Emerging niche clustering results from both competition and predation.

Understanding species coexistence has been a central question in ecology for decades, and the notion that competing species need to differ in their ecological niche for stable coexistence has dominated. Recent theoretical and empirical work suggests differently. Species can also escape competitive exclusion by being similar, leading to clusters of species with similar traits. This theory has so far only been explored under competition. By combining mathematical and numerical analyses, we reveal that competition and predation are equally capable to promote clusters of similar species in prey-predator communities, their relative importance being modulated by resource availability. We further show that predation has a stabilizing effect on clustering patterns, making the clusters more diverse. Our results merge different ecological theories and bring new light to the emergent neutrality theory by adding the perspective of trophic interactions. These results open new perspectives to the study of trait distributions in ecological interaction networks.

Quality matters: Stoichiometry of resources modulates spatial feedbacks in aquatic-terrestrial meta-ecosystems.

Species dispersal and resource spatial flows greatly affect the dynamics of connected ecosystems. So far, research on meta-ecosystems has mainly focused on the quantitative effect of subsidy flows. Yet, resource exchanges at heterotrophic-autotrophic (e.g. aquatic-terrestrial) ecotones display a stoichiometric asymmetry that likely matters for functioning. Here, we joined ecological stoichiometry and the meta-ecosystem framework to understand how subsidy stoichiometry mediates the response of the meta-ecosystem to subsidy flows. Our model results demonstrate that resource flows between ecosystems can induce a positive spatial feedback loop, leading to higher production at the meta-ecosystem scale by relaxing local ecosystem limitations ('spatial complementarity'). Furthermore, we show that spatial flows can also have an unexpected negative impact on production when accentuating the stoichiometric mismatch between local resources and basal species needs. This study paves the way for studies on the interdependency of ecosystems at the landscape extent.

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.

Predicting effects of multiple interacting global change drivers across trophic levels.

Global change encompasses many co-occurring anthropogenic drivers, which can act synergistically or antagonistically on ecological systems. Predicting how different global change drivers simultaneously contribute to observed biodiversity change is a key challenge for ecology and conservation. However, we lack the mechanistic understanding of how multiple global change drivers influence the vital rates of multiple interacting species. We propose that reaction norms, the relationships between a driver and vital rates like growth, mortality, and consumption, provide insights to the underlying mechanisms of community responses to multiple drivers. Understanding how multiple drivers interact to affect demographic rates using a reaction-norm perspective can improve our ability to make predictions of interactions at higher levels of organization-that is, community and food web. Building on the framework of consumer-resource interactions and widely studied thermal performance curves, we illustrate how joint driver impacts can be scaled up from the population to the community level. A simple proof-of-concept model demonstrates how reaction norms of vital rates predict the prevalence of driver interactions at the community level. A literature search suggests that our proposed approach is not yet used in multiple driver research. We outline how realistic response surfaces (i.e., multidimensional reaction norms) can be inferred by parametric and nonparametric approaches. Response surfaces have the potential to strengthen our understanding of how multiple drivers affect communities as well as improve our ability to predict when interactive effects emerge, two of the major challenges of ecology today.

Native and exotic plants play different roles in urban pollination networks across seasons.

Urban areas often host exotic plant species, whether managed or spontaneous. These plants are suspected of affecting pollinator diversity and the structure of pollination networks. However, in dense cityscapes, exotic plants also provide additional flower resources during periods of scarcity, and the consequences for the seasonal dynamics of networks still need to be investigated. For two consecutive years, we monitored monthly plant-pollinator networks in 12 green spaces in Paris, France. We focused on seasonal variations in the availability and attractiveness of flower resources, comparing native and exotic plants at both the species and community levels. We also considered their respective contributions to network properties over time (specialization and nestedness). Exotic plants provided more abundant and diverse flower resources than native plants, especially from late summer on. However, native plants received more visits and attracted more pollinator species at the community level; and during certain times of the year at the species level as well. Exotic plants were involved in more generalist interactions, increasingly so over the seasons. In addition, they contributed more to network nestedness than native plants. These results show that exotic plants are major components of plant-pollinator interactions in a dense urban landscape, even though they are less attractive than natives. They constitute a core of generalist interactions that increase nestedness and can participate in the overall stability of the network. However, most exotic species were seldom visited by insects. Pollinator communities may benefit from including more native species when managing urban green spaces.

