Systematic distributions of interaction strengths across tree interaction networks yield positive diversity-productivity relationships.

Understanding the mechanisms underlying diversity-productivity relationships (DPRs) is crucial to mitigating the effects of forest biodiversity loss. Tree-tree interactions in diverse communities are fundamental in driving growth rates, potentially shaping the emergent DPRs, yet remain poorly explored. Here, using data from a large-scale forest biodiversity experiment in subtropical China, we demonstrated that changes in individual tree productivity were driven by species-specific pairwise interactions, with higher positive net pairwise interaction effects on trees in more diverse neighbourhoods. By perturbing the interactions strength from empirical data in simulations, we revealed that the positive differences between inter- and intra-specific interactions were the critical determinant for the emergence of positive DPRs. Surprisingly, the condition for positive DPRs corresponded to the condition for coexistence. Our results thus provide a novel insight into how pairwise tree interactions regulate DPRs, with implications for identifying the tree mixtures with maximized productivity to guide forest restoration and reforestation efforts.

Reciprocal inhibition and competitive hierarchy cause negative biodiversity-ecosystem function relationships.

The relationship between biodiversity and ecosystem function (BEF) captivates ecologists, but the factors responsible for the direction of this relationship remain unclear. While higher ecosystem functioning at higher biodiversity levels ('positive BEF') is not universal in nature, negative BEF relationships seem puzzlingly rare. Here, we develop a dynamical consumer-resource model inspired by microbial decomposer communities in pitcher plant leaves to investigate BEF. We manipulate microbial diversity via controlled colonization and measure their function as total ammonia production. We test how niche partitioning among bacteria and other ecological processes influence BEF in the leaves. We find that a negative BEF can emerge from reciprocal interspecific inhibition in ammonia production causing a negative complementarity effect, or from competitive hierarchies causing a negative selection effect. Absent these factors, a positive BEF was the typical outcome. Our findings provide a potential explanation for the rarity of negative BEF in empirical data.

More is not always better: peat moss mixtures slightly enhance peatland stability.

Terrestrial wetland ecosystems challenge biodiversity-ecosystem function theory, which generally links high species diversity to stable ecosystem functions. An open question in ecosystem ecology is whether assemblages of co-occurring peat mosses contribute to the stability of peatland ecosystem processes. We conducted a two-species (Sphagnum cuspidatum, Sphagnum medium) replacement series mesocosm experiment to evaluate the resistance, resilience, and recovery rates of net ecosystem CO2 exchange (NEE) under mild and deep water table drawdown. Our results show a positive effect of mild water table drawdown on NEE with no apparent role for peat moss mixture. Our study indicates that the carbon uptake capacity by peat moss mixtures is rather resilient to mild water table drawdown, but seriously affected by deeper drought conditions. Co-occurring peat moss species seem to enhance the resilience of the carbon uptake function (i.e. ability of NEE to return to pre-perturbation levels) of peat moss mixtures only slightly. These findings suggest that assemblages of co-occurring Sphagnum mosses do only marginally contribute to the stability of ecosystem functions in peatlands under drought conditions. Above all, our results highlight that predicted severe droughts can gravely affect the sink capacity of peatlands, with only a small extenuating role for peat moss mixtures.

Mycorrhizal associations modify tree diversity-productivity relationships across experimental tree plantations.

Decades of studies have demonstrated links between biodiversity and ecosystem functioning, yet the generality of the relationships and the underlying mechanisms remain unclear, especially for forest ecosystems. Using 11 tree-diversity experiments, we tested tree species richness-community productivity relationships and the role of arbuscular (AM) or ectomycorrhizal (ECM) fungal-associated tree species in these relationships. Tree species richness had a positive effect on community productivity across experiments, modified by the diversity of tree mycorrhizal associations. In communities with both AM and ECM trees, species richness showed positive effects on community productivity, which could have resulted from complementarity between AM and ECM trees. Moreover, both AM and ECM trees were more productive in mixed communities with both AM and ECM trees than in communities assembled by their own mycorrhizal type of trees. In communities containing only ECM trees, species richness had a significant positive effect on productivity, whereas species richness did not show any significant effects on productivity in communities containing only AM trees. Our study provides novel explanations for variations in diversity-productivity relationships by suggesting that tree-mycorrhiza interactions can shape productivity in mixed-species forest ecosystems.

