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

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.

Covering approximately 40% of land surfaces, grasslands provide critical ecosystem services that rely on soil organisms. However, the global determinants of soil biodiversity and functioning remain underexplored. In this study, we investigate the drivers of soil microbial and detritivore activity in grasslands across a wide range of climatic conditions on five continents. We apply standardized treatments of nutrient addition and herbivore reduction, allowing us to disentangle the regional and local drivers of soil organism activity. We use structural equation modeling to assess the direct and indirect effects of local and regional drivers on soil biological activities. Microbial and detritivore activities are positively correlated across global grasslands. These correlations are shaped more by global climatic factors than by local treatments, with annual precipitation and soil water content explaining the majority of the variation. Nutrient addition tends to reduce microbial activity by enhancing plant growth, while herbivore reduction typically increases microbial and detritivore activity through increased soil moisture. Our findings emphasize soil moisture as a key driver of soil biological activity, highlighting the potential impacts of climate change, altered grazing pressure, and eutrophication on nutrient cycling and decomposition within grassland ecosystems. A global grassland study reveals soil moisture, not local treatments, drives microbial and detritivore activity, highlighting the potential influence of climate change and human interventions on grassland ecosystems. [ABSTRACT FROM AUTHOR]

Eutrophication usually impacts grassland biodiversity, community composition, and biomass production, but its impact on the stability of these community aspects is unclear. One challenge is that stability has many facets that can be tightly correlated (low dimensionality) or highly disparate (high dimensionality). Using standardized experiments in 55 grassland sites from a globally distributed experiment (NutNet), we quantify the effects of nutrient addition on five facets of stability (temporal invariability, resistance during dry and wet growing seasons, recovery after dry and wet growing seasons), measured on three community aspects (aboveground biomass, community composition, and species richness). Nutrient addition reduces the temporal invariability and resistance of species richness and community composition during dry and wet growing seasons, but does not affect those of biomass. Different stability measures are largely uncorrelated under both ambient and eutrophic conditions, indicating consistently high dimensionality. Harnessing the dimensionality of ecological stability provides insights for predicting grassland responses to global environmental change. Anthropogenic eutrophication is a driver of plant community shifts in many grassland ecosystems. Here, the authors use data from a globally distributed experiment to assess how nutrient addition affects multiple facets of grassland ecological stability and their correlations. [ABSTRACT FROM AUTHOR]

Soil net nitrogen (N) mineralization is a key biogeochemical process influencing plant available N and net primary productivity in terrestrial ecosystems. However, the spatial variations and controlling factors of soil net N mineralization (RPNM) in arid and semi‐arid grasslands are less studied and unclear.In this study, we investigated the soil RPNM by performing a laboratory incubation experiment. Soil samples were collected from 30 sites in three east–west transects on the Inner Mongolia Plateau (MP), Loess Plateau (LP) and Tibetan Plateau (TP) along a 3200 km arid and semi‐arid grassland gradient, with each transect containing three different grassland types (meadow steppe [MS], typical steppe [TS] and desert steppe [DS], respectively).Results showed that the average RPNM values ranged from −0.37 to 1.29 mg N kg−1 day−1, with a significantly lower RPNM found in the DS (0.08 ± 0.01 mg N kg−1 day−1) compared with those in the MS (0.30 ± 0.03 mg N kg−1 day−1) and in the TS (0.33 ± 0.03 mg N kg−1 day−1) in the MP and LP transects (p < 0.05). This difference could be explained by variations in climatic and soil factors, such as hydrothermal index (HT), the soil pH, soil organic matter (SOM) and precipitation. However, no significant differences in RPNM were found among different grassland types in the TP transect, possibly due to the similarly low microbial activity, as indicated by the microbial biomass carbon values. Across all three grassland transects, HT, SOM and microbial variables were the major factors controlling RPNM, which together explained 20.7% of the variation in RPNM. Further structural equation model analysis indicated HT was an integral predictor of RPNM, directly or indirectly via SOM, under different conditions of precipitation and temperature.Our findings provide field evidence and parameters for biogeochemical cycling to better predict future N transformation processes under changing precipitation and temperature regimes across a wide range of arid and semi‐arid grassland ecosystems. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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.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.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.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.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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Anthropogenic nutrient enrichment is driving global biodiversity decline and modifying ecosystem functions. Theory suggests that plant functional types that fix atmospheric nitrogen have a competitive advantage in nitrogen-poor soils, but lose this advantage with increasing nitrogen supply. By contrast, the addition of phosphorus, potassium, and other nutrients may benefit such species in lownutrient environments by enhancing their nitrogen-fixing capacity. We present a global-scale experiment confirming these predictions for nitrogen-fixing legumes (Fabaceae) across 45 grasslands on six continents. Nitrogen addition reduced legume cover, richness, and biomass, particularly in nitrogen-poor soils, while cover of non-nitrogenfixing plants increased. The addition of phosphorous, potassium, and other nutrients enhanced legume abundance, but did not mitigate the negative effects of nitrogen addition. Increasing nitrogen supply thus has the potential to decrease the diversity and abundance of grassland legumes worldwide regardless of the availability of other nutrients, with consequences for biodiversity, food webs, ecosystem resilience, and genetic improvement of protein-rich agricultural plant species. [ABSTRACT FROM AUTHOR]

