Dr. Martin Schütz

Aims: We introduce ReSurveyEurope — a new data source of resurveyed vegetation plots in Europe, compiled by a collaborative network of vegetation scientists. We describe the scope of this initiative, provide an overview of currently available data, governance, data contribution rules, and accessibility. In addition, we outline further steps, including potential research questions.
Results: ReSurveyEurope includes resurveyed vegetation plots from all habitats. Version 1.0 of ReSurveyEurope contains 283,135 observations (i.e., individual surveys of each plot) from 79,190 plots sampled in 449 independent resurvey projects. Of these, 62,139 (78%) are permanent plots, that is, marked in situ, or located with GPS, which allow for high spatial accuracy in resurvey. The remaining 17,051 (22%) plots are from studies in which plots from the initial survey could not be exactly relocated. Four data sets, which together account for 28,470 (36%) plots, provide only presence/absence information on plant species, while the remaining 50,720 (64%) plots contain abundance information (e.g., percentage cover or cover–abundance classes such as variants of the Braun-Blanquet scale). The oldest plots were sampled in 1911 in the Swiss Alps, while most plots were sampled between 1950 and 2020.
Conclusions: ReSurveyEurope is a new resource to address a wide range of research questions on fine-scale changes in European vegetation. The initiative is devoted to an inclusive and transparent governance and data usage approach, based on slightly adapted rules of the well-established European Vegetation Archive (EVA). ReSurveyEurope data are ready for use, and proposals for analyses of the data set can be submitted at any time to the coordinators. Still, further data contributions are highly welcome.

Topsoil removal, among other restoration measures, has been recognized as one of the most successful methods to restore biodiversity and ecosystem functioning in European grasslands. However, knowledge about how removal as well as other restoration methods influence interactions between plant and microbial communities is very limited. The aims of the current study were to understand the impact of topsoil removal on plant-microorganism interactions and on soil nitrogen (N) mineralization, as one example of ecosystem functioning. We examined how three different grassland restoration methods, namely ‘Harvest only’, ‘Topsoil removal’ and ‘Topsoil removal + Propagules (plant seed addition)’, affected i) the interactions between plants and soil microorganisms, ii) soil microbial community assembly processes, and iii) soil N mineralization. We compared the outcome of these three restoration methods to initial degraded and target semi-natural grasslands in the Canton of Zurich, Switzerland. We were able to show that ‘Topsoil removal’ and ‘Topsoil removal + Propagules’, but not ‘Harvest only’, reduced the soil total N pool and available N concentration, but increased soil N mineralization and strengthened the plant-microorganism interactions. Microbial community assembly processes shifted towards more deterministic after both topsoil removal treatments. These shifts could be attributed to an increase in dispersal limitation and selection due to stronger interactions between plants and soil microorganisms. The negative relationship between soil N mineralization and microbial community stochasticity indicated that microbial assembly processes, to some extent, can be incorporated into model predictions of soil functions. Overall, the results suggest that topsoil removal may change the microbial assembly processes and thus the functioning of grassland ecosystems by enhancing the interaction between plants and soil microorganisms.

Ecological theory posits that temporal stability patterns in plant populations are associated with differences in species' ecological strategies. However, empirical evidence is lacking about which traits, or trade-offs, underlie species stability, especially across different biomes. We compiled a worldwide collection of long-term permanent vegetation records (greater than 7000 plots from 78 datasets) from a large range of habitats which we combined with existing trait databases. We tested whether the observed inter-annual variability in species abundance (coefficient of variation) was related to multiple individual traits. We found that populations with greater leaf dry matter content and seed mass were more stable over time. Despite the variability explained by these traits being low, their effect was consistent across different datasets. Other traits played a significant, albeit weaker, role in species stability, and the inclusion of multi-variate axes or phylogeny did not substantially modify nor improve predictions. These results provide empirical evidence and highlight the relevance of specific ecological trade-offs, i.e. in different resource-use and dispersal strategies, for plant populations stability across multiple biomes. Further research is, however, necessary to integrate and evaluate the role of other specific traits, often not available in databases, and intraspecific trait variability in modulating species stability.

