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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.

Human impacts such as habitat loss, climate change and biological invasions are radically altering biodiversity, with greater effects projected into the future. Evidence suggests human impacts may differ substantially between terrestrial and freshwater ecosystems, but the reasons for these differences are poorly understood. We propose an integrative approach to explain these differences by linking impacts to four fundamental processes that structure communities: dispersal, speciation, species-level selection and ecological drift. Our goal is to provide process-based insights into why human impacts, and responses to impacts, may differ across ecosystem types using a mechanistic, eco-evolutionary comparative framework. To enable these insights, we review and synthesise (i) how the four processes influence diversity and dynamics in terrestrial versus freshwater communities, specifically whether the relative importance of each process differs among ecosystems, and (ii) the pathways by which human impacts can produce divergent responses across ecosystems, due to differences in the strength of processes among ecosystems we identify. Finally, we highlight research gaps and next steps, and discuss how this approach can provide new insights for conservation. By focusing on the processes that shape diversity in communities, we aim to mechanistically link human impacts to ongoing and future changes in ecosystems., Ecology Letters, 26 (2), ISSN:1461-023X, ISSN:1461-0248

Lakes are often described as sentinels of global change. Phenomena like lake eutrophication, algal blooms, or reorganization in community composition belong to the most studied ecosystem regime shifts. However, although regime shifts have been well documented in several lakes, a global assessment of the prevalence of regime shifts is still missing, and, more in general, of the factors altering stability in lake status, is missing. Here, we provide a first global assessment of regime shifts and stability in the productivity of 1,015 lakes worldwide using trophic state index (TSI) time series derived from satellite imagery. We find that 12.8% of the lakes studied show regime shifts whose signatures are compatible with tipping points, while the number of detected regime shifts from low to high TSI has increased over time. Although our results suggest an overall stable picture for global lake dynamics, the limited instability signatures do not mean that lakes are insensitive to global change. Modeling the interaction between lake climatic, geophysical, and socioeconomic features and their stability properties, we find that the probability of a lake experiencing a tipping point increases with human population density in its catchment, while it decreases as the gross domestic product of that population increases. Our results show how quantifying lake productivity dynamics at a global scale highlights socioeconomic inequalities in conserving natural environments., Proceedings of the National Academy of Sciences, 119 (35), ISSN:0027-8424, ISSN:1091-6490

Competition for limited resources is a major force in structuring ecological communities. Species minimum resource requirements (R*s) can predict competitive outcomes and evolve under selection in simple communities under controlled conditions. However, whether R*s predict competitive outcomes or demonstrate adaptive evolution in naturally complex communities is unknown. We subjected natural phytoplankton communities to three types of resource limitation (nitrogen, phosphorus, light) in outdoor mesocosms over 10 weeks. We examined the community composition weekly and isolated 21 phytoplankton strains from seven species to quantify responses to the selection of R* for these resources. We investigated the evolutionary change in R*s in the dominant species, Desmodesmus armatus. R*s were good predictors of species changes in relative abundance, though this was largely driven by the success of D. armatus across several treatments. This species also demonstrated an evolutionary change in R*s under resource limitation, supporting the potential for adaptive trait change to modify competitive outcomes in natural communities., Ecology Letters, 25 (11), ISSN:1461-023X, ISSN:1461-0248

Biodiversity loss and climate warming are occurring in concert, with potentially profound impacts on ecosystem functioning. We currently know very little about the combined effects of these changes on the links between the community structure, dynamics and the resulting in situ CO2 concentrations in freshwater ecosystems. Here we aimed to determine both individual and combined effects of temperature and non-resource diversity (species inedible for a given consumer) on CO2 concentration. Our analysis further aimed to establish both direct effects on CO2 concentrations and potential indirect effects that occur via changes to the phytoplankton and zooplankton biomasses. Our results showed that there were no interactive effects of changes in temperature and diversity on CO2 concentration in the water. Instead, independent increases in either temperature or non-resource diversity resulted in a substantial reduction in CO2 concentrations, particularly at the highest non-resource diversity. The effects of non-resource diversity and warming on CO2 were indirect, resulting largely from the positive impacts on total biomass of primary producers. Our study is the first to experimentally partition the impacts of temperature and diversity on the consumer–resource dynamics and associated changes to CO2 concentrations. It provides new mechanistic insights into the role of diverse plankton communities for ecosystem functioning and their importance in regulating CO2 dynamics under ongoing climate warming.

Eutrophication is a persistent threat to aquatic ecosystems worldwide. Foundation species, namely those that play a central role in the structuring of communities and functioning of ecosystems, are likely important for the resilience of aquatic ecosystems in the face of disturbance. However, little is known about how interactions among such species influence ecosystem responses to nutrient perturbation. Here, using an array (N = 20) of outdoor experimental pond ecosystems (15,000 L), we manipulated the presence of two foundation species, the macrophyte Myriophyllum spicatum and the mussel Dreissena polymorpha, and quantified ecosystem responses to multiple nutrient disturbances, spread over two years. In the first year, we added five nutrient pulses, ramping up from 10 to 50 μg P/L over a 10-week period from mid-July to mid-October, and in the second year, we added a single large pulse of 50 μg P/L in mid-October. We used automated sondes to measure multiple ecosystems properties at high frequency (15-minute intervals), including phytoplankton and dissolved organic matter fluorescence, and to model whole-ecosystem metabolism. Overall, both foundation species strongly affected the ecosystem responses to nutrient perturbation, and, as expected, initially suppressed the increase in phytoplankton abundance following nutrient additions. However, when both species were present, phytoplankton biomass increased substantially relative to other treatment combinations: non-additivity was evident for multiple ecosystem metrics following the nutrient perturbations in both years but was diminished in the intervening months between our perturbations. Overall, these results demonstrate how interactions between foundation species can cause surprisingly strong deviations from the expected responses of aquatic ecosystems to perturbations such as nutrient additions.