Microclimate modulation: An overlooked mechanism influencing the impact of plant diversity on ecosystem functioning.

Changes in climate and biodiversity are widely recognized as primary global change drivers of ecosystem structure and functioning, also affecting ecosystem services provided to human populations. Increasing plant diversity not only enhances ecosystem functioning and stability but also mitigates climate change effects and buffers extreme weather conditions, yet the underlying mechanisms remain largely unclear. Recent studies have shown that plant diversity can mitigate climate change (e.g. reduce temperature fluctuations or drought through microclimatic effects) in different compartments of the focal ecosystem, which as such may contribute to the effect of plant diversity on ecosystem properties and functioning. However, these potential plant diversity-induced microclimate effects are not sufficiently understood. Here, we explored the consequences of climate modulation through microclimate modification by plant diversity for ecosystem functioning as a potential mechanism contributing to the widely documented biodiversity-ecosystem functioning (BEF) relationships, using a combination of theoretical and simulation approaches. We focused on a diverse set of response variables at various levels of integration ranging from ecosystem-level carbon exchange to soil enzyme activity, including population dynamics and the activity of specific organisms. Here, we demonstrated that a vegetation layer composed of many plant species has the potential to influence ecosystem functioning and stability through the modification of microclimatic conditions, thus mitigating the negative impacts of climate extremes on ecosystem functioning. Integrating microclimatic processes (e.g. temperature, humidity and light modulation) as a mechanism contributing to the BEF relationships is a promising avenue to improve our understanding of the effects of climate change on ecosystem functioning and to better predict future ecosystem structure, functioning and services. In addition, microclimate management and monitoring should be seen as a potential tool by practitioners to adapt ecosystems to climate change.

The north-south divide? Macroalgal functional trait diversity and redundancy varies with intertidal aspect.

BACKGROUND AND AIMS: Marine macroalgae ('seaweeds') are critical to coastal ecosystem structure and function, but also vulnerable to the many environmental changes associated with anthropogenic climate change (ACC). The local habitat conditions underpinning observed and predicted ACC-driven changes in intertidal macroalgal communities are complex and probably site-specific and operate in addition to more commonly reported regional factors such as sea surface temperatures. METHODS: We examined how the composition and functional trait expression of macroalgal communities in SW England varied with aspect (i.e. north-south orientation) at four sites with opposing Equator- (EF) and Pole-facing (PF) surfaces. Previous work at these sites had established that average annual (low tide) temperatures vary by 1.6 °C and that EF-surfaces experience six-fold more frequent extremes (i.e. >30 °C). KEY RESULTS: PF macroalgal communities were consistently more taxon rich; 11 taxa were unique to PF habitats, with only one restricted to EF. Likewise, functional richness and dispersion were greater on PF-surfaces (dominated by algae with traits linked to rapid resource capture and utilization, but low desiccation tolerance), although differences in both taxon and functional richness were probably driven by the fact that less diverse EF-surfaces were dominated by desiccation-tolerant fucoids. CONCLUSIONS: Although we cannot disentangle the influence of temperature variation on algal ecophysiology from the indirect effects of aspect on species interactions (niche pre-emption, competition, grazing, etc.), our study system provides an excellent model for understanding how environmental variation at local scales affects community composition and functioning. By virtue of enhanced taxonomic diversity, PF-aspects supported higher functional diversity and, consequently, greater effective functional redundancy. These differences may imbue PF-aspects with resilience against environmental perturbation, but if predicted increases in global temperatures are realized, some PF-sites may shift to a depauperate, desiccation-tolerant seaweed community with a concomitant loss of functional diversity and redundancy.

Aquatic biodiversity enhances multiple nutritional benefits to humans.