Interaction fidelity is less common than expected in plant-pollinator communities.

Pairs of plants and pollinators species sometimes consistently interact throughout time and across space. Such consistency can be interpreted as a sign of interaction fidelity, that is a consistent interaction between two species when they co-occur in the same place. But how common interaction fidelity is and what determines interaction fidelity in plant-pollinator communities remain open questions. We aim to assess how frequent is interaction fidelity between plants and their pollinators and what drives interaction fidelity across plant-pollinator communities. Using a dataset of 141 networks around the world, we quantify whether the interaction between pairs of plant and pollinator species happens more ('interaction fidelity') or less ('interaction avoidance') often than expected by chance given the structure of the networks in which they co-occur. We also explore the relationship between interaction fidelity and species' degree (i.e. number of interactions), and the taxonomy of the species involved in the interaction. Our findings reveal that most plant-pollinator interactions do not differ from random expectations, in other words show neither fidelity nor avoidance. Out of the total 44,814 co-occurring species pairs we found 7,877 unique pair interactions (18%). Only 551 (7%) of the 7,877 plant-pollinator interactions did show significant interaction fidelity, meaning that these pairs interact in a consistent and non-random way across networks. We also find that 39 (0.09%) out of 44,814 plant-pollinator pairs showed significant interaction avoidance. Our results suggest that interactions involving specialist species have a high probability to show interaction fidelity and a low probability of interaction avoidance. In addition, we find that particular associations between plant and insect orders, as for example interactions between Hymenoptera and Fabales, showed high fidelity and low avoidance. Although niche and neutral processes simultaneously influence patterns of interaction in ecological communities, our findings suggest that it is rather neutral processes that are shaping the patterns of interactions in plant-pollinator networks.

Phenological traits foster persistence of mutualistic networks by promoting facilitation.

Morphological and phenological traits are key determinants of the structure of mutualistic networks. Both traits create forbidden links, but phenological traits can also decouple interaction in time. While such difference likely affects the indirect effects among species and consequently network persistence, it remains overlooked. Here, using a dynamic model, we show that networks structured by phenology favour facilitation over competition within guilds of pollinators and plants, thereby increasing network persistence, while the contrary holds for networks structured by morphology. We further show that such buffering of competition by phenological traits mostly beneficiate to specialists, the most vulnerable species otherwise, which propagate the most positive effects within guilds and promote nestedness. Our results indicate that beyond trophic mismatch, phenological shifts such as those induced by climate change are likely to affect indirect effects within mutualistic assemblages, with consequences for biodiversity.

Species richness and food-web structure jointly drive community biomass and its temporal stability in fish communities.

Biodiversity-ecosystem functioning and food-web complexity-stability relationships are central to ecology. However, they remain largely untested in natural contexts. Here, we estimated the links among environmental conditions, richness, food-web structure, annual biomass and its temporal stability using a standardised monitoring dataset of 99 stream fish communities spanning from 1995 to 2018. We first revealed that both richness and average trophic level are positively related to annual biomass, with effects of similar strength. Second, we found that community stability is fostered by mean trophic level, while contrary to expectation, it is decreased by species richness. Finally, we found that environmental conditions affect both biomass and its stability mainly via effects on richness and network structure. Strikingly, the effect of species richness on community stability was mediated by population stability rather than synchrony, which contrasts with results from single trophic communities. We discuss the hypothesis that it could be a characteristic of multi-trophic communities.

Small freshwater ecosystems with dissimilar microbial communities exhibit similar temporal patterns.

