Temperature effects on fish production across a natural thermal gradient.

Global warming is widely predicted to reduce the biomass production of top predators, or even result in species loss. Several exceptions to this expectation have been identified, however, and it is vital that we understand the underlying mechanisms if we are to improve our ability to predict future trends. Here, we used a natural warming experiment in Iceland and quantitative theoretical predictions to investigate the success of brown trout as top predators across a stream temperature gradient (4-25 °C). Brown trout are at the northern limit of their geographic distribution in this system, with ambient stream temperatures below their optimum for maximal growth, and above it in the warmest streams. A five-month mark-recapture study revealed that population abundance, biomass, growth rate, and production of trout all increased with stream temperature. We identified two mechanisms that contributed to these responses: (1) trout became more selective in their diet as stream temperature increased, feeding higher in the food web and increasing in trophic position; and (2) trophic transfer through the food web was more efficient in the warmer streams. We found little evidence to support a third potential mechanism: that external subsidies would play a more important role in the diet of trout with increasing stream temperature. Resource availability was also amplified through the trophic levels with warming, as predicted by metabolic theory in nutrient-replete systems. These results highlight circumstances in which top predators can thrive in warmer environments and contribute to our knowledge of warming impacts on natural communities and ecosystem functioning.

Light and temperature controls of aquatic plant photosynthesis downstream of a hydropower plant and the effect of plant removal.

Many rivers worldwide are regulated, and the altered hydrology can lead to mass development of aquatic plants. Plant invasions are often seen as a nuisance for human activities leading to costly remedial actions with uncertain implications for aquatic biodiversity and ecosystem functioning. Mechanical harvesting is often used to remove aquatic plants and knowledge of plant growth rate could improve management decisions. Here, we used a simple light-temperature theoretical model to make a priori prediction of aquatic plant photosynthesis. These predictions were assessed through an open-channel diel change in O2 mass balance approach. A Michaelis-Menten type model was fitted to observed gross primary production (GPP) standardised at 10 °C using a temperature dependence from thermodynamic theory of enzyme kinetics. The model explained 87 % of the variability in GPP of a submerged aquatic plant (Juncus bulbosus L.) throughout an annual cycle in the River Otra, Norway. The annual net plant production was about 2.4 (1.0-3.8) times the standing biomass of J. bulbosus. This suggests a high continuous mass loss due to hydraulic stress and natural mechanical breakage of stems, as the biomass of J. bulbosus remained relatively constant throughout the year. J. bulbosus was predicted to be resilient to mechanical harvesting with photosynthetic capacity recovered within two years following 50-85 % plant removal. The predicted recovery was confirmed through a field experiment where 72 % of J. bulbosus biomass was mechanically removed. We emphasise the value of using a theoretical approach, like metabolic theory, over statistical models where a posteriori results are not always easy to interpret. Finally, the ability to predict ecosystem resilience of aquatic photosynthesis in response to varying management scenarios offers a valuable tool for estimating aquatic ecosystem services, such as carbon regulation. This tool can benefit the EU Biodiversity Strategy and UN Sustainable Development Goals.

Estimation of ecosystem respiration and photosynthesis in supersaturated stream water downstream of a hydropower plant.

