Intraspecific demographic and trait responses to environmental change drivers are linked in two species of ciliate.

BACKGROUND: Over the past decade, theory and observations have suggested intraspecific variation, trait-based differences within species, as a buffer against biodiversity loss from multiple environmental changes. This buffering effect can only occur when different populations of the same species respond differently to environmental change. More specifically, variation of demographic responses fosters buffering of demography, while variation of trait responses fosters buffering of functioning. Understanding how both responses are related is important for predicting biodiversity loss and its consequences. In this study, we aimed to empirically assess whether population-level trait responses to multiple environmental change drivers are related to the demographic response to these drivers. To this end, we measured demographic and trait responses in microcosm experiments with two species of ciliated protists. For three clonal strains of each species, we measured responses to two environmental change drivers (climate change and pollution) and their combination. We also examined if relationships between demographic and trait responses existed across treatments and strains. RESULTS: We found different demographic responses across strains of the same species but hardly any interactive effects between the two environmental change drivers. Also, trait responses (summarized in a survival strategy index) varied among strains within a species, again with no driver interactions. Demographic and trait responses were related across all strains of both species tested in this study: Increasing intrinsic growth and self-limitation were associated with a shift in survival strategy from sit-and-wait towards flee. CONCLUSIONS: Our results support the existence of a link between a population's demographic and trait responses to environmental change drivers in two species of ciliate. Future work could dive deeper into the specifics of phenotypical trait values, and changes therein, related to specific life strategies in different species of ciliate and other zooplankton grazers.

Relationships of temperature and biodiversity with stability of natural aquatic food webs.

Temperature and biodiversity changes occur in concert, but their joint effects on ecological stability of natural food webs are unknown. Here, we assess these relationships in 19 planktonic food webs. We estimate stability as structural stability (using the volume contraction rate) and temporal stability (using the temporal variation of species abundances). Warmer temperatures were associated with lower structural and temporal stability, while biodiversity had no consistent effects on either stability property. While species richness was associated with lower structural stability and higher temporal stability, Simpson diversity was associated with higher temporal stability. The responses of structural stability were linked to disproportionate contributions from two trophic groups (predators and consumers), while the responses of temporal stability were linked both to synchrony of all species within the food web and distinctive contributions from three trophic groups (predators, consumers, and producers). Our results suggest that, in natural ecosystems, warmer temperatures can erode ecosystem stability, while biodiversity changes may not have consistent effects.

Mean species responses predict effects of environmental change on coexistence.

Environmental change research is plagued by the curse of dimensionality: the number of communities at risk and the number of environmental drivers are both large. This raises the pressing question if a general understanding of ecological effects is achievable. Here, we show evidence that this is indeed possible. Using theoretical and simulation-based evidence for bi- and tritrophic communities, we show that environmental change effects on coexistence are proportional to mean species responses and depend on how trophic levels on average interact prior to environmental change. We then benchmark our findings using relevant cases of environmental change, showing that means of temperature optima and of species sensitivities to pollution predict concomitant effects on coexistence. Finally, we demonstrate how to apply our theory to the analysis of field data, finding support for effects of land use change on coexistence in natural invertebrate communities.

Density declines, richness increases, and composition shifts in stream macroinvertebrates.

Documenting trends of stream macroinvertebrate biodiversity is challenging because biomonitoring often has limited spatial, temporal, and taxonomic scopes. We analyzed biodiversity and composition of assemblages of >500 genera, spanning 27 years, and 6131 stream sites across forested, grassland, urban, and agricultural land uses throughout the United States. In this dataset, macroinvertebrate density declined by 11% and richness increased by 12.2%, and insect density and richness declined by 23.3 and 6.8%, respectively, over 27 years. In addition, differences in richness and composition between urban and agricultural versus forested and grassland streams have increased over time. Urban and agricultural streams lost the few disturbance-sensitive taxa they once had and gained disturbance-tolerant taxa. These results suggest that current efforts to protect and restore streams are not sufficient to mitigate anthropogenic effects.

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.

Dispersal syndromes in challenging environments: A cross-species experiment.

Dispersal is a central biological process tightly integrated into life-histories, morphology, physiology and behaviour. Such associations, or syndromes, are anticipated to impact the eco-evolutionary dynamics of spatially structured populations, and cascade into ecosystem processes. As for dispersal on its own, these syndromes are likely neither fixed nor random, but conditional on the experienced environment. We experimentally studied how dispersal propensity varies with individuals' phenotype and local environmental harshness using 15 species ranging from protists to vertebrates. We reveal a general phenotypic dispersal syndrome across studied species, with dispersers being larger, more active and having a marked locomotion-oriented morphology and a strengthening of the link between dispersal and some phenotypic traits with environmental harshness. Our proof-of-concept metacommunity model further reveals cascading effects of context-dependent syndromes on the local and regional organisation of functional diversity. Our study opens new avenues to advance our understanding of the functioning of spatially structured populations, communities and ecosystems.

