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Clonal perennial grasses are the dominant species in almost all natural grasslands, however their seed production is typically low. The reasons why seed set is so low remains unclear. We studied a rhizomatous grass (Leymus chinensis) using 13 C tracing the different photosynthetic organs to investigate carbon fixation and allocation during the seed-filling stage. We found that the vegetative ramet leaves are the largest (81%) source for total plant fixed carbon, whereas almost all carbon is allocated to vegetative reproduction. The spike is the largest (54%) carbon source for the seeds. However, the spike produced carbon only allocated 37% to the seeds, with the majority allocated to vegetative reproduction. This preferential carbon allocation to vegetative reproduction limits sexual reproduction. Nitrogen application significantly increased assimilated carbon. However, nearly all increased carbon accumulated in the vegetative reproduction rather than in the seeds. Only the carbon produced by the spike increased its allocation to the seeds by 13%. Taken together, we conclude that the predominance of vegetative reproduction, combined with self-incompatibility, results in low ovule fertilization and very weak seed sink strength for carbon competition, suggests that the weak seed sink strength is the key reason causing low seed set in L. chinensis., (© 2024 John Wiley & Sons Ltd.)

Plants display a range of temporal patterns of inter-annual reproduction, from relatively constant seed production to "mast seeding," the synchronized and highly variable interannual seed production of plants within a population. Previous efforts have compiled global records of seed production in long-lived plants to gain insight into seed production, forest and animal population dynamics, and the effects of global change on masting. Existing datasets focus on seed production dynamics at the population scale but are limited in their ability to examine community-level mast seeding dynamics across different plant species at the continental scale. We harmonized decades of plant reproduction data for 141 woody plant species across nine Long-Term Ecological Research (LTER) or long-term ecological monitoring sites from a wide range of habitats across the United States. Plant reproduction data are reported annually between 1957 and 2021 and based on either seed traps or seed and/or cone counts on individual trees. A wide range of woody plant species including trees, shrubs, and lianas are represented within sites allowing for direct community-level comparisons among species. We share code for filtering of data that enables the comparison of plot and individual tree data across sites. For each species, we compiled relevant life history attributes (e.g., seed mass, dispersal syndrome, seed longevity, sexual system) that may serve as important predictors of mast seeding in future analyses. To aid in phylogenetically informed analyses, we also share a phylogeny and phylogenetic distance matrix for all species in the dataset. These data can be used to investigate continent-scale ecological properties of seed production, including individual and population variability, synchrony within and across species, and how these properties of seed production vary in relation to plant species traits and environmental conditions. In addition, these data can be used to assess how annual variability in seed production is associated with climate conditions and how that varies across populations, species, and regions. The dataset is released under a CC0 1.0 Universal public domain license., (© 2024 The Author(s). Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Global change is associated with variable shifts in the annual production of aboveground plant biomass, suggesting localized sensitivities with unclear causal origins. Combining remotely sensed normalized difference vegetation index data since the 1980s with contemporary field data from 84 grasslands on 6 continents, we show a widening divergence in site-level biomass ranging from +51% to -34% globally. Biomass generally increased in warmer, wetter and species-rich sites with longer growing seasons and declined in species-poor arid areas. Phenological changes were widespread, revealing substantive transitions in grassland seasonal cycling. Grazing, nitrogen deposition and plant invasion were prevalent in some regions but did not predict overall trends. Grasslands are undergoing sizable changes in production, with implications for food security, biodiversity and carbon storage especially in arid regions where declines are accelerating., (© 2024. The Author(s), under exclusive licence to Springer Nature Limited.)

The fundamental trade-off between current and future reproduction has long been considered to result in a tendency for species that can grow large to begin reproduction at a larger size. Due to the prolonged time required to reach maturity, estimates of tree maturation size remain very rare and we lack a global view on the generality and the shape of this trade-off. Using seed production from five continents, we estimate tree maturation sizes for 486 tree species spanning tropical to boreal climates. Results show that a species' maturation size increases with maximum size, but in a non-proportional way: the largest species begin reproduction at smaller sizes than would be expected if maturation were simply proportional to maximum size. Furthermore, the decrease in relative maturation size is steepest in cold climates. These findings on maturation size drivers are key to accurately represent forests' responses to disturbance and climate change., (© 2024 John Wiley & Sons Ltd.)