Competition for limiting resources is among the most fundamental ecological interactions and has long been considered a key driver of species coexistence and biodiversity. Species' minimum resource requirements, their R *s, are key traits that link individual physiological demands to the outcome of competition. However, a major question remains unanswered—to what extent are species’ competitive traits able to evolve in response to resource limitation? To address this knowledge gap, we performed an evolution experiment in which we exposed Chlamydomonas reinhardtii for approximately 285 generations to seven environments in chemostats that differed in resource supply ratios (including nitrogen, phosphorus and light limitation) and salt stress. We then grew the ancestors and descendants in a common garden and quantified their competitive abilities for essential resources. We investigated constraints on trait evolution by testing whether changes in resource requirements for different resources were correlated. Competitive abilities for phosphorus improved in all populations, while competitive abilities for nitrogen and light increased in some populations and decreased in others. In contrast to the common assumption that there are trade-offs between competitive abilities for different resources, we found that improvements in competitive ability for a resource came at no detectable cost. Instead, improvements in competitive ability for multiple resources were either positively correlated or not significantly correlated. Using resource competition theory, we then demonstrated that rapid adaptation in competitive traits altered the predicted outcomes of competition. These results highlight the need to incorporate contemporary evolutionary change into predictions of competitive community dynamics over environmental gradients. This article is part of the theme issue ‘Conceptual challenges in microbial community ecology’.

Evolution is a fundamental ecosystem process. The study of genomic variation of organisms can not only improve our understanding of evolutionary processes, but also of contemporary and future ecosystem dynamics. We argue that integrative research between the fields of genomics and ecosystem ecology could generate new insights. Specifically, studies of biodiversity and ecosystem functioning, evolutionary rescue, and eco-evolutionary dynamics could all benefit from information about variation in genome structure and the genetic architecture of traits, whereas genomic studies could benefit from information about the ecological context of evolutionary dynamics. We propose new ways to help link research on functional genomic diversity with (reciprocal) interactions between phenotypic evolution and ecosystem change. Despite numerous challenges, we anticipate that the wealth of genomic data being collected on natural populations will improve our understanding of ecosystems.

The research of a generation of ecologists was catalysed by the recognition that the number and identity of species in communities influences the functioning of ecosystems. The relationship between biodiversity and ecosystem functioning (BEF) is most often examined by controlling species richness and randomising community composition. In natural systems, biodiversity changes are often part of a bigger community assembly dynamic. Therefore, focusing on community assembly and the functioning of ecosystems (CAFE), by integrating both species richness and composition through species gains, losses and changes in abundance, will better reveal how community changes affect ecosystem function. We synthesise the BEF and CAFE perspectives using an ecological application of the Price equation, which partitions the contributions of richness and composition to function. Using empirical examples, we show how the CAFE approach reveals important contributions of composition to function. These examples show how changes in species richness and composition driven by environmental perturbations can work in concert or antagonistically to influence ecosystem function. Considering how communities change in an integrative fashion, rather than focusing on one axis of community structure at a time, will improve our ability to anticipate and predict changes in ecosystem function. ISSN:1461-023X ISSN:1461-0248

For algal biofuels to be economically sustainable and avoid exacerbating nutrient pollution, algal cultivation and processing must maximize rates of biofuel production while simultaneously minimizing the consumption of nitrogen (N) and phosphorus (P) fertilizers. We experimentally tested whether algal polycultures could be engineered to improve N and P nutrient-use efficiency compared to monocultures by balancing trade-offs in nutrient-use efficiency and biocrude production. We analyzed the flows of N and P through the processes of cultivation, biocrude production through hydrothermal liquefaction, and nutrient recycling in a laboratory-scale system. None of the six species we examined exhibited high N efficiency, P efficiency, and biocrude production simultaneously; each had poor performance in at least one function (i.e.,25th percentile). Polycultures of two to six species did not outperform the best species in any single function, but some polycultures exhibited more balanced performance and maintained all three functions at higher levels simultaneously than any of the monocultures (i.e.,67th percentile). Moreover, certain polycultures came closer to optimizing all three functions than any of the monocultures. By balancing trade-offs between N and P efficiency and biocrude production, polycultures could be used to simultaneously reduce the demand for both N and P fertilizers by up to 85%.

We compared several measures of quality for biomass and biocrude oil produced via hydrothermal liquefaction for monocultures and polycultures of the freshwater microalgae Ankistrodesmus falcatus, Chlorella sorokiniana, Pediastrum duplex, Scenedesmus acuminatus, Scenedesmus ecornis, and Selenastrum capricornutum. On average, the 2-species cultures provided product quality comparable to that of the monocultures, while that of the average 4- and 6-species polycultures was lower. No single monoculture or polyculture performed the best with respect to all quality metrics considered, including biomass fatty acid content and biocrude elemental content and higher heating value. However, for each measure of quality, some polycultures did outperform or match the performance of the best monoculture for each respective metric. Numerous polycultures performed outside the range of product quality exhibited by their constituent species (transgressive overyielding), and polycultures with P. duplex did so to the greatest extent. Microalgal biochemical composition was correlated with biocrude properties, however including species identity as an additional correlation variable further explained the observed variation. Finally, we highlight several polycultures that offer potentially compelling trade-offs between product quantity and quality compared to the most productive monoculture.