Seminatural grasslands are important biodiversity hotspots, but they are increasingly degraded by intensive agriculture. Grassland restoration is considered to be promising in halting the ongoing loss of biodiversity, but this evaluation is mostly based on plant communities. Insect herbivores contribute substantially to grassland biodiversity and to the provisioning of a variety of ecosystem functions. However, it is unclear how they respond to different measures that are commonly used to restore seminatural grasslands from intensively used agricultural land. We studied the long‐term success of different restoration techniques, which were originally targeted at reestablishing seminatural grassland plant communities, for herbivorous insect communities on taxonomic as well as functional level. Therefore, we sampled insect communities 22 yr after the establishment of restoration measures. These measures ranged from harvest and removal of biomass to removal of the topsoil layer and subsequent seeding of plant propagules. We found that insect communities in restored grasslands had higher taxonomic and functional diversity compared to intensively managed agricultural grasslands and were more similar in composition to target grasslands. Restoration measures including topsoil removal proved to be more effective, in particular in restoring species characterized by functional traits susceptible to intensive agriculture (e.g., large‐bodied species). Our study shows that long‐term success in the restoration of herbivorous insect communities of seminatural grasslands can be achieved by different restoration measures and that more invasive approaches that involve the removal of the topsoil layer are more effective. We attribute these restoration successes to accompanying changes in the plant community, resulting in bottom‐up control of the herbivore community. Our results are of critical importance for management decisions aiming to restore multi‐trophic communities, their functional composition and consequently the proliferation of ecosystem functions. [ABSTRACT FROM AUTHOR]

Microbial processing of aggregate‐unprotected organic matter inputs is key for soil fertility, long‐term ecosystem carbon and nutrient sequestration and sustainable agriculture. We investigated the effects of adding multiple nutrients (nitrogen, phosphorus and potassium plus nine essential macro‐ and micro‐nutrients) on decomposition and biochemical transformation of standard plant materials buried in 21 grasslands from four continents. Addition of multiple nutrients weakly but consistently increased decomposition and biochemical transformation of plant remains during the peak‐season, concurrent with changes in microbial exoenzymatic activity. Higher mean annual precipitation and lower mean annual temperature were the main climatic drivers of higher decomposition rates, while biochemical transformation of plant remains was negatively related to temperature of the wettest quarter. Nutrients enhanced decomposition most at cool, high rainfall sites, indicating that in a warmer and drier future fertilized grassland soils will have an even more limited potential for microbial processing of plant remains. [ABSTRACT FROM AUTHOR]