The estimation of whole stream metabolism, as determined by photosynthesis and respiration, is critical to our understanding of carbon cycling and carbon subsidies to aquatic food-webs. The mass development of aquatic plants is a worldwide problem for human activities and often occurs in regulated rivers, altering biodiversity and ecosystem functions. Hydropower plants supersaturate water with gases and prevent the use of common whole stream metabolism models to estimate ecosystem respiration. Here we used the inert noble gas argon to parse out biological from physical processes in stream metabolism calculations. We coupled the O2:Ar ratio determined by gas chromatography in grab samples with in-situ oxygen concentrations measured by an optode to estimate aquatic plant photosynthesis and ecosystem respiration during supersaturation events through a parsimonious approach. The results compared well with a more complicated two-station model based on O2 mass balances in non-supersatured water, and with associated changes in dissolved CO2 (or dissolved inorganic carbon). This new method provides an independent approach to evaluate alternative corrections of dissolved oxygen data (e.g. through the use of total dissolved gases) in long term studies. The use of photosynthesis-irradiance models allows the determination of light parameters such as the onset of light saturation or low light use efficiency, which could be used for inverse modelling. The use of the O2:Ar approach to correct for oversaturation may become more applicable with the emergence of portable mass inlet mass spectrometers (MIMS). Photosynthesis was modest (2.9-5.8 g O2 m2 day-1) compared to other rivers with submerged vegetation, likely indicating nutrient co-limitations (CO2, inorganic N and P). Respiration was very low (-2.1 to -3.9 g O2 m2 day-1) likely due to a lack of allochthonous carbon supply and sandy sediment.

Effects of pollution-induced changes in oxygen conditions scaling up from individuals to ecosystems in a tropical river network.

Anthropogenic inputs of nutrients and organic matter are common in tropical lowland rivers while little is known about the pollution-induced changes in oxygen availability and respiratory performance of ectotherms in these high temperature systems. We investigated the effects of agriculture and urban land-use on river water oxygen levels (diel measurements), decomposition rates (Wettex) and macroinvertebrate assemblages (field studies), as well as the oxy-regulatory capacity of eight riverine macroinvertebrate taxa (laboratory study) from a tropical lowland river network in Myanmar. The highest decomposition rates (0.1-5.5 mg Wettex degree day-1) and oxygen stress (≤91% saturation deficits) were found in reaches draining degraded catchments with elevated concentrations of nutrients. All individual macroinvertebrate taxa investigated were to some extent able to regulate their respiration when placed under oxygen stress in the laboratory (regulation value of 0.74-0.89). The oxy-regulation capacity of macroinvertebrate assemblages in the river network were, as predicted, inversely related to diel oxygen stress (maximum deficit; lm, R2 = 0.69), where taxonomic richness and pollution sensitivity (ASPT metric) also declined sharply (lm, R2 ≥ 0.79). Our study shows that eutrophication and organic pollution induce oxygen deficits in tropical rivers but stimulate decomposition rates, which may further deplete oxygen levels. Furthermore, macroinvertebrate oxy-regulatory capacity predicts assemblage composition along gradients in oxygen stress at the ecosystem level. Our findings suggest that tropical lowland river systems could be highly sensitive to pollution by nutrients and organic matter leading to substantial impacts on ectotherm community composition and ecosystem functioning.

Mechanical removal of macrophytes in freshwater ecosystems: Implications for ecosystem structure and function.

Macrophytes are generally considered a nuisance when they interfere with human activities. To combat perceived nuisance, macrophytes are removed, and considerable resources are spent every year worldwide on this practice. Macrophyte removal can, however, have severe negative impacts on ecosystem structure and functioning and interfere with management goals of healthy freshwater ecosystems. Here, we reviewed the existing literature on mechanical macrophyte removal and summarised current information from 98 studies on short- and long-term consequences for ecosystem structure and functioning. In general, the majority of studies were conducted in rivers and streams and evaluated short-term effects of removal on single ecosystem properties. Moreover, most studies did not address the interrelationships between ecosystem properties and the underlying mechanisms. Contrasting effects of removal on ecosystem structure and function were found and these discrepancies were highly dependent on the context of each study, making meaningful quantitative comparisons across studies very difficult. We illustrated how a Bayesian network (BN) approach can be used to assess the implications of macrophyte removal on interrelated ecosystem properties across a wide range of environmental conditions. The BN approach could also help engage a conversation with stakeholders on the management of freshwater ecosystems.

Juncus Bulbosus Tissue Nutrient Concentrations and Stoichiometry in Oligotrophic Ecosystems: Variability with Seasons, Growth Forms, Organs and Habitats.