Alien plant species regularly and simultaneously invade agricultural landscapes and ecosystems; however, the effects of co-invasion on crop production and native biodiversity have rarely been studied. Secondary metabolites produced by alien plants may be allelopathic; if they enter the soil, they may be transported by agricultural activities, negatively affecting crop yield and biodiversity. It is unknown whether substances from different alien species in combination have a greater impact on crops and wild plants than if they are from only one of the alien species. In this study, we used a set of common garden experiments to test the hypothesis that mixed extracts from two common invasive species have synergistic effects on crops and weeds (defined as all non-crop plants) in European agricultural fields compared to single-species extracts. We found that both the combined and individual extracts had detrimental effects on the seed germination, seedling growth, biomass, and photosynthetic performance of both crops and weeds. We found that the negative effect of mixed extracts was not additive and that crop plants were more strongly affected by invasive species extracts than the weeds. Our results are important for managing invasive species in unique ecosystems on agricultural land and preventing economic losses in yield production., (© 2023. The Author(s).)

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., (© 2023. The Author(s).)

Little is currently known about how climate modulates the relationship between plant diversity and soil organic carbon and the mechanisms involved. Yet, this knowledge is of crucial importance in times of climate change and biodiversity loss. Here, we show that plant diversity is positively correlated with soil carbon content and soil carbon-to-nitrogen ratio across 84 grasslands on six continents that span wide climate gradients. The relationships between plant diversity and soil carbon as well as plant diversity and soil organic matter quality (carbon-to-nitrogen ratio) are particularly strong in warm and arid climates. While plant biomass is positively correlated with soil carbon, plant biomass is not significantly correlated with plant diversity. Our results indicate that plant diversity influences soil carbon storage not via the quantity of organic matter (plant biomass) inputs to soil, but through the quality of organic matter. The study implies that ecosystem management that restores plant diversity likely enhances soil carbon sequestration, particularly in warm and arid climates., (© 2023. Springer Nature Limited.)

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., (© 2023. Springer Nature Limited.)

Over the last 40 years, a burrowing mammal eradication policy has been prevalent on the Qinghai-Tibetan Plateau (QTP). This policy is based on similar burrowing mammal eradication programs in other areas and is justified on the assumptions that burrowing mammals compete with livestock for forage and contribute to grassland degradation. However, there is no clear theoretical or experimental evidence supporting these assumptions. This paper synthesizes the ecological functioning of small burrowing mammals in natural grasslands and discusses the irrationality and consequences of burrowing mammal eradication for sustainable livestock grazing and grassland degradation. Past burrowing mammal eradication efforts have failed because increased food availability for the remaining rodents and reduced predator populations led to rapid population rebounds. Herbivores differ in diet, and there is clear evidence that burrowing mammals, especially plateau zokors Myospalax baileyi, have a different diet than livestock. In QTP meadows, burrowing mammal eradication induces a shift towards plant communities with fewer species preferred by livestock and more species preferred by burrowing mammals. Thus, eradicating burrowing mammals has the opposite effect, a reduction in livestock preferred vegetation. We suggest that the policy of poisoning burrowing mammals needs to be reconsidered and revoked as soon as possible. We argue that incorporating density-dependent factors such as predation and food availability are essential for maintaining a low burrowing mammal density. For degraded grasslands, we suggest that the optimal sustainable approach is to decrease the intensity of livestock grazing. Lower grazing induces changes in vegetation structure and plant species composition that increases predation on burrowing mammals and decreases the abundance of plants preferred by burrowing mammals. Such a nature-based grassland management system maintains the density of burrowing mammals at a low stable density while minimizing human management and interventions., (© 2023 The Authors. Journal of Animal Ecology © 2023 British Ecological Society.)

