Search

Global changes will modify future nutrient availability with implications for grassland biogeochemistry. Soil organic matter (SOM) is central to grasslands for both provision of nutrients and climate mitigation through carbon (C) storage. While we know that C and nitrogen (N) in SOM can be influenced by greater nutrient availability, we lack understanding of nutrient effects on C and N coupling and stability in soil. Different SOM fractions have different functional relevance and mean residence times, i.e., mineral-associated organic matter (MAOM) has a higher mean residence time than particulate organic matter (POM). By separating effects of nutrient supply on the different SOM fractions, we can better evaluate changes in soil C and N coupling and stability and associated mechanisms. To this end, we studied responses of C and N ratios and distributions across POM and MAOM to 6–10 years of N, phosphorus (P), potassium and micronutrients (K +µ), and combined NPK +µ additions at 11 grassland sites spanning 3 continents and globally relevant environmental gradients in climate, plant growth, soil texture, and nutrient availability. We found addition of N and NPK +µ generally reduced C:N in MAOM and POM. However, at low fertility and at warm, sandy sites, nutrient addition promoted higher MAOM and POM C:N, respectively. Addition of NPK +µ also promoted C storage in POM relative to MAOM, and this was consistent across sites. Our results suggest that addition of macro- and micronutrients consistently decrease SOM stabilization, whereas responses of soil C:N stoichiometry were contingent on SOM fraction and environmental conditions.

SummaryEutrophication impacts grassland diversity such as species richness, functional trait diversity and composition, but whether and how these changes affect ecosystem functional stability under increasing climate extremes is unknown. We quantify the direct and diversity-mediated effects of nutrient addition on functional stability of productivity as measured by resistance during and recovery after dry and wet growing seasons, and temporal invariability across 19 grasslands spanning four continents. We use leaf traits measured in the field to construct slow-fast leaf economic spectrum and calculate functional trait composition and diversity of the slow-fast trait. Nutrient addition increased resistance during dry growing seasons mainly through its direct effects. It also increased resistance during wet growing seasons directly, but these positive effects were weakened by indirect negative effects through reducing slow-fast functional diversity. Nutrient addition had weak effects on recovery after dry growing seasons, but it decreased recovery after wet growing seasons mainly through its direct effects. Nutrient addition increased temporal invariability through resistance, but decreased it through recovery and reducing species richness, leading to overall weak net effects. Thus, processes and influencing pathways independent of plant diversity should be carefully taken into account in understanding ecosystem stability in response to increasing global changes.

The effects of altered nutrient supplies and herbivore density on species diversity vary with spatial scale, because coexistence mechanisms are scale dependent. This scale dependence may alter the shape of the species–area relationship (SAR), which can be described by changes in species richness (S) as a power function of the sample area (A): S = cAz, where c and z are constants. We analysed the effects of experimental manipulations of nutrient supply and herbivore density on species richness across a range of scales (0.01–75 m²) at 30 grasslands in 10 countries. We found that nutrient addition reduced the number of species that could co-occur locally, indicated by the SAR intercepts (log c), but did not affect the SAR slopes (z). As a result, proportional species loss due to nutrient enrichment was largely unchanged across sampling scales, whereas total species loss increased over threefold across our range of sampling scales.

Plant‐soil feedbacks (PSFs) have been shown to strongly affect plant performance under controlled conditions, and PSFs are thought to have far reaching consequences for plant population dynamics and the structuring of plant communities. However, thus far the relationship between PSF and plant species abundance in the field is not consistent. Here, we synthesize PSF experiments from tropical forests to semiarid grasslands, and test for a positive relationship between plant abundance in the field and PSFs estimated from controlled bioassays. We meta‐analyzed results from 22 PSF experiments and found an overall positive correlation (0.12 ≤ r¯ ≤ 0.32) between plant abundance in the field and PSFs across plant functional types (herbaceous and woody plants) but also variation by plant functional type. Thus, our analysis provides quantitative support that plant abundance has a general albeit weak positive relationship with PSFs across ecosystems. Overall, our results suggest that harmful soil biota tend to accumulate around and disproportionately impact species that are rare. However, data for the herbaceous species, which are most common in the literature, had no significant abundance‐PSFs relationship. Therefore, we conclude that further work is needed within and across biomes, succession stages and plant types, both under controlled and field conditions, while separating PSF effects from other drivers (e.g., herbivory, competition, disturbance) of plant abundance to tease apart the role of soil biota in causing patterns of plant rarity versus commonness., Across studies, we detected a small but significant positive correlation between plant‐soil feedbacks measured during controlled experiments and plant field abundance. Plants that are rare are seemingly burdened more by the negative effects of harmful soil biota than common plants.

