Cumulative nitrogen enrichment alters the drivers of grassland overyielding.

Effects of plant diversity on grassland productivity, or overyielding, are found to be robust to nutrient enrichment. However, the impact of cumulative nitrogen (N) addition (total N added over time) on overyielding and its drivers are underexplored. Synthesizing data from 15 multi-year grassland biodiversity experiments with N addition, we found that N addition decreases complementarity effects and increases selection effects proportionately, resulting in no overall change in overyielding regardless of N addition rate. However, we observed a convex relationship between overyielding and cumulative N addition, driven by a shift from complementarity to selection effects. This shift suggests diminishing positive interactions and an increasing contribution of a few dominant species with increasing N accumulation. Recognizing the importance of cumulative N addition is vital for understanding its impacts on grassland overyielding, contributing essential insights for biodiversity conservation and ecosystem resilience in the face of increasing N deposition.

Biodiversity: Net primary productivity relationships are eliminated by invasive species dominance.

Experiments often find that net primary productivity (NPP) increases with species richness when native species are considered. However, relationships may be altered by exotic (non-native) species, which are hypothesized to reduce richness but increase productivity (i.e., 'invasion-diversity-productivity paradox'). We compared richness-NPP relationships using a comparison of exotic versus native-dominated sites across the central USA, and two experiments under common environments. Aboveground NPP was measured using peak biomass clipping in all three studies, and belowground NPP was measured in one study with root ingrowth cores using root-free soil. In all studies, there was a significantly positive relationship between NPP and richness across native species-dominated sites and plots, but no relationship across exotic-dominated ones. These results indicate that relationships between NPP and richness depend on whether native or exotic species are dominant, and that exotic species are 'breaking the rules', altering richness-productivity and richness-C stock relationships after invasion.

Experimental impacts of grazing on grassland biodiversity and function are explained by aridity.

Grazing by domestic herbivores is the most widespread land use on the planet, and also a major global change driver in grasslands. Yet, experimental evidence on the long-term impacts of livestock grazing on biodiversity and function is largely lacking. Here, we report results from a network of 10 experimental sites from paired grazed and ungrazed grasslands across an aridity gradient, including some of the largest remaining native grasslands on the planet. We show that aridity partly explains the responses of biodiversity and multifunctionality to long-term livestock grazing. Grazing greatly reduced biodiversity and multifunctionality in steppes with higher aridity, while had no effects in steppes with relatively lower aridity. Moreover, we found that long-term grazing further changed the capacity of above- and below-ground biodiversity to explain multifunctionality. Thus, while plant diversity was positively correlated with multifunctionality across grasslands with excluded livestock, soil biodiversity was positively correlated with multifunctionality across grazed grasslands. Together, our cross-site experiment reveals that the impacts of long-term grazing on biodiversity and function depend on aridity levels, with the more arid sites experiencing more negative impacts on biodiversity and ecosystem multifunctionality. We also highlight the fundamental importance of conserving soil biodiversity for protecting multifunctionality in widespread grazed grasslands.

Giving fair credits to efforts in science and policy.

International collaborations aim to solve global environmental issues. Academic work and science-policy interfaces are instrumental in this pursuit, although scholars often overlook their significance. There is a need for fair credit distribution, transparency, and diversity in academia and policy reports. Recognizing these factors can enhance inclusivity and equity, driving solutions.

Assessing the importance of species and their assemblages for the biodiversity-ecosystem multifunctionality relationship.

Biodiversity changes, such as decline in species richness and biotic homogenization, can have grave consequences for ecosystem functionality. Careful investigation of biodiversity-ecosystem multifunctionality linkages with due consideration of conceptual and technical challenges is required to make the knowledge practically useful in managing social-ecological systems. In this paper, we introduced different methods to assess perspectives regarding the issue of diversity-multifunctionality, including a possible multifunctional redundancy/uniqueness, and the influences of the number and identity of functions on multifunctionality. In particular, we aimed to align methods with detecting the mechanisms underpinning diversity-multifunctional relationships that are free from statistical biases. Based on a set of novel methods that excluded analytical biases resulting from differences in the number and identities of multiple functions considered, we found that a substantial portion of species disproportionately supported ecosystem functions and that the diversity effects on multifunctionality were more markedly observed when more functions were considered. These results jointly emphasize that individual species are, to some extent, both functionally unique as well as redundant, highlighting the complexity and necessity for managed assemblages to retain high levels of diversity. We also observed that the relative magnitude of uniqueness or redundancy can differ between species and functions and therefore should be defined in a multifunctional context. We further found that only a small subset of species was identified as significantly less important, especially at low levels of multifunctionality. Taken together, given the low level of multifunctional redundancy we identified, we stress that unraveling the hierarchical roles of biodiversity at different levels, such as individual species and their assemblages, should be a high research priority, in both theory and practice.

