Soil microorganisms are critical to ecosystem functioning and the maintenance of soil fertility. However, despite global increases in the inputs of nitrogen (N) and phosphorus (P) to ecosystems due ...to human activities, we lack a predictive understanding of how microbial communities respond to elevated nutrient inputs across environmental gradients. Here we used high-throughput sequencing of marker genes to elucidate the responses of soil fungal, archaeal, and bacterial communities using an N and P addition experiment replicated at 25 globally distributed grassland sites. We also sequenced metagenomes from a subset of the sites to determine how the functional attributes of bacterial communities change in response to elevated nutrients. Despite strong compositional differences across sites, microbial communities shifted in a consistent manner with N or P additions, and the magnitude of these shifts was related to the magnitude of plant community responses to nutrient inputs. Mycorrhizal fungi and methanogenic archaea decreased in relative abundance with nutrient additions, as did the relative abundances of oligotrophic bacterial taxa. The metagenomic data provided additional evidence for this shift in bacterial life history strategies because nutrient additions decreased the average genome sizes of the bacterial community members and elicited changes in the relative abundances of representative functional genes. Our results suggest that elevated N and P inputs lead to predictable shifts in the taxonomic and functional traits of soil microbial communities, including increases in the relative abundances of faster-growing, copiotrophic bacterial taxa, with these shifts likely to impact belowground ecosystems worldwide.
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.
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.
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).
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.
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.
Droughts can strongly affect grassland productivity and biodiversity, but responses differ widely. Nutrient availability may be a critical factor explaining this variation, but is often ignored in ...analyses of drought responses. Here, we used a standardized nutrient addition experiment covering 10 European grasslands to test if full‐factorial nitrogen, phosphorus, and potassium addition affected plant community responses to inter‐annual variation in drought stress and to the extreme summer drought of 2018 in Europe. We found that nutrient addition amplified detrimental drought effects on community aboveground biomass production. Drought effects also differed between functional groups, with a negative effect on graminoid but not forb biomass production. Our results imply that eutrophication in grasslands, which promotes dominance of drought‐sensitive graminoids over forbs, amplifies detrimental drought effects. In terms of climate change adaptation, agricultural management would benefit from taking into account differential drought impacts on fertilized versus unfertilized grasslands, which differ in ecosystem services they provide to society.
Drought impacts on grasslands vary widely. Nutrient availability may be a critical factor explaining this variation. We used a standardized nutrient addition experiment covering 10 European grasslands to test if nitrogen, phosphorus, and potassium addition affected responses to inter‐annual variation in drought stress and to the extreme summer drought of 2018. We found that nutrient addition amplified detrimental drought effects on biomass production. Drought effects differed between functional groups, with a negative effect on graminoids but not forbs. For climate change adaptation, agricultural management would benefit from taking into account differential drought impacts on fertilized versus unfertilized grasslands, which differ in ecosystem service provisioning.
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.
Summary statement
Plants are potentially exposed to many different herbivores simultaneously, but we often study the effect of specific herbivore guilds or species on plant performance. In this review, we unpack and explore differences between vertebrate and invertebrate focussed studies which examined how plant traits respond to and affect herbivory. We examined overall trends and biases, differences in the types of traits examined and the relationships between plant traits and herbivory.
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.
All multicellular organisms host a diverse microbiome composed of microbial pathogens, mutualists, and commensals, and changes in microbiome diversity or composition can alter host fitness and ...function. Nonetheless, we lack a general understanding of the drivers of microbiome diversity, in part because it is regulated by concurrent processes spanning scales from global to local. Global-scale environmental gradients can determine variation in microbiome diversity among sites, however an individual host's microbiome also may reflect its local micro-environment. We fill this knowledge gap by experimentally manipulating two potential mediators of plant microbiome diversity (soil nutrient supply and herbivore density) at 23 grassland sites spanning global-scale gradients in soil nutrients, climate, and plant biomass. Here we show that leaf-scale microbiome diversity in unmanipulated plots depended on the total microbiome diversity at each site, which was highest at sites with high soil nutrients and plant biomass. We also found that experimentally adding soil nutrients and excluding herbivores produced concordant results across sites, increasing microbiome diversity by increasing plant biomass, which created a shaded microclimate. This demonstration of consistent responses of microbiome diversity across a wide range of host species and environmental conditions suggests the possibility of a general, predictive understanding of microbiome diversity.