Almost 50 years ago, Michael Rosenzweig pointed out that nutrient addition can destabilise food webs, leading to loss of species and reduced ecosystem function through the paradox of enrichment. Around the same time, David Tilman demonstrated that increased nutrient loading would also be expected to cause competitive exclusion leading to deleterious changes in food web diversity. While both concepts have greatly illuminated general diversity-stability theory, we currently lack a coherent framework to predict how nutrients influence food web stability across a landscape. This is a vitally important gap in our understanding, given mounting evidence of serious ecological disruption arising from anthropogenic displacement of resources and organisms. Here, we combine contemporary theory on food webs and meta-ecosystems to show that nutrient additions are indeed expected to drive loss in stability and function in human-impacted regions. Our models suggest that destabilisation is more likely to be caused by the complete loss of an equilibrium due to edible plant species being competitively excluded. In highly modified landscapes, spatial nutrient transport theory suggests that such instabilities can be amplified over vast distances from the sites of nutrient addition. Consistent with this theoretical synthesis, the empirical frequency of these distant propagating ecosystem imbalances appears to be growing. This synthesis of theory and empirical data suggests that human modification of the Earth is strongly connecting distantly separated ecosystems, causing rapid, expansive and costly nutrient-driven instabilities over vast areas of the planet. Similar to existing food web theory, the corollary to this spatial nutrient theory is that slowing down spatial nutrient pathways can be a potent means of stabilising degraded ecosystems.

Aim: Climate variability threatens to destabilize production in many ecosystems. Asynchronous species dynamics may buffer against such variability when a decrease in performance by some species is offset by an increase in performance of others. However, high climatic variability can eliminate species through stochastic extinctions or cause similar stress responses among species that reduce buffering. Local conditions, such as soil nutrients, can also alter production stability directly or by influencing asynchrony. We test these hypotheses using a globally distributed sampling experiment. Location: Grasslands in North America, Europe and Australia. Time period: Annual surveys over 5 year intervals occurring between 2007 and 2014. Major taxa studied: Herbaceous plants. Methods: We sampled annually the per species cover and aboveground community biomass [net primary productivity (NPP)], plus soil chemical properties, in 29 grasslands. We tested how soil conditions, combined with variability in precipitation and temperature, affect species richness, asynchrony and temporal stability of primary productivity. We used bivariate relationships and structural equation modelling to examine proximate and ultimate relationships. Results: Climate variability strongly predicted asynchrony, whereas NPP stability was more related to soil conditions. Species richness was structured by both climate variability and soils and, in turn, increased asynchrony. Variability in temperature and precipitation caused a unimodal asynchrony response, with asynchrony being lowest at low and high climate variability. Climate impacted stability indirectly, through its effect on asynchrony, with stability increasing at higher asynchrony owing to lower inter-annual variability in NPP. Soil conditions had no detectable effect on asynchrony but increased stability by increasing the mean NPP, especially when soil organic matter was high. Main conclusions: We found globally consistent evidence that climate modulates species asynchrony but that the direct effect on stability is low relative to local soil conditions. Nonetheless, our observed unimodal responses to variability in temperature and precipitation suggest asynchrony thresholds, beyond which there are detectable destabilizing impacts of climate on primary productivity.

Nutrient enrichment can simultaneously increase and destabilise plant biomass production, with co-limitation by multiple nutrients potentially intensifying these effects. Here, we test how factorial additions of nitrogen (N), phosphorus (P) and potassium with essential nutrients (K+) affect the stability (mean/standard deviation) of aboveground biomass in 34 grasslands over 7 years. Destabilisation with fertilisation was prevalent but was driven by single nutrients, not synergistic nutrient interactions. On average, N-based treatments increased mean biomass production by 21-51% but increased its standard deviation by 40-68% and so consistently reduced stability. Adding P increased interannual variability and reduced stability without altering mean biomass, while K+ had no general effects. Declines in stability were largest in the most nutrient-limited grasslands, or where nutrients reduced species richness or intensified species synchrony. We show that nutrients can differentially impact the stability of biomass production, with N and P in particular disproportionately increasing its interannual variability.