Summary Phenotypic variation controls the species interactions which determine whether or not species coexist. Long-standing hypotheses in ecology and evolution posit that phenotypic differentiation enables coexistence by increasing the size of niche differentiation. This hypothesis has only been tested using macroscopic traits to date, but niche differentiation, particularly of microscopic organisms, also occurs at the molecular and metabolic level. We examined how phenotypic variation that arises at the level of gene expression over evolutionary time affects phytoplankton species interactions and coexistence. We predicted that similarity in gene expression among species would decline with phylogenetic distance, and that reduced similarity in gene expression would weaken competition, increase facilitation and promote coexistence. To test this, we grew eight species of freshwater green algae in monocultures and bicultures for 46 days in a laboratory microcosm experiment. We quantified the strength of species interactions by: (i) fitting Lotka–Volterra models to time-series densities and estimating interaction coefficients, and (ii) calculating relative densities that compare species’ steady-state densities in biculture to those in monoculture. We used Illumina high throughput sequencing to quantify the expression of 1253 families of homologous genes, including a set of 17 candidate genes that we hypothesized a priori to be involved in competition or facilitation. Synthesis. We found that closely related species had greater similarity in gene expression than did distantly related species, but as gene expression became more similar, species experienced weaker competition or greater facilitation, and were more likely to coexist. We identified gene functional categories that were uniquely differentially regulated in association with particular species interaction types. Contrary to common thinking in ecology and evolution, similarity in gene expression, and not differentiation, was associated with weaker competition, facilitation and coexistence.

A major challenge in ecology is to understand determinants of ecosystem functioning and stability in the face of disturbance. Some important species can strongly shape community structure and ecosystem functioning, but their impacts and interactions on ecosystem-level responses to disturbance are less well known. Shallow ponds provide a model system in which to study the effects of such species because some taxa mitigate transitions between alternative ecosystem states caused by eutrophication. We performed pond experiments to test how two foundation species (a macrophyte and a mussel) affected the biomass of planktonic primary producers and its stability in response to nutrient additions. Individually, each species reduced phytoplankton biomass and tended to increase rates of recovery from disturbance, but together the species reversed these effects, particularly with larger nutrient additions. This reversal was mediated by high cyanobacterial dominance of the community and a resulting loss of trait evenness. Effects of the foundation species on primary producer biomass were associated with effects on other ecosystem properties, including turbidity and dissolved oxygen. Our work highlights the important role of foundation species and their interactive effects in determining responses of ecosystem functioning to disturbance.

International audience; Species interactions lie at the heart of many theories of macroevolution, from adaptive radiation to the Red Queen. Although some theories describe the imprint that interactions will have over long timescales, we are still missing a comprehensive understanding of the effects of interactions on macroevolution. Current research shows strong evidence for the impact of interactions on macroevolutionary patterns of trait evolution and diversification, yet many macroevolutionary studies have only a tenuous relationship to ecological studies of interactions over shorter timescales. We review current research in this area, highlighting approaches that explicitly model species interactions and connect them to broad-scale macroevolutionary patterns. We also suggest that progress has been made by taking an integrative interdisciplinary look at individual clades. We focus on African cichlids as a case study of how this approach can be fruitful. Overall, although the evidence for species interactions shaping macroevolution is strong, further work using integrative and model-based approaches is needed to spur progress towards understanding the complex dynamics that structure communities over time and space.

Biodiversity loss and climate warming are occurring in concert, with potentially profound impacts on ecosystem functioning. We currently know very little about the combined effects of these changes on the links between the community structure, dynamics and the resulting in situ CO2 concentrations in freshwater ecosystems. Here we aimed to determine both individual and combined effects of temperature and non-resource diversity (species inedible for a given consumer) on CO2 concentration. Our analysis further aimed to establish both direct effects on CO2 concentrations and potential indirect effects that occur via changes to the phytoplankton and zooplankton biomasses. Our results showed that there were no interactive effects of changes in temperature and diversity on CO2 concentration in the water. Instead, independent increases in either temperature or non-resource diversity resulted in a substantial reduction in CO2 concentrations, particularly at the highest non-resource diversity. The effects of non-resource diversity and warming on CO2 were indirect, resulting largely from the positive impacts on total biomass of primary producers. Our study is the first to experimentally partition the impacts of temperature and diversity on the consumer–resource dynamics and associated changes to CO2 concentrations. It provides new mechanistic insights into the role of diverse plankton communities for ecosystem functioning and their importance in regulating CO2 dynamics under ongoing climate warming. Electronic supplementary material The online version of this article (10.1007/s00442-020-04593-0) contains supplementary material, which is available to authorized users.

Resource competition theory is a conceptual framework that provides mechanistic insights into competition and community assembly of species with different resource requirements. However, there has been little exploration of how resource requirements depend on other environmental factors, including temperature. Changes in resource requirements as influenced by environmental temperature would imply that climate warming can alter the outcomes of competition and community assembly.We experimentally demonstrate that environmental temperature alters the minimum light and nitrogen requirements – as well as other growth parameters – of six widespread phytoplankton species from distinct taxonomic groups. We found that species require the most nitrogen at the highest temperatures while light requirements tend to be lowest at intermediate temperatures, although there are substantial interspecific differences in the exact shape of this relationship.We also experimentally parameterize two competition models, which we use to illustrate how temperature, through its effects on species’ traits, alters competitive hierarchies in multispecies assemblages, determining community dynamics.Developing a mechanistic understanding of how temperature influences the ability to compete for limiting resources is a critical step towards improving forecasts of community dynamics under climate warming.

