Climate change vulnerability of Arctic char across Scandinavia.

Climate change is anticipated to cause species to shift their ranges upward and poleward, yet space for tracking suitable habitat conditions may be limited for range-restricted species at the highest elevations and latitudes of the globe. Consequently, range-restricted species inhabiting Arctic freshwater ecosystems, where global warming is most pronounced, face the challenge of coping with changing abiotic and biotic conditions or risk extinction. Here, we use an extensive fish community and environmental dataset for 1762 lakes sampled across Scandinavia (mid-1990s) to evaluate the climate vulnerability of Arctic char (Salvelinus alpinus), the world's most cold-adapted and northernly distributed freshwater fish. Machine learning models show that abiotic and biotic factors strongly predict the occurrence of Arctic char across the region with an overall accuracy of 89 percent. Arctic char is less likely to occur in lakes with warm summer temperatures, high dissolved organic carbon levels (i.e., browning), and presence of northern pike (Esox lucius). Importantly, climate warming impacts are moderated by habitat (i.e., lake area) and amplified by the presence of competitors and/or predators (i.e., northern pike). Climate warming projections under the RCP8.5 emission scenario indicate that 81% of extant populations are at high risk of extirpation by 2080. Highly vulnerable populations occur across their range, particularly near the southern range limit and at lower elevations, with potential refugia found in some mountainous and coastal regions. Our findings highlight that range shifts may give way to range contractions for this cold-water specialist, indicating the need for pro-active conservation and mitigation efforts to avoid the loss of Arctic freshwater biodiversity.

Species interactions, environmental gradients and body size shape population niche width.

Competition for shared resources is commonly assumed to restrict population-level niche width of coexisting species. However, the identity and abundance of coexisting species, the prevailing environmental conditions, and the individual body size may shape the effects of interspecific interactions on species' niche width. Here we study the effects of interspecific and intraspecific interactions, lake area and altitude, and fish body size on the trophic niche width and resource use of a generalist predator, the littoral-dwelling large, sparsely rakered morph of European whitefish (Coregonus lavaretus; hereafter LSR whitefish). We use stable isotope, diet and survey fishing data from 14 subarctic lakes along an environmental gradient in northern Norway. The isotopic niche width of LSR whitefish showed a humped-shaped relationship with increasing relative abundance of sympatric competitors, suggesting widest population niche at intermediate intensity of interspecific interactions. The isotopic niche width of LSR whitefish tended to decrease with increasing altitude, suggesting reduced niche in colder, less productive lakes. LSR whitefish typically shifted to a higher trophic position and increased reliance on littoral food resources with increasing body size, although between-lake differences in ontogenetic niche shifts were evident. In most lakes, LSR whitefish relied less on littoral food resources than coexisting fishes and the niche overlap between sympatric competitors was most evident among relatively large individuals (>250 mm). Individual niche variation was highest among >200 mm long LSR whitefish, which likely have escaped the predation window of sympatric predators. We demonstrate that intermediate intensity of interspecific interactions may broaden species' niche width, whereas strong competition for limited resources and high predation risk may suppress niche width in less productive environments. Acknowledging potential humped-shaped relationships between population niche width and interspecific interactions can help us understand species' responses to environmental disturbance (e.g. climate change and species invasions) as well as the driving forces of niche specialization.

Open Data Practices among Users of Primary Biodiversity Data.

Presence-only biodiversity data are increasingly relied on in biodiversity, ecology, and conservation research, driven by growing digital infrastructures that support open data sharing and reuse. Recent reviews of open biodiversity data have clearly documented the value of data sharing, but the extent to which the biodiversity research community has adopted open data practices remains unclear. We address this question by reviewing applications of presence-only primary biodiversity data, drawn from a variety of sources beyond open databases, in the indexed literature. We characterize how frequently researchers access open data relative to data from other sources, how often they share newly generated or collated data, and trends in metadata documentation and data citation. Our results indicate that biodiversity research commonly relies on presence-only data that are not openly available and neglects to make such data available. Improved data sharing and documentation will increase the value, reusability, and reproducibility of biodiversity research.

Community structure influences species' abundance along environmental gradients.