Despite plants realistically being affected by vertebrate and invertebrate herbivores simultaneously, fundamental differences in the ecology and evolution of these two herbivore guilds often means their impacts on plants are studied separately. A synthesis of the literature is needed to understand the types of plant traits examined and their response to, and effect on (in terms of forage selection) vertebrate and invertebrate herbivory, and to identify associated knowledge gaps. Focusing on grassland systems and species, we found 138 articles that met our criteria: 39 invertebrate, 97 vertebrate and 2 focussed on both vertebrate and invertebrate herbivores. Our study identified invertebrate focussed research, research conducted in the Southern Hemisphere and research on nondomesticated herbivores was significantly underrepresented based on our search and should be a focus of future research. Differences in study focus (trait response or trait effect), along with differences in the types of traits examined, led to limited opportunity for comparison between the two herbivore guilds. This review therefore predominantly discusses the response and effect of plant traits to each herbivore guild separately. In future studies, we suggest this review be used as a guide for trait selection, to improve comparability and the broader significance of results.

Aim: The microbial metabolic quotient (MMQ; mg CO₂-C/mg MBC/h), defined as the amount of microbial CO₂ respired (MR; mg CO₂-C/kg soil/h) per unit of microbial biomass C (MBC; mg C/kg soil), is a key parameter for understanding the microbial regulation of the carbon (C) cycle, including soil C sequestration. Here, we experimentally tested hypotheses about the individual and interactive effects of multiple nutrient addition (nitrogen + phosphorus + potassium + micronutrients) and herbivore exclusion on MR, MBC and MMQ across 23 sites (five continents). Our sites encompassed a wide range of edaphoclimatic conditions; thus, we assessed which edaphoclimatic variables affected MMQ the most and how they interacted with our treatments.
Location: Australia, Asia, Europe, North/South America.
Time period: 2015–2016.
Major taxa: Soil microbes.
Methods: Soils were collected from plots with established experimental treatments. MR was assessed in a 5-week laboratory incubation without glucose addition, MBC via substrate-induced respiration. MMQ was calculated as MR/MBC and corrected for soil temperatures (MMQsoil). Using linear mixed effects models (LMMs) and structural equation models (SEMs), we analysed how edaphoclimatic characteristics and treatments interactively affected MMQsoil.
Results: MMQsoil was higher in locations with higher mean annual temperature, lower water holding capacity and lower soil organic C concentration, but did not respond to our treatments across sites as neither MR nor MBC changed. We attributed this relative homeostasis to our treatments to the modulating influence of edaphoclimatic variables. For example, herbivore exclusion, regardless of fertilization, led to greater MMQsoil only at sites with lower soil organic C (< 1.7%).
Main conclusions: Our results pinpoint the main variables related to MMQsoil across grasslands and emphasize the importance of the local edaphoclimatic conditions in controlling the response of the C cycle to anthropogenic stressors. By testing hypotheses about MMQsoil across global edaphoclimatic gradients, this work also helps to align the conflicting results of prior studies.

Nutrients and herbivores are well-known drivers of grassland diversity and stability in local communities. However, whether they interact to impact the stability of aboveground biomass and whether these effects depend on spatial scales remain unknown. It is also unclear whether nutrients and herbivores impact stability via different facets of plant diversity including species richness, evenness, and changes in community composition through time and space. We used a replicated experiment adding nutrients and excluding herbivores for 5 years in 34 global grasslands to explore these questions. We found that both nutrient addition and herbivore exclusion alone reduced stability at the larger spatial scale (aggregated local communities; gamma stability), but through different pathways. Nutrient addition reduced gamma stability primarily by increasing changes in local community composition over time, which was mainly driven by species replacement. Herbivore exclusion reduced gamma stability primarily by decreasing asynchronous dynamics among local communities (spatial asynchrony). Their interaction weakly increased gamma stability by increasing spatial asynchrony. Our findings indicate that disentangling the processes operating at different spatial scales may improve conservation and management aiming at maintaining the ability of ecosystems to reliably provide functions and services for humanity.