Fertilisation experiments have demonstrated that nutrient availability is a key determinant of biomass production and carbon sequestration in grasslands. However, the influence of nutrients in explaining spatial variation in grassland biomass production has rarely been assessed. Using a global dataset comprising 72 sites on six continents, we investigated which of 16 soil factors that shape nutrient availability associate most strongly with variation in grassland aboveground biomass. Climate and N deposition were also considered. Based on theory-driven structural equation modelling, we found that soil micronutrients (particularly Zn and Fe) were important predictors of biomass and, together with soil physicochemical properties and C:N, they explained more unique variation (32%) than climate and N deposition (24%). However, the association between micronutrients and biomass was absent in grasslands limited by NP. These results highlight soil properties as key predictors of global grassland biomass production and point to serial co-limitation by NP and micronutrients.

Grassland ecosystems cover around 37% of the ice-free land surface on Earth and have critical socioeconomic importance globally. As in many terrestrial ecosystems, biological dinitrogen (N2) fixation represents an essential natural source of nitrogen (N). The ability to fix atmospheric N2 is limited to diazotrophs, a diverse guild of bacteria and archaea. To elucidate the abiotic (climatic, edaphic), biotic (vegetation), and spatial factors that govern diazotrophic community composition in global grassland soils, amplicon sequencing of the dinitrogenase reductase gene—nifH—was performed on samples from a replicated standardized nutrient [N, phosphorus (P)] addition experiment in 23 grassland sites spanning four continents. Sites harbored distinct and diverse diazotrophic communities, with most of reads assigned to diazotrophic taxa within the Alphaproteobacteria (e.g., Rhizobiales), Cyanobacteria (e.g., Nostocales), and Deltaproteobacteria (e.g., Desulforomonadales) groups. Likely because of the wide range of climatic and edaphic conditions and spatial distance among sampling sites, only a few of the taxa were present at all sites. The best model describing the variation among soil diazotrophic communities at the OTU level combined climate seasonality (temperature in the wettest quarter and precipitation in the warmest quarter) with edaphic (C:N ratio, soil texture) and vegetation factors (various perennial plant covers). Additionally, spatial variables (geographic distance) correlated with diazotrophic community variation, suggesting an interplay of environmental variables and spatial distance. The diazotrophic communities appeared to be resilient to elevated nutrient levels, as 2–4 years of chronic N and P additions had little effect on the community composition. However, it remains to be seen, whether changes in the community composition occur after exposure to long-term, chronic fertilization regimes.

Spatial rarity is often used to predict extinction risk, but rarity can also occur temporally. Perhaps more relevant in the context of global change is whether a species is core to a community (persistent) or transient (intermittently present), with transient species often susceptible to human activities that reduce niche space. Using 5–12 yr of data on 1,447 plant species from 49 grasslands on five continents, we show that local abundance and species persistence under ambient conditions are both effective predictors of local extinction risk following experimental exclusion of grazers or addition of nutrients; persistence was a more powerful predictor than local abundance. While perturbations increased the risk of exclusion for low persistence and abundance species, transient but abundant species were also highly likely to be excluded from a perturbed plot relative to ambient conditions. Moreover, low persistence and low abundance species that were not excluded from perturbed plots tended to have a modest increase in abundance following perturbance. Last, even core species with high abundances had large decreases in persistence and increased losses in perturbed plots, threatening the long-term stability of these grasslands. Our results demonstrate that expanding the concept of rarity to include temporal dynamics, in addition to local abundance, more effectively predicts extinction risk in response to environmental change than either rarity axis predicts alone. © 2021 by the Ecological Society of America

Arctic plants are adapted to climatic variability, but their long-term responses to warming remain unclear. Responses may occur by range shifts, phenological adjustments in growth and reproduction, or both. Here, we compare distribution and phenology of 83 arctic and boreal mountain species, sampled identically in the early 20th (1917–1919) and 21st centuries (2017–2018) from a region of northern Sweden that has warmed significantly. We test two compensatory hypotheses to high-latitude warming—upward shifts in distribution, and earlier or extended growth and reproduction. For distribution, we show dramatic upward migration by 69% of species, averaging 6.1 m per decade, especially boreal woodland taxa whose upward expansion has reduced arctic montane habitat by 30%. Twenty percent of summit species showed distributional shifts but downward, especially moisture-associated snowbed flora. For phenology, we detected wide inter-annual variability in the onset of leafing and flowering in both eras. However, there was no detectable change in growing-season length, relating to two mechanisms. First, plot-level snow melt data starting in 1917 demonstrated that melt date, rather than vernal temperatures, better predicts plant emergence, with snow melt influenced by warmer years having greater snowfall—warmer springs did not always result in earlier emergence because snowbeds can persist longer. Second, the onset of reproductive senescence between eras was similar, even when plant emergence was earlier by a month, possibly due to intensified summer heat stress or hard-wired ‘canalization’ where senescence occurs regardless of summer temperature. Migrations in this system have possibly buffered arctic species against displacement by boreal expansion and warming, but ongoing temperature increases, woody plant invasion, and a potential lack of flexibility in timing of senescence may foreshadow challenges.