Species' response to abiotic environmental variation can be influenced by local community structure and interspecific interactions, particularly in restricted habitats such as islands and lakes. In temperate lakes, future increase in water temperature and run-off of terrestrial (allochthonous) dissolved organic carbon (DOC) are predicted to alter community composition and the overall ecosystem productivity. However, little is known about how the present community structure and abiotic environmental variation interact to affect the abundance of native fish populations. We used a space-for-time approach to study how local community structure interact with lake morphometric and climatic characteristics (i.e. temperature and catchment productivity) to affect brown trout (Salmo trutta L.) yield in 283 Norwegian lakes located in different biogeographical regions. Brown trout yield (based on data from standardized survey gill net fishing; g 100 m(-2) gill net night(-1)) was generally lower in lakes where other fish species were present than in lakes with brown trout only. The yield showed an overall negative relationship with increasing temperature and a positive relationship with lake shoreline complexity. Brown trout yield was also negatively correlated with DOC load (measured using Normalized Difference Vegetation Index as a proxy) and lake size and depth (measured using terrain slope as a proxy), but only in lakes where other fish species were present. The observed negative response of brown trout yield to increasing DOC load and proportion of the pelagic open-water area is likely due to restricted (littoral) niche availability and competitive dominance of more pelagic fishes such as Arctic charr (Salvelinus alpinus (L.)). Our study highlights that, through competitive interactions, the local community structure can influence the response of a species' abundance to variation in abiotic conditions. Changes in biomass and niche use of top predators (such as the brown trout), associated with predicted changes in direct and indirect climatic factors, may have further influences on the structure and function of temperate lake ecosystems.

Unimodal response of fish yield to dissolved organic carbon.

Here, we demonstrate a contrasting effect of terrestrial coloured dissolved organic material on the secondary production of boreal nutrient poor lakes. Using fish yield from standardised brown trout gill-net catches as a proxy, we show a unimodal response of lake secondary productivity to dissolved organic carbon (DOC). This suggests a trade-off between positive and negative effects, where the initial increase may hinge upon several factors such as energy subsidising, screening of UV-radiation or P and N load being associated with organic carbon. The subsequent decline in production with further increase in DOC is likely associated with light limitations of primary production. We also show that shallow lakes switch from positive to negative effects at higher carbon loads than deeper lakes. These results underpin the major role of organic carbon for structuring productivity of boreal lake ecosystems.

Using mass scaling of movement cost and resource encounter rate to predict animal body size-population density relationships.

The negative relationship between body mass and population abundance was documented decades ago and forms one of the most fundamental scaling-laws in ecology. However, current theory fails to capture observed variations and the subject continues to raise controversy. Here we unify empirically observed size-abundance relationships with theory, by incorporating allometries in resource encounter rate and metabolic costs of movements. Fractal geometry is used to quantify the underlying resources distributions. Our model predicts that in environments packed with resources, body mass to population abundance relationships is less negative than the commonly assumed -3/4 power law. When resources are more patchily distributed, we predict a more negative exponent. These predictions are consistent with empirical observations. The current research provides an important step towards synthesizing metabolism, resource distribution and the global scaling of animal abundance, explaining why size-abundance relationships vary among feeding guilds and ecosystems.

Effects of temperature and food quality on age and size at maturity in ectotherms: an experimental test with Atlantic salmon.

The reaction norm between growth rate, age and size at maturity in ectotherms is widely debated in ecological literature. It has been proposed that the effect depends on whether growth is affected by food quality or temperature (called the Berrigan-Charnov puzzle). The present experiment tested this for Atlantic salmon (Salmo salar). We enhanced growth rates by increasing temperature and ratio of lipids to proteins in the food for groups of Atlantic salmon. Both treatments gave higher percentages of early mature and therefore smaller adults in contrast to the proposed Berrigan-Charnov puzzle. There was a difference between sexes in that males could attain maturity 1 year younger than females when reared under similar environmental conditions. Males that matured during the first year in sea water were smaller than similar aged immature males. The probability of that Atlantic salmon attained maturity for the first time during their second year in sea increased with growth rate during the preceding winter and if fed a high-lipid diet. Increased summer temperature exhibited no additional effect. Similar aged fish reared at elevated temperature and fed high-lipid diet attained maturity at a larger body mass and exhibited higher mass-length-ratios than those reared at natural temperature and fed a low-lipid diet, indicating that structural growth has priority over lipid deposits. Increased growth rate before the onset of maturation, whether this is owing to enhanced lipid content in food or increased water temperature, decreased age and therefore size at maturity. Enhanced lipid relative to protein content in food, but not temperature, had an additive positive effect on early maturation probability, likely due to increased amounts of reserve energy. These results may be general for ectotherm organisms.