Aquatic plant nutrient concentrations provide important information to characterise their role in nutrient retention and turnover in aquatic ecosystems. While large standing biomass of aquatic plants is typically found in nutrient-rich localities, it may also occur in oligotrophic ecosystems. Juncus bulbosus is able to form massive stands even in very nutrient-dilute waters. Here we show that this may be achieved by tissues with very high carbon-to-nutrient ratios combined with perennial (slow) growth and a poor food source for grazers inferred from plant stoichiometry and tissue nutrient thresholds. We also show that the C, N, P and C:N:P stoichiometric ratios of Juncus bulbosus vary with the time of year, habitats (lakes versus rivers) and organs (roots versus shoots). We found no differences between growth forms (notably in P, inferred as the most limiting nutrient) corresponding to small and large plant stands. The mass development of J. bulbosus requires C, N and P, whatever the ecosystem (lake or river), and not just CO2 and NH4, as suggested in previous studies. Since macrophytes inhabiting oligotrophic aquatic ecosystems are dominated by isoetids (perennial plants with a high root/shoot ratio), attention should be paid to quantifying the role of roots in aquatic plant stoichiometry, nutrient turnover and nutrient retention.

Rethinking Phage Ecology by Rooting it Within an Established Plant Framework.

Despite the abundance and significance of bacteriophages to microbial ecosystems, no broad ecological frameworks exist within which to determine "bacteriophage types" that reflect their ecological strategies and ways in which they interact with bacterial cells. To address this, we repurposed the well-established Grime's triangular CSR framework, which classifies plants according to three axes: competitiveness (C), ability to tolerate stress (S), and capacity to cope with disturbance (R). This framework is distinguished from other accepted schemes, as it seeks to identify individual characteristics of plants to understand their biological strategies and roles within an ecosystem. Our repurposing of the CSR triangle is based on phage transcription and the observation that typically phages have three major distinguishable transcription phases: early, middle, and late. We hypothesize that the proportion of genes expressed in these phases reflects key information about the phage "ecological strategy," namely the C, S, and R strategies, allowing us to examine phages in a similar way to how plants are projected onto the triangle. In the "phage version" of this scheme, we suggest: (1) that some phages prioritize the early phase of transcription that shuts off host defense mechanisms, which reflects competitiveness; (2) other phages prioritize tuning resource management mechanisms in the cell such as nucleotide metabolism during their "mid" expression profile to tolerate stress; and (3) a further subset of phages (termed Ruderals) survive disturbance by investing significant resources into regeneration so they express a higher proportion of their genes during late infection. We examined 42 published phage transcriptomes and show that they fall into discrete CSR categories according to their expression profiles. We discuss these positions in the context of their biology, which is largely consistent with our predictions of specific phage characteristics. In this opinion article, we suggest a starting point to ascribe phages into different functional types and thus understand them in an ecological framework. We suggest that this may have far-reaching implications for the application of phages in therapy and their exploitation to manipulate bacterial communities. We invite further use of this framework via our online tool; www.PhageCSR.ml.

Distribution of aquatic macrophytes in contrasting river systems: a critique of compositional-based assessment of water quality.

A brief summary of the historical developments relating to plant distribution and aquatic macrophyte-nutrient indices provided a means of assessing the general context and validity of previous assumptions. This has particular current relevance because of the prominent use of bioindicators for defining nutrient enrichment. A survey of 161 sites distributed across two broadly contrasting groups of rivers (circum-neutral versus alkaline) recorded 110 species of aquatic macrophytes and these have been statistically analyzed to (i) rank and separate the individual effects of local environmental conditions and spatial isolation on species distribution in the two contrasting groups of sites; (ii) calculate a macrophyte index based on plant cover and species indicator values (Mean Trophic Rank, MTR); and finally (iii) investigate the implications for biomonitoring. Chemical, physical and hydrological site attributes together with spatial isolation, each explained a significant and at least partially independent influence over plant species distribution. It was extremely difficult, however, to separate the single effects of different site attributes on plant distribution. While some plant species are more restricted to certain environmental conditions, many appeared indifferent to the range of those being tested. The role played by nutrients (nitrogen (N) and phosphorus (P)) were either mostly indistinguishable from other site attributes (e.g., nitrate from conductivity) or subordinate (e.g., soluble reactive phosphorus, ammonium). It is therefore very unlikely that macrophyte species composition could provide a reliable bioindicator of the surrounding nutrient (N, P) status. The calculation of the plant index illustrated this unreliability by showing that strong correlations existed with many environmental variables, not just inorganic N and P.