Plant traits are commonly used to predict ecosystem-level processes, but the validity of such predictions is dependent on the assumption that trait variability between species is greater than trait variability within a species—the robustness assumption. Here, we compare leaf trait intraspecific and interspecific variability depending on geographical differences between sites and 5 years of experimental herbivore exclusion in two vegetation types of subalpine grasslands in Switzerland. Four leaf traits were measured from eight herbaceous species common to all 18 sites. Intraspecific trait variability differed significantly depending on site and herbivory. However, the amount and structure of variability depended on the trait measured and whether considering leaf traits separately or multiple leaf traits simultaneously. Leaf phosphorus concentration showed the highest intraspecific variability, while specific leaf area showed the highest interspecific variability and displayed intraspecific variability only in response to herbivore exclusion. Species identity based on multiple traits was not predictable. We find intraspecific variability is an essential consideration when using plant functional traits as a common currency not just species mean traits. This is particularly true for leaf nutrient concentrations, which showed high intraspecific variability in response to site differences and herbivore exclusion, a finding which suggests that the robustness assumption does not always hold. [ABSTRACT FROM AUTHOR]

Mammalian grazing induces changes in vegetation properties in grasslands, which can affect a wide variety of other animals including many arthropods. However, the impacts may depend on the type and body size of these mammals. Furthermore, how mammals influence functional trait syndromes of arthropod communities is not well known. We progressively excluded large (e.g. red deer, chamois), medium (e.g. alpine marmot, mountain hare), and small (e.g. mice) mammals using size‐selective fences in two vegetation types (short‐ and tall‐grass vegetation) of subalpine grasslands. We then assessed how these exclusions affected the community composition and functional traits of ground beetles (Coleoptera, Carabidae), and which vegetation characteristic mediated the observed effects. Total carabid biomass, the activity densities of carabids with specific traits (i.e. small eyes, short wings), the richness of small‐eyed species and the richness of herbivorous species were significantly higher when certain mammals were excluded compared to when all mammals had access, regardless of vegetation type. Excluding large and medium mammals increased the activity density of herbivorous carabid species, but only in short‐grass vegetation. Similarly, excluding large mammals (ungulates) altered carabid species composition in the short‐, but not in the tall‐grass vegetation. All these responses were related to aboveground plant biomass, but not to plant Shannon diversity or vegetation structural heterogeneity. Our results indicate that changes in aboveground plant biomass are key drivers of mammalian grazers’ influence on carabids, suggesting that bottom–up forces are important in subalpine grassland systems. The exclusion of ungulates provoked the strongest carabid response. Our results, however, also highlight the ecological significance of smaller herbivorous mammals. Our study furthermore shows that mammalian grazing not only altered carabid community composition, but also caused community‐wide functional trait shifts, which could potentially have a wider impact on species interactions and ecosystem functioning. [ABSTRACT FROM AUTHOR]

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

Ungulates, smaller mammals, and invertebrates can each affect soil biota through their influence on vegetation and soil characteristics. However, direct and indirect effects of the aboveground biota on soil food webs remain to be unraveled. We assessed effects of progressively excluding aboveground large-, medium- and small-sized mammals as well as invertebrates on soil nematode diversity and feeding type abundances in two subalpine grassland types: short- and tall-grass vegetation. We explored pathways that link exclusions of aboveground biota to nematode feeding type abundances via changes in plants, soil environment, soil microbial biomass, and soil nutrients. In both vegetation types, exclusions caused a similar shift toward higher abundance of all nematode feeding types, except plant feeders, lower Shannon diversity, and lower evenness. These effects were strongest when small mammals, or both small mammals and invertebrates were excluded in addition to excluding larger mammals. Exclusions resulted in a changed abiotic soil environment that only affected nematodes in the short-grass vegetation. In each vegetation type, exclusion effects on nematode abundances were mediated by different drivers related to plant quantity and quality. In the short-grass vegetation, not all exclusion effects on omni-carnivorous nematodes were mediated by the abundance of lower trophic level nematodes, suggesting that omni-carnivores also depended on other prey than nematodes. We conclude that small aboveground herbivores have major impacts on the soil food web of subalpine short- and tall-grass ecosystems. Excluding aboveground animals caused similar shifts in soil nematode assemblages in both subalpine vegetation types, however, mechanisms turned out to be system-specific. [ABSTRACT FROM AUTHOR]