Spatial predictors and temporal forecast of total organic carbon levels in boreal lakes.

Browning of Fennoscandian boreal lakes is raising concerns for negative ecosystem impacts as well as reduced drinking water quality. Declined sulfur deposition and warmer climate, along with afforestation, other climate impacts and less outfield grazing, have resulted in increased fluxes of Total Organic Carbon (TOC) from catchments to freshwater, and subsequently to coastal waters. This study assesses the major governing factors for increased TOC levels among several catchment characteristics in almost 5000 Fennoscandian lakes and catchments. Normalized Difference Vegetation Index (NDVI), a proxy for plant biomass, and the proportions of peatland in the catchment, along with surface runoff intensity and nitrogen deposition loading, were identified as the main spatial predictors for lake TOC concentrations. A multiple linear model, based on these explanatory variables, was used to simulate future TOC concentration in surface runoff from coastal drainage basins in 2050 and 2100, using the forecasts of climatic variables in two of the Shared Socio-economic Pathways (SSP): 1-2.6 (+2 °C) and 3-7.0 (+4,5 °C). These scenarios yield contrasting effects. SSP 1-2.6 predicts an overall decrease of TOC export to coastal waters, while SSP 3-7.0 in contrast leads to an increase in TOC export.

Recognizability bias in citizen science photographs.

Citizen science and automated collection methods increasingly depend on image recognition to provide the amounts of observational data research and management needs. Recognition models, meanwhile, also require large amounts of data from these sources, creating a feedback loop between the methods and tools. Species that are harder to recognize, both for humans and machine learning algorithms, are likely to be under-reported, and thus be less prevalent in the training data. As a result, the feedback loop may hamper training mostly for species that already pose the greatest challenge. In this study, we trained recognition models for various taxa, and found evidence for a 'recognizability bias', where species that are more readily identified by humans and recognition models alike are more prevalent in the available image data. This pattern is present across multiple taxa, and does not appear to relate to differences in picture quality, biological traits or data collection metrics other than recognizability. This has implications for the expected performance of future models trained with more data, including such challenging species.

Direct and indirect climatic drivers of biotic interactions: ice-cover and carbon runoff shaping Arctic char Salvelinus alpinus and brown trout Salmo trutta competitive asymmetries.

One of the major challenges in ecological climate change impact science is to untangle the climatic effects on biological interactions and indirect cascading effects through different ecosystems. Here, we test for direct and indirect climatic drivers on competitive impact of Arctic char (Salvelinus alpinus L.) on brown trout (Salmo trutta L.) along a climate gradient in central Scandinavia, spanning from coastal to high-alpine environments. As a measure of competitive impact, trout food consumption was measured using (137)Cs tracer methodology both during the ice-covered and ice-free periods, and contrasted between lakes with or without char coexistence along the climate gradient. Variation in food consumption between lakes was best described by a linear mixed effect model including a three-way interaction between the presence/absence of Arctic char, season and Secchi depth. The latter is proxy for terrestrial dissolved organic carbon run-off, strongly governed by climatic properties of the catchment. The presence of Arctic char had a negative impact on trout food consumption. However, this effect was stronger during ice-cover and in lakes receiving high carbon load from the catchment, whereas no effect of water temperature was evident. In conclusion, the length of the ice-covered period and the export of allochthonous material from the catchment are likely major, but contrasting, climatic drivers of the competitive interaction between two freshwater lake top predators. While future climatic scenarios predict shorter ice-cover duration, they also predict increased carbon run-off. The present study therefore emphasizes the complexity of cascading ecosystem effects in future effects of climate change on freshwater ecosystems.

Combining counts and incidence data: an efficient approach for estimating the log-normal species abundance distribution and diversity indices.