1. Plant damage by invertebrate herbivores and pathogens influences the dynamics of grassland ecosystems, but anthropogenic changes in nitrogen and phosphorus availability can modify these relationships.
2. Using a globally distributed experiment, we describe leaf damage on 153 plant taxa from 27 grasslands worldwide, under ambient conditions and with experimentally elevated nitrogen and phosphorus.
3. Invertebrate damage significantly increased with nitrogen addition, especially in grasses and non-leguminous forbs. Pathogen damage increased with nitrogen in grasses and legumes but not forbs. Effects of phosphorus were generally weaker. Damage was higher in grasslands with more precipitation, but climatic conditions did not change effects of nutrients on leaf damage. On average, invertebrate damage was relatively higher on legumes and pathogen damage was relatively higher on grasses. Community-weighted mean damage reflected these functional group patterns, with no effects of N on community-weighted pathogen damage (due to opposing responses of grasses and forbs) but stronger effects of N on community-weighted invertebrate damage (due to consistent responses of grasses and forbs).
4. Synthesis. As human-induced inputs of nitrogen and phosphorus continue to increase, understanding their impacts on invertebrate and pathogen damage becomes increasingly important. Our results demonstrate that eutrophication frequently increases plant damage and that damage increases with precipitation across a wide array of grasslands. Invertebrate and pathogen damage in grasslands is likely to increase in the future, with potential consequences for plant, invertebrate and pathogen communities, as well as the transfer of energy and nutrients across trophic levels.

1. 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
2. We determined the effects of addition of experimental nitrogen (N), phosphorus (P) and potassium plus micronutrient (K_μ) 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.
3. 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%.
4. 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.

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.

Ecological models predict that the effects of mammalian herbivore exclusion on plant diversity depend on resource availability and plant exposure to ungulate grazing over evolutionary time. Using an experiment replicated in 57 grasslands on six continents, with contrasting evolutionary history of grazing, we tested how resources (mean annual precipitation and soil nutrients) determine herbivore exclusion effects on plant diversity, richness and evenness. Here we show that at sites with a long history of ungulate grazing, herbivore exclusion reduced plant diversity by reducing both richness and evenness and the responses of richness and diversity to herbivore exclusion decreased with mean annual precipitation. At sites with a short history of grazing, the effects of herbivore exclusion were not related to precipitation but differed for native and exotic plant richness. Thus, plant species' evolutionary history of grazing continues to shape the response of the world’s grasslands to changing mammalian herbivory.

1. Evaluation of restoration activities is indispensable to assess the extent to which targets have been reached. Usually, the main goal of ecological restoration is to restore biodiversity and ecosystem functioning, but validation is often based on a single indicator, which may or may not cope with whole-ecosystem dynamics. Network analyses are, however, powerful tools, allowing to examine both the recovery of various biotic and abiotic properties and the integrated response at community and ecosystem level.
2. We used restoration sites where topsoil was removed from former intensively managed grassland and seeds were added. These sites were between 3 and 32 years old. We assessed how plants, soil biota, soil properties and correlation-based interactions between biotic communities and their abiotic environment developed over time and compared the results with (i) intensively managed (not restored), and (ii) well-preserved targeted semi-natural grasslands.
3. Plant, nematode, fungal and prokaryotic diversity and community structures of the restored grasslands revealed clear successional patterns and followed similar trajectories towards targeted semi-natural grasslands. All biotic communities reached targeted diversity levels no later than 18 years post-restoration.
4. Ecological networks of intensively managed and short-term (~4 years) restored grasslands were less tightly connected compared to those found in mid- and long-term (~18–30 years) restored and target grasslands. Restoration specifically enhanced interactions among biotic communities, but reduced interactions between biotic communities and their abiotic environment as well as interactions among abiotic properties in the short- and mid-term.
5. Synthesis and applications: Overall, our study demonstrated that topsoil removal and seed addition were successful in restoring diverse, tightly coupled and well-connected biotic communities above- and below-ground similar to those found in the semi-natural grasslands that were restoration targets. Network analyses proved to be powerful in examining the long-term re-establishment of functionally connected biotic communities in restored ecosystems. Thus, we provide an approach to holistically assess restoration activities by notably considering the complexity of ecosystems, much in contrast to most traditional approaches.