Vapocoolant sprays are convenient forms of cold temperature analgesia. These sprays may not be suitable for all patients with particular concern for patients with sickle cell disease. To prevent any further cases from occurring, we propose adding a more specific cautionary statement to the manufacturer guidelines. We also hope that medical personnel can help patients with sickle cell avoid topical and environmental cold temperature triggers for sickle vaso-occlusive pain and reduce the suffering in this rare disease.

Abandoned agricultural lands often accumulate soil carbon (C) following depletion of soil C by cultivation. The potential for this recovery to provide significant C storage benefits depends on the rate of soil C accumulation, which, in turn, may depend on nutrient supply rates. We tracked soil C for almost four decades following intensive agricultural soil disturbance along an experimentally imposed gradient in nitrogen (N) added annually in combination with other macro- and micro-nutrients. Soil %C accumulated over the course of the study in unfertilized control plots leading to a gain of 6.1 Mg C ha-1 in the top 20 cm of soil. Nutrient addition increased soil %C accumulation leading to a gain of 17.8 Mg C ha-1 in fertilized plots, nearly a threefold increase over the control plots. These results demonstrate that substantial increases in soil C in successional grasslands following agricultural abandonment occur over decadal timescales, and that C gain is increased by high supply rates of soil nutrients. In addition, soil %C continued to increase for decades under elevated nutrient supply, suggesting that short-term nutrient addition experiments underestimate the effects of soil nutrients on soil C accumulation.

Nutrients and herbivores have independent effects on the temporal stability of aboveground biomass in grasslands; however, their joint effects may not be additive and may also depend on spatial scales. In an experiment adding nutrients and excluding herbivores in 34 globally distributed grasslands, we found that nutrients and herbivores mainly had additive effects. Nutrient addition consistently reduced stability at the local and larger spatial scales (aggregated local communities), while herbivore exclusion weakly reduced stability at these scales. Moreover, nutrient addition reduced stability primarily by causing changes in local community composition over time and by reducing local species richness and evenness. In contrast, herbivore exclusion weakly reduced stability at the larger scale mainly by decreasing asynchronous dynamics among local communities, but also by weakly decreasing local species richness. Our findings indicate disentangling the influences of processes operating at different spatial scales may improve conservation and management in stabilizing grassland biomass.

Soil organic carbon (SOC) is strongly affected by farm cropping, which covers >10% of the earth's surface. Land retirement of marginal fields, now a global initiative, can increase SOC storage but reported accumulation rates are variable. Here, we quantify SOC in crop fields and retired marginal land in an intensely farmed 10,000 km 2 region of central North America, testing nutrients, soil texture and management as drivers of SOC storage. Overwhelmingly, SOC was associated with farm management with among-farm differences varying >fourfold (17.4–81 t ha −1) in the top 15 cm. Total farm SOC averaged 502.2 t farm −1 but again ranged widely (216–1611 t farm −1). Farm-specific SOC was often, but not always, higher on farms with N-rich silt-clay soils, and lower on sandy soils with higher P relating to former tobacco production. In contrast, within-farm SOC between crop fields and retired land did not significantly differ with time. Low SOC on retired lands was associated with persistently high soil N and P and elevated microbial respiration. Retired soils did possess substantially larger pools of lignin-rich root biomass to depths of 60 cm, which may signify eventual SOC accumulation possibly as nutrient legacies diminish. Our work shows that management legacy, interacting with soil texture and nutrients, predicts SOC more than short-term retirement . Indeed, crop fields averaged 67% of farm SOC because they represented up to 94% of total farm area – SOC retention on cropland remains a management priority, above and beyond gains with retirement. Interestingly, the largest per-volume SOC levels were in remnant forest that contained 25% of farm SOC despite only averaging 11% of farm area. Maintaining SOC stocks in farm landscapes may be more quickly attained by protecting remnant forest, with retired lands needing time to re-build SOC stocks.