Despite small freshwater ecosystems being biodiversity reservoirs and contributing significantly to greenhouse fluxes, their microbial communities remain largely understudied. Yet, microorganisms intervene in biogeochemical cycling and impact water quality. Because of their small size, these ecosystems are in principle more sensitive to disturbances, seasonal variation and pluri-annual climate change. However, how microbial community composition varies over space and time, and whether archaeal, bacterial and microbial eukaryote communities behave similarly remain unanswered. Here, we aim to unravel the composition and intra/interannual temporal dynamic patterns for archaea, bacteria and microbial eukaryotes in a set of small freshwater ecosystems. We monitored archaeal and bacterial community composition during 24 consecutive months in four ponds and one brook from northwestern France by 16S rRNA gene amplicon sequencing (microbial eukaryotes were previously investigated for the same systems). Unexpectedly for oxic environments, bacterial Candidate Phyla Radiation (CPR) were highly diverse and locally abundant. Our results suggest that microbial community structure is mainly driven by environmental conditions acting over space (ecosystems) and time (seasons). A low proportion of operational taxonomic units (OTUs) (<1%) was shared by the five ecosystems despite their geographical proximity (2-9 km away), making microbial communities almost unique in each ecosystem and highlighting the strong selective influence of local environmental conditions. Marked and similar seasonality patterns were observed for archaea, bacteria and microbial eukaryotes in all ecosystems despite strong turnovers of rare OTUs. Over the 2-year survey, microbial community composition varied despite relatively stable environmental parameters. This suggests that biotic associations play an important role in interannual community assembly.

Relative effects of anthropogenic pressures, climate, and sampling design on the structure of pollination networks at the global scale.

Pollinators provide crucial ecosystem services that underpin to wild plant reproduction and yields of insect-pollinated crops. Understanding the relative impacts of anthropogenic pressures and climate on the structure of plant-pollinator interaction networks is vital considering ongoing global change and pollinator decline. Our ability to predict the consequences of global change for pollinator assemblages worldwide requires global syntheses, but these analytical approaches may be hindered by variable methods among studies that either invalidate comparisons or mask biological phenomena. Here we conducted a synthetic analysis that assesses the relative impact of anthropogenic pressures and climatic variability, and accounts for heterogeneity in sampling methodology to reveal network responses at the global scale. We analyzed an extensive dataset, comprising 295 networks over 123 locations all over the world, and reporting over 50,000 interactions between flowering plant species and their insect visitors. Our study revealed that anthropogenic pressures correlate with an increase in generalism in pollination networks while pollinator richness and taxonomic composition are more related to climatic variables with an increase in dipteran pollinator richness associated with cooler temperatures. The contrasting response of species richness and generalism of the plant-pollinator networks stresses the importance of considering interaction network structure alongside diversity in ecological monitoring. In addition, differences in sampling design explained more variation than anthropogenic pressures or climate on both pollination networks richness and generalism, highlighting the crucial need to report and incorporate sampling design in macroecological comparative studies of pollination networks. As a whole, our study reveals a potential human impact on pollination networks at a global scale. However, further research is needed to evaluate potential consequences of loss of specialist species and their unique ecological interactions and evolutionary pathways on the ecosystem pollination function at a global scale.

Trophic redundancy reduces vulnerability to extinction cascades.

Current species extinction rates are at unprecedentedly high levels. While human activities can be the direct cause of some extinctions, it is becoming increasingly clear that species extinctions themselves can be the cause of further extinctions, since species affect each other through the network of ecological interactions among them. There is concern that the simplification of ecosystems, due to the loss of species and ecological interactions, increases their vulnerability to such secondary extinctions. It is predicted that more complex food webs will be less vulnerable to secondary extinctions due to greater trophic redundancy that can buffer against the effects of species loss. Here, we demonstrate in a field experiment with replicated plant-insect communities, that the probability of secondary extinctions is indeed smaller in food webs that include trophic redundancy. Harvesting one species of parasitoid wasp led to secondary extinctions of other, indirectly linked, species at the same trophic level. This effect was markedly stronger in simple communities than for the same species within a more complex food web. We show that this is due to functional redundancy in the more complex food webs and confirm this mechanism with a food web simulation model by highlighting the importance of the presence and strength of trophic links providing redundancy to those links that were lost. Our results demonstrate that biodiversity loss, leading to a reduction in redundant interactions, can increase the vulnerability of ecosystems to secondary extinctions, which, when they occur, can then lead to further simplification and run-away extinction cascades.

