How plant traits respond to and affect vertebrate and invertebrate herbivores-Are measurements comparable across herbivore types?

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.

Impacts of Myrtle Rust Induced Tree Mortality on Species and Functional Richness within Seedling Communities of a Wet Sclerophyll Forest in Eastern Australia.

Austropuccinia psidii is an introduced plant pathogen known to have caused significant declines in populations of several Australian native Myrtaceae species. However, limited research has focused on the impacts of the pathogen on plant communities in the aftermath of its invasion. This study investigated the relationship between disease impact level, plant species diversity, and functional richness in seedling communities in a wet sclerophyll forest in southeast Queensland. A clear shift was found from early colonizer Myrtaceae species in the mid- and understory to a more diverse non-Myrtaceae seedling community indicative of secondary succession. Comparisons of key Myrtaceae species and the seedling community suggest that there may also be a shift towards species that produce drupes and larger seeds, and overall, a current reduction in fruit availability due to the dramatic loss of previously dominant species. Seedling diversity showed no significant correlation with tree mortality, possibly due to favorable rainfall conditions during the study period. The more subtle changes in forest composition, such as changes in fruit type and availability due to myrtle rust, however, could affect the visitation of local bird species in the short term and certainly reduce the store of early colonizing native shrub and tree species.

The community capacity curve applied to reforestation: a framework to support success.

Community involvement is critical for the success of many interventions designed to promote reforestation. To secure this involvement, it helps to recognize that communities are heterogenous both within and among themselves and possess diverse mixes of livelihood assets required to implement reforestation. We explore the relationship between livelihood assets and reforestation success and outline a conceptual model that we call the community capacity curve (CCC) applied to reforestation. We argue that the shape of the CCC is sigmoidal. Importantly, communities at the lower end of the CCC have limited capacity to implement reforestation projects without substantial and ongoing capacity building and other sorts of support, including through livelihood projects that improve food security and provide cash benefits. Communities at the higher part of the CCC have greater capacity to implement reforestation projects, especially projects focused on biodiversity and environmental services. The CCC can help design, implement, monitor and assess reforestation projects, select appropriate livelihood activities and types of reforestation, select communities suited to a reforestation project, guide implementation and understand projects' successes and failure. The CCC also provides a framework to engage with policy makers and funding bodies to explore the types of support for communities to reforest successfully. This article is part of the theme issue 'Understanding forest landscape restoration: reinforcing scientific foundations for the UN Decade on Ecosystem Restoration'.

Impact of roadside burning on genetic diversity in a high-biomass invasive grass.

The invasive grass-fire cycle is a widely documented feedback phenomenon in which invasive grasses increase vegetation flammability and fire frequency, resulting in further invasion and compounded effects on fire regimes. Few studies have examined the role of short-term adaptation in driving the invasive grass-fire cycle, despite invasive species often thriving after introduction to new environments. We used a replicated (nine locations), paired sampling design (burn vs unburnt sites) to test the hypothesis that roadside burning increases genetic diversity and thus adaptive potential in the invasive, high-biomass grass Cenchrus ciliaris. Between four and five samples per site (n = 93) were genotyped using the DArTseq platform, and we filtered the data to produce panels of 15,965 neutral and 5030 non-neutral single nucleotide polymorphism (SNP) markers. Using fastSTRUCTURE, we detected three distinct genetic clusters with extremely high F ST values among them (0.94-0.97) suggesting three different cultivars. We found high rates of asexual reproduction, possibly related to clonality or apomixis common in this species. At three locations, burnt and unburnt sites were genetically different, but genetic structure was not consistently related to fire management across the study region. Burning was associated with high genetic diversity and sexual reproduction in one genetic cluster, but with low genetic diversity and clonality in another. Individual SNPs were associated with longitude and genetic clustering, but not with recent fire management. Overall, we found limited evidence that roadside burning consistently increased genetic diversity and adaptive potential in C. ciliaris; evolutionary and breeding history more strongly shaped genetic structure. Roadside burning could therefore continue to be used for managing biomass in this species, with continued monitoring. Our study provides a framework for detecting fire-related changes on a genetic level-a process that could be used as an early warning system to detect the invasive grass-fire cycle in future.

The contribution of insects to global forest deadwood decomposition.