Vertebrate and invertebrate herbivores alter plant communities directly by selectively consuming plant species; and indirectly by inducing morphological and physiological changes to plant traits that provide competitive or survivorship advantages to some life forms over others. Progressively excluding aboveground herbivore communities (ungulates, medium and small sized mammals, invertebrates) over five growing seasons, we explored how leaf morphology (specific leaf area or SLA) and nutrition (nitrogen, carbon, phosphorous, potassium, sodium, and calcium) of different plant life forms (forbs, legumes, grasses, sedges) correlated with their dominance. We experimented in two subalpine grassland types with different land-use histories: (1) heavily grazed, nutrient-rich, short-grass vegetation and (2) lightly grazed, lower nutrient tall-grass vegetation. We found differences in leaf traits between treatments where either all herbivores were excluded or all herbivores were present, showing the importance of considering the impacts of both vertebrates and invertebrates on the leaf traits of plant species. Life forms responses to the progressive exclusion of herbivores were captured by six possible combinations: (1) increased leaf size and resource use efficiency (leaf area/nutrients) where lower nutrient levels are invested in leaf construction, but a reduction in the number of leaves, for example, forbs in both vegetation types, (2) increased leaf size and resource use efficiency, for example, legumes in short grass, (3) increased leaf size but a reduction in the number of leaves, for example, legumes in the tall grass, (4) increased number of leaves produced and increased resource use efficiency, for example, grasses in the short grass, (5) increased resource use efficiency of leaves only, for example, grasses and sedges in the tall grass, and (6) no response in terms of leaf construction or dominance, for example, sedges in the short grass. Although we found multiple possible responses by life forms to progressive exclusion of herbivores, we also found some important generalities. Changes in leaf traits of legumes and grasses correlated with their increasing dominance in the short-grass vegetation and plants were more efficient at constructing photosynthetic tissue when herbivores are present with few exceptions. These results demonstrate that vertebrate and invertebrate herbivores are essential to maintain plant species richness and resource-use efficiency. [ABSTRACT FROM AUTHOR]

The comparable and integrated nature of plant functional types and advances in high-spectral-resolution remote sensing techniques (i.e. imaging spectroscopy) make their combination highly interesting for spatially continuous and repeatable large-scale ecosystem monitoring. Depending on physical environment and stress, plants invest in covarying biochemical and structural traits, influencing spectral characteristics of vegetation. These traits are assumed to bear a more direct causal relationship to plant functional types than to plant life/growth forms. However, the connection between a vegetation community's functional and spectral response remains to be established., We assessed the correlation structure between (i) biochemical and structural vegetation traits (biomass, dry matter content, nitrogen content, neutral detergent fibre content), (ii) plant life/growth forms and (iii) seven plant functional types of two categories (strategy types, indicator values) collected in heterogeneous alpine grassland. We then used airborne imaging spectroscopy data from the same area to model and predict plant life/growth forms and plant functional types at the vegetation community level using partial least squares regression and validated our models based on an independent data set., We found high correlations between many of the biochemical and structural vegetation traits, plant life/growth forms and plant functional types tested. Using airborne imaging spectroscopy data, we successfully modelled and predicted most plant life/growth forms ( R2 max. = 0·56) and all plant functional types ( R2 max. = 0·62). However, model performance for plant life/growth forms decreased substantially during external validation and overall model consistency was low (average change in R2 = 72%), while plant functional type models were much more consistent (average change in R2 = 20%). Based on our findings, we developed a conceptual framework using the theory and methodology of vegetation ecology and imaging spectroscopy to link the vegetation community's functional to its spectral signature., Our results encourage the use of plant functional types in imaging spectroscopy in order to aid the large-scale monitoring of ecosystems, which is particularly important given the increased availability of airborne data and the prospective launches of spaceborne instruments in the near future. [ABSTRACT FROM AUTHOR]