Long-term effects of global change on occupancy and flight period of wild bees in Belgium.

Global change affects species by modifying their abundance, spatial distribution, and activity period. The challenge is now to identify the respective drivers of those responses and to understand how those responses combine to affect species assemblages and ecosystem functioning. Here we correlate changes in occupancy and mean flight date of 205 wild bee species in Belgium with temporal changes in temperature trend and interannual variation, agricultural intensification, and urbanization. Over the last 70 years, bee occupancy decreased on average by 33%, most likely because of agricultural intensification, and flight period of bees advanced on average by 4 days, most likely because of interannual temperature changes. Those responses resulted in a synergistic effect because species which increased in occupancy tend to be those that have shifted their phenologies earlier in the season. This leads to an overall advancement and shortening of the pollination season by 9 and 15 days respectively, with lower species richness and abundance compared to historical pollinator assemblages, except at the early start of the season. Our results thus suggest a strong decline in pollination function and services.

Advancing our understanding of ecological stability.

The concept of ecological stability occupies a prominent place in both fundamental and applied ecological research. We review decades of work on the topic and examine how our understanding has progressed. We show that our understanding of stability has remained fragmented and is limited largely to simple or simplified systems. There has been a profusion of metrics proposed to quantify stability, of which only a handful are used commonly. Furthermore, studies typically quantify one to two metrics of stability at a time and in response to a single perturbation, with some of the main environmental pressures of today being the least studied. We argue that we need to build on the existing consensus and strong theoretical foundation of the stability concept to better understand its multidimensionality and the interdependencies between metrics, levels of organisation and types of perturbations. Only by doing so can we make progress in the quantification of stability in theory and in practice, and eventually build a more comprehensive understanding of how ecosystems will respond to ongoing environmental change.

How plants connect pollination and herbivory networks and their contribution to community stability.

Pollination and herbivory networks have mainly been studied separately, highlighting their distinct structural characteristics and the related processes and dynamics. However, most plants interact with both pollinators and herbivores, and there is evidence that both types of interaction affect each other. Here we investigated the way plants connect these mutualistic and antagonistic networks together, and the consequences for community stability. Using an empirical data set, we show that the way plants connect pollination and herbivory networks is not random and promotes community stability. Analyses of the structure of binary and quantitative networks show different results: the plants' generalism with regard to pollinators is positively correlated to their generalism with regard to herbivores when considering binary interactions, but not when considering quantitative interactions. We also show that plants that share the same pollinators do not share the same herbivores. However, the way plants connect pollination and herbivory networks promotes stability for both binary and quantitative networks. Our results highlight the relevance of considering the diversity of interaction types in ecological communities, and stress the need to better quantify the costs and benefits of interactions, as well as to develop new metrics characterizing the way different interaction types are combined within ecological networks.

How plants connect pollination and herbivory networks and their contribution to community stability.

Pollination and herbivory networks have mainly been studied separately, highlighting their distinct structural characteristics and the related processes and dynamics. However, most plants interact with both pollinators and herbivores, and there is evidence that both types of interaction affect each other. Here we investigated the way plants connect these mutualistic and antagonistic networks together, and the consequences for community stability. Using an empirical data set, we show that the way plants connect pollination and herbivory networks is not random and promotes community stability. Analyses of the structure of binary and quantitative networks show different results: the plants' generalism with regard to pollinators is positively correlated to their generalism with regard to herbivores when considering binary interactions, but not when considering quantitative interactions. We also show that plants that share the same pollinators do not share the same herbivores. However, the way plants connect pollination and herbivory networks promotes stability for both binary and quantitative networks. Our results highlight the relevance of considering the diversity of interaction types in ecological communities, and stress the need to better quantify the costs and benefits of interactions, as well as to develop new metrics characterizing the way different interaction types are combined within ecological networks.

Soil food web properties explain ecosystem services across European land use systems.