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

Biodiversity—both above‐ and belowground—influences multiple functions in terrestrial ecosystems. Yet, it is unclear whether differences in above‐ and belowground species composition (β‐diversity) are associated with differences in multiple ecosystem functions (e.g., spatial turnover in ecosystem function). Here, we partitioned the contributions of above‐ and belowground β‐diversity and abiotic factors (geographic distance, differences in environments) on the spatial turnover of multiple grassland ecosystem functions. We compiled a dataset of plant and soil microbial communities and six indicators of grassland ecosystem functions (i.e., plant aboveground live biomass, plant nitrogen [N], plant phosphorus [P], root biomass, soil total N, and soil extractable P) from 18 grassland sites on four continents contributing to the Nutrient Network experiment. We used Mantel tests and structural equation models to disentangle the relationship between above‐ and belowground β‐diversity and spatial turnover in grassland ecosystem functions. We found that the effects of abiotic factors on the spatial turnover of ecosystem functions were largely indirect through their influences on above‐ and belowground β‐diversity, and that spatial turnover of ecosystem function was more strongly associated with plant β‐diversity than with soil microbial β‐diversity. These results indicate that changes in above‐ and belowground species composition are one mechanism that interacts with environmental change to determine variability in multiple ecosystem functions across spatial scales. As grasslands face global threats from shrub encroachment, conversion to agriculture, or are lost to development, the functions and services they provide will more strongly converge with increased aboveground community homogenization than with soil microbial community homogenization.

Soil nitrogen (N) availability is critical for grassland functioning. However, human activities have increased the supply of biologically limiting nutrients, and changed the density and identity of mammalian herbivores. These anthropogenic changes may alter net soil N mineralization (soil net Nmin), that is, the net balance between N mineralization and immobilization, which could severely impact grassland structure and functioning. Yet, to date, little is known about how fertilization and herbivore removal individually, or jointly, affect soil net Nmin across a wide range of grasslands that vary in soil and climatic properties. Here we collected data from 22 grasslands on five continents, all part of a globally replicated experiment, to assess how fertilization and herbivore removal affected potential (laboratory‐based) and realized (field‐based) soil net Nmin. Herbivore removal in the absence of fertilization did not alter potential and realized soil net Nmin. However, fertilization alone and in combination with herbivore removal consistently increased potential soil net Nmin. Realized soil net Nmin, in contrast, significantly decreased in fertilized plots where herbivores were removed. Treatment effects on potential and realized soil net Nmin were contingent on site‐specific soil and climatic properties. Fertilization effects on potential soil net Nmin were larger at sites with higher mean annual precipitation (MAP) and temperature of the wettest quarter (T.q.wet). Reciprocally, realized soil net Nmin declined most strongly with fertilization and herbivore removal at sites with lower MAP and higher T.q.wet. In summary, our findings show that anthropogenic nutrient enrichment, herbivore exclusion and alterations in future climatic conditions can negatively impact soil net Nmin across global grasslands under realistic field conditions. This is an important context‐dependent knowledge for grassland management worldwide.

AimNutrient enrichment is associated with plant invasions and biodiversity loss. Functional trait advantages may predict the ascendancy of invasive plants following nutrient enrichment but this is rarely tested. Here, we investigate (a) whether dominant native and non‐native plants differ in important morphological and physiological leaf traits, (b) how their traits respond to nutrient addition, and (c) whether responses are consistent across functional groups.LocationAustralia, Europe, North America and South Africa.Time period2007–2014.Major taxa studiedGraminoids and forbs.MethodsWe focused on two types of leaf traits connected to resource acquisition: morphological features relating to light‐foraging surfaces and investment in tissue (specific leaf area, SLA) and physiological features relating to internal leaf chemistry as the basis for producing and utilizing photosynthate. We measured these traits on 503 leaves from 151 dominant species across 27 grasslands on four continents. We used an identical nutrient addition treatment of nitrogen (N), phosphorus (P) and potassium (K) at all sites. Sites represented a broad range of grasslands that varied widely in climatic and edaphic conditions.ResultsWe found evidence that non‐native graminoids invest in leaves with higher nutrient concentrations than native graminoids, particularly at sites where native and non‐native species both dominate. We found little evidence that native and non‐native forbs differed in the measured leaf traits. These results were consistent in natural soil fertility levels and nutrient‐enriched conditions, with dominant species responding similarly to nutrient addition regardless of whether they were native or non‐native.Main conclusionsOur work identifies the inherent physiological trait advantages that can be used to predict non‐native graminoid establishment, potentially because of higher efficiency at taking up crucial nutrients into their leaves. Most importantly, these inherent advantages are already present at natural soil fertility levels and are maintained following nutrient enrichment.