The amount of carbon stored in deadwood is equivalent to about 8 per cent of the global forest carbon stocks1. The decomposition of deadwood is largely governed by climate2-5 with decomposer groups-such as microorganisms and insects-contributing to variations in the decomposition rates2,6,7. At the global scale, the contribution of insects to the decomposition of deadwood and carbon release remains poorly understood7. Here we present a field experiment of wood decomposition across 55 forest sites and 6 continents. We find that the deadwood decomposition rates increase with temperature, and the strongest temperature effect is found at high precipitation levels. Precipitation affects the decomposition rates negatively at low temperatures and positively at high temperatures. As a net effect-including the direct consumption by insects and indirect effects through interactions with microorganisms-insects accelerate the decomposition in tropical forests (3.9% median mass loss per year). In temperate and boreal forests, we find weak positive and negative effects with a median mass loss of 0.9 per cent and -0.1 per cent per year, respectively. Furthermore, we apply the experimentally derived decomposition function to a global map of deadwood carbon synthesized from empirical and remote-sensing data, obtaining an estimate of 10.9 ± 3.2 petagram of carbon per year released from deadwood globally, with 93 per cent originating from tropical forests. Globally, the net effect of insects may account for 29 per cent of the carbon flux from deadwood, which suggests a functional importance of insects in the decomposition of deadwood and the carbon cycle.

Negative effects of nitrogen override positive effects of phosphorus on grassland legumes worldwide.

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 low-nutrient 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-nitrogen-fixing 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.

Temporal rarity is a better predictor of local extinction risk than spatial rarity.

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

Non-Native Eragrostis curvula Impacts Diversity of Pastures in South-Eastern Australia Even When Native Themeda triandra Remains Co-Dominant.

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.

Inter-reef Halimeda algal habitats within the Great Barrier Reef support a distinct biotic community and high biodiversity.

Tropical marine biodiversity studies have been biased towards more accessible coastal habitats and shallow coral reefs, while deeper inter-reef habitats are less studied due to different survey challenges. One such inter-reef habitat is the 'bioherms' dominated by the calcareous Halimeda macroalgae. In the northern section of Australia's Great Barrier Reef, Halimeda algal bioherms occupy >6,000 km2 of the inter-reef seabed, more than twice the area of adjacent shallow coral reefs. Here, we describe the biodiversity of the plant, vertebrate and invertebrate communities inhabiting Halimeda bioherms. By combining previous spatial mapping with legacy benthic biodiversity datasets, we find that Halimeda bioherms are a critically important complex habitat that hosts higher average species richness and diversity for both plants and invertebrates than the surrounding inter-reef (non-coral reef) seascape. Furthermore, at the community level, the structure of the bioherm-associated biotic assemblage is distinct from the non-bioherm community, with 40% of Halimeda bioherm-associated species not recorded at any non-bioherm sites. These findings improve estimates of the biodiversity of the Great Barrier Reef and elevate Halimeda bioherms as a critically important inter-reef habitat. Regular long-term monitoring is required to detect potential impacts to inter-reef biodiversity and ecosystem structure and function under future climate change scenarios.

Seedling diversity in actively and passively restored tropical forest understories.

Alternative methods for restoring tropical forests influence the ecological processes that shape recruitment of understory species. In turn, the traits of species recruited will influence the ecological processes the forests provide now and over the long term. We assess the phylogenetic and functional structure of seedlings beneath monoculture plantations, mixed-species plantations (both active restoration) and regenerating selectively logged native forests (passive restoration), considering traits of specific leaf area (SLA, including within-species variation), leaf nitrogen and phosphorus content, life-form, potential plant height, and dispersal type. Monoculture plantations comprised seedlings that were more closely related then would be expected by chance (i.e., phylogenetically clustered), and regenerating forest contained species more distantly related then would be expected by chance (i.e., phylogenetically overdispersed). This suggests that seedlings beneath monocultures assemble through environmental filtering and through the dispersal limitation of predictable functional guilds. However, dispersal limitation is frequently overcome by human-assisted dispersal, increasing trait diversity. Comparing SLA values revealed that regenerating forests recruit seedlings with both high and low mean and variation of SLA, leading to higher overall diversity. Regenerating forest seedlings showed signs of environmental filtering, only based on within-species variation of SLA. Regenerating forest understories appear to favor species that show a high intraspecific variation in SLA values (e.g., Pterocarpus indicus Willd.) and at the same time provided habitat for later successional seedlings that show a lower intraspecific variation in SLA (e.g., Canarium luzonicum (Blume) A.Gray). This trait diversity suggests limiting similarity or competitive exclusion may be reduced because of niche differences, allowing species with different traits to coexist. Phylogenetic and functionally distinct species are restricted in their regeneration capacity, many of which are of conservation significance (under the IUCN Red List). Reforestation projects should maximize desired ecological services (including conservation value) by actively managing for the recruitment of species that are phylogenetically and functionally (including intraspecifically) distinct. This management aim will increase the probability of fulfilling a wider array of niche spaces and potentially increase the diversity of ecosystem services provided.