Humanity depends on biodiversity for health, well-being, and a stable environment. As biodiversity change accelerates, we are still discovering the full range of consequences for human health and well-being. Here, we test the hypothesis-derived from biodiversity-ecosystem functioning theory-that species richness and ecological functional diversity allow seafood diets to fulfill multiple nutritional requirements, a condition necessary for human health. We analyzed a newly synthesized dataset of 7,245 observations of nutrient and contaminant concentrations in 801 aquatic animal taxa and found that species with different ecological traits have distinct and complementary micronutrient profiles but little difference in protein content. The same complementarity mechanisms that generate positive biodiversity effects on ecosystem functioning in terrestrial ecosystems also operate in seafood assemblages, allowing more diverse diets to yield increased nutritional benefits independent of total biomass consumed. Notably, nutritional metrics that capture multiple micronutrients and fatty acids essential for human well-being depend more strongly on biodiversity than common ecological measures of function such as productivity, typically reported for grasslands and forests. Furthermore, we found that increasing species richness did not increase the amount of protein in seafood diets and also increased concentrations of toxic metal contaminants in the diet. Seafood-derived micronutrients and fatty acids are important for human health and are a pillar of global food and nutrition security. By drawing upon biodiversity-ecosystem functioning theory, we demonstrate that ecological concepts of biodiversity can deepen our understanding of nature's benefits to people and unite sustainability goals for biodiversity and human well-being.

Phytoplankton biodiversity is more important for ecosystem functioning in highly variable thermal environments.

The 21st century has seen an acceleration of anthropogenic climate change and biodiversity loss, with both stressors deemed to affect ecosystem functioning. However, we know little about the interactive effects of both stressors and in particular about the interaction of increased climatic variability and biodiversity loss on ecosystem functioning. This should be remedied because larger climatic variability is one of the main features of climate change. Here, we demonstrated that temperature fluctuations led to changes in the importance of biodiversity for ecosystem functioning. We used microcosm communities of different phytoplankton species richness and exposed them to a constant, mild, and severe temperature-fluctuating environment. Wider temperature fluctuations led to steeper biodiversity-ecosystem functioning slopes, meaning that species loss had a stronger negative effect on ecosystem functioning in more fluctuating environments. For severe temperature fluctuations, the slope increased through time due to a decrease of the productivity of species-poor communities over time. We developed a theoretical competition model to better understand our experimental results and showed that larger differences in thermal tolerances across species led to steeper biodiversity-ecosystem functioning slopes. Species-rich communities maintained their ecosystem functioning with increased fluctuation as they contained species able to resist the thermally fluctuating environments, while this was on average not the case in species-poor communities. Our results highlight the importance of biodiversity for maintaining ecosystem functions and services in the context of increased climatic variability under climate change.

Animal and plant space-use drive plant diversity-productivity relationships.

Plant community productivity generally increases with biodiversity, but the strength of this relationship exhibits strong empirical variation. In meta-food-web simulations, we addressed if the spatial overlap in plants' resource access and animal space-use can explain such variability. We found that spatial overlap of plant resource access is a prerequisite for positive diversity-productivity relationships, but causes exploitative competition that can lead to competitive exclusion. Space-use of herbivores causes apparent competition among plants, resulting in negative relationships. However, space-use of larger top predators integrates sub-food webs composed of smaller species, offsetting the negative effects of exploitative and apparent competition and leading to strongly positive diversity-productivity relationships. Overall, our results show that spatial overlap of plants' resource access and animal space-use can greatly alter the strength and sign of such relationships. In particular, the scaling of animal space-use effects opens new perspectives for linking landscape processes without effects on biodiversity to productivity patterns.

A novel proxy to examine interspecific phosphorus facilitation between plant species.