The evolution of trait variance creates a tension between species diversity and functional diversity.

It seems intuitively obvious that species diversity promotes functional diversity: communities with more plant species imply more varied plant leaf chemistry, more species of crops provide more kinds of food, etc. Recent literature has nuanced this view, showing how the relationship between the two can be modulated along latitudinal or environmental gradients. Here we show that even without such effects, the evolution of functional trait variance can erase or even reverse the expected positive relationship between species- and functional diversity. We present theory showing that trait-based eco-evolutionary processes force species to evolve narrower trait breadths in more tightly packed, species-rich communities, in their effort to avoid competition with neighboring species. This effect is so strong that it leads to an overall reduction in trait space coverage whenever a new species establishes. Empirical data from land snail communities on the Galápagos Islands are consistent with this claim. The finding that the relationship between species- and functional diversity can be negative implies that trait data from species-poor communities may misjudge functional diversity in species-rich ones, and vice versa.

Interactive effects of multiple stressors vary with consumer interactions, stressor dynamics and magnitude.

Predicting the impacts of multiple stressors is important for informing ecosystem management but is impeded by a lack of a general framework for predicting whether stressors interact synergistically, additively or antagonistically. Here, we use process-based models to study how interactions generalise across three levels of biological organisation (physiological, population and consumer-resource) for a two-stressor experiment on a seagrass model system. We found that the same underlying processes could result in synergistic, additive or antagonistic interactions, with interaction type depending on initial conditions, experiment duration, stressor dynamics and consumer presence. Our results help explain why meta-analyses of multiple stressor experimental results have struggled to identify predictors of consistently non-additive interactions in the natural environment. Experiments run over extended temporal scales, with treatments across gradients of stressor magnitude, are needed to identify the processes that underpin how stressors interact and provide useful predictions to management.

Biodiversity promotes ecosystem functioning despite environmental change.

Three decades of research have demonstrated that biodiversity can promote the functioning of ecosystems. Yet, it is unclear whether the positive effects of biodiversity on ecosystem functioning will persist under various types of global environmental change drivers. We conducted a meta-analysis of 46 factorial experiments manipulating both species richness and the environment to test how global change drivers (i.e. warming, drought, nutrient addition or CO2 enrichment) modulated the effect of biodiversity on multiple ecosystem functions across three taxonomic groups (microbes, phytoplankton and plants). We found that biodiversity increased ecosystem functioning in both ambient and manipulated environments, but often not to the same degree. In particular, biodiversity effects on ecosystem functioning were larger in stressful environments induced by global change drivers, indicating that high-diversity communities were more resistant to environmental change. Using a subset of studies, we also found that the positive effects of biodiversity were mainly driven by interspecific complementarity and that these effects increased over time in both ambient and manipulated environments. Our findings support biodiversity conservation as a key strategy for sustainable ecosystem management in the face of global environmental change.

Stressor richness intensifies productivity loss but mitigates biodiversity loss.

Ecosystems are subject to a multitude of anthropogenic environmental changes. Experimental research in the field of multiple stressors has typically involved varying the number of stressors, here termed stressor richness, but without controlling for total stressor intensity. Mistaking stressor intensity effects for stressor richness effects can misinform management decisions when there is a trade-off between mitigating these two factors. We incorporate multiple stressors into three community models and show that, at a fixed total stressor intensity, increasing stressor richness aggravates joint stressor effects on ecosystem functioning, but reduces effects on species persistence and composition. In addition, stressor richness weakens the positive selection and negative complementarity effects on ecosystem function. We identify the among-species variation of stressor effects on traits as a key determinant of the resulting community-level stressor effects. Taken together, our results unravel the mechanisms linking multiple environmental changes to biodiversity and ecosystem function.

Species richness increases fitness differences, but does not affect niche differences.

A key question in ecology is what limits species richness. Modern coexistence theory presents the persistence of species as a balance between niche differences and fitness differences that favour and hamper coexistence, respectively. With most applications focusing on species pairs, however, we know little about if and how this balance changes with species richness. Here, we apply recently developed definitions of niche and fitness differences, based on invasion analysis, to multispecies communities. We present the first mathematical proof that, for invariant average interaction strengths, the average fitness difference among species increases with richness, while the average niche difference stays constant. Extensive simulations with more complex models and analyses of empirical data confirmed these mathematical results. Combined, our work suggests that, as species accumulate in ecosystems, ever-increasing fitness differences will at some point exceed constant niche differences, limiting species richness. Our results contribute to a better understanding of coexistence multispecies communities.