Opposing community assembly patterns for dominant and nondominant plant species in herbaceous ecosystems globally.

Biotic and abiotic factors interact with dominant plants-the locally most frequent or with the largest coverage-and nondominant plants differently, partially because dominant plants modify the environment where nondominant plants grow. For instance, if dominant plants compete strongly, they will deplete most resources, forcing nondominant plants into a narrower niche space. Conversely, if dominant plants are constrained by the environment, they might not exhaust available resources but instead may ameliorate environmental stressors that usually limit nondominants. Hence, the nature of interactions among nondominant species could be modified by dominant species. Furthermore, these differences could translate into a disparity in the phylogenetic relatedness among dominants compared to the relatedness among nondominants. By estimating phylogenetic dispersion in 78 grasslands across five continents, we found that dominant species were clustered (e.g., co-dominant grasses), suggesting dominant species are likely organized by environmental filtering, and that nondominant species were either randomly assembled or overdispersed. Traits showed similar trends for those sites (<50%) with sufficient trait data. Furthermore, several lineages scattered in the phylogeny had more nondominant species than expected at random, suggesting that traits common in nondominants are phylogenetically conserved and have evolved multiple times. We also explored environmental drivers of the dominant/nondominant disparity. We found different assembly patterns for dominants and nondominants, consistent with asymmetries in assembly mechanisms. Among the different postulated mechanisms, our results suggest two complementary hypotheses seldom explored: (1) Nondominant species include lineages adapted to thrive in the environment generated by dominant species. (2) Even when dominant species reduce resources to nondominant ones, dominant species could have a stronger positive effect on some nondominants by ameliorating environmental stressors affecting them, than by depleting resources and increasing the environmental stress to those nondominants. These results show that the dominant/nondominant asymmetry has ecological and evolutionary consequences fundamental to understand plant communities.

Increasing effects of chronic nutrient enrichment on plant diversity loss and ecosystem productivity over time.

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.

Ranking buffel: Comparative risk and mitigation costs of key environmental and socio-cultural threats in central Australia.

Changed fire regimes and the introduction of rabbits, cats, foxes, and large exotic herbivores have driven widespread ecological catastrophe in Australian arid and semi-arid zones, which encompass over two-thirds of the continent. These threats have caused the highest global mammal extinction rates in the last 200 years, as well as significantly undermining social, economic, and cultural practices of Aboriginal peoples of this region. However, a new and potentially more serious threat is emerging. Buffel grass (Cenchrus ciliaris L.) is a globally significant invader now widespread across central Australia, but the threat this ecological transformer species poses to biodiversity, ecosystem function, and culture has received relatively little attention. Our analyses suggest threats from buffel grass in arid and semi-arid areas of Australia are at least equivalent in magnitude to those posed by invasive animals and possibly higher, because unlike these more recognized threats, buffel has yet to occupy its potential distribution. Buffel infestation also increases the intensity and frequency of wildfires that affect biodiversity, cultural pursuits, and productivity. We compare the logistical and financial challenges of creating and maintaining areas free of buffel for the protection of biodiversity and cultural values, with the creation and maintenance of refuges from introduced mammals or from large-scale fire in natural habitats. The scale and expense of projected buffel management costs highlight the urgent policy, research, and financing initiatives essential to safeguard threatened species, ecosystems, and cultural values of Aboriginal people in central Australia.

Global impacts of fertilization and herbivore removal on soil net nitrogen mineralization are modulated by local climate and soil properties.

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

Rapid recovery of tropical forest diversity and structure after shifting cultivation in the Philippines uplands.