Resource complementarity can contribute to enhanced ecosystem functioning in diverse plant communities, but the role of facilitation in the enhanced complementarity is poorly understood. Here, we use leaf manganese concentration ([Mn]) as a proxy for rhizosheath carboxylate concentration to explore novel mechanisms of complementarity mediated by phosphorus (P) facilitation. In pot experiments, we showed that mixtures involving Carex korshinskyi, an efficient P-mobilizing species, exhibited greater biomass and relative complementarity effect than combinations without C. korshinskyi on P-deficient soils. Compared with monocultures, leaf [Mn] and [P] of species that are inefficient at P mobilization increased by 27% and 21% when grown with C. korshinskyi (i.e. interspecific P facilitation via carboxylates) rather than next to another inefficient P-mobilizing species. This experimental result was supported by a meta-analysis including a range of efficient P-mobilizing species. Phosphorus facilitation enhanced the relative complementarity effect in low-P environments, related to a greater change in several facilitated species of their root morphological traits relative to those in monoculture. Using leaf [Mn] as a proxy, we highlight a vital mechanism of interspecific P facilitation via belowground processes and provide evidence for the pivotal role of P facilitation mediated by the plasticity of root traits in biodiversity research.

The essential role of biodiversity in the key axes of ecosystem function.

Biodiversity is essential for maintaining the terrestrial ecosystem multifunctionality (EMF). Recent studies have revealed that the variations in terrestrial ecosystem functions are captured by three key axes: the maximum productivity, water use efficiency, and carbon use efficiency of the ecosystem. However, the role of biodiversity in supporting these three key axes has not yet been explored. In this study, we combined the (i) data collected from more than 840 vegetation plots across a large climatic gradient in China using standard protocols, (ii) data on plant traits and phylogenetic information for more than 2,500 plant species, and (iii) soil nutrient data measured in each plot. These data were used to systematically assess the contribution of environmental factors, species richness, functional and phylogenetic diversity, and community-weighted mean (CWM) and ecosystem traits (i.e., traits intensity normalized per unit land area) to EMF via hierarchical partitioning and Bayesian structural equation modeling. Multiple biodiversity attributes accounted for 70% of the influence of all the variables on EMF, and ecosystems with high functional diversity had high resource use efficiency. Our study is the first to systematically explore the role of different biodiversity attributes, including species richness, phylogenetic and functional diversity, and CWM and ecosystem traits, in the key axes of ecosystem functions. Our findings underscore that biodiversity conservation is critical for sustaining EMF and ultimately ensuring human well-being.

Interactive effects of global change drivers as determinants of the link between soil biodiversity and ecosystem functioning.

Biodiversity, both aboveground and belowground, is negatively affected by global changes such as drought or warming. This loss of biodiversity impacts Earth's ecosystems, as there is a positive relationship between biodiversity and ecosystem functioning (BEF). Even though soils host a large fraction of biodiversity that underlies major ecosystem functions, studies exploring the relationship between soil biodiversity and ecosystem functioning (sBEF) as influenced by global change drivers (GCDs) remain scarce. Here we highlight the need to decipher sBEF relationships under the effect of interactive GCDs that are intimately connected in a changing world. We first state that sBEF relationships depend on the type of function (e.g., C cycling or decomposition) and biodiversity facet (e.g., abundance, species richness, or biomass) considered. Then, we shed light on the impact of single and interactive GCDs on soil biodiversity and sBEF and show that results from scarce studies studying interactive effects range from antagonistic to additive to synergistic when two individual GCDs cooccur. This indicates the need for studies quantitatively accounting for the impacts of interactive GCDs on sBEF relationships. Finally, we provide guidelines for optimized methodological and experimental approaches to study sBEF in a changing world that will provide more valuable information on the real impact of (interactive) GCDs on sBEF. Together, we highlight the need to decipher the sBEF relationship in soils to better understand soil functioning under ongoing global changes, as changes in sBEF are of immediate importance for ecosystem functioning.

Why are biodiversity-ecosystem functioning relationships so elusive? Trophic interactions may amplify ecosystem function variability.