Nitrogen-Induced Hysteresis in Grassland Biodiversity: A Theoretical Test of Litter-Mediated Mechanisms.

AbstractThe global rise in anthropogenic reactive nitrogen and the negative impacts of N deposition on terrestrial plant diversity are well documented. The R* theory of resource competition predicts reversible decreases in plant diversity in response to N loading. However, empirical evidence for the reversibility of N-induced biodiversity loss is mixed. In a long-term N-enrichment experiment in Minnesota, a low-diversity state that emerged during N addition has persisted for decades after additions ceased. Hypothesized mechanisms preventing recovery of biodiversity include nutrient recycling, insufficient external seed supply, and litter inhibition of plant growth. Here, we present an ordinary differential equation model that unifies these mechanisms, produces bistability at intermediate N inputs, and qualitatively matches the observed hysteresis at Cedar Creek. Key features of the model, including native species' growth advantage in low-N conditions and limitation by litter accumulation, generalize from Cedar Creek to North American grasslands. Our results suggest that effective biodiversity restoration in these systems may require management beyond reducing N inputs, such as burning, grazing, haying, and seed additions. By coupling resource competition with an additional interspecific inhibitory process, the model also illustrates a general mechanism for bistability and hysteresis that may occur in multiple ecosystem types.

A Conceptual Framework to Integrate Biodiversity, Ecosystem Function, and Ecosystem Service Models.

Global biodiversity and ecosystem service models typically operate independently. Ecosystem service projections may therefore be overly optimistic because they do not always account for the role of biodiversity in maintaining ecological functions. We review models used in recent global model intercomparison projects and develop a novel model integration framework to more fully account for the role of biodiversity in ecosystem function, a key gap for linking biodiversity changes to ecosystem services. We propose two integration pathways. The first uses empirical data on biodiversity-ecosystem function relationships to bridge biodiversity and ecosystem function models and could currently be implemented globally for systems and taxa with sufficient data. We also propose a trait-based approach involving greater incorporation of biodiversity into ecosystem function models. Pursuing both approaches will provide greater insight into biodiversity and ecosystem services projections. Integrating biodiversity, ecosystem function, and ecosystem service modeling will enhance policy development to meet global sustainability goals.

Plant genetic diversity affects multiple trophic levels and trophic interactions.

Intraspecific genetic diversity is an important component of biodiversity. A substantial body of evidence has demonstrated positive effects of plant genetic diversity on plant performance. However, it has remained unclear whether plant genetic diversity generally increases plant performance by reducing the pressure of plant antagonists across trophic levels for different plant life forms, ecosystems and climatic zones. Here, we analyse 4702 effect sizes reported in 413 studies that consider effects of plant genetic diversity on trophic groups and their interactions. We found that that increasing plant genetic diversity decreased the performance of plant antagonists including invertebrate herbivores, weeds, plant-feeding nematodes and plant diseases, while increasing the performance of plants and natural enemies of herbivores. Structural equation modelling indicated that plant genetic diversity increased plant performance partly by reducing plant antagonist pressure. These results reveal that plant genetic diversity often influences multiple trophic levels in ways that enhance natural pest control in managed ecosystems and consumer control of plants in natural ecosystems for sustainable plant production.

Grand challenges in biodiversity-ecosystem functioning research in the era of science-policy platforms require explicit consideration of feedbacks.

Feedbacks are an essential feature of resilient socio-economic systems, yet the feedbacks between biodiversity, ecosystem services and human wellbeing are not fully accounted for in global policy efforts that consider future scenarios for human activities and their consequences for nature. Failure to integrate feedbacks in our knowledge frameworks exacerbates uncertainty in future projections and potentially prevents us from realizing the full benefits of actions we can take to enhance sustainability. We identify six scientific research challenges that, if addressed, could allow future policy, conservation and monitoring efforts to quantitatively account for ecosystem and societal consequences of biodiversity change. Placing feedbacks prominently in our frameworks would lead to (i) coordinated observation of biodiversity change, ecosystem functions and human actions, (ii) joint experiment and observation programmes, (iii) more effective use of emerging technologies in biodiversity science and policy, and (iv) a more inclusive and integrated global community of biodiversity observers. To meet these challenges, we outline a five-point action plan for collaboration and connection among scientists and policymakers that emphasizes diversity, inclusion and open access. Efforts to protect biodiversity require the best possible scientific understanding of human activities, biodiversity trends, ecosystem functions and-critically-the feedbacks among them.

Diversifying bioenergy crops increases yield and yield stability by reducing weed abundance.