The benefits of masting (volatile, quasi-synchronous seed production at lagged intervals) include satiation of seed predators, but these benefits come with a cost to mutualist pollen and seed dispersers. If the evolution of masting represents a balance between these benefits and costs, we expect mast avoidance in species that are heavily reliant on mutualist dispersers. These effects play out in the context of variable climate and site fertility among species that vary widely in nutrient demand. Meta-analyses of published data have focused on variation at the population scale, thus omitting periodicity within trees and synchronicity between trees. From raw data on 12 million tree-years worldwide, we quantified three components of masting that have not previously been analysed together: (i) volatility, defined as the frequency-weighted year-to-year variation; (ii) periodicity, representing the lag between high-seed years; and (iii) synchronicity, indicating the tree-to-tree correlation. Results show that mast avoidance (low volatility and low synchronicity) by species dependent on mutualist dispersers explains more variation than any other effect. Nutrient-demanding species have low volatility, and species that are most common on nutrient-rich and warm/wet sites exhibit short periods. The prevalence of masting in cold/dry sites coincides with climatic conditions where dependence on vertebrate dispersers is less common than in the wet tropics. Mutualist dispersers neutralize the benefits of masting for predator satiation, further balancing the effects of climate, site fertility and nutrient demands., (© 2023. The Author(s), under exclusive licence to Springer Nature Limited.)

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., (© 2023. The Author(s).)

Causal effects of biodiversity on ecosystem functions can be estimated using experimental or observational designs - designs that pose a tradeoff between drawing credible causal inferences from correlations and drawing generalizable inferences. Here, we develop a design that reduces this tradeoff and revisits the question of how plant species diversity affects productivity. Our design leverages longitudinal data from 43 grasslands in 11 countries and approaches borrowed from fields outside of ecology to draw causal inferences from observational data. Contrary to many prior studies, we estimate that increases in plot-level species richness caused productivity to decline: a 10% increase in richness decreased productivity by 2.4%, 95% CI [-4.1, -0.74]. This contradiction stems from two sources. First, prior observational studies incompletely control for confounding factors. Second, most experiments plant fewer rare and non-native species than exist in nature. Although increases in native, dominant species increased productivity, increases in rare and non-native species decreased productivity, making the average effect negative in our study. By reducing the tradeoff between experimental and observational designs, our study demonstrates how observational studies can complement prior ecological experiments and inform future ones., (© 2023. The Author(s).)

Plant productivity varies due to environmental heterogeneity, and theory suggests that plant diversity can reduce this variation. While there is strong evidence of diversity effects on temporal variability of productivity, whether this mechanism extends to variability across space remains elusive. Here we determine the relationship between plant diversity and spatial variability of productivity in 83 grasslands, and quantify the effect of experimentally increased spatial heterogeneity in environmental conditions on this relationship. We found that communities with higher plant species richness (alpha and gamma diversity) have lower spatial variability of productivity as reduced abundance of some species can be compensated for by increased abundance of other species. In contrast, high species dissimilarity among local communities (beta diversity) is positively associated with spatial variability of productivity, suggesting that changes in species composition can scale up to affect productivity. Experimentally increased spatial environmental heterogeneity weakens the effect of plant alpha and gamma diversity, and reveals that beta diversity can simultaneously decrease and increase spatial variability of productivity. Our findings unveil the generality of the diversity-stability theory across space, and suggest that reduced local diversity and biotic homogenization can affect the spatial reliability of key ecosystem functions., (© 2023. The Author(s).)