The relationship between biodiversity and ecosystem functions (BEFs) has attracted great interest. Studies on BEF have so far focused on the average trend of ecosystem function as species diversity increases. A tantalizing but rarely addressed question is why large variations in ecosystem functions are often observed across systems with similar species diversity, likely obscuring observed BEFs. Here we use a multi-trophic food web model in combination with empirical data to examine the relationships between species richness and the variation in ecosystem functions (VEFs) including biomass, metabolism, decomposition, and primary and secondary production. We then probe the mechanisms underlying these relationships, focusing on the role of trophic interactions. While our results reinforce the previously documented positive BEF relationships, we found that ecosystem functions exhibit significant variation within each level of species richness and the magnitude of this variation displays a hump-shaped relationship with species richness. Our analyses demonstrate that VEFs is reduced when consumer diversity increases through elevated nonlinearity in trophic interactions, and/or when the diversity of basal species such as producers and decomposers decreases. This explanation is supported by a 34-year empirical food web time series from the Gulf of Riga ecosystem. Our work suggests that biodiversity loss may not only result in ecosystem function decline, but also reduce the predictability of functions by generating greater function variability among ecosystems. It thus helps to reconcile the debate on the generality of positive BEF relationships and to disentangle the drivers of ecosystem stability. The role of trophic interactions and the variation in their strengths mediated by functional responses in shaping ecosystem function variation warrants further investigations and better incorporation into biodiversity-ecosystem functioning research.

The hidden role of multi-trophic interactions in driving diversity-productivity relationships.

Resource-use complementarity of producer species is often invoked to explain the generally positive diversity-productivity relationships. Additionally, multi-trophic interactions that link processes across trophic levels have received increasing attention as a possible key driver. Given that both are integral to natural ecosystems, their interactive effect should be evident but has remained hidden. We address this issue by analysing diversity-productivity relationships in a simulation experiment of producer communities nested within complex food-webs, manipulating resource-use complementarity and multi-trophic animal richness. We show that these two mechanisms interactively create diverse communities of complementary producer species. This shapes diversity-productivity relationships such that their joint contribution generally exceeds their individual effects. Specifically, multi-trophic interactions in animal-rich ecosystems facilitate producer coexistence by preventing competitive exclusion despite overlaps in resource-use, which increases the realised complementarity. The interdependence of food-webs and producer complementarity in creating biodiversity-productivity relationships highlights the importance to adopt a multi-trophic perspective on biodiversity-ecosystem functioning relationships.

No complementarity no gain-Net diversity effects on tree productivity occur once complementarity emerges during early stand development.

Although there is compelling evidence that tree diversity has an overall positive effect on forest productivity, there are important divergences among studies on the nature and strength of these diversity effects and their timing during forest stand development. To clarify conflicting results related to stand developmental stage, we explored how diversity effects on productivity change through time in a diversity experiment spanning 11 years. We show that the strength of diversity effects on productivity progressively increases through time, becoming significantly positive after 9 years. Moreover, we demonstrate that the strengthening of diversity effects is driven primarily by gradual increases in complementarity. We also show that mixing species with contrasting resource-acquisition strategies, and the dominance of deciduous, fast-developing species, promote positive diversity effects on productivity. Our results suggest that the canopy closure and subsequent stem exclusion phase are key for promoting niche complementarity in diverse tree communities.

Failures to disagree are essential for environmental science to effectively influence policy development.

While environmental science, and ecology in particular, is working to provide better understanding to base sustainable decisions on, the way scientific understanding is developed can at times be detrimental to this cause. Locked-in debates are often unnecessarily polarised and can compromise any common goals of the opposing camps. The present paper is inspired by a resolved debate from an unrelated field of psychology where Nobel laureate David Kahneman and Garry Klein turned what seemed to be a locked-in debate into a constructive process for their fields. The present paper is also motivated by previous discourses regarding the role of thresholds in natural systems for management and governance, but its scope of analysis targets the scientific process within complex social-ecological systems in general. We identified four features of environmental science that appear to predispose for locked-in debates: (1) The strongly context-dependent behaviour of ecological systems. (2) The dominant role of single hypothesis testing. (3) The high prominence given to theory demonstration compared investigation. (4) The effect of urgent demands to inform and steer policy. This fertile ground is further cultivated by human psychological aspects as well as the structure of funding and publication systems.

Marine heatwaves of differing intensities lead to distinct patterns in seafloor functioning.

