistle), a perennial, non-native thistle in coastal California grasslands presently dominated by non-native annual grasses, may alter rates of ecosystem CO₂ exchange and evapotranspiration (ET). During spring and summer 2006, we compared midday maximum net ecosystem CO₂ exchange (NEE) and ET among adjacent grassland plots where Cynara was present and where it was absent. Measurements of NEE supported the prediction that deeply-rooted Cynara increase midday ecosystem C-assimilation. Cynara-mediated shifts in NEE were associated with increases in ecosystem photosynthesis rather than changes in ecosystem respiration. Furthermore, the presence of Cynara was associated with increased ET during the growing season. An increase in aboveground live biomass (a proxy for leaf area) associated with Cynara invasion may underlie shifts in ecosystem CO₂ and water vapor exchange. Following mid-growing season sampling during April, we removed Cynara from half of the Cynara-containing plots with spot applications of herbicide. Three weeks later, midday fluxes in removal plots were indistinguishable from those in plots where Cynara was never present suggesting a lack of biogeochemical legacy effects. Similar to woody-encroachment in some semi-arid ecosystems, Cynara invasion increases midday ecosystem CO₂ assimilation and evapotranspiration rates and has the potential to increase C-storage in California coastal grasslands.

Declines in global biodiversity have inspired a generation of studies that seek to characterize relationships between biodiversity and ecosystem functioning. The metrics for complementarity and selection effects derived by Loreau and Hector in 2001 remain some of the most influential and widely used statistics for studying these relationships. These metrics quantify the degree to which the effect of biodiversity on a given ecosystem function depends on only a few species that perform well in monoculture and in mixture (the selection effect) or if the effect of biodiversity on a given ecosystem function is independent of monoculture performance (the complementarity effect). This distinction may be useful in determining the consequences of the loss of rare versus common or dominant species in natural systems. However, because these metrics require observations of all species in a community in monoculture, applications in natural systems have been limited.Here, we derive a statistical augmentation of the original partition, which can be applied to incomplete random samples of species drawn from a larger pool. This augmentation controls for the bias introduced by using only a subsample of species in monocultures rather than having monocultures of all species.Using simulated and empirical examples, we demonstrate the robustness of these metrics, and provide source code for calculating them. We find that these augmentations provide a reliable estimate of complementarity and selection effects as long as approximately 50% of the species present in mixture are present in monoculture and these species represent a random subset of the mixture.We foresee two primary applications for this method: (a) estimating complementarity and selection effects for experimentally assembled communities where monoculture data are lacking for some species, and (b) extrapolating results from biodiversity experiments to diverse natural systems.

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.

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.

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.