Increased nutrient inputs due to anthropogenic activity are expected to increase primary productivity across terrestrial ecosystems, but changes in allocation aboveground versus belowground with nutrient addition have different implications for soil carbon (C) storage. Thus, given that roots are major contributors to soil C storage, understanding belowground net primary productivity (BNPP) and biomass responses to changes in nutrient availability is essential to predicting carbon-climate feedbacks in the context of interacting global environmental changes. To address this knowledge gap, we tested whether a decade of nitrogen (N) and phosphorus (P) fertilization consistently influenced aboveground and belowground biomass and productivity at nine grassland sites spanning a wide range of climatic and edaphic conditions in the continental United States. Fertilization effects were strong aboveground, with both N and P addition stimulating aboveground biomass at nearly all sites (by 30% and 36%, respectively, on average). P addition consistently increased root production (by 15% on average), whereas other belowground responses to fertilization were more variable, ranging from positive to negative across sites. Site-specific responses to P were not predicted by the measured covariates. Atmospheric N deposition mediated the effect of N fertilization on root biomass and turnover. Specifically, atmospheric N deposition was positively correlated with root turnover rates, and this relationship was amplified with N addition. Nitrogen addition increased root biomass at sites with low N deposition but decreased it at sites with high N deposition. Overall, these results suggest that the effects of nutrient supply on belowground plant properties are context dependent, particularly with regard to background N supply rates, demonstrating that site conditions must be considered when predicting how grassland ecosystems will respond to increased nutrient loading from anthropogenic activity., (© 2022 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Effective application of functional trait approaches to ecological questions requires understanding the patterns of trait variation within species as well as between them. However, few studies address the potential for intraspecific variation to occur on a temporal basis and, thus, for trait-based findings to be contingent upon sampling year. To quantify annual variation in the functional traits of grassland plant species, we measured specific leaf area, leaf dry matter content, plant height, and chlorophyll content in 12 shortgrass prairie plant species. We repeated these measurements across 4 years, both in long-term nitrogen addition plots and in corresponding control plots. Three of the four traits showed significant year-to-year variation in a linear mixed model analysis, generally following a pattern of more acquisitive leaf economics spectrum traits in higher rainfall years. Furthermore, two of the measured traits responded interactively to nitrogen addition and sampling year, although only one, leaf dry matter content, showed the expected pattern of stronger nitrogen responses in high rainfall years. For leaf dry matter content and specific leaf area, trait responses to sampling year were larger than responses to the nitrogen addition treatment. These findings illustrate that species' functional traits can respond strongly to environmental changes across years, and thus that trait variation in a species or community is likely to extend beyond the values and patterns observed in any single year., (© 2022 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Ontogenic changes in soybean radiation use efficiency (RUE) have been attributed to variation in specific leaf nitrogen (SLN) based only on data collected during seed filling. We evaluated this hypothesis using data on leaf area, absorbed radiation (ARAD), aboveground dry matter (ADM), and plant nitrogen (N) concentration collected during the entire crop season from seven field experiments conducted in a stress-free environment. Each experiment included a full-N treatment that received ample N fertilizer and a zero-N treatment that relied on N fixation and soil N mineralization. We estimated RUE based on changes in ADM between sampling times and associated ARAD, accounting for changes in biomass composition. The RUE and SLN exhibited different seasonal patterns: a bell-shaped pattern with a peak around the beginning of seed filling, and a convex pattern followed by an abrupt decline during late seed filling, respectively. Changes in SLN explained the decline in RUE during seed filling but failed to predict changes in RUE in earlier stages and underestimated the maximum RUE observed during pod setting. Comparison between observed and simulated RUE using a process-based crop simulation model revealed similar discrepancies. The decoupling between RUE and SLN during early crop stages suggests that leaf N is above that needed to maximize crop growth but may play a role in storing N that can be used in later reproductive stages to meet the large seed N demand associated with high-yielding crops., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Organic farming supports higher biodiversity than conventional farming, but at the cost of lower yields. We conducted a meta-analysis quantifying the trade-off between biodiversity and yield, comparing conventional and organic farming. We developed a compatibility index to assess whether biodiversity gains from organic farming exceed yield losses, and a substitution index to assess whether organic farming would increase biodiversity in an area if maintaining total production under organic farming would require cultivating more land at the expense of nature. Overall, organic farming had 23% gain in biodiversity with a similar cost of yield decline. Biodiversity gain is negatively correlated to yield loss for microbes and plants, but no correlation was found for other taxa. The biodiversity and yield trade-off varies under different contexts of organic farming. The overall compatibility index value was close to zero, with negative values for cereal crops, positive for non-cereal crops, and varies across taxa. Our results indicate that, on average, the proportion of biodiversity gain is similar to the proportion of yield loss for paired field studies. For some taxa in non-cereal crops, switching to organic farming can lead to a biodiversity gain without yield loss. We calculated the overall value of substitution index and further discussed the application of this index to evaluate when the biodiversity of less intensified farming system is advantageous., (© 2022 John Wiley & Sons Ltd.)