Forest ecosystems sequester approximately half of the world's organic carbon (C), most of it in the soil. The amount of soil C stored depends on the input and decomposition rate of soil organic matter (OM), which is controlled by the abundance and composition of the microbial and invertebrate communities, soil physico-chemical properties, and (micro)-climatic conditions. Although many studies have assessed how these site-specific climatic and soil properties affect the decomposition of fresh OM, differences in the type and quality of the OM substrate used, make it difficult to compare and extrapolate results across larger scales. Here, we used standard wood stakes made from aspen (Populus tremuloides Michx.) and loblolly pine (Pinus taeda L.) to explore how climate and abiotic soil properties affect wood decomposition across 44 unharvested forest stands located across the northern hemisphere. Stakes were placed in three locations: (i) on top of the surface organic horizons (surface), (ii) at the interface between the surface organic horizons and mineral soil (interface), and (iii) into the mineral soil (mineral). Decomposition rates of both wood species was greatest for mineral stakes and lowest for stakes placed on the surface organic horizons, but aspen stakes decomposed faster than pine stakes. Our models explained 44 and 36% of the total variation in decomposition for aspen surface and interface stakes, but only 0.1% (surface), 12% (interface), 7% (mineral) for pine, and 7% for mineral aspen stakes. Generally, air temperature was positively, precipitation negatively related to wood stake decomposition. Climatic variables were stronger predictors of decomposition than soil properties (surface C:nitrogen ratio, mineral C concentration, and pH), regardless of stake location or wood species. However, climate-only models failed in explaining wood decomposition, pointing toward the importance of including local-site properties when predicting wood decomposition. The difficulties we had in explaining the variability in wood decomposition, especially for pine and mineral soil stakes, highlight the need to continue assessing drivers of decomposition across large global scales to better understand and estimate surface and belowground C cycling, and understand the drivers and mechanisms that affect C pools, CO₂ emissions, and nutrient cycles.

Increasing evidence suggests that remotely sensed spectral diversity is linked to plant species richness. However, a conflicting spectral diversity–biodiversity relationship in grasslands has been found in previous studies. In particular, it remains unclear how well the spectral diversity–biodiversity relationship holds in naturally assembled species-rich grasslands. To address the linkage between spectral diversity and plant species richness in a species-rich alpine grassland ecosystem, we investigated (i) the trade-off between spectral and spatial resolution in remote sensing data; (ii) the suitability of three different spectral metrics to describe spectral diversity (coefficient of variation, convex hull volume and spectral species richness) and (iii) the importance of confounding effects of live plant biomass, dead plant biomass and plant life forms on the spectral diversity–biodiversity relationship. We addressed these questions using remote sensing data collected with consumer-grade cameras with four spectral bands and 10 cm spatial resolution on an unmanned aerial vehicle (UAV), airborne imaging spectrometer data (AVIRIS-NG) with 372 bands and 2.5 m spatial resolution, and a fused data product of both datasets. Our findings suggest that a fused dataset can cope with the requirement of both high spatial- and spectral resolution to remotely measure biodiversity. However, in contrast to several previous studies, we found a negative correlation between plant species richness and spectral metrics based on the spectral information content (i.e. spectral complexity). The spectral diversity calculated based on the spectral complexity was sensitive to live and dead plant biomass. Overall, our results suggest that remote sensing of plant species diversity requires a high spatial resolution, the use of classification-based spectral metrics, such as spectral species richness, and awareness of confounding factors (e.g. plant biomass), which may be ecosystem specific.