Human‐caused loss of vertebrate and invertebrate animals, defaunation, is increasing, and potentially affects plant community structure of diverse grassland ecosystems world‐wide.
We experimentally ...simulated defaunation using size‐selective fences to progressively exclude large‐, medium‐ and small‐sized mammals, and invertebrates from two subalpine vegetation types in the Swiss National Park (SNP): intensively grazed short‐grass and moderately grazed tall‐grass vegetation. We assessed plant community properties yearly from 2009 to 2013, and examined treatment effects on plant community structure in the two grassland types.
In the short‐grass vegetation, the exclusion of large mammals increased total plant biomass, while the exclusion of large and medium‐sized mammals increased total, grass and forb biomass compared to when all animals had access. These increases became stronger when invertebrates were also excluded. The exclusion of all mammals and invertebrates increased biomass of grasses by 205%, forbs by 100% and total plant biomass by 118% compared to when all animals had access, hence enhancing relative biomass of grasses from 43.6% to 60%, changing plant species composition and lowering richness of forbs by 16%, the number of plant families by 13% and family‐level Shannon diversity by 23%. In contrast to these significant community‐level responses found in the short‐grass vegetation, there was no evidence that the size‐selective exclusion of animals altered the plant community structure of the tall‐grass vegetation. The contrasting results were due to the difference in plant community composition prior to our experiment, which were related to differences in quantity and quality of forage and in grazing intensities of herbivores between the two grassland types.
Synthesis. Our results showed that different‐sized animals, in particular large mammals and invertebrates, contributed to maintain the plant community structure in the short‐grass vegetation, highlighting the importance of multiple, functionally different animal groups for ecosystem functioning and stability. In contrast to the short‐grass vegetation, we could not detect such a top‐down control by animals in the tall‐grass vegetation. Our results suggest that potential defaunation effects on grassland plant community structure depend on the degree of grazing pressure release and grassland vegetation type.
Different‐sized animals, in particular large mammals and invertebrates, contribute to maintain the plant community structure in the short‐grass vegetation, highlighting the importance of multiple, functionally different animal groups for ecosystem functioning and stability. However, we cannot detect such a top‐down control by animals in the tall‐grass vegetation. Our results suggest that potential defaunation effects on grassland plant community structure depend on the degree of grazing pressure release and grassland vegetation type.
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.
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 m2 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.
Niche dimensionality provides a general theoretical explanation for biodiversity-more niches, defined by more limiting factors, allow for more ways that species can coexist. Because plant species ...compete for the same set of limiting resources, theory predicts that addition of a limiting resource eliminates potential trade-offs, reducing the number of species that can coexist. Multiple nutrient limitation of plant production is common and therefore fertilization may reduce diversity by reducing the number or dimensionality of belowground limiting factors. At the same time, nutrient addition, by increasing biomass, should ultimately shift competition from belowground nutrients towards a one-dimensional competitive trade-off for light. Here we show that plant species diversity decreased when a greater number of limiting nutrients were added across 45 grassland sites from a multi-continent experimental network. The number of added nutrients predicted diversity loss, even after controlling for effects of plant biomass, and even where biomass production was not nutrient-limited. We found that elevated resource supply reduced niche dimensionality and diversity and increased both productivity and compositional turnover. Our results point to the importance of understanding dimensionality in ecological systems that are undergoing diversity loss in response to multiple global change factors.
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