The relationships that control seed production in trees are fundamental to understanding the evolution of forest species and their capacity to recover from increasing losses to drought, fire, and harvest. A synthesis of fecundity data from 714 species worldwide allowed us to examine hypotheses that are central to quantifying reproduction, a foundation for assessing fitness in forest trees. Four major findings emerged. First, seed production is not constrained by a strict trade-off between seed size and numbers. Instead, seed numbers vary over ten orders of magnitude, with species that invest in large seeds producing more seeds than expected from the 1:1 trade-off. Second, gymnosperms have lower seed production than angiosperms, potentially due to their extra investments in protective woody cones. Third, nutrient-demanding species, indicated by high foliar phosphorus concentrations, have low seed production. Finally, sensitivity of individual species to soil fertility varies widely, limiting the response of community seed production to fertility gradients. In combination, these findings can inform models of forest response that need to incorporate reproductive potential., (© 2022. The Author(s).)

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., (© 2022 John Wiley & Sons Ltd.)

Despite the long history of the study of the biodiversity-ecosystem function relationship, uncertainty remains about the relationship of natural grassland ecosystems under stressful conditions. Recently, trait- and phylogenetic-based tests provide a powerful way to detect the relationship in different spaces but have seldom been applied to stressful zones on a large spatial scale. We selected Qinghai-Tibetan as the study area and collected a grassland community database involving 581 communities. We calculated biomass and species', functional, and phylogenetic diversity of each community and examined their relationships by using linear and non-linear regression models. Results showed an overall positive biodiversity-productivity relationship in species', functional and phylogenetic space. The relationship, however, was non-linear, in which biodiversity explained better the variation in community biomass when species diversity was more than a threshold, showing a weak effect of biodiversity on ecosystem function in low species diversity communities. We also found a filled triangle for the limit of the relationship between species and functional diversity, implying that functional diversity differs significantly among communities when their species diversity is low but finally converges to be a constant with increasing communities' species diversity. Our study suggests that multiple niche processes may structure the grassland communities, and their forces tend to balance in high-biodiversity communities., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Qi, Kang, Knops, Jiang, Abuman and Du.)

Tree fecundity and recruitment have not yet been quantified at scales needed to anticipate biogeographic shifts in response to climate change. By separating their responses, this study shows coherence across species and communities, offering the strongest support to date that migration is in progress with regional limitations on rates. The southeastern continent emerges as a fecundity hotspot, but it is situated south of population centers where high seed production could contribute to poleward population spread. By contrast, seedling success is highest in the West and North, serving to partially offset limited seed production near poleward frontiers. The evidence of fecundity and recruitment control on tree migration can inform conservation planning for the expected long-term disequilibrium between climate and forest distribution., Competing Interests: Competing interest statement: M.B., W.D.K., and I.S.P. are collaborators on a 2020 review with T.W., (Copyright © 2021 the Author(s). Published by PNAS.)

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., (© 2021 John Wiley & Sons Ltd.)

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., Competing Interests: None declared., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

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., (© 2020 by the Ecological Society of America.)

Biological invasions are a major human induced global change that is threatening global biodiversity by homogenizing the world's fauna and flora. Species spread because humans have moved species across geographical boundaries and have changed ecological factors that structure ecosystems, such as nitrogen deposition, disturbance, etc. Many biological invasions are caused accidentally, as a byproduct of human travel and commerce driven product shipping. However, humans also have spread many species intentionally because of perceived benefits. Of interest is the role of the recent exponential growth in information exchange via internet social media in driving biological invasions. To date, this has not been examined. Here, we show that for one such invasive species, goldenrod, social networks spread misleading and incomplete information that is enhancing the spread of goldenrod invasions into new environments. We show that the notion of goldenrod honey as a "superfood" with unsupported healing properties is driving a demand that leads beekeepers to produce goldenrod honey. Social networks provide a forum for such information exchange and this is leading to further spread of goldenrod in many countries where goldenrod is not native, such as Poland. However, this informal social information exchange ignores laws that focus on preventing the further spread of invasive species and the strong negative effects that goldenrod has on native ecosystems, including floral resources that negatively impact honeybee performance. Thus, scientifically unsupported information on "superfoods" such as goldenrod honey that is disseminated through social internet networks has real world consequences such as increased goldenrod invasions into novel geographical regions which decreases native biodiversity., (© 2020 John Wiley & Sons Ltd.)