Reinterpreting the relationship between number of species and number of links connects community structure and stability.

For 50 years, ecologists have examined how the number of interactions (links) scales with the number of species in ecological networks. Here, we show that the way the number of links varies when species are sequentially removed from a community is fully defined by a single parameter identifiable from empirical data. We mathematically demonstrate that this parameter is network-specific and connects local stability and robustness, establishing a formal connection between community structure and two prime stability concepts. Importantly, this connection highlights a local stability-robustness trade-off, which is stronger in mutualistic than in trophic networks. Analysis of 435 empirical networks confirmed these results. We finally show how our network-specific approach relates to the classical across-network approach found in literature. Taken together, our results elucidate one of the intricate relationships between network structure and stability in community networks.

The topology and drivers of ant-symbiont networks across Europe.

Intimate associations between different species drive community composition across ecosystems. Understanding the ecological and evolutionary drivers of these symbiotic associations is challenging because their structure eventually determines stability and resilience of the entire species network. Here, we compiled a detailed database on naturally occurring ant-symbiont networks in Europe to identify factors that affect symbiont network topology. These networks host an unrivalled diversity of macrosymbiotic associations, spanning the entire mutualism-antagonism continuum, including: (i) myrmecophiles - commensalistic and parasitic arthropods; (ii) trophobionts - mutualistic aphids, scale insects, planthoppers and caterpillars; (iii) social parasites - parasitic ant species; (iv) parasitic helminths; and (v) parasitic fungi. We dissected network topology to investigate what determines host specificity, symbiont species richness, and the capacity of different symbiont types to switch hosts. We found 722 macrosymbionts (multicellular symbionts) associated with European ants. Symbiont type explained host specificity and the average relatedness of the host species. Social parasites were associated with few hosts that were phylogenetically highly related, whereas the other symbiont types interacted with a larger number of hosts across a wider taxonomic distribution. The hosts of trophobionts were the least phylogenetically related across all symbiont types. Colony size, host range and habitat type predicted total symbiont richness: ant hosts with larger colony size, a larger distribution range or with a wider habitat range contained more symbiont species. However, we found that different sets of host factors affected diversity in the different types of symbionts. Ecological factors, such as colony size, host range and niche width predominantly determined myrmecophile species richness, whereas host phylogeny was the most important predictor of mutualistic trophobiont, social parasite and parasitic helminth species richness. Lastly, we found that hosts with a common biogeographic history support a more similar community of symbionts. Phylogenetically related hosts also shared more trophobionts, social parasites and helminths, but not myrmecophiles. Taken together, these results suggest that ecological and evolutionary processes structure host specificity and symbiont richness in large-scale ant-symbiont networks, but these drivers may shift in importance depending on the type of symbiosis. Our findings highlight the potential of well-characterized bipartite networks composed of different types of symbioses to identify candidate processes driving community composition.

Biodiversity mediates the effects of stressors but not nutrients on litter decomposition.

Understanding the consequences of ongoing biodiversity changes for ecosystems is a pressing challenge. Controlled biodiversity-ecosystem function experiments with random biodiversity loss scenarios have demonstrated that more diverse communities usually provide higher levels of ecosystem functioning. However, it is not clear if these results predict the ecosystem consequences of environmental changes that cause non-random alterations in biodiversity and community composition. We synthesized 69 independent studies reporting 660 observations of the impacts of two pervasive drivers of global change (chemical stressors and nutrient enrichment) on animal and microbial decomposer diversity and litter decomposition. Using meta-analysis and structural equation modeling, we show that declines in decomposer diversity and abundance explain reduced litter decomposition in response to stressors but not to nutrients. While chemical stressors generally reduced biodiversity and ecosystem functioning, detrimental effects of nutrients occurred only at high levels of nutrient inputs. Thus, more intense environmental change does not always result in stronger responses, illustrating the complexity of ecosystem consequences of biodiversity change. Overall, these findings provide strong evidence that the consequences of observed biodiversity change for ecosystems depend on the kind of environmental change, and are especially significant when human activities decrease biodiversity.