Drawing insights from multiple disciplines is essential for finding integrative solutions that are required to tackle complex environmental problems. Human activities are causing unprecedented influence on global ecosystems, culminating in the loss of species and fundamental changes in the selective environments of organisms across the tree of life. Our collective understanding about biological evolution can help identify and mitigate many of the environmental problems in the Anthropocene. To this end, we propose a stronger integration of environmental sciences with evolutionary biology., Graphical abstract Unlabelled Image, Highlights • Evolutionary theory is not widely used to address environmental problems. • Evolutionary processes can both ameliorate and exacerbate environmental problems. • Evolution can be informative about disease dynamics, biofuel production, adaptation to pollution, and harmful algal blooms. • We propose a stronger integration of evolutionary biology with environmental sciences to help solve environmental problems.

Algal biofuels have the potential to curb the emissions of greenhouse gases from fossil fuels, but current growing methods fail to produce fuels that meet the multiple standards necessary for economical industrial use. For example, algae grown as monocultures for biofuel production have not simultaneously and economically achieved high yields of the high-quality lipid-rich biomass desired for the industrial-scale production of bio-oil. Decades of study in the field of ecology have demonstrated that simultaneous increases in multiple functions, such as the quantity and quality of biomass, can occur in natural ecosystems by increasing biological diversity. Here, we show that species consortia of algae can improve the production of bio-oil, which benefits from both a high biomass yield and a high quality of biomass rich in fatty acids. We explain the underlying causes of increased quantity and quality of algal biomass among species consortia by showing that, relative to monocultures, species consortia can differentially regulate lipid metabolism genes while growing to higher levels of biomass, in part due to a greater utilization of nutrient resources. We identify multiple genes involved in lipid biosynthesis that are frequently upregulated in bicultures and further show that these elevated levels of gene expression are highly predictive of the elevated levels in biculture relative to that in monoculture of multiple quality metrics of algal biomass. These results show that interactions between species can alter the expression of lipid metabolism genes and further demonstrate that our understanding of diversity-function relationships from natural ecosystems can be harnessed to improve the production of bio-oil. IMPORTANCE Algal biofuels are one of the more promising forms of renewable energy. In our study, we investigate whether ecological interactions between species of microalgae regulate two important factors in cultivation—the biomass of the crop produced and the quality of the biomass that is produced. We found that species interactions often improved production yields, especially the fatty acid content of the algal biomass, and that differentially expressed genes involved in fatty acid metabolism are predictive of improved quality metrics of bio-oil. Other studies have found that diversity often improves productivity and stability in agricultural and natural ecosystems. Our results provide further evidence that growing multispecies crops of microalgae may improve the production of high-quality biomass for bio-oil.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Recent meta-analyses indicate that an increase in the number of natural enemy species raises the overall effectiveness of biological control of insect pests in agricultural settings. Although economic valuations of biological pest control in agriculture show an impressive cost savings from this ecosystem service, no direct estimate of the value of biodiversity for biological control has been attempted. We apply a basic microeconomic model for estimating the value of changes in the richness of arthropod natural enemies or in functional diversity for biological pest control for two sample crops using experimental results reported in the ecological literature. Market-based outcomes were driven by changes in crop yields associated with experimental reductions in natural enemy species richness, and modified by supply shifts and price elasticities. We show how our simple model differs from common approaches used in the ecological literature, and explain why this alternative model more accurately estimates societal well-being for consumers and producers participating in these crop markets. We conclude by discussing the additional research and data needed to make economic valuation of ecosystem services in agricultural settings more feasible, rigorous, and realistic in the future.

Hundreds of experiments have now manipulated species richness of various groups of organisms and examined how this aspect of biological diversity influences ecosystem functioning. Ecologists have recently expanded this field to look at whether genetic diversity among species, often quantified as the degree of evolutionary divergence on a molecular phylogeny, also predicts ecological function. Some have hypothesized that phylogenetic divergence should be a superior predictor of ecological function than species richness because evolutionary relatedness represents the degree of ecological and functional differentiation among species. But studies to date have provided mixed support for this hypothesis. Here, we re-analyze data from 16 experiments that have manipulated plant species richness in grassland ecosystems and examined the impact on aboveground biomass production over multiple time points. Using a new molecular phylogeny of the plant species used in these experiments, we quantified how the phylogenetic diversity of plants impacts average community biomass production as well as the stability of community biomass production through time. Using four complementary analyses we show that, after statistically controlling for variation in species richness, phylogenetic diversity (the sum of branches in a molecular phylogenetic tree connecting all species in a community) is neither related to mean community biomass nor to the temporal stability of biomass. These results run counter to past claims. After controlling for species richness, phylogenetic diversity was positively related to variation in community biomass over time via an increase in individual species variances but this relationship was not strong enough to influence community stability. In contrast to the non-significant relationships between phylogenetic diversity, biomass, and stability, after controlling for species richness, our analyses show that species richness tends to increase the mean biomass production of plant communities, after controlling for phylogenetic diversity. The relationship between species richness and temporal variation in community biomass was either non-significant or negative depending on which analysis was used. However, the increases in community biomass with species richness, independent of phylogenetic diversity, always led to increased stability. These results suggest that phylogenetic diversity is no better as a predictor of ecosystem functioning than species richness. Synthesis. Our study on grasslands offers a cautionary tale when trying to relate phylogenetic diversity to ecosystem functioning as there may be trait and functional variation among species that cannot be explained by their evolutionary relatedness. Management recommendations suggesting the conservation of evolutionarily distinct species will lead to more productive and more stable communities under changing environmental conditions are not well supported by the data explored in this study. Fil: Venail, Patrick. University of Michigan. School of Natural Resources and Environment; Estados Unidos. University of Geneva. Institute F.-A. Forel. Section of Earth and Environmental Sciences; Suiza Fil: Gross, Kevin. North Carolina State University. Statistics Department; Estados Unidos Fil: Oakley, Todd H.. University of California. Department of Ecology, Evolution and Marine Biology; Estados Unidos Fil: Narwani, Anita. University of Michigan. School of Natural Resources and Environment; Estados Unidos. Swiss Federal Institute of Aquatic Science and Technology. Aquatic Ecology; Suiza Fil: Allan, Eric. University of Bern. Institute of Plant Sciences; Estados Unidos Fil: Flombaum, Pedro. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinacion Administrativa Ciudad Universitaria. Centro de Investigaciones del Mar y la Atmósfera; Argentina Fil: Isbell, Forest. University of Georgia. Department of Plant Biology; Estados Unidos Fil: Joshi, Jasmin. University of Potsdam. Biodiversity Research/Systematic Botany. Institute of Biochemistry and Biology; Alemania. Berlin-Brandenburg Institute of Advanced Biodiversity Research; Alemania Fil: Reich, Peter B.. University of Minnesota. Department of Forest Resources; Estados Unidos. University of Western Sydney. Hawkesbury Institute for the Environment; Australia Fil: Tilman, David. University of Minnesota. College of Biological Sciences; Estados Unidos. University of California. Bren School of Environmental Science and Management; Estados Unidos Fil: Van Ruijven, Jasper. Wageningen University. Department of Nature Conservation and Plant Ecology; Países Bajos Fil: Cardinale, Bradley J.. University of Michigan. School of Natural Resources and Environment; Estados Unidos