Shifting cultivation is a widespread land-use in the tropics that is considered a major threat to rainforest diversity and structure. In the Philippines, a country with rich biodiversity and high rates of species endemism, shifting cultivation, locally termed as kaingin, is a major land-use and has been for centuries. Despite the potential impact of shifting cultivation on forests and its importance to many people, it is not clear how biodiversity and forest structure recover after kaingin abandonment in the country, and how well these post-kaingin secondary forests can complement the old-growth forests. We investigated parameters of forest diversity and structure along a fallow age gradient in secondary forests regenerating after kaingin abandonment in Leyte Island, the Philippines (elevation range: 445-650 m asl). We first measured the tree diversity and forest structure indices in regenerating secondary forests and old-growth forest. We then measured the recovery of tree diversity and forest structure parameters in relation to the old-growth forest. Finally, using linear mixed effect models (LMM), we assessed the effect of different environmental variables on the recovery of forest diversity and structure. We found significantly higher species density in the oldest fallow sites, while Shannon's index, species evenness, stem number, basal area, and leaf area index were higher in the old-growth forest. A homogeneous species composition was found across the sites of older fallow age. Multivariate analysis revealed patch size as a strong predictor of tree diversity and forest structure recovery after shifting cultivation. Our study suggests that, secondary forests regenerating after shifting cultivation abandonment can recover rapidly. Although recovery of forest structure was not as rapid as the tree diversity, our older fallow sites contained a similar number of species as the old-growth forest. Many of these species are also endemic to the Philippines. Novel and emerging ecosystems like tropical secondary forests are of high conservation importance and can act as a refuge for dwindling tropical forest biodiversity.

Rapidly assessing cobenefits to advance threat-management alliances.

Conservation strategies aimed at reducing threats to biodiversity can have significant implications for multiple sectors in a socioeconomic system, but these cobenefits are often poorly understood. For example, many of the threats to native species also impede agricultural production, yet agriculture is typically perceived as in competition with conservation objectives. Although a comprehensive, multiobjective decision analysis is usually beyond the scope and capacity of conservation decision makers, failing to incorporate key socioeconomic costs and benefits into conservation decision-making processes can result in missed opportunities for diversifying outcomes and creating cost-sharing multisectoral partnerships. We devised a straightforward and readily interpretable approach to incorporate cobenefits into a threat-management prioritization approach. We used it to analyze the agricultural cobenefits of implementing 9 invasive animal management strategies designed to ensure the persistence of 148 threatened species across Australia's Lake Eyre Basin over 50 years. A structured elicitation process with 24 participants (scientists, land managers, agriculturalists, and other stakeholders) was used to collect information on each strategy, including costs, technical and social feasibility, benefits to native threatened species, and cobenefits to agricultural production systems. The costs of targeted invasive animal management to save threatened species across the basin (AU$33 million/year) outweighed the overall benefits to the agricultural industry (estimated AU$226 million/year). The return on investment for these management strategies varied substantially when agricultural cobenefits were considered alongside threatened species benefits and showed synergies and challenges. Our approach demonstrates the value of incorporating cobenefits of conservation actions into cost-effectiveness analyses to guide potential investment and partnerships and to diversify implementation pathways.

Evaluación Rápida de los Cobeneficios para Promover Alianzas de Manejo de Amenazas Resumen Las estrategias de conservación enfocadas en la reducción de las amenazas para la biodiversidad pueden tener implicaciones importantes para muchos sectores de un sistema socioeconómico, pero existe un entendimiento reducido de estos cobeneficios. Por ejemplo, muchas de las amenazas para las especies nativas también impiden la producción agrícola y a pesar de esto, comúnmente se percibe a la agricultura como una competencia para los objetivos de conservación. Aunque un análisis completo de decisiones con objetivos múltiples está usualmente más allá del enfoque y la capacidad del órgano decisorio, no incluir costos y beneficios socioeconómicos importantes dentro del proceso de toma de decisiones puede resultar en oportunidades perdidas para la diversificación de resultados y la creación de colaboraciones multisectoriales con reparto de costes. Diseñamos una estrategia directa y de fácil interpretación para incorporar los cobeneficios dentro de una estrategia de priorización de manejo de amenazas. Usamos esta estrategia para analizar los cobeneficios agrícolas de la implementación de nueve estrategias de manejo de animales invasores diseñadas para asegurar la persistencia de 148 especies amenazadas en la cuenca del Lago Eyre en Australia durante 50 años. Usamos un proceso estructurado de extracción con 24 participantes (científicos, administradores de tierras, agricultores y otros actores) para recolectar información sobre cada estrategia, incluyendo los costos, viabilidad técnica y social, beneficios para las especies nativas amenazadas y los cobeneficios para los sistemas de producción agrícola. Los costos del manejo enfocado en animales invasores para salvar a las especies amenazadas de la cuenca (AU$33 millones al año) superaron a los beneficios generales para la industria agrícola (estimados en AU$226 millones al año). El rendimiento de la inversión para estas estrategias de manejo varió sustancialmente cuando los cobeneficios agrícolas estuvieron considerados junto con los beneficios para las especies amenazadas y mostró retos y sinergias. Nuestra estrategia demuestra la importancia de la incorporación de los cobeneficios de las acciones de conservación dentro de los análisis de rentabilidad para guiar la inversión potencial y las alianzas y para diversificar las vías de implementación.