Ecosystems are at their healthiest when they have a high level of biodiversity ­ that is, a wide variety of different species all living together. But human activity is changing the environment and putting ecosystems under pressure. One of the places this is most evident is in the communities of organisms responsible for breaking down dead plants. These organisms ­ called decomposers ­ are highly sensitive to pesticides, metals and other chemical stressors, as well as excess nutrients, such as nitrogen, released by industry and farming. Exposing decomposers to these chemicals can change both the number of individuals of each species and the number of different species that are present. In other words, these chemicals can, respectively, alter both the abundance and diversity of decomposer communities. Controlled experiments in simplified conditions suggest that these changes in biodiversity affect ecosystem health. But, it remained unclear to what extent these results applied to real-world scenarios of environmental change. To test the findings of controlled experiments, Beaumelle et al. investigated how chemical stressors and excess nutrients affect the breakdown of leaf litter ­ the debris of decomposing leaves that forms on top of soil. Previous studies suggest that the reduced biodiversity caused by chemicals should result in leaf litter breaking down more slowly. Whereas the loss in biodiversity caused by nutrients will increase the number of some decomposer species, causing leaf litter to break down faster or slower, depending on local conditions. Beaumelle et al. tested these predictions by gathering the results from 69 independent studies conducted across the globe. The results showed that stressors caused the diversity and abundance of decomposers to decline, which reduced the breakdown of leaf litter, as expected. But, the outcomes of excess nutrients were more varied. Low levels of excess nutrients increased the breakdown of leaf litter, but at high levels slowed down the rate leaves decomposed. Furthermore, the effect excess nutrients had on biodiversity in decomposer communities changed according to the types of organisms in the ecosystem. This suggests that variations in biodiversity can impact ecosystems differently depending on the type of environmental change. The breakdown of leaf litter plays a critical role in carbon balance, and this has knock-on effects for the Earth's climate. This work suggests that improving biodiversity is crucial to maintain the health of ecosystems, but successful strategies will have to be adjusted depending on the type of human impact (for example, chemical stressors or nutrient additions). These findings could help researchers design better approaches for boosting ecosystem health in the future.

Community composition modifies direct and indirect effects of pesticides in freshwater food webs.

For environmental risk assessment, the effects of pesticides on aquatic ecosystems are often assessed based on single species tests, disregarding the potential influence of community composition. We, therefore, studied the influence of changing the horizontal (the number of species within trophic levels) and vertical composition (number of trophic levels) on the ecological effects of the herbicide linuron and the insecticide chlorpyrifos, targeting producers and herbivores, respectively. We tested how adding, to a single primary producer, 4 selected competing producer species, 0-1-4 selected herbivore species, and one selected predator species resulting in 1, 2 and 3 trophic levels, changes the effects of the two pesticides. Linuron decreased producer biovolume less (17%) when the 4 producers were added, because insensitive producers compensated for the loss of sensitive producers. However, linuron decreased producer biovolume 42% and 32% more as we increased the number of herbivore species from 0 to 4 and as we increased trophic levels from 1 to 3, respectively. The indirect negative effect of linuron on herbivore biovolume was 11% and 15% lower when more producer and herbivores were added, respectively. Adding a predator increased this indirect negative effect by 22%. Chlorpyrifos decreased herbivore biovolume about 10% less when adding multiple herbivore or producer species. However, adding a predator magnified the direct negative impact on herbivores (13%). Increasing the number of producer, herbivore species and adding trophic levels increased the indirect positive impact on producer biovolume (between 10% and 35%). Our study shows that changing horizontal composition can both increase and decrease the effects of the selected pesticides, while changing vertical composition by adding number of trophic levels always increased these effects. Therefore, single species sensitivity will not always represent a worst case estimate of ecological effects. Protecting the most sensitive species may not ensure protection of ecosystems.

Potential impact of chemical stress on freshwater invertebrates: A sensitivity assessment on continental and national scale based on distribution patterns, biological traits, and relatedness.

Current chemical risk assessment approaches rely on a standard suite of test species to assess toxicity to environmental species. Assessment factors are used to extrapolate from single species to communities and ecosystem effects. This approach is pragmatic, but lacks resolution in biological and environmental parameters. Novel modelling approaches can help improve the biological resolution of assessments by using mechanistic information to identify priority species and priority regions that are potentially most impacted by chemical stressors. In this study we developed predictive sensitivity models by combining species-specific information on acute chemical sensitivity (LC50 and EC50), traits, and taxonomic relatedness. These models were applied at two spatial scales to reveal spatial differences in the sensitivity of species assemblages towards two chemical modes of action (MOA): narcosis and acetylcholinesterase (AChE) inhibition. We found that on a relative scale, 46% and 33% of European species were ranked as more sensitive towards narcosis and AChE inhibition, respectively. These more sensitive species were distributed with higher occurrences in the south and north-eastern regions, reflecting known continental patterns of endemic macroinvertebrate biodiversity. We found contradicting sensitivity patterns depending on the MOA for UK scenarios, with more species displaying relative sensitivity to narcotic MOA in north and north-western regions, and more species with relative sensitivity to AChE inhibition MOA in south and south-western regions. Overall, we identified hotspots of species sensitive to chemical stressors at two spatial scales, and discuss data gaps and crucial technological advances required for the successful application of the proposed methodology to invertebrate scenarios, which remain underrepresented in global conservation priorities.