Eutrophication is a widespread environmental change that usually reduces the stabilizing effect of plant diversity on productivity in local communities. Whether this effect is scale dependent remains to be elucidated. Here, we determine the relationship between plant diversity and temporal stability of productivity for 243 plant communities from 42 grasslands across the globe and quantify the effect of chronic fertilization on these relationships. Unfertilized local communities with more plant species exhibit greater asynchronous dynamics among species in response to natural environmental fluctuations, resulting in greater local stability (alpha stability). Moreover, neighborhood communities that have greater spatial variation in plant species composition within sites (higher beta diversity) have greater spatial asynchrony of productivity among communities, resulting in greater stability at the larger scale (gamma stability). Importantly, fertilization consistently weakens the contribution of plant diversity to both of these stabilizing mechanisms, thus diminishing the positive effect of biodiversity on stability at differing spatial scales. Our findings suggest that preserving grassland functional stability requires conservation of plant diversity within and among ecological communities., Eutrophication has been shown to weaken diversity-stability relationships in grasslands, but it is unclear whether the effect depends on scale. Analysing a globally distributed network of grassland sites, the authors show a positive role of beta diversity and spatial asynchrony as drivers of stability but find that nitrogen enrichment weakens the diversity-stability relationships at different spatial scales.

Currently, either highly multiplexed genetic manipulations can be delivered to mammalian cells all at once, or extensive engineering of gene regulatory sequences can be used to conditionally activate a few manipulations. Here, we provide proof-of-principle for a new system enabling multiple genetic manipulations to be executed as a preprogrammed cascade of events. The system leverages the programmability of the S. pyogenes Cas9 and is based on flexible arrangements of individual modules of activity. The basic module consists of an inactive single guide RNA (sgRNA) -like component that is converted to an active state through the effects of another sgRNA. Modules can be arranged to bring about an algorithmic program of sequential genetic manipulations without the need for engineering cell type specific promoters or gene regulatory sequences. With the expanding diversity of available tools that utilize spCas9, this sgRNA-based system provides multiple levels of interfacing with mammalian cell biology.

SUMMARYGenetic manipulation of mammalian cells is instrumental to modern biomedical research but is currently limited by poor capabilities of sequentially controlling multiple manipulations in cells. Currently, either highly multiplexed manipulations can be delivered to populations of cells all at one time, or gene regulatory sequences can be engineered to conditionally activate a few manipulations within individual cells. Here, we provide proof-of-principle for a new system enabling multiple genetic manipulations to be executed as a preprogrammed cascade of events. The system leverages the programmability of the S. pyogenes Cas9 RNA-guided nuclease and is based on flexible arrangements of individual modules of activity. The basic module consists of an inactive single guide RNA (sgRNA) - like component that is converted to an active state through the effects of another sgRNA. Modules can be arranged to bring about an algorithmic program of genetic manipulations without the need for engineering cell type specific promoters or gene regulatory sequences. With the expanding diversity of available tools that utilize spCas9 to edit, repress or activate genes, this sgRNA-based system provides multiple levels for interfacing with host cell biology. In addition, ability of the system to progress through multiple modules from episomal plasmid DNA makes it suitable for applications sensitive to the presence of heterologous genomic DNA sequences and broadly applicable to biomedical research and mammalian cell engineering.