Author Correction: Leaf nutrients, not specific leaf area, are consistent indicators of elevated nutrient inputs.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Nutrient availability controls the impact of mammalian herbivores on soil carbon and nitrogen pools in grasslands.

Grasslands are subject to considerable alteration due to human activities globally, including widespread changes in populations and composition of large mammalian herbivores and elevated supply of nutrients. Grassland soils remain important reservoirs of carbon (C) and nitrogen (N). Herbivores may affect both C and N pools and these changes likely interact with increases in soil nutrient availability. Given the scale of grassland soil fluxes, such changes can have striking consequences for atmospheric C concentrations and the climate. Here, we use the Nutrient Network experiment to examine the responses of soil C and N pools to mammalian herbivore exclusion across 22 grasslands, under ambient and elevated nutrient availabilities (fertilized with NPK + micronutrients). We show that the impact of herbivore exclusion on soil C and N pools depends on fertilization. Under ambient nutrient conditions, we observed no effect of herbivore exclusion, but under elevated nutrient supply, pools are smaller upon herbivore exclusion. The highest mean soil C and N pools were found in grazed and fertilized plots. The decrease in soil C and N upon herbivore exclusion in combination with fertilization correlated with a decrease in aboveground plant biomass and microbial activity, indicating a reduced storage of organic matter and microbial residues as soil C and N. The response of soil C and N pools to herbivore exclusion was contingent on temperature - herbivores likely cause losses of C and N in colder sites and increases in warmer sites. Additionally, grasslands that contain mammalian herbivores have the potential to sequester more N under increased temperature variability and nutrient enrichment than ungrazed grasslands. Our study highlights the importance of conserving mammalian herbivore populations in grasslands worldwide. We need to incorporate local-scale herbivory, and its interaction with nutrient enrichment and climate, within global-scale models to better predict land-atmosphere interactions under future climate change.

Global change effects on plant communities are magnified by time and the number of global change factors imposed.

Global change drivers (GCDs) are expected to alter community structure and consequently, the services that ecosystems provide. Yet, few experimental investigations have examined effects of GCDs on plant community structure across multiple ecosystem types, and those that do exist present conflicting patterns. In an unprecedented global synthesis of over 100 experiments that manipulated factors linked to GCDs, we show that herbaceous plant community responses depend on experimental manipulation length and number of factors manipulated. We found that plant communities are fairly resistant to experimentally manipulated GCDs in the short term (<10 y). In contrast, long-term (≥10 y) experiments show increasing community divergence of treatments from control conditions. Surprisingly, these community responses occurred with similar frequency across the GCD types manipulated in our database. However, community responses were more common when 3 or more GCDs were simultaneously manipulated, suggesting the emergence of additive or synergistic effects of multiple drivers, particularly over long time periods. In half of the cases, GCD manipulations caused a difference in community composition without a corresponding species richness difference, indicating that species reordering or replacement is an important mechanism of community responses to GCDs and should be given greater consideration when examining consequences of GCDs for the biodiversity-ecosystem function relationship. Human activities are currently driving unparalleled global changes worldwide. Our analyses provide the most comprehensive evidence to date that these human activities may have widespread impacts on plant community composition globally, which will increase in frequency over time and be greater in areas where communities face multiple GCDs simultaneously.