Functional biogeography: Stoichiometry and thresholds for interpreting nutrient limitation in aquatic plants.

Atmospheric N pollution may shift nutrient limitations in aquatic autotrophs from N to P or cause an intensification of P limitation in formerly pristine areas. Small changes in nutrient supply in oligotrophic lakes and rivers could lead to large changes in relative plant growth and yield with possible knock on effects on ecosystem carbon cycling through changes in the decomposition rate of their tissue. Previous biogeographical studies have shown inconsistent responses of plant nutrient tissue content and stoichiometry (functional traits) to external nutrient availability. Here we used a single species, Juncus bulbosus, to test the interplay between plant tissue nutrient (content and stoichiometry) and external environmental factors (local and catchment scale). We developed a comparative approach applicable globally to assess the thresholds for nutrient limitation in aquatic plants in the wild. Phosphorus in Juncus bulbosus tissue was negatively related to sediment organic matter (Fe root plaque limiting P uptake) and catchment vegetation cover (less P leaching to lakes). Our comparative approach revealed that the lack of increase in N plant tissue along the strong gradient in external N concentration may be explained by P limitation and strict plant tissue N:P ratio. Our comparative approach further showed that the nutrient content and stoichiometry of Juncus bulbosus was similar to other submerged aquatic plants growing in nutrient poor aquatic ecosystems. In southern Norway, mass development of Juncus bulbosus may be primarily triggered by changes in P availability, rather than CO2 or inorganic N, as previously thought, although co-limitations are also possible. If so, the mass development of Juncus bulbosus in oligotrophic aquatic ecosystems could be an early indicator of increasing P fluxes through these ecosystems which are less limited by N due to high atmospheric N deposition.

Detection of an invasive aquatic plant in natural water bodies using environmental DNA.

The ability to detect founding populations of invasive species or rare species with low number of individuals is important for aquatic ecosystem management. Traditional approaches use historical data, knowledge of the species' ecology and time-consuming surveys. Within the past decade, environmental DNA (eDNA) has emerged as a powerful additional tracking tool. While much work has been done with animals, comparatively very little has been done with aquatic plants. Here we investigated the transportation and seasonal changes in eDNA concentrations for an invasive aquatic species, Elodea canadensis, in Norway. A specific probe assay was developed using chloroplast DNA to study the fate of the targeted eDNA through space and time. The spatial study used a known source of Elodea canadensis within Lake Nordbytjern 400 m away from the lake outlet flowing into the stream Tveia. The rate of disappearance of E. canadensis eDNA was an order of magnitude loss over about 230 m in the lake and 1550 m in the stream. The time series study was performed monthly from May to October in lake Steinsfjorden harbouring E. canadensis, showing that eDNA concentrations varied by up to three orders of magnitude, peaking during fall. In both studies, the presence of suspended clay or turbidity for some samples did not hamper eDNA analysis. This study shows how efficient eDNA tools may be for tracking aquatic plants in the environment and provides key spatial and temporal information on the fate of eDNA.

Aquatic macrophytes as bioindicators of carbon dioxide in groundwater fed rivers.