Obtaining accurate estimates of diversity indices is difficult because the number of species encountered in a sample increases with sampling intensity. We introduce a novel method that requires that the presence of species in a sample to be assessed while the counts of the number of individuals per species are only required for just a small part of the sample. To account for species included as incidence data in the species abundance distribution, we modify the likelihood function of the classical Poisson log-normal distribution. Using simulated community assemblages, we contrast diversity estimates based on a community sample, a subsample randomly extracted from the community sample, and a mixture sample where incidence data are added to a subsample. We show that the mixture sampling approach provides more accurate estimates than the subsample and at little extra cost. Diversity indices estimated from a freshwater zooplankton community sampled using the mixture approach show the same pattern of results as the simulation study. Our method efficiently increases the accuracy of diversity estimates and comprehension of the left tail of the species abundance distribution. We show how to choose the scale of sample size needed for a compromise between information gained, accuracy of the estimates and cost expended when assessing biological diversity. The sample size estimates are obtained from key community characteristics, such as the expected number of species in the community, the expected number of individuals in a sample and the evenness of the community.

Clavis: An open and versatile identification key format.

The skills and knowledge needed to recognize and classify taxa are becoming increasingly scarce in the scientific community. At the same time, it is clear that these skills are strongly needed in biodiversity monitoring for management and conservation, especially when carried out by citizen scientists. Formalizing the required knowledge in the form of digital identification keys is one way of making such knowledge more available for professional and amateur observers of biodiversity. In this paper we describe Clavis, an open and versatile data format for capturing the knowledge required for taxon identification through digital keys, allowing for a level of detail beyond that of any current key format. We present the format independently from any particular implementation, as our aim is for Clavis to serve as a basis for interoperable tools and interfaces serving different needs and actors.

Maximizing citizen scientists' contribution to automated species recognition.

Technological advances and data availability have enabled artificial intelligence-driven tools that can increasingly successfully assist in identifying species from images. Especially within citizen science, an emerging source of information filling the knowledge gaps needed to solve the biodiversity crisis, such tools can allow participants to recognize and report more poorly known species. This can be an important tool in addressing the substantial taxonomic bias in biodiversity data, where broadly recognized, charismatic species are highly over-represented. Meanwhile, the recognition models are trained using the same biased data, so it is important to consider what additional images are needed to improve recognition models. In this study, we investigated how the amount of training data influenced the performance of species recognition models for various taxa. We utilized a large citizen science dataset collected in Norway, where images are added independently from identification. We demonstrate that while adding images of currently under-represented taxa will generally improve recognition models more, there are important deviations from this general pattern. Thus, a more focused prioritization of data collection beyond the basic paradigm that "more is better" is likely to significantly improve species recognition models and advance the representativeness of biodiversity data.

Impacts of piscicide-induced fish removal on resource use and trophic diversity of lake invertebrates.

Chemical eradication of non-native species has become a widely used method to mitigate the potential negative impacts of altered competitive or predatory dynamics on biodiversity and natural ecosystem processes. However, the responses of non-target species can vary from rapid full recovery to delayed or absent recolonization, and little is known about the potential shifts in resource use and trophic diversity of native species following chemical treatments. We used a before-after-control-impact approach to study the effects of rotenone piscicide treatment on abundance and trophic niche of benthic invertebrates in three untreated and three treated lakes in central Norway, the latter group hosting non-native roach (Rutilus rutilus) and pike (Esox lucius) prior to rotenone treatment. Based on community composition data, the relative abundance of invertebrate grazers and collectors decreased while that of predators increased following fish removal in the treated lakes. The stable isotope data indicated minor shifts in resource use of, and trophic diversity among, benthic invertebrate communities. While the predatory dragonfly larvae (Odonata) and grazer snails (Lymnaeidae) showed increased δ13C values indicating increased reliance on littoral benthic algae, the collector mayfly larvae (Leptophlebia) showed decreased δ13C values following fish removal in treated lakes. Grazer snails also showed a shift to a lower trophic position, while the predatory dragonflies and collector mayflies showed no changes in δ15N values following fish removal. The community-level isotopic niches of benthic invertebrates showed no consistent changes, although the sample-size corrected and Bayesian estimates of standard ellipse areas (SEAC and SEAB) slightly increased in two of the three treated lakes due to an increased range in δ15N. In conclusion, our study findings indicate some changes in species assemblages but minor shifts in the resource use and trophic diversity of benthic invertebrate communities following fish removal in rotenone treated lakes.

A regionally coherent ecological fingerprint of climate change, evidenced from natural history collections.