Relationships between species diversity, productivity, temporal stability of productivity, and plant invasion have been well documented in grasslands, and these relationships could translate to improved agricultural sustainability. However, few studies have explored these relationships in agricultural contexts where fertility and weeds are managed. Using 7 years of biomass yield and species composition data from 12 species mixture treatments varying in native species diversity, we found that species richness increased yield and interannual yield stability by reducing weed abundance. Stability was driven by yield as opposed to temporal variability of yield. Nitrogen fertilization increased yield but at the expense of yield stability. We show how relationships between diversity, species asynchrony, invasion, productivity, and stability observed in natural grasslands can extend into managed agricultural systems. Increasing bioenergy crop diversity can improve farmer economics via increased yield, reduced yield variability, and reduced inputs for weed control, thus promoting perennial vegetation on agricultural lands.

Consistent functional clusters explain the effects of biodiversity on ecosystem productivity in a long-term experiment.

Biomass production in ecosystems is a complex process regulated by several facets of biodiversity and species identity, but also species interactions such as competition or complementarity between species. For studying these different facets separately, ecosystem biomass is generally partitioned in two biodiversity effects. The composition effect is a simple, linear effect, and the interaction effect is a more subtle, nonlinear effect. Here we used a clustering approach (1) to separately and comprehensively capture all linear and nonlinear effects induced by both biodiversity effects on ecosystem functioning, and (2) to determine the functional composition at the origin of each biodiversity effect. We used data from the long-term Cedar Creek BioDIV experiment carried out over 22 yr, and we partitioned multiplicatively the biomass in composition and interaction effects. Both biodiversity effects were weakly correlated. Our clustering approach accurately explains and predicts each diversity effect over time: each one is modeled by a different functional composition. Even if environmental conditions and the strength of interaction effect strongly varied over time, the functional clusters of species that govern the interaction effect do not change over the 22 yr of the experiment. The functional composition governing the interaction effect is therefore very robust. In contrast, the functional clusters of species that govern the composition effect are less robust and change with environmental conditions. Understanding ecosystem functioning therefore requires that ecological properties are first partitioned by type, then each type of property is analyzed and modeled separately. Approaches without a priori groupings of species, such as functional clustering, appear particularly efficient and robust to unravel the web of species interactions, and identify the role played by species on biodiversity effects.

Determinants of community compositional change are equally affected by global change.

Global change is impacting plant community composition, but the mechanisms underlying these changes are unclear. Using a dataset of 58 global change experiments, we tested the five fundamental mechanisms of community change: changes in evenness and richness, reordering, species gains and losses. We found 71% of communities were impacted by global change treatments, and 88% of communities that were exposed to two or more global change drivers were impacted. Further, all mechanisms of change were equally likely to be affected by global change treatments-species losses and changes in richness were just as common as species gains and reordering. We also found no evidence of a progression of community changes, for example, reordering and changes in evenness did not precede species gains and losses. We demonstrate that all processes underlying plant community composition changes are equally affected by treatments and often occur simultaneously, necessitating a wholistic approach to quantifying community changes.

Biodiversity as insurance: from concept to measurement and application.

Biological insurance theory predicts that, in a variable environment, aggregate ecosystem properties will vary less in more diverse communities because declines in the performance or abundance of some species or phenotypes will be offset, at least partly, by smoother declines or increases in others. During the past two decades, ecology has accumulated strong evidence for the stabilising effect of biodiversity on ecosystem functioning. As biological insurance is reaching the stage of a mature theory, it is critical to revisit and clarify its conceptual foundations to guide future developments, applications and measurements. In this review, we first clarify the connections between the insurance and portfolio concepts that have been used in ecology and the economic concepts that inspired them. Doing so points to gaps and mismatches between ecology and economics that could be filled profitably by new theoretical developments and new management applications. Second, we discuss some fundamental issues in biological insurance theory that have remained unnoticed so far and that emerge from some of its recent applications. In particular, we draw a clear distinction between the two effects embedded in biological insurance theory, i.e. the effects of biodiversity on the mean and variability of ecosystem properties. This distinction allows explicit consideration of trade-offs between the mean and stability of ecosystem processes and services. We also review applications of biological insurance theory in ecosystem management. Finally, we provide a synthetic conceptual framework that unifies the various approaches across disciplines, and we suggest new ways in which biological insurance theory could be extended to address new issues in ecology and ecosystem management. Exciting future challenges include linking the effects of biodiversity on ecosystem functioning and stability, incorporating multiple functions and feedbacks, developing new approaches to partition biodiversity effects across scales, extending biological insurance theory to complex interaction networks, and developing new applications to biodiversity and ecosystem management.

Disturbance type and species life history predict mammal responses to humans.