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 N min ), 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 N min 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 N min . Herbivore removal in the absence of fertilization did not alter potential and realized soil net N min . However, fertilization alone and in combination with herbivore removal consistently increased potential soil net N min. Realized soil net N min , in contrast, significantly decreased in fertilized plots where herbivores were removed. Treatment effects on potential and realized soil net N min were contingent on site-specific soil and climatic properties. Fertilization effects on potential soil net N min were larger at sites with higher mean annual precipitation (MAP) and temperature of the wettest quarter (T.q.wet). Reciprocally, realized soil net N min 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 N min across global grasslands under realistic field conditions. This is an important context-dependent knowledge for grassland management worldwide., (© 2020 John Wiley & Sons Ltd.)

Degradation of permafrost with a thin overlying active layer can greatly affect vegetation via changes in the soil water and nutrient regimes within the active layer, while little is known about the presence or absence of such effects in areas with a deep active layer. Here, we selected the northeastern Qinghai-Tibet Plateau as the study area. We examined the vegetation communities and biomass along an active layer thickness (ALT) gradient from 0.6 to 3.5 m. Our results showed that plant cover, below-ground biomass, species richness, and relative sedge cover declined with the deepening active layer, while the evenness, and relative forb cover showed a contrary trend. The vegetation indices and the dissimilarity of vegetation composition exhibited significant changes when the ALT was greater than 2.0 m. The vegetation indices (plant cover, below-ground biomass, evenness index, relative forb cover and relative sedge cover) were closely associated with soil water content, soil pH, texture and nutrient content. Soil water content played a key role in the ALT-vegetation relationship, especially at depths of 30-40 cm. Our results suggest that when the ALT is greater than 2.0 m, the presence of underlying permafrost still benefits vegetation growth via maintaining adequate soil water contents at 30-40 cm depth. Furthermore, the degradation of permafrost may lead to declines of vegetation cover and below-ground biomass with a shift in vegetation species.

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

Nitrogen (N) supply can limit the yields of soybean [Glycine max (L.) Merr.] in highly productive environments. To explore the physiological mechanisms underlying this limitation, seasonal changes in N dynamics, aboveground dry matter (ADM) accumulation, leaf area index (LAI) and fraction of absorbed radiation (fAPAR) were compared in crops relying only on biological N 2 fixation and available soil N (zero-N treatment) versus crops receiving N fertilizer (full-N treatment). Experiments were conducted in seven high-yield environments without water limitation, where crops received optimal management. In the zero-N treatment, biological N 2 fixation was not sufficient to meet the N demand of the growing crop from early in the season up to beginning of seed filling. As a result, crop LAI, growth, N accumulation, radiation-use efficiency and fAPAR were consistently higher in the full-N than in the zero-N treatment, leading to improved seed set and yield. Similarly, plants in the full-N treatment had heavier seeds with higher N concentration because of greater N mobilization from vegetative organs to seeds. Future yield gains in high-yield soybean production systems will require an increase in biological N 2 fixation, greater supply of N from soil or fertilizer, or alleviation of the trade-off between these two sources of N in order to meet the plant demand., (© 2020 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Grasslands worldwide are expected to experience an increase in extreme events such as drought, along with simultaneous increases in mineral nutrient inputs as a result of human industrial activities. These changes are likely to interact because elevated nutrient inputs may alter plant diversity and increase the sensitivity to droughts. Dividing a system's sensitivity to drought into resistance to change during the drought and rate of recovery after the drought generates insights into different dimensions of the system's resilience in the face of drought. Here, we examine the effects of experimental nutrient fertilization and the resulting diversity loss on the resistance to and recovery from severe regional droughts. We do this at 13 North American sites spanning gradients of aridity, five annual grasslands in California, and eight perennial grasslands in the Great Plains. We measured rate of resistance as the change in annual aboveground biomass (ANPP) per unit change in growing season precipitation as conditions declined from normal to drought. We measured recovery as the change in ANPP during the postdrought period and the return to normal precipitation. Resistance and recovery did not vary across the 400-mm range of mean growing season precipitation spanned by our sites in the Great Plains. However, chronic nutrient fertilization in the Great Plains reduced drought resistance and increased drought recovery. In the California annual grasslands, arid sites had a greater recovery postdrought than mesic sites, and nutrient addition had no consistent effects on resistance or recovery. Across all study sites, we found that predrought species richness in natural grasslands was not consistently associated with rates of resistance to or recovery from the drought, in contrast to earlier findings from experimentally assembled grassland communities. Taken together, these results suggest that human-induced eutrophication may destabilize grassland primary production, but the effects of this may vary across regions and flora, especially between perennial and annual-dominated grasslands., (Published 2020. This article has been contributed to by US Government employees and their work is in the public domain in the USA.)