Marine heatwaves (MHWs) are increasing in frequency and intensity due to climate change. Several well-documented effects of heatwaves on community structure exist, but examples of their effect on functioning of species, communities or ecosystems remain scarce. We tested the effects of short-term, moderate and strong MHWs on macrofauna bioturbation and associated solute fluxes as examples of ecosystem functioning. We also measured macrofaunal excretion rates to assess effects of temperature on macrofauna metabolism. For this experiment, we used unmanipulated sediment cores with natural animal communities collected from a muddy location at 32 m depth in the northern Baltic Sea. Despite the mechanistic effect of bioturbation remaining unchanged between the treatments, there were significant differences in oxygen consumption, solute fluxes and excretion. Biogeochemical and biological processes were boosted by the moderate heatwave, whereas biogeochemical cycling seemed to decrease under a strong heatwave. A prolonged, moderate heatwave could possibly lead to resource depletion if primary production cannot meet the demands of benthic consumption. By contrast, decreased degradation activities under strong heatwaves could lead to a build-up of organic material and potentially hypoxia. The strong variability and the complexity of the response highlight the context dependency of these processes complicating future predictions.

The presence of multiple parasitoids decreases host survival under warming, but parasitoid performance also decreases.

Current global changes are reshaping ecological communities and modifying environmental conditions. We need to recognize the combined impact of these biotic and abiotic factors on species interactions, community dynamics and ecosystem functioning. Specifically, the strength of predator-prey interactions often depends on the presence of other natural enemies: it weakens with competition and interference or strengthens with facilitation. Such effects of multiple predators on prey are likely to be affected by changes in the abiotic environment, altering top-down control, a key structuring force in natural and agricultural ecosystems. Here, we investigated how warming alters the effects of multiple predators on prey suppression using a dynamic model coupled with empirical laboratory experiments with Drosophila-parasitoid communities. While multiple parasitoids enhanced top-down control under warming, parasitoid performance generally declined when another parasitoid was present owing to competitive interactions. This could reduce top-down control over multiple generations. Our study highlights the importance of accounting for interactive effects between abiotic and biotic factors to better predict community dynamics in a rapidly changing world and thus better preserve ecosystem functioning and services such as biological control.

The role of diversity, body size and climate in dung removal: A correlative and experimental approach.

The mechanisms by which climatic changes influence ecosystem functions, that is, by a direct climatic control of ecosystem processes or by modifying richness and trait compositions of species communities, remain unresolved. This study is a contribution to this discourse by elucidating the linkages between climate, land use, biodiversity, body size and ecosystem functions. We disentangled direct climatic from biodiversity-mediated effects by using dung removal by dung beetles as a model system and by combining correlative field data and exclosure experiments along an extensive elevational gradient on Mt. Kilimanjaro, Tanzania. Dung removal declined with increasing elevation, being associated with a strong reduction in the richness and body size traits of dung beetle communities. Climate influenced dung removal rates by modifying biodiversity rather than by direct effects. The biodiversity-ecosystem effect was driven by a change in the mean body size of dung beetles. Dung removal rates were strongly reduced when large dung beetles were experimentally excluded. This study underscores that climate influences ecosystem functions mainly by modifying biodiversity and underpins the important role of body size for dung removal.

Avoiding artifacts when varying the number of species in ecological models.

Ecological theory recognizes the importance of the variety of species for maintaining the functioning of ecosystems and their derived services. We assert that when studying the effects of shifts in biodiversity levels using mathematical models, their dynamics must be sensitive to the variety of species traits but not to raw species numbers, a property that we call order-invariance. We present a testing procedure for verifying order-invariance of ecological network models -with or without trait adaptation- expressed as ODEs. Furthermore, we applied our test to several influential models used for evaluating biodiversity effects on ecosystem functioning. In most of the surveyed studies the equations failed our test. This raises doubts about the validity of previous results and calls for revisiting the theory derived from these studies. Our results foster the creation of artifact-free models, a necessary step towards building a more robust theory of biodiversity-driven ecosystem functioning.