Summary 1. A long-standing hypothesis in ecology and evolutionary biology is that closely related species are more ecologically similar to each other and therefore compete more strongly than distant relatives do. A recent hypothesis posits that evolutionary relatedness may also explain the prevalence of mutualisms, with facilitative interactions being more common among distantly related species. Despite the importance of these hypotheses for understanding the structure and function of ecological communities, experimental tests to determine how evolutionary relatedness influences competition and facilitation are still somewhat rare. 2. Here, we report results of a laboratory experiment in which we assessed how competitive and facilitative interactions among eight species of freshwater green algae are influenced by their relatedness. We measured the prevalence of competition and facilitation among 28 pairs of freshwater green algal species that were chosen to span a large gradient of phylogenetic distances. For each species, we first measured its invasion success when introduced into a steady-state population of another resident species. Then, we compared its growth rate when grown alone in monoculture to its growth rate when introduced as an invader. The change in the species’ population growth rate as an invader (sensitivity) is used as a measure of the strength of its interaction with the resident species. A reduced growth rate in the presence of another species indicates competition, whereas an increased growth rate indicates facilitation. 3. Although competition between species was more frequent (75% of interactions), facilitation was common (the other 25% of interactions). We found no significant relationship between the phylogenetic distance separating two interacting species and the success of invasion, nor the prevalence or strength of either competition or facilitation. Interspecific interactions depended more on the identity of the species, with certain taxa consistently acting as good or bad competitors/facilitators. These species were not predictable a priori from their positions on a phylogeny. 4. Synthesis. The phylogenetic relatedness of the green algae species used here did not predict the prevalence of competitive and facilitative interactions, rejecting the hypothesis that close relatives compete strongly and contesting recent evidence that facilitation is likely to occur between distant relatives.

The coexistence of competing species depends on the balance between their fitness differences, which determine their competitive inequalities, and their niche differences, which stabilise their competitive interactions. Darwin proposed that evolution causes species’ niches to diverge, but the influence of evolution on relative fitness differences, and the importance of both niche and fitness differences in determining coexistence have not yet been studied together. We tested whether the phylogenetic distances between species of green freshwater algae determined their abilities to coexist in a microcosm experiment. We found that niche differences were more important in explaining coexistence than relative fitness differences, and that phylogenetic distance had no effect on either coexistence or on the sizes of niche and fitness differences. These results were corroborated by an analysis of the frequency of the co-occurrence of 325 pairwise combinations of algal taxa in > 1100 lakes across North America. Phylogenetic distance may not explain the coexistence of freshwater green algae.

Phenotypic variation is vital for microbial populations to survive environmental perturbations. Both genetic and non-genetic factors contribute to an organism’s phenotypic variation and therefore its fitness. To investigate the correlation between genetic diversity and phenotypic variation, we applied our recently developed mass spectrometry method that allows for the simultaneous measurement of more than 25 different lipids and pigments with high throughput in the unicellular microalga Chlamydomonas reinhardtii. We monitored the impact of nitrogen limitation on a genetically diverse wild-type strain CC-1690 and two isoclonal isolates from CC-1690 named ANC3 and ANC5. Measuring molecular composition of thousands of single cells at different time points of the experiment allowed us to capture a dynamic picture of the phenotypic composition and adaptation of the populations over time. Although the genetically diverse population maintained phenotypic variation over the whole time course of the experiment, the isoclonal cultures showed higher synchronicity in their phenotypic response. Furthermore, the genetically diverse population showed equal or greater phenotypic variation over the whole time range in multidimensional trait space compared with isoclonal populations. However, along individual trait axes non-genetic variance was higher in isoclonal populations.