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., (© 2020 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Background: The Eurasian steppe is an important vegetation type characterized by cold, arid and nitrogen poor conditions. At the Eastern edge, including in the Songnen grassland, the vegetation is dominated by Leymus chinensis (henceforth L. chinensis) and is increasing threatened by elevated anthropogenic nitrogen deposition. L. chinensis is a perennial grass that mainly reproduces vegetatively and its sexual reproduction is limited. However, sexual reproduction plays an important role influencing colonization after large disturbances. To develop an understanding of how elevated nitrogen deposition changes the plant community structure and functioning we need a better understanding how sexual reproduction of L. chinensis changes with nitrogen enrichment. Here we report on a field experiment where we added 10 g N m - 2  yr - 1 and examined changes in seed traits, seed germination and early seedling growth., Results: Nitrogen addition increased seed production by 79%, contributing to this seed increases were a 28% increase in flowering plant density, a 40% increase in seed number per plant and a 11% increase in seed weight. Seed size increased with a 42% increase in large seeds and a 49% decrease in the smallest seed size category. Seed germination success improved by 10% for small seeds and 18% for large seeds. Combined, the increased in seed production and improved seed quality doubled the potential seed germination. Subsequent seedling above and below-ground biomass also significantly increased., Conclusions: All aspects of L. chinensis sexual reproduction increased with nitrogen addition. Thus, L. chinensis competitive ability may increase when atmospheric nitrogen deposition increases, which may further reduce overall plant diversity in the low diversity Songnen grasslands.

Plant traits-the morphological, anatomical, physiological, biochemical and phenological characteristics of plants-determine how plants respond to environmental factors, affect other trophic levels, and influence ecosystem properties and their benefits and detriments to people. Plant trait data thus represent the basis for a vast area of research spanning from evolutionary biology, community and functional ecology, to biodiversity conservation, ecosystem and landscape management, restoration, biogeography and earth system modelling. Since its foundation in 2007, the TRY database of plant traits has grown continuously. It now provides unprecedented data coverage under an open access data policy and is the main plant trait database used by the research community worldwide. Increasingly, the TRY database also supports new frontiers of trait-based plant research, including the identification of data gaps and the subsequent mobilization or measurement of new data. To support this development, in this article we evaluate the extent of the trait data compiled in TRY and analyse emerging patterns of data coverage and representativeness. Best species coverage is achieved for categorical traits-almost complete coverage for 'plant growth form'. However, most traits relevant for ecology and vegetation modelling are characterized by continuous intraspecific variation and trait-environmental relationships. These traits have to be measured on individual plants in their respective environment. Despite unprecedented data coverage, we observe a humbling lack of completeness and representativeness of these continuous traits in many aspects. We, therefore, conclude that reducing data gaps and biases in the TRY database remains a key challenge and requires a coordinated approach to data mobilization and trait measurements. This can only be achieved in collaboration with other initiatives., (© 2019 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Soil nitrogen mineralisation (N min ), the conversion of organic into inorganic N, is important for productivity and nutrient cycling. The balance between mineralisation and immobilisation (net N min ) varies with soil properties and climate. However, because most global-scale assessments of net N min are laboratory-based, its regulation under field-conditions and implications for real-world soil functioning remain uncertain. Here, we explore the drivers of realised (field) and potential (laboratory) soil net N min across 30 grasslands worldwide. We find that realised N min is largely explained by temperature of the wettest quarter, microbial biomass, clay content and bulk density. Potential N min only weakly correlates with realised N min , but contributes to explain realised net N min when combined with soil and climatic variables. We provide novel insights of global realised soil net N min and show that potential soil net N min data available in the literature could be parameterised with soil and climate data to better predict realised N min .