Soil stores approximately twice as much carbon as the atmosphere and fluctuations in the size of the soil carbon pool directly influence climate conditions. We used the Nutrient Network global change experiment to examine how anthropogenic nutrient enrichment might influence grassland soil carbon storage at a global scale. In isolation, enrichment of nitrogen and phosphorous had minimal impacts on soil carbon storage. However, when these nutrients were added in combination with potassium and micronutrients, soil carbon stocks changed considerably, with an average increase of 0.04 KgCm 2 year 1 (standard deviation 0.18 KgCm 2 year 1 ). These effects did not correlate with changes in primary productivity, suggesting that soil carbon decomposition may have been restricted. Although nutrient enrichment caused soil carbon gains most dry, sandy regions, considerable absolute losses of soil carbon may occur in high-latitude regions that store the majority of the world’s soil carbon. These mechanistic insights into the sensitivity of grassland carbon stocks to nutrient enrichment can facilitate biochemical modelling efforts to project carbon cycling under future climate scenarios. Fil: Crowther, Thomas W.. Univerität strasse; Suiza Fil: Riggs, C.. University of Minnesota; Estados Unidos Fil: Lind, E. M.. University of Minnesota; Estados Unidos Fil: Borer, Elizabeth. University of Minnesota; Estados Unidos Fil: Seabloom, Eric. University of Minnesota; Estados Unidos Fil: Hobbie, S. E.. University of Minnesota; Estados Unidos Fil: Wubs, E. R. Jasper. Netherlands Institute of Ecology; Países Bajos Fil: Adler, Peter. University of Utah; Estados Unidos Fil: Firn, Jennifer. Queensland University of Technology; Australia Fil: Gherardi, Laureano. Arizona State University; Estados Unidos Fil: Hagenah, N.. University of Pretoria; Sudáfrica Fil: Hofmockel, K. S.. University of Iowa; Estados Unidos Fil: Knops, Johannes M. H.. Earth and Biological Sciences Directorate. Pacific Northwest National Laboratory; Estados Unidos Fil: McCulley, R. L.. University of Kentucky; Estados Unidos Fil: MacDougall, A. S.. University of Guelph; Canadá Fil: Peri, Pablo Luis. Instituto Nacional de Tecnología Agropecuaria; Argentina. Universidad Tecnológica Nacional; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Prober, S.M.. CSIRO Land and Water; Australia Fil: Stevens, C. J.. Lancaster University; Reino Unido Fil: Routh, D.. Univerität strasse; Suiza

Almost 50 years ago, Michael Rosenzweig pointed out that nutrient addition can destabilize food webs, leading to loss of species and reduced ecosystem function through the paradox of enrichment. Around the same time, David Tilman demonstrated that increased nutrient loading would also be expected to cause competitive exclusion leading to deleterious changes in food web diversity. While both concepts have greatly illuminated general diversity-stability theory, we currently lack a coherent framework to predict how nutrients influence food web stability across a landscape. This is a vitally important gap in our understanding, given mounting evidence of serious ecological disruption arising from anthropogenic displacement of resources and organisms. Here, we combine contemporary theory on food webs and meta-ecosystems to show that nutrient additions are indeed expected to drive loss in stability and function in human-impacted regions. However, this loss in stability occurs not just from wild oscillations in population abundance, but more frequently from the complete loss of an equilibrium due to edible plant species being competitively excluded. In highly modified landscapes, spatial nutrient transport theory suggests that such instabilities can be amplified over vast distances from the sites of nutrient addition. Consistent with this theoretical synthesis, the empirical frequency of these distant propagating ecosystem imbalances appears to be growing. This synthesis of theory and empirical data suggests that human modification of the Earth’s ecological connectivity is “entangling” once distantly separated ecosystems, causing rapid, expansive, and costly nutrient-driven instabilities over vast areas of the planet. The corollary to this spatial nutrient theory, though -- akin to weak interaction theory from food web networks -- is that slow spatial nutrient pathways can be potent stabilizers by moderating flows across a landscape.

The paradigmatic hypothesis for the effect of fertilisation on plant diversity represents a one-dimensional trade-off for plants competing for below-ground nutrients (generically) and above-ground light: fertilisation reduces competition for nutrients while increasing biomass and thereby shifts competition for depleted available light. The essential problem of this simple paradigm is that it misses both the multivariate and mechanistic nature of the factors that determine biodiversity as well as their causal relationships. We agree that light limitation, as DeMalach and Kadmon argue, can indeed be an important factor associated with diversity loss, and we presented it as an integral part of our tests of the niche dimension hypothesis. We disagree with DeMalach and Kadmon that light is the ‘main’ factor explaining diversity, because this misrepresents the causal structure represented in the design of our experiment in which multiple nutrient addition was the ultimate causal driver of a suite of correlated responses that included diversity and light, and especially live and dead biomass, which are the factors that control light depletion. Our findings highlight that multiple nutrient limitations can structure plant diversity and composition independently of changes in light and biomass. For example, approximately one-third of our sites showed no significant increase in biomass with greater number of added nutrients yet still lost diversity when nutrients were added. The important message is that while light limitation can be an important contributor to diversity loss, it is not a necessary mechanism. Fil: Harpole, W. Stanley. Helmholtz Center for Environmental Research; Alemania Fil: Sullivan, Lauren L.. University of Minnesota; Estados Unidos Fil: Lind, Eric M.. University of Minnesota; Estados Unidos Fil: Firn, Jennifer. Queensland University of Technology; Australia Fil: Adler, Peter B.. State University of Utah; Estados Unidos Fil: Borer, Elizabeth T.. University of Minnesota; Estados Unidos Fil: Chase, Jonathan. Martin Luther University Halle Wittenberg; Alemania Fil: Fay, Philip A.. USDA-ARS Grassland Soil and Water Research Lab; Estados Unidos Fil: Hautier, Yann. Utrecht University; Países Bajos Fil: Hillebrand, Helmut. University of Oldenburg; Alemania Fil: MacDougall, Andrew S.. University of Guelph; Canadá Fil: Seabloom, Eric W.. University of Minnesota; Estados Unidos Fil: Bakker, Jonathan D.. University of Washington; Estados Unidos Fil: Cadotte, Marc W.. University of Toronto; Canadá Fil: Chaneton, Enrique Jose. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura. Universidad de Buenos Aires. Facultad de Agronomía; Argentina Fil: Chu, Chengjin. Sun Yat-sen University; China Fil: Hagenah, Nicole. University of KwaZulu-Natal; Sudáfrica Fil: Kirkman, Kevin. University of KwaZulu-Natal; Sudáfrica Fil: La Pierre, Kimberly J.. Smithsonian Environmental Research Center; Estados Unidos Fil: Moore, Joslin L.. Monash University; Australia Fil: Morgan, John W.. La Trobe University; Australia Fil: Prober, Suzanne M.. CSIRO Land and Water; Australia Fil: Risch, Anita C.. Swiss Federal Institute for Forest, Snow and Landscape Research; Suiza Fil: Schuetz, Martin. Swiss Federal Institute for Forest, Snow and Landscape Research; Suiza Fil: Stevens, Carly J.. Lancaster University; Reino Unido