Algae-derived biocrude oil is a possible renewable energy alternative to fossil fuel based crude oil. Outdoor cultivation in raceway ponds is estimated to provide a better return on energy invested than closed photobioreactor systems. However, in these open systems, algal crops are subjected to environmental variation in temperature and irradiance, as well as biotic invasions which can cause costly crop instabilities. In this paper, we used an experimental approach to investigate the ability of species richness to maximize and stabilize biocrude production in the face of weekly temperature fluctuations between 17 and 27 °C, relative to a constant-temperature control. We hypothesized that species richness would lead to higher mean biocrude production and greater stability of biocrude production over time in the variable temperature environment. Counter to our hypothesis, species richness tended to cause a decline in mean biocrude production, regardless of environmental temperature variation. However, biodiversity did have stabilizing effects on biocrude production over time in the variable temperature environment and not in the constant temperature environment. Altogether, our results suggest that when the most productive and stable monoculture is unknown, inoculating raceway ponds with a diverse mixture of algae will tend to ensure stable harvests over time., Environmental Science & Technology, 50 (23), ISSN:0013-936X, ISSN:1520-5851

The aim of this study was to determine if polycultures of algae could enhance tolerance to aqueous-phase coproduct (ACP) from hydrothermal liquefaction (HTL) of algal biomass to produce biocrude. The growth of algal monocultures and polycultures was characterized across a range ACP concentrations and sources. All of the monocultures were either killed or inhibited by 2% ACP, but polycultures of the same species were viable at up to 10%. The addition of ACP increased the growth rate (up to 25%) and biomass production (53%) of polycultures, several of which were more productive in ACP than any monoculture was in the presence or absence of ACP. These results suggest that a cultivation process that applies biodiversity to nutrient recycling could produce more algae with less fertilizer consumption.

Summary 1. The importance of species diversity for the stability of populations, communities and ecosystem functions is a central question in ecology. 2. Biodiversity experiments have shown that diversity can impact both the average and variability of stocks and rates at these levels of ecological organization in single trophic-level ecosystems. Whether these impacts hold in food webs and across trophic levels is still unclear. 3. We asked whether resource species diversity, community composition and consumer feeding selectivity in planktonic food webs impact the stability of resource or consumer populations, community biomass and ecosystem functions. We also tested the relative importance of resource diversity and community composition. 4. We found that resource diversity negatively affected resource population stability, but had no effect on consumer population stability, regardless of the consumer’s feeding selectivity. Resource diversity had positive effects on most ecosystem functions and their stability, including primary production, resource biomass and particulate carbon, nitrogen and phosphorus concentrations. 5. Community composition, however, generally explained more variance in population, community and ecosystem properties than species diversity per se. This result points to the importance of the outcomes of particular species interactions and individual species’ effect traits in determining food web properties and stability. 6. Among the stabilizing mechanisms tested, an increase in the average resource community biomass with increasing resource diversity had the greatest positive impact on stability. 7. Our results indicate that resource diversity and composition are generally important for the functioning and stability of whole food webs, but do not have straightforward impacts on consumer populations.

At present, the disciplines of evolutionary biology and ecosystem science are weakly integrated. As a result, we have a poor understanding of how the ecological and evolutionary processes that create, maintain, and change biological diversity affect the flux of energy and materials in global biogeochemical cycles. The goal of this article was to review several research fields at the interfaces between ecosystem science, community ecology and evolutionary biology, and suggest new ways to integrate evolutionary biology and ecosystem science. In particular, we focus on how phenotypic evolution by natural selection can influence ecosystem functions by affecting processes at the environmental, population and community scale of ecosystem organization. We develop an eco-evolutionary model to illustrate linkages between evolutionary change (e. g. phenotypic evolution of producer), ecological interactions (e. g. consumer grazing) and ecosystem processes (e. g. nutrient cycling). We conclude by proposing experiments to test the ecosystem consequences of evolutionary changes.

The effect of species diversity on ecosystem function is commonly studied within a single trophic level, but less is known about how resource diversity affects species interactions between trophic levels. We conducted a grazing experiment to determine how resource species diversity affects rates of consumption by three species of freshwater zooplankton consumers. We measured the effect of resource diversity on rates of consumption for several resource community compositions. These compositions varied in terms of palatability for the consumers. The effect of resource diversity on consumption rates depended on the diet breadth of the consumer species (from specialist to generalist) and the community composition of resources. Overall, high resource diversity commonly caused a decline in consumption rates of consumers. The most selective grazer showed reduced consumption for nearly all community compositions, whereas the most generalist grazer showed accelerated consumption when all resource species were palatable. Our results demonstrate that resource species diversity can modulate rates of consumption through the action of multiple different mechanisms.

The competitive exclusion principle poses the pressing question of how biodiversity is maintained in nature. Many mechanisms have been proposed to explain diversity and to resolve what has become known as the “paradox of the plankton”. We propose a dichotomy among these mechanisms in order to enable empiricists to begin testing their relative importance. Specifically, the mechanisms can be categorized as being internally generated or as depending on forces external to the competitive community. Here we tested whether the internal competitive dynamics of a phytoplankton assemblage or externally generated resource variability (a disturbance) were more effective at maintaining species diversity over time. We also tested whether the species composition of assemblages was important in determining the persistence of species diversity. We employed controlled microcosm experiments in which we either imposed exogenous variability in nutrient availability via serial dilution or allowed the communities to ...