Aquatic plants have been used as hydrological tracers in groundwater fed river systems. In nature, patterns in plant distribution have been attributed to ammonium (NH(4)) toxicity and phosphate (PO(4)) limitation, while some laboratory studies have focused on the role of the partial pressure of CO(2) (pCO(2)). The aims of this study were (i) to test whether plant distribution was more related to pCO(2) than NH(4) and PO(4) in nature, (ii) to develop and test the predictive power of new plant indices for pCO(2), NH(4) and PO(4), and (iii) to test the potential causality of the relationships using species eco-physiological traits. These tests were carried out with field data from the Rhine, Rhône and Danube river basins. Species composition was best related to the effect of pCO(2). The pCO(2) plant index was well calibrated (r(2)=0.73) and had the best predictive power (r(2)=0.47) of the three indices tested on independent datasets. The plant-pCO(2) relationship was supported by a biological mechanism: the ability of strictly submerged species of aquatic vascular plants to use HCO(3) under low pCO(2). This was not the whole story: the effects of pCO(2), NH(4) and PO(4) on plant distribution were partially confounded and interacted all together with temperature. However, neither NH(4) toxicity nor P limitation could be asserted using species eco-physiological traits. Moreover, the predictive power of the NH(4) and PO(4) plant indices was not as strong as pCO(2), at r(2)=0.24 and r(2)=0.27, respectively. Other potentially confounding variables such as spatial structure, biotic and physical factors were unlikely to confound the findings of this study.

Attribute-based classification of European hydrophytes and its relationship to habitat utilization.

1Here we classify selected European hydrophytes into 'attribute groups' based on the possession of homogenous sets of characteristics, and explore the correspondence between these attribute groups, or individual attributes, and habitat use.2Non-hierarchical clustering was used to assign 120 species to twenty groups based on a matrix of categorical scores for literature- and field-derived information covering seventeen intrinsic morphological and life-history traits. Subdivision of some of these traits produced a total of 58 attributes (i.e. modalities). The robustness of this classification was confirmed by a high rate of reclassification (92%) under multiple discriminant analysis (MDA). The phylogenetic contribution was explored using ordination methods with taxonomy at family level acting as a covariable.3Our approach differed from earlier classifications based on growth or life form because we regarded growth form plasticity as a property of the species and its range of growing conditions, rather than of each individual population, and we considered additional (e.g. regenerative) traits. However, some conventional life form groups were preserved (i.e. utricularids, isoetids, hydrocharids and lemnids).4Some parallels existed with established theory on terrestrial plant growth strategies, but we used strictly intrinsic attributes relevant specifically to hydrophytes and our groups could not be decomposed into three or four primary strategies. Only finer levels of partitioning appear to be of fundamental and applied ecological relevance in hydrophytes.5A principal components analysis ordination based on 26 attributes related to physical habitat utilization separated species and their attribute groups along axes relating to: (a) flow, substratum grade and organic matter content, scour frequency, and sedimentation; and (b) depth, water level stability and biotic disturbance. A MDA applied to species ordination scores indicated only a modest overall correspondence between attribute groups and habitat use (54% correct reclassification). Poor reclassification was the result of intergroup overlap (indicating alternative sets of attributes for a given habitat) or high intragroup variance in habitat utilization (indicating commonality of attributes between different habitats). These results are interpreted in terms of trade-offs between resistance and resilience traits, 'functional plasticity' in traits, phylogenetic dependence in some groups and methodological constraints. The predictive potential of hydrophyte groups and their limitations are discussed.6Redundancy analysis revealed a highly significant correlation between traits and habitat use (P < 0.01). Our attribute matrix explained 72% of variation in physical habitat use with eight attributes (i.e. turions, anchored emergent leaves, high or low body flexibility, high root:shoot biomass ratio, free-floating surface or free-floating submerged growth form, and annual life history) explaining half of this variation.7Most attributes were mapped in accordance with habitat template predictions, although tests were confounded by the underlying correlation between spatial and temporal heterogeneity. The main features were: (a) a trade-off between resistance-type traits (related to stream lining, flexibility and anchorage) in more spatially heterogenous riverine and littoral zone habitats, and resilience type traits (i.e. turions, very small body size and free-floating growth forms) in spatially simple, rarely disturbed habitats, such as backwaters and canals; and (b) a shift from high investment competitive traits with a low reproductive output in deep stable habitats to classically ruderal and desiccation resistance traits in shallow fluctuating habitats.