Climate change has dramatic impacts on ecological systems, affecting a range of ecological factors including phenology, species abundance, diversity, and distribution. The breadth of climate change impacts on ecological systems leads to the occurrence of fingerprints of climate change. However, climate fingerprints are usually identified across broad geographical scales and are potentially influenced by publication biases. In this study, we used natural history collections spanning over 250 years, to quantify a range of ecological responses to climate change, including phenology, abundance, diversity, and distributions, across a range of taxa, including vertebrates, invertebrates, plants, and fungi, within a single region, Central Norway. We tested the hypotheses that ecological responses to climate change are apparent and coherent at a regional scale, that longer time series show stronger trends over time and in relation to temperature, and that ecological responses change in trajectory at the same time as shifts in temperature. We identified a clear regional coherence in climate signal, with decreasing abundances of limnic zooplankton (on average by 7691 individuals m-3 °C-1) and boreal forest breeding birds (on average by 1.94 territories km-2 °C-1), and earlier plant flowering phenology (on average 2 days °C-1) for every degree of temperature increase. In contrast, regional-scale species distributions and species diversity were largely stable. Surprisingly, the effect size of ecological response did not increase with study duration, and shifts in responses did not occur at the same time as shifts in temperature. This may be as the long-term studies include both periods of warming and temperature stability, and that ecological responses lag behind warming. Our findings demonstrate a regional climate fingerprint across a long timescale. We contend that natural history collections provide a unique window on a broad spectrum of ecological responses at timescales beyond most ecological monitoring programs. Natural history collections are thus an essential source for long-term ecological research.

Ice-cover effects on competitive interactions between two fish species.

1. Variations in the strength of ecological interactions between seasons have received little attention, despite an increased focus on climate alterations on ecosystems. Particularly, the winter situation is often neglected when studying competitive interactions. In northern temperate freshwaters, winter implies low temperatures and reduced food availability, but also strong reduction in ambient light because of ice and snow cover. Here, we study how brown trout [Salmo trutta (L.)] respond to variations in ice-cover duration and competition with Arctic charr [Salvelinus alpinus (L.)], by linking laboratory-derived physiological performance and field data on variation in abundance among and within natural brown trout populations. 2. Both Arctic charr and brown trout reduced resting metabolic rate under simulated ice-cover (darkness) in the laboratory, compared to no ice (6-h daylight). However, in contrast to brown trout, Arctic charr was able to obtain positive growth rate in darkness and had higher food intake in tank experiments than brown trout. Arctic charr also performed better (lower energy loss) under simulated ice-cover in a semi-natural environment with natural food supply. 3. When comparing brown trout biomass across 190 Norwegian lakes along a climate gradient, longer ice-covered duration decreased the biomass only in lakes where brown trout lived together with Arctic charr. We were not able to detect any effect of ice-cover on brown trout biomass in lakes where brown trout was the only fish species. 4. Similarly, a 25-year time series from a lake with both brown trout and Arctic charr showed that brown trout population growth rate depended on the interaction between ice breakup date and Arctic charr abundance. High charr abundance was correlated with low trout population growth rate only in combination with long winters. 5. In conclusion, the two species differed in performance under ice, and the observed outcome of competition in natural populations was strongly dependent on duration of the ice-covered period. Our study shows that changes in ice phenology may alter species interactions in Northern aquatic systems. Increased knowledge of how adaptations to winter conditions differ among coexisting species is therefore vital for our understanding of ecological impacts of climate change.

Population consequences of climate change through effects on functional traits of lentic brown trout in the sub-Arctic.

Climate-induced plasticity in functional traits has received recent attention due to the immense importance phenotypic variation plays in population level responses. Here, we explore the effect of different climate-change scenarios on lentic populations of a freshwater ectotherm, the brown trout (Salmo trutta L.), through climate effects on functional traits. We first parameterize models of climate variables on growth, spawning probability and fecundity. The models are utilized to inform a dynamic age-structured projection matrix, enabling long-term population viability projections under climate and population density variation. Ambient temperature and winter conditions had a substantial effect on population growth rate. In general, warmer summer temperatures resulted in faster growth rates for young fish but ended in smaller size at age as fish got older. Increasing summer temperatures also induced maturation at younger age and smaller size. In addition, we found effects of first-year growth on later growth trajectories for a fish, indicating that environmental conditions experienced the first year will also influence size at age later in life. At the population level, increasing temperatures average (up to 4 °C increase in areas with mean summer temperature at approximately 12 °C) resulted in a positive effect on population growth rate (i.e. smaller but more fish) during climate simulations including increasing and more variable temperatures.