Aboveground herbivores have strong effects on grassland nitrogen (N) cycling. They can accelerate or slow down soil net N mineralization depending on ecosystem productivity and grazing intensity. Yet, most studies only consider either ungulates or invertebrate herbivores, but not the combined effect of several functionally different vertebrate and invertebrate herbivore species or guilds. We assessed how a diverse herbivore community affects net N mineralization in subalpine grasslands. By using size-selective fences, we progressively excluded large, medium, and small mammals, as well as invertebrates from two vegetation types, and assessed how the exclosure types (ET) affected net N mineralization. The two vegetation types differed in long-term management (centuries), forage quality, and grazing history and intensity. To gain a more mechanistic understanding of how herbivores affect net N mineralization, we linked mineralization to soil abiotic (temperature; moisture; NO3- NH44+, and total inorganic N concentrations/pools; C, N, P concentrations; pH; bulk density), soil biotic (microbial biomass; abundance of collembolans, mites, and nematodes) and plant (shoot and root biomass; consumption; plant C, N, and fiber content; plant N pool) properties. Net N mineralization differed between ET, but not between vegetation types. Thus, short-term changes in herbivore community composition and, therefore, in grazing intensity had a stronger effect on net N mineralization than long-term management and grazing history. We found highest N mineralization values when only invertebrates were present, suggesting that mammals had a negative effect on net N mineralization. Of the variables included in our analyses, only mite abundance and aboveground plant biomass explained variation in net N mineralization among ET. Abundances of both mites and leaf-sucking invertebrates were positively correlated with aboveground plant biomass, and biomass increased with progressive exclusion. The negative impact of mammals on net N mineralization may be related partially to (1) differences in the amount of plant material (litter) returned to the belowground subsystem, which induced a positive bottom-up effect on mite abundance, and (2) alterations in the amount and/or distribution of dung, urine, and food waste. Thus, our results clearly show that short-term alterations of the aboveground herbivore community can strongly impact nutrient cycling within ecosystems independent of long-term management and grazing history. [ABSTRACT FROM AUTHOR]

Recognition is growing that besides ungulates, small vertebrate and invertebrate herbivores are important drivers of grassland functioning. Even though soil microarthropods play key roles in several soil processes, effects of herbivores—especially those of smaller body size—on their communities are not well understood. Therefore, we progressively excluded large, medium and small vertebrate and invertebrate herbivores for three growing seasons using size-selective fences in two vegetation types in subalpine grasslands; short-grass and tall-grass vegetation generated by high and low historical levels of ungulate grazing. Herbivore exclusions generally had few effects on microarthropod communities, but exclusion of all herbivore groups resulted in decreased total springtail and Poduromorpha richness compared with exclusion of only ungulates and medium-sized mammals, regardless of vegetation type. The tall-grass vegetation had a higher total springtail richness and mesostigmatid mite abundance than the short-grass vegetation and a different oribatid mite community composition. Although several biotic and abiotic variables differed between the exclusion treatments and vegetation types, effects on soil microarthropods were best explained by differences in nutrient and fibre content of the previous year’s vegetation, a proxy for litter quality, and to a lesser extent soil temperature. After three growing seasons, smaller herbivores had a stronger impact on these functionally important soil microarthropod communities than large herbivores. Over longer time-scales, however, large grazers created two different vegetation types and thereby influenced microarthropod communities bottom-up, e.g. by altering resource quality. Hence, both short- and long-term consequences of herbivory affected the structure of the soil microarthropod community. [ABSTRACT FROM AUTHOR]