Human activity and land use change impact every landscape on Earth, driving declines in many animal species while benefiting others. Species ecological and life history traits may predict success in human-dominated landscapes such that only species with "winning" combinations of traits will persist in disturbed environments. However, this link between species traits and successful coexistence with humans remains obscured by the complexity of anthropogenic disturbances and variability among study systems. We compiled detection data for 24 mammal species from 61 populations across North America to quantify the effects of (1) the direct presence of people and (2) the human footprint (landscape modification) on mammal occurrence and activity levels. Thirty-three percent of mammal species exhibited a net negative response (i.e., reduced occurrence or activity) to increasing human presence and/or footprint across populations, whereas 58% of species were positively associated with increasing disturbance. However, apparent benefits of human presence and footprint tended to decrease or disappear at higher disturbance levels, indicative of thresholds in mammal species' capacity to tolerate disturbance or exploit human-dominated landscapes. Species ecological and life history traits were strong predictors of their responses to human footprint, with increasing footprint favoring smaller, less carnivorous, faster-reproducing species. The positive and negative effects of human presence were distributed more randomly with respect to species trait values, with apparent winners and losers across a range of body sizes and dietary guilds. Differential responses by some species to human presence and human footprint highlight the importance of considering these two forms of human disturbance separately when estimating anthropogenic impacts on wildlife. Our approach provides insights into the complex mechanisms through which human activities shape mammal communities globally, revealing the drivers of the loss of larger predators in human-modified landscapes.

How complementarity and selection affect the relationship between ecosystem functioning and stability.

The biotic mechanisms underlying ecosystem functioning and stability have been extensively-but separately-explored in the literature, making it difficult to understand the relationship between functioning and stability. In this study, we used community models to examine how complementarity and selection, the two major biodiversity mechanisms known to enhance ecosystem biomass production, affect ecosystem stability. Our analytic and simulation results show that although complementarity promotes stability, selection impairs it. The negative effects of selection on stability operate through weakening portfolio effects and selecting species that have high productivity but low tolerance to perturbations ("risk-prone" species). In contrast, complementarity enhances stability by increasing portfolio effects and reducing the relative abundance of risk-prone species. Consequently, ecosystem functioning and stability exhibit either a synergy, if complementarity effects prevail, or trade-off, if selection effects prevail. Across species richness levels, ecosystem functioning and stability tend to be positively related, but negative relationships can occur when selection co-varies with richness. Our findings provide novel insights for understanding the functioning-stability relationship, with potential implications for both ecological research and ecosystem management.

Biotic homogenization destabilizes ecosystem functioning by decreasing spatial asynchrony.

Our planet is facing significant changes of biodiversity across spatial scales. Although the negative effects of local biodiversity (α diversity) loss on ecosystem stability are well documented, the consequences of biodiversity changes at larger spatial scales, in particular biotic homogenization, that is, reduced species turnover across space (ß diversity), remain poorly known. Using data from 39 grassland biodiversity experiments, we examine the effects of ß diversity on the stability of simulated landscapes while controlling for potentially confounding biotic and abiotic factors. Our results show that higher ß diversity generates more asynchronous dynamics among local communities and thereby contributes to the stability of ecosystem productivity at larger spatial scales. We further quantify the relative contributions of α and ß diversity to ecosystem stability and find a relatively stronger effect of α diversity, possibly due to the limited spatial scale of our experiments. The stabilizing effects of both α and ß diversity lead to a positive diversity-stability relationship at the landscape scale. Our findings demonstrate the destabilizing effect of biotic homogenization and suggest that biodiversity should be conserved at multiple spatial scales to maintain the stability of ecosystem functions and services.

Behaviorally-mediated trophic cascade attenuated by prey use of risky places at safe times.

The mere threat of predation may incite behavioral changes in prey that lead to community-wide impacts on productivity, biodiversity, and nutrient cycling. The paucity of experimental manipulations, however, has contributed to controversy over the strength of this pathway in wide-ranging vertebrate systems. We investigated whether simulated gray wolf (Canis lupus) presence can induce behaviorally-mediated trophic cascades, specifically, whether the 'fear' of wolf olfactory cues alone can change deer foraging behavior in ways that affect plants and soils. Wolves were recently removed from the Cedar Creek Ecosystem Science Reserve (Minnesota, USA), such that consumptively mediated predator effects were negligible. At 32 experimental plots, we crossed two nested treatments: wolf urine application and herbivore exclosures. We deployed camera traps to quantify how white-tailed deer (Odocoileus virginianus) adjusted their spatiotemporal habitat use, foraging, and vigilance in response to wolf cues and how these behavioral changes affected plant productivity, plant communities, and soil nutrients. Weekly applications of wolf urine significantly altered deer behavior, but deer responses did not cascade to affect plant or soil properties. Deer substantially reduced crepuscular activity at wolf-simulated sites compared to control locations. As wolves in this area predominantly hunted during mornings and evenings, this response potentially allows deer to maximize landscape use by accessing dangerous areas when temporal threat is low. Our experiment suggests that prey may be sensitive to 'dynamic' predation risk that is structured across both space and time and, consequentially, prey use of risky areas during safe times may attenuate behaviorally-mediated trophic cascades at the predator-prey interface.