Intensive land use reduces the diversity and abundance of many soil biota, with consequences for the processes that they govern and the ecosystem services that these processes underpin. Relationships between soil biota and ecosystem processes have mostly been found in laboratory experiments and rarely are found in the field. Here, we quantified, across four countries of contrasting climatic and soil conditions in Europe, how differences in soil food web composition resulting from land use systems (intensive wheat rotation, extensive rotation, and permanent grassland) influence the functioning of soils and the ecosystem services that they deliver. Intensive wheat rotation consistently reduced the biomass of all components of the soil food web across all countries. Soil food web properties strongly and consistently predicted processes of C and N cycling across land use systems and geographic locations, and they were a better predictor of these processes than land use. Processes of carbon loss increased with soil food web properties that correlated with soil C content, such as earthworm biomass and fungal/bacterial energy channel ratio, and were greatest in permanent grassland. In contrast, processes of N cycling were explained by soil food web properties independent of land use, such as arbuscular mycorrhizal fungi and bacterial channel biomass. Our quantification of the contribution of soil organisms to processes of C and N cycling across land use systems and geographic locations shows that soil biota need to be included in C and N cycling models and highlights the need to map and conserve soil biodiversity across the world.

Intensive agriculture reduces soil biodiversity across Europe.

Soil biodiversity plays a key role in regulating the processes that underpin the delivery of ecosystem goods and services in terrestrial ecosystems. Agricultural intensification is known to change the diversity of individual groups of soil biota, but less is known about how intensification affects biodiversity of the soil food web as a whole, and whether or not these effects may be generalized across regions. We examined biodiversity in soil food webs from grasslands, extensive, and intensive rotations in four agricultural regions across Europe: in Sweden, the UK, the Czech Republic and Greece. Effects of land-use intensity were quantified based on structure and diversity among functional groups in the soil food web, as well as on community-weighted mean body mass of soil fauna. We also elucidate land-use intensity effects on diversity of taxonomic units within taxonomic groups of soil fauna. We found that between regions soil food web diversity measures were variable, but that increasing land-use intensity caused highly consistent responses. In particular, land-use intensification reduced the complexity in the soil food webs, as well as the community-weighted mean body mass of soil fauna. In all regions across Europe, species richness of earthworms, Collembolans, and oribatid mites was negatively affected by increased land-use intensity. The taxonomic distinctness, which is a measure of taxonomic relatedness of species in a community that is independent of species richness, was also reduced by land-use intensification. We conclude that intensive agriculture reduces soil biodiversity, making soil food webs less diverse and composed of smaller bodied organisms. Land-use intensification results in fewer functional groups of soil biota with fewer and taxonomically more closely related species. We discuss how these changes in soil biodiversity due to land-use intensification may threaten the functioning of soil in agricultural production systems.

Functionally and phylogenetically diverse plant communities key to soil biota.

Recent studies assessing the role of biological diversity for ecosystem functioning indicate that the diversity of functional traits and the evolutionary history of species in a community, not the number of taxonomic units, ultimately drives the biodiversity--ecosystem-function relationship. Here, we simultaneously assessed the importance of plant functional trait and phylogenetic diversity as predictors of major trophic groups of soil biota (abundance and diversity), six years from the onset of a grassland biodiversity experiment. Plant functional and phylogenetic diversity were generally better predictors of soil biota than the traditionally used species or functional group richness. Functional diversity was a reliable predictor for most biota, with the exception of soil microorganisms, which were better predicted by phylogenetic diversity. These results provide empirical support for the idea that the diversity of plant functional traits and the diversity of evolutionary lineages in a community are important for maintaining higher abundances and diversity of soil communities.

A bioenergetic framework for aboveground terrestrial food webs.

Bioenergetic approaches have been greatly influential for understanding community functioning and stability and predicting effects of environmental changes on biodiversity. These approaches use allometric relationships to establish species' trophic interactions and consumption rates and have been successfully applied to aquatic ecosystems. Terrestrial ecosystems, where body mass is less predictive of plant-consumer interactions, present inherent challenges that these models have yet to meet. Here, we discuss the processes governing terrestrial plant-consumer interactions and develop a bioenergetic framework integrating those processes. Our framework integrates bioenergetics specific to terrestrial plants and their consumers within a food web approach while also considering mutualistic interactions. Such a framework is poised to advance our understanding of terrestrial food webs and to predict their responses to environmental changes.