Plant growth can be limited by resource acquisition and defence against consumers, leading to contrasting trade-off possibilities. The competition-defence hypothesis posits a trade-off between competitive ability and defence against enemies (e.g. herbivores and pathogens). The growth-defence hypothesis suggests that strong competitors for nutrients are also defended against enemies, at a cost to growth rate. We tested these hypotheses using observations of 706 plant populations of over 500 species before and following identical fertilisation and fencing treatments at 39 grassland sites worldwide. Strong positive covariance in species responses to both treatments provided support for a growth-defence trade-off: populations that increased with the removal of nutrient limitation (poor competitors) also increased following removal of consumers. This result held globally across 4 years within plant life-history groups and within the majority of individual sites. Thus, a growth-defence trade-off appears to be the norm, and mechanisms maintaining grassland biodiversity may operate within this constraint. [ABSTRACT FROM AUTHOR]

. 1. Grasslands cover approximately 40% of the Earth's terrestrial landscape, supporting large communities of vertebrate and invertebrate herbivores. Orthoptera play an important role, consuming relatively large amounts of biomass. Their occurrence can be strongly affected by habitat diversity and structure, which can be shaped by large herbivores. Several studies have focused on the impact of livestock on Orthoptera communities, but little is known about how wild ungulates influence the abundance and diversity of these insects in grassland ecosystems. 2. We studied Orthoptera abundance and diversity in subalpine grasslands in the Swiss Alps, where grazing by red deer and chamois has created a mosaic of short- and tall-grass patches. Data on vegetation structure, habitat diversity and plant nitrogen (N) content allowed us to consider how these parameters affected the occurrence of Orthoptera at our study sites. 3. We found a total of nine Orthoptera species with an average density of 2.6 individuals m−2. Neither Orthoptera abundance nor diversity differed between short- and tall-grass patches created by large ungulates. Both Orthoptera abundance and diversity were, however, positively influenced by increasing vegetation height, but negatively by increasing habitat diversity within patches. Increasing plant N content promoted a more even spread of species within the insect assemblage on short- but not on tall-grass patches. 4. Large-scale habitat alteration by wild ungulates had no direct effect on the abundance and diversity of Orthoptera. However, we observed that they indirectly affected Orthoptera abundance and diversity by altering plant N content and the structure of the habitat at small scales. [ABSTRACT FROM AUTHOR]

A major obstacle for predicting the effects of climate and land use changes on global soil carbon (C) stores is the very limited knowledge about the long timescale dynamics of the relatively stable fraction of soil C, which represents the bulk of soil C and the primary determinant of the long-term C balance of terrestrial ecosystems. In this study, we examined how variable topo-edaphic conditions and herds of native migratory ungulates influenced turnover of the stable pool (total minus active fraction) of soil C in grasslands of Yellowstone National Park (YNP). Soil C properties were determined for grasslands located inside and outside long-term ungulate exclosures established 1958-1962 at seven variable topographic positions. Active C pool sizes, estimated with soil laboratory incubations, and soil radiocarbon measures were used to parameterize a process-based model to determine turnover of the stable C pool at the sites. Stable C turnover ranged 37-653 and 89-869 years for 0-10 and 0-20 cm soils, respectively. Among ungrazed communities, there was a trend for stable soil C turnover to slow along topographic gradients of increasing soil moisture, soil C content, and shoot biomass from hilltop to slope-bottom positions. This was likely a result of an increasing amount of support tissue resulting in greater concentrations of lignin and cellulose as shoot biomass increased down slope. In contrast, across the grazed landscape, stable C turnover sped up from hilltop to slope-bottom positions, which was likely a consequence of grazer effects on plant species composition along the topographic gradient. These findings indicated that despite topography playing the primary role in controlling such important site characteristics as soil moisture, soil C content, and plant production in YNP grassland, the long-term turnover of the stable C pool was determined by herbivores. The results demonstrate the important regulatory role of herbivores in controlling the C balance of this semiarid grassland ecosystem. [ABSTRACT FROM AUTHOR]