Analysing temporal patterns in plant communities is extremely important to quantify the extent and the consequences of ecological changes, especially considering the current biodiversity crisis. Long-term data collected through the regular sampling of permanent plots represent the most accurate resource to study ecological succession, analyse the stability of a community over time and understand the mechanisms driving vegetation change. We hereby present the LOng-Term Vegetation Sampling (LOTVS) initiative, a global collection of vegetation time-series derived from the regular monitoring of plant species in permanent plots. With 79 data sets from five continents and 7,789 vegetation time-series monitored for at least 6 years and mostly on an annual basis, LOTVS possibly represents the largest collection of temporally fine-grained vegetation time-series derived from permanent plots and made accessible to the research community. As such, it has an outstanding potential to support innovative research in the fields of vegetation science, plant ecology and temporal ecology.

Background and aims The amount of nitrogen (N) derived from symbiotic N₂ fixation by legumes in grasslands might be affected by anthropogenic N and phosphorus (P) inputs, but the underlying mechanisms are not known.
Methods We evaluated symbiotic N₂ fixation in 17 natural and semi-natural grasslands on four continents that are subjected to the same full-factorial N and P addition experiment, using the ¹⁵N natural abundance method.
Results N as well as combined N and P (NP) addition reduced aboveground legume biomass by 65% and 45%, respectively, compared to the control, whereas P addition had no significant impact. Addition of N and/or P had no significant effect on the symbiotic N₂ fixation per unit legume biomass. In consequence, the amount of N fixed annually per grassland area was less than half in the N addition treatments compared to control and P addition, irrespective of whether the dominant legumes were annuals or perennials.
Conclusion Our results reveal that N addition mainly impacts symbiotic N₂ fixation via reduced biomass of legumes rather than changes in N₂ fixation per unit legume biomass. The results show that soil N enrichment by anthropogenic activities significantly reduces N₂ fixation in grasslands, and these effects cannot be reversed by additional P amendment.

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.

It is generally assumed that restoring biodiversity will enhance diversity and ecosystem functioning. However, to date, it has rarely been evaluated whether and how restoration efforts manage to rebuild biodiversity and multiple ecosystem functions (ecosystem multifunctionality) simultaneously. Here, we quantified how three restoration methods of increasing intervention intensity (harvest only < topsoil removal < topsoil removal + propagule addition) affected grassland ecosystem multifunctionality 22 yr after the restoration event. We compared restored with intensively managed and targeted seminatural grasslands based on 13 biotic and abiotic, above- and belowground properties. We found that all three restoration methods improved ecosystem multifunctionality compared to intensively managed grasslands and developed toward the targeted seminatural grasslands. However, whereas higher levels of intervention intensity reached ecosystem multifunctionality of targeted seminatural grasslands after 22 yr, lower intervention missed this target. Moreover, we found that topsoil removal with and without seed addition accelerated the recovery of biotic and aboveground properties, and we found no negative long-term effects on abiotic or belowground properties despite removing the top layer of the soil. We also evaluated which ecosystem properties were the best indicators for restoration success in terms of accuracy and cost efficiency. Overall, we demonstrated that low-cost measures explained relatively more variation of ecosystem multifunctionality compared to high-cost measures. Plant species richness was the most accurate individual property in describing ecosystem multifunctionality, as it accounted for 54% of ecosystem multifunctionality at only 4% of the costs of our comprehensive multifunctionality approach. Plant species richness is the property that typically is used in restoration monitoring by conservation agencies. Vegetation structure, soil carbon storage and water-holding capacity together explained 70% of ecosystem multifunctionality at only twice the costs (8%) of plant species richness, which is, in our opinion, worth considering in future restoration monitoring projects. Hence, our findings provide a guideline for land managers how they could obtain an accurate estimate of aboveground-belowground ecosystem multifunctionality and restoration success in a highly cost-efficient way.