We investigated how the history of local disturbances in a watershed can influence the regional coherence of ecosystem properties in lakes that have similar morphometry and climatic conditions. We measured sedimentary δ13C, δ15N, C:N and %BSiO2 in Sooke Lake Reservoir (SOL) and Shawnigan Lake (SHL), which are located within 4 km of each other on Vancouver Island, Canada. SOL is an impounded lake whose watershed has been fully protected over the last century, although the lake level has been raised 3 times via impoundment during this period. SHL has a similar limnological regime, but the surrounding watershed has been developed extensively for residential uses. We investigated how a pulse disturbance regime in SOL (i.e. repeated dam raising) and a press disturbance regime in SHL (i.e. persistent development) influenced the variability of paleoindicators in each system over time. We found that these contrasting disturbance regimes reduced the regional temporal coherence of aquatic productivity between the two lakes (indicated by %BSiO2), but did not influence the regional coherence of nutrient status or the main carbon sources of the lakes (indicated by %C, %N and δ13C). In contrast, an indicator of the sources and cycling of nitrogen (δ15N) showed increased coherence. Local disturbances also affected the variability of the paleoindicators within each system over time. In SOL, impoundments led to both declines (%N, δ15N) and increases (δ13C) in the variability of paleoindicators. In SHL, persistent watershed development led to lower variability of two paleoindicators (%N, %BSiO2). Overall, our data suggest that local disturbances can influence the %BSiO2 and C:N ratio of lake sediments, but are less likely to alter the regional coherence of %C, %N and δ13C between lakes.

Phytoplankton species traits have been used to successfully predict the outcome of competition, but these traits are notoriously laborious to measure. If these traits display a phylogenetic signal, phylogenetic distance (PD) can be used as a proxy for trait variation. We provide the first investigation of the degree of phylogenetic signal in traits related to competition in freshwater green phytoplankton. We measured 17 traits related to competition and tested whether they displayed a phylogenetic signal across a molecular phylogeny of 59 species of green algae. We also assessed the fit of five models of trait evolution to trait variation across the phylogeny. There was no significant phylogenetic signal for 13 out of 17 ecological traits. For 7 traits, a non-phylogenetic model provided the best fit. For another 7 traits, a phylogenetic model was selected, but parameter values indicated that trait variation evolved recently, diminishing the importance of common ancestry. This study suggests that traits related to competition in freshwater green algae are not generally well-predicted by patterns of common ancestry. We discuss the mechanisms by which the link between phylogenetic distance and phenotypic differentiation may be broken., PLoS ONE, 10 (9), ISSN:1932-6203

Interspecific competition between phytophagous insects can occur when plant responses induced by an early-season herbivore alter host quality for later colonizers. Recent evidence for specificity in the elicitation of induced plant responses by different attackers suggests that dynamics of host use in the field may be more complex than previously anticipated, because host suitability for colonizing herbivores may depend on which herbivore species has initially damaged a plant. In each of two years, we manipulated the first herbivore to attack Solanum dulcamara plants in an experimental population using several different arthropod species and subsequently monitored colonization by natural herbivores over the course of the growing season. We additionally performed weekly herbivore counts in wild S. dulcamara populations following natural variation in herbivore arrival. Plant-mediated interactions occurred primarily between two leaf-feeding beetles, Psylliodes affinis and Plagiometriona clavata. In both manipulative and observational experiments, P. clavata oviposition was reduced on plants initially damaged by P. affinis (or a third leaf-feeding beetle, Lema trilinea) relative to plants that were initially undamaged. Lowered P. clavata occurrence continued through subsequent life-history stages, resulting in decreased emergence of second-generation P. clavata adults on these plants. The occurrence of P. affinis was also lowered on plants damaged by conspecifics in both manipulative and observational experiments. Resistance against P. affinis also followed applications of jasmonic acid, an elicitor of plant defensive responses. Conversely, early-season damage by P. clavata did not influence plant quality for either later conspecifics or P. affinis. Initial herbivory by the spittlebug Aphrophora saratogensis or generalist taildropper slugs (Prophysaon sp.) likewise had no influence on P. clavata and P. affinis colonization, whereas L. trilinea damage did not affect later arriving P. affinis. Hence, only a subset of early-season damagers influenced herbivore occurrence on S. dulcamara. Preference tests examining P. affinis feeding and P. clavata oviposition confirmed that specificity in elicitation of induced plant responses produced the divergent herbivore occurrence patterns observed in the field. Overall, the existence of plant-mediated competitive asymmetry between herbivore species on S. dulcamara highlights the dynamic nature of plant resistance and its potential role in organizing herbivore communities.

Species richness (SR) and phylogenetic diversity (PD) are highly correlated measures of plant diversity. Each, by itself, is significantly associated with plant community biomass in biodiversity experiments. As presented by Cadotte (2015) and as we present below, reasonable but alternative analyses that attempt to control for this correlation in different ways provide contradictory or inconclusive support for the hypothesis that PD is superior to SR as a predictor of community biomass. In Venail et al. (2015), we re-analysed data from 16 experimental manipulations of grassland SR to look at how SR and PD influence variation in plant community biomass through time. Using four types of analyses, we showed that, after statistically controlling for variation in SR, PD was not related to community biomass or to the temporal stability of biomass. We did, however, find that SR tends to increase the biomass production of plant communities after controlling for PD. In his comment, Cadotte expressed two concerns about our analyses. One is that we used non-random subsets of experiments, rather than the full data set, for some of our analyses (types 2, 3). We were clear in stating these analyses were based on non-random subsets that were specifically chosen to minimize the SR–PD correlation and avoid problems associated with multicollinearity. We acknowledge that our tests are conservative, a cost of which is that they sacrifice statistical power while, at the same time, minimizing the chance of drawing an incorrect conclusion. But we disagree with Cadotte’s suggestion that our use of non-random data subsets led to ‘biased’ conclusions, and demonstrate later in this response that his claim of bias is unsubstantiated. Cadotte’s second concern was that our analyses did not account for differences in biomass across studies. This is an important criticism to consider; we made a mistake by not controlling for variation in biomass. To address this issue, Cadotte used mixed models where study was included as a random effect, and ran analyses that standardized biomass among sites. Collectively, these led Cadotte to conclude ‘All analyses strongly support previous literature claims about the value of PD and I further show that: (i) PD provides a more powerful explanation of variation in biomass production than species richness; (ii) PD explains variation in biomass production after controlling for richness; and (iii) the use of data subsets inadvertently biased the conclusions’. We have two concerns with Cadotte’s re-analysis. First, Cadotte’s approach largely ignores the concerns we raised about multicollinearity. When two or more predictors