Towards a unified study of multiple stressors: divisions and common goals across research disciplines.

Anthropogenic environmental changes, or 'stressors', increasingly threaten biodiversity and ecosystem functioning worldwide. Multiple-stressor research is a rapidly expanding field of science that seeks to understand and ultimately predict the interactions between stressors. Reviews and meta-analyses of the primary scientific literature have largely been specific to either freshwater, marine or terrestrial ecology, or ecotoxicology. In this cross-disciplinary study, we review the state of knowledge within and among these disciplines to highlight commonality and division in multiple-stressor research. Our review goes beyond a description of previous research by using quantitative bibliometric analysis to identify the division between disciplines and link previously disconnected research communities. Towards a unified research framework, we discuss the shared goal of increased realism through both ecological and temporal complexity, with the overarching aim of improving predictive power. In a rapidly changing world, advancing our understanding of the cumulative ecological impacts of multiple stressors is critical for biodiversity conservation and ecosystem management. Identifying and overcoming the barriers to interdisciplinary knowledge exchange is necessary in rising to this challenge. Division between ecosystem types and disciplines is largely a human creation. Species and stressors cross these borders and so should the scientists who study them.

Intuitive and broadly applicable definitions of niche and fitness differences.

Explaining nature's biodiversity is a key challenge for science. To persist, populations must be able to grow faster when rare, a feature called negative frequency dependence and quantified as 'niche differences' ( N ) in modern coexistence theory. Here, we first show that available definitions of N differ in how N link to species interactions, are difficult to interpret and often apply to specific community types only. We then present a new definition of N that is intuitive and applicable to a broader set of (modelled and empirical) communities than is currently the case, filling a main gap in the literature. Given N , we also redefine fitness differences ( F ) and illustrate how N and F determine coexistence. Finally, we demonstrate how to apply our definitions to theoretical models and experimental data, and provide ideas on how they can facilitate comparison and synthesis in community ecology.

Quantifying the primary biotic resource use by fisheries: A global assessment.

In this paper, the specific primary production required (SPPR expressed as kg-NPP/kg-fish in wet weight) of more than 1700 marine species were calculated directly from 96 published food web models using the newly developed SPPR calculation framework. The relationship between SPPR and other ecological factors were then statistically analyzed. Among- and within-species variability of SPPR were found to be both explained by trophic level (TL), suggesting similar mechanisms underpinning both sources of variability. Among species, we found that harvesting species at higher mean trophic levels (MTL) increases the mean SPPR by a factor of 19 per 1 unit increase in MTL. Based on our empirical relationship, the mean SPPR of more than 9000 marine species were predicted and subsequently used to assess the primary production required (PPR) to support fisheries in five major fishing countries in Europe. The results indicated that conventional approach to estimating PPR, which neglects food web ecology, can underestimate PPR by up to a factor of 5. Within species, we found that harvesting populations occupying a higher TL leads to a higher SPPR. For example, the SPPR of Atlantic cod in the Celtic Sea (TL = 4.75) was 5 times higher than in the Gilbert Bay (TL = 3.3). Our results, which are based on large amounts of field data, highlight the importance of properly accounting for ecological factors during the impact assessment of fisheries.

Food Web Uncertainties Influence Predictions of Climate Change Effects on Soil Carbon Sequestration in Heathlands.

Carbon cycling models consider soil carbon sequestration a key process for climate change mitigation. However, these models mostly focus on abiotic soil processes and, despite its recognized critical mechanistic role, do not explicitly include interacting soil organisms. Here, we use a literature study to show that even a relatively simple soil community (heathland soils) contains large uncertainties in temporal and spatial food web structure. Next, we used a Lotka-Volterra-based food web model to demonstrate that, due to these uncertainties, climate change can either increase or decrease soil carbon sequestration to varying extents. Both the strength and direction of changes strongly depend on (1) the main consumer's (enchytraeid worms) feeding preferences and (2) whether decomposers (fungi) or enchytraeid worms are more sensitive to stress. Hence, even for a soil community with a few dominant functional groups and a simulation model with a few parameters, filling these knowledge gaps is a critical first step towards the explicit integration of soil food web dynamics into carbon cycling models in order to better assess the role soils play in climate change mitigation.