The increasing availability of remote sensing data allows the quantification of biodiversity in space and time. In particular, spectral diversity, defined as the variability of electromagnetic radiation reflected from plants, can be assessed with remote sensing. Plant traits vary diurnally and seasonally due to plant phenology and land management. This results in strong temporal variation of spectral diversity, which cannot be accurately represented by remotely sensed data collected at a single point in time. However, knowledge of how datasets sampled at multiple points in time should best be used to quantify spectral diversity is scarce. To address this issue, we first introduced a new approach using spatio-temporal spectral diversity based on the dissimilarity measure Rao's quadratic entropy index (RaoQ). Thereby, we demonstrated how RaoQ can be used to partition the total spectral diversity of a region (γSD) into additive alpha (αSD, within communities) and spatio-temporal beta (βSD; between communities) components, allowing the calculation of βSD from community mean spectral features, independent from αSD. Second, we illustrated our methodological approach with a case study in which βSD is calculated from Sentinel-2 satellite data at high temporal resolution for managed grasslands which differ across a large gradient of environmental properties. We were able to show differences in βSD and separate its components into phenological and management effects. Furthermore, the contribution of different plant communities to βSD was assessed, and the results were validated against a dataset of in-situ measured β diversity from plant surveys. Compared to spatial dissimilarities from distinct stages of the growing season, using spatio-temporal dissimilarities between communities produced a more accurate estimation of the uniqueness of a community. This study shows how to account for temporal variations in the spectral diversity of plant communities and demonstrates that this improves the estimation of plant biodiversity through remote sensing. Spectral diversity in space and time makes it possible to assess mechanisms that drive biodiversity and identify plant communities relevant for conservation purposes.

Lowland grassy woodlands in Australia’s south-east face reductions in native plant diversity because of invasion by non-native plants. We compared the relative abundance and diversity of plant species among sites dominated by the native Kangaroo grass (KG) Themeda triandra with sites co-dominated by the non-native African lovegrass (ALG) Eragrostis curvula and KG. We found significant differences in plant species composition depending on the dominant species. Furthermore, our results revealed differences in several diversity parameters such as a lower species richness and forb diversity on sites co-dominated by ALG and KG. This was the case despite the functional similarity of both ALG and KG—both C₄ perennial tussock grasses of a similar height. Therefore, our results highlight the critical function of the native KG in maintaining and enhancing the target plant species composition and diversity within these grassy woodlands. Herbivore grazing potentially impacts on the abundance of the dominant grass and forb species in various ways, but its impact likely differs depending on their evolutionary origin. Therefore, disentangling the role of individual herbivore groups (native-, non-native mammals, and invertebrates) on the plant community composition of the lowland grassy woodlands is essential to find appropriate grazing regimes for ALG management in these ecosystems.

Human activities are enriching many of Earth's ecosystems with biologically limiting mineral nutrients such as nitrogen (N) and phosphorus (P). In grasslands, this enrichment generally reduces plant diversity and increases productivity. The widely demonstrated positive effect of diversity on productivity suggests a potential negative feedback, whereby nutrient-induced declines in diversity reduce the initial gains in productivity arising from nutrient enrichment. In addition, plant productivity and diversity can be inhibited by accumulations of dead biomass, which may be altered by nutrient enrichment. Over longer time frames, nutrient addition may increase soil fertility by increasing soil organic matter and nutrient pools. We examined the effects of 5–11 yr of nutrient addition at 47 grasslands in 12 countries. Nutrient enrichment increased aboveground live biomass and reduced plant diversity at nearly all sites, and these effects became stronger over time. We did not find evidence that nutrient-induced losses of diversity reduced the positive effects of nutrients on biomass; however, nutrient effects on live biomass increased more slowly at sites where litter was also increasing, regardless of plant diversity. This work suggests that short-term experiments may underestimate the long-term nutrient enrichment effects on global grassland ecosystems.