There is a long tradition in ecology of trying to understand community assembly processes by making inferences from patterns of community structure (Diamond 1975; Connor Simberloff 1979). Inferring process from pattern is appealing because the latter is more easily observed and quantified especially when manipulative controlled experiments are infeasible. In such cases our hope (albeit naïve at times) is that patterns of community structure bear some signature of the processes that generated them by assuming that a particular pattern is overwhelmingly the outcome of a single dominant process. However multiple processes can often generate the same pattern. A now well known example of the inference of processes from patterns is Joseph Connell's ‘Ghost of Competition Past’ (1980) which highlighted that although coexisting competitors may tend to be niche differentiated on average this pattern in and of itself is not proof that the observed niche differences are the result of divergent selection experienced by coevolving competitors. Connell emphasized that direct empirical support for the notion of character displacement enabling coexistence was virtually non existent. Another classic example of inferring process from pattern relates to the intermediate disturbance hypothesis (IDH) (Connell 1978) which suggests that intermediate levels of disturbance promote the highest levels of diversity (pattern) by preventing competitive exclusion by either good competitors in undisturbed environments or rapid growers in heavily disturbed environments (process). There have been very few attempts to directly test the process leading to this pattern and it has more recently been recognized that two alternative processes (i.e. competitive relative nonlinearity and storage effects) are capable of generating the same pattern (Chesson 2000; Shea Roxburgh Rauschert 2004). Such examples are important reminders that understanding community assembly requires careful understanding of how pattern and process are linked. Critical evaluation of existing research approaches an important role to play here by pointing the way towards more productive approaches and research agendas. For instance in response to the criticisms of Connell (1980) and others Schluter McPhail (1992) developed a checklist of criteria that must be satisfied in order to demonstrate character displacement. Recent reviews highlight which items on the checklist rarely are checked off thereby identifying productive directions for future work (Beans 2014; Stuart et al. 2014).

The most unique feature of Earth is the existence of life, and the most extraordinary feature of life is its diversity. Approximately 9 million types of plants, animals, protists and fungi inhabit the Earth. So, too, do 7 billion people. Two decades ago, at the first Earth Summit, the vast majority of the world's nations declared that human actions were dismantling the Earth's ecosystems, eliminating genes, species and biological traits at an alarming rate. This observation led to the question of how such loss of biological diversity will alter the functioning of ecosystems and their ability to provide society with the goods and services needed to prosper.

1. The importance of species diversity for the stability of populations, communities and ecosystem functions is a central question in ecology. 2. Biodiversity experiments have shown that diversity can impact both the average and variability of stocks and rates at these levels of ecological organization in single trophic-level ecosystems. Whether these impacts hold in food webs and across trophic levels is still unclear. 3. We asked whether resource species diversity, community composition and consumer feeding selectivity in planktonic food webs impact the stability of resource or consumer populations, community biomass and ecosystem functions. We also tested the relative importance of resource diversity and community composition. 4. We found that resource diversity negatively affected resource population stability, but had no effect on consumer population stability, regardless of the consumer's feeding selectivity. Resource diversity had positive effects on most ecosystem functions and their stability, including primary production, resource biomass and particulate carbon, nitrogen and phosphorus concentrations. 5. Community composition, however, generally explained more variance in population, community and ecosystem properties than species diversity per se. This result points to the importance of the outcomes of particular species interactions and individual species' effect traits in determining food web properties and stability. 6. Among the stabilizing mechanisms tested, an increase in the average resource community biomass with increasing resource diversity had the greatest positive impact on stability. 7. Our results indicate that resource diversity and composition are generally important for the functioning and stability of whole food webs, but do not have straightforward impacts on consumer populations.

At present, the disciplines of evolutionary biology and ecosystem science are weakly integrated. As a result, we have a poor understanding of how the ecological and evolutionary processes that create, maintain, and change biological diversity affect the flux of energy and materials in global biogeochemical cycles. The goal of this article was to review several research fields at the interfaces between ecosystem science, community ecology and evolutionary biology, and suggest new ways to integrate evolutionary biology and ecosystem science. In particular, we focus on how phenotypic evolution by natural selection can influence ecosystem functions by affecting processes at the environmental, population and community scale of ecosystem organization. We develop an eco-evolutionary model to illustrate linkages between evolutionary change (e.g. phenotypic evolution of producer), ecological interactions (e.g. consumer grazing) and ecosystem processes (e.g. nutrient cycling). We conclude by proposing experiments to test the ecosystem consequences of evolutionary changes.

Nature 486, 59–67 (2012); doi:10.1038/nature11148 In Table 1 and Supplementary Table 2 of this Review, under the ‘Category of service’ called ‘Regulating’, the first two ‘Measures of service provision’ related to ‘Biocontrol’ should read ‘Abundance of herbivorous pests’ instead of ‘Control of herbivorous pests’.