Summary Clarifying what species are being consumed at what times can improve our understanding of how anthropogenic change affects food web dynamics, with implications for community assembly including restoration. This includes human-based changes to plant communities via species introductions, which can interact with consumer feeding preferences to indirectly alter assembly outcomes including reduced restoration success if planted species are preferentially targeted. We used DNA barcoding of plant material in rodent scat, combined with field-based feeding trials, rodent trapping, and rodent exclosures to test for dietary preferences and assembly impacts of native rodents on a restored and regionally rare tallgrass prairie of central North America. We examined whether native rodents preferred non-targeted and mostly non-native oldfield plants that are more locally abundant, thus protecting the rarer native planted species from consumption, or if rodents preferred native plants regardless of abundance. Our results supported the latter outcome. Barcoding revealed that native rodents consumed mostly non-planted oldfield species for ten months of the year (92% of the diet). The exceptions were August–September, when planted prairie species accounted for 87% of plants consumed and coincided with their peak seed production. Cafeteria trials suggested diet seasonality to be explained by food limitation – native prairie seeds were consumed year-round when made available. Non-targeted oldfield species thus appeared to be of sufficient quality to support rodent populations (peaking at 23 individuals ha−1 in summer), but without rescuing rarer prairie species from targeted granivory and resulting in reduced prairie diversity outside of experimental exclosures. Synthesis and applications. Our work illustrates how anthropogenic-based changes to producer communities may affect feeding pathways in grassland food webs, potentially facilitating establishment by non-target species via indirect consumer effects that can be difficult to detect. These findings suggest that rodents may reduce the restoration success of tallgrass prairie in our region, with some planted species likely to suffer future recruitment difficulties due to granivory. Managers may need to consider multiple trophic levels with restoration, not just the commonly planted producer communities but also the consumers and predators associated with them. This article is protected by copyright. All rights reserved.

Protected areas are a cornerstone of the global strategy for conserving biodiversity, and yet their efficacy in comparison to unprotected areas is rarely tested. In the highly fragmented forests of temperate regions, landscape context and forest history may be more important than protection status for plant species diversity. To determine whether there are differences in plant diversity between protected areas and private lands while controlling for landscape context, forest age, and other important factors. We used a database of 156 one-hectare forest plots distributed over 120,000 km2 in the fragmented forests of southern Ontario to test whether protected areas and private forests differed in native species richness, relative abundance of exotic species, and the probability of finding species of conservation concern. Plots with more forest on the surrounding landscape had higher native species richness, lower abundance of exotic species, and greater probability of supporting at least one species of conservation concern. Young forests tended to have higher abundance of exotics, and were less likely to support species of conservation concern. Surprisingly, privately owned forests had greater native species richness and were more likely to support species of conservation concern once these other factors were accounted for. In addition, there were significant interactions between ownership type, forest history, and landscape context. Our results highlight the importance of privately owned forests in this region, and the need to consider forest history and landscape context when comparing the efficacy of protected areas versus private land for sustaining biodiversity.