Grazing and seasonal variation in precipitation and temperature are important controls of soil and plant processes in grasslands. As these ecosystems store up to 30% of the world’s belowground carbon (C), it is important to understand how this variability affects mineral soil C pools/fluxes, and how C cycling might be affected by changes in precipitation and temperature, due to climate change. The aim of this study was to investigate the effects of grazing and differences in soil temperature and moisture on standard organic matter (OM) decomposition rates (cotton cloth) incubated in the top 10 cm soil of grasslands with variable topography in Yellowstone National Park (YNP) during the 2004 growing season. Grazing did not affect soil temperature, moisture, cotton cloth decomposition rates, soil bulk density, soil C and N concentrations, or soil C:N ratios. However, a large spatio-temporal variability in decomposition was observed: cotton cloth decomposition was positively related to soil moisture and soil C and N concentrations, and negatively to soil temperature. Highest decomposition rates were found in wetter slope bottom soils [season averages of decomposition given as rate of decomposition (cotton rotting rate = CRR) = 23–26%] and lower rates in drier, hill-top soils (season averages, CRR = 20%). Significantly higher decomposition rates were recorded in spring, early summer and early fall when soils were moist and cool (spring, CRR = 25%; early summer, CRR = 26%; fall, CRR = 20%) compared to mid-summer (CRR = 18%) when soils were dry and warm. Our findings suggest that climate-change related decreases in precipitation and increases in temperature predicted for North American grasslands would decrease soil OM decomposition in YNP, which contrasts the general assumption that increases in temperature would accelerate OM decomposition rates. [ABSTRACT FROM AUTHOR]

We examined the role of red deer ( Cervus elaphus L.) in translocating phosphorus (P) from their preferred grazing sites (short-grass vegetation on subalpine grasslands) to their wider home range in a subalpine grassland ecosystem in the Central European Alps. Phosphorus was used because it is the limiting nutrient in these grasslands. When we compared P removal of aboveground biomass due to grazing with P input due to the deposit of feces on a grid of 268 cells (20 m × 20 m) covering the entire grassland, we detected distinct spatial patterns: the proportion of heavily grazed short-grass vegetation increased with increasing soil-P pool, suggesting that red deer preferably grazed on grid cells with a higher soil-P pool. Biomass consumption related to increased proportion of short-grass vegetation, and therefore P removal, increased with increasing soil-P pool. However, within the two vegetation types (short-grass and tall-grass), consumption was independent from soil-P pool. In addition, P input rates from defecation increased with increasing soil-P pool, resulting in a constant mean net P loss of 0.083 kg ha−1 y−1 (0.03%–0.07% of soil-P pool) independent of both soil-P pool and vegetation type. Thus, there was no P translocation between grid cells with different soil-P pools or between short-grass and tall-grass vegetation. Based on these results, it is likely that the net rate of P loss is too small to explain the observed changes in vegetation composition from tall-herb/meadow communities to short-grass and from tall-grass to short-grass on the grassland since 1917. Instead, we suggest that the grazing patterns of red deer directly induced succession from tall-herb/meadow communities to short-grass vegetation. Yet, it is also possible that long-term net soil-P losses indirectly drive plant succession from short-grass to tall-grass vegetation, because nutrient depletion could reduce grazing pressure in short-grass vegetation and enable the characteristic tall-grass species Carex sempervirens Vill. to establish. [ABSTRACT FROM AUTHOR]

Landscape position, grazing, and seasonal variation in precipitation and temperature create spatial and temporal variability in soil processes, and plant biomass and composition in grasslands. However, it is unclear how this variation in plant and soil properties affects carbon dioxide (CO2) fluxes. The aim of this study is to explore the effect of grazing, topographic position, and seasonal variation in soil moisture and temperature on plant assimilation, shoot and soil respiration, and net ecosystem CO2 exchange (NEE). Carbon dioxide fluxes, vegetation, and environmental variables were measured once a month inside and outside long-term ungulate exclosures in hilltop (dry) to slope bottom (mesic) grassland throughout the 2004 growing season in Yellowstone National Park. There was no difference in vegetation properties and CO2 fluxes between the grazed and the ungrazed sites. The spatial and temporal variability in CO2 fluxes were related to differences in aboveground biomass and total shoot nitrogen content, which were both related to variability in soil moisture. All sites were CO2 sinks (NEE>0) for all our measurments taken throughout the growing season; but CO2 fluxes were four- to fivefold higher at sites supporting the most aboveground biomass located at slope bottoms, compared to the sites with low biomass located at hilltops or slopes. The dry sites assimilated more CO2 per gram aboveground biomass and stored proportionally more of the gross-assimilated CO2 in the soil, compared to wet sites. These results indicate large spatio-temporal variability of CO2 fluxes and suggest factors that control the variability in Yellowstone National Park. [ABSTRACT FROM AUTHOR]

The spatial grazing patterns of red deer (Cervus elaphus L.) in a subalpine grassland ecosystem of the Swiss National Park were analysed with reference to the phosphorus content in the topsoil and to the former agricultural management system. Changes in the composition of the vegetation and in plant species richness were studied on permanent plots set up between 1917 and 1945 in areas which today are heavily grazed by red deer, and were related to the development of the red deer population.The spatial grazing patterns of red deer are, 80 years after the Park’s foundation, still determined by former agricultural use. Red deer prefer the most phosphorus-rich, formerly irrigated parts for grazing.Grazing by red deer was found to be the driving force behind temporal changes in the vegetation observed on the subalpine grasslands. Plant succession has proceeded faster in the preferred grazing areas and the vegetation has adapted to the grazing pressure with a decrease in tall-growing plant species. At the same time, the abundance of unpalatable species has increased due to (i) morphological defences, (ii) spatial avoidance, i.e. by growing close to the ground and (iii) temporal avoidance because they have a short life-span (annuals). Changes in vegetation composition have coincided with a sharp increase in species richness. Between 1917 and 1999, the number of plant species has doubled on average. We found a highly significant correlation between the changes in the numbers of plant species and red deer, with species richness increasing with increasing deer density. [Copyright &y& Elsevier]

Increasing evidence suggests that ecosystem functions are strongly linked to morphological plant traits, like specific leaf area (SLA) and its variability, which serve as a proxy of functional diversity (FD). Functional diversity is rarely studied at regional scales, and its scale dependence is poorly understood. Capturing trait variations at distinct spatial scales and in differently managed grasslands remains challenging, mainly because a limited number of trait measurements are available and field campaigns are time-consuming. Here, we derived α- and β-FD indices based on SLA measured in the field and estimated from optical satellite data by using molecular absorption profiles of leaves in canopies. We inverted the 1-D columnar radiative transfer model PROSAIL using Sentinel-2 reflectance data at canopy level. From the inversion we were able to distinguish different alpine management types based on retrieved SLA. Model uncertainties were mainly related to the different local plant communities, here represented by functional diversity indices and community-weighted means of traits. Thus, successful PROSAIL application was affected by management type. Management categories displaying lower α-FD, like mowed and fertilized, delivered the most reliable results. Further, we compared FD (i.e., richness, evenness, divergence) from local to regional scales. Locally, management determines the magnitude of FD, whereas on a regional scale, parcel size and the uniformity of agricultural practices control trait diversity. Our results highlight the importance of quantifying β-FD from space as it delivers additional information on the impact of management types, differing from locally measured α-FD values. • CWM of SLA estimable from Sentinel-2 datasets with PROSAIL. • Alpha functional diversity influences remotely sensed plant trait retrieval. • It is crucial to quantify an area-wide beta functional diversity. • Management types impact spatial components of functional diversity differently. [ABSTRACT FROM AUTHOR]