Adaptive winter survival strategies: defended energy levels in juvenile Atlantic salmon along a latitudinal gradient.

Current knowledge suggests that patterns of energy storage and depletion in animals are governed by behavioural trade-offs between risks associated with feeding and future energy demands. However, the length of adverse periods varies over geographical or climatic gradients. To explore the potential for genotypic sources of variation in behavioural trade-offs, we compared the winter energy-depletion patterns among 13 wild populations of juvenile Atlantic salmon (Salmo salar L.) along a latitudinal gradient (58-70 degrees N) and performed common-environment experiments of energy-state-dependent feeding. In the wild, winter lipid-depletion rates were lower for northern than for southern populations. The variation in spring lipid levels among the population was lower than autumn variation, with storage lipid levels clustered close to critical limits for survival. In semi-natural stream channels with natural food supply, hatchery-reared fish originating from northern populations showed a positive scaling of feeding activity with decreasing energy levels, whereas southern populations did not. In conclusion, juvenile Atlantic salmon from northern populations defend their energy levels more strongly than fish from southern populations. Adaptive variation in feeding activity appears important for this difference. Thus, the present study shows a link between geographical patterns in storage energy trajectories and adaptive differences in state-dependent feeding motivation.

Spatial distribution of limited resources and local density regulation in juvenile Atlantic salmon.

1. Spatial heterogeneity of resources may influence competition among individuals and thus have a fundamental role in shaping population dynamics and carrying capacity. In the present study, we identify shelter opportunities as a limiting resource for juvenile Atlantic salmon (Salmo salar L.). Experimental and field studies are combined in order to demonstrate how the spatial distribution of shelters may influence population dynamics on both within and among population scales. 2. In closed experimental streams, fish performance scaled negatively with decreasing shelter availability and increasing densities. In contrast, the fish in open stream channels dispersed according to shelter availability and performance of fish remaining in the streams did not depend on initial density or shelters. 3. The field study confirmed that spatial variation in densities of 1-year-old juveniles was governed both by initial recruit density and shelter availability. Strength of density-dependent population regulation, measured as carrying capacity, increased with decreasing number of shelters. 4. Nine rivers were surveyed for spatial variation in shelter availability and increased shelter heterogeneity tended to decrease maximum observed population size (measured using catch statistics of adult salmon as a proxy). 5. Our studies highlight the importance of small-scale within-population spatial structure in population dynamics and demonstrate that not only the absolute amount of limiting resources but also their spatial arrangement can be an important factor influencing population carrying capacity.

Hydropower impacts on reservoir fish populations are modified by environmental variation.

Global transition towards renewable energy production has increased the demand for new and more flexible hydropower operations. Before management and stakeholders can make informed choices on potential mitigations, it is essential to understand how the hydropower reservoir ecosystems respond to water level regulation (WLR) impacts that are likely modified by the reservoirs' abiotic and biotic characteristics. Yet, most reservoir studies have been case-specific, which hampers large-scale planning, evaluation and mitigation actions across various reservoir ecosystems. Here, we investigated how the effect of the magnitude, frequency and duration of WLR on fish populations varies along environmental gradients. We used biomass, density, size, condition and maturation of brown trout (Salmo trutta L.) in Norwegian hydropower reservoirs as a measure of ecosystem response, and tested for interacting effects of WLR and lake morphometry, climatic conditions and fish community structure. Our results showed that environmental drivers modified the responses of brown trout populations to different WLR patterns. Specifically, brown trout biomass and density increased with WLR magnitude particularly in large and complex-shaped reservoirs, but the positive relationships were only evident in reservoirs with no other fish species. Moreover, increasing WLR frequency was associated with increased brown trout density but decreased condition of individuals within the populations. WLR duration had no significant impacts on brown trout, and the mean weight and maturation length of brown trout showed no significant response to any WLR metrics. Our study demonstrates that local environmental characteristics and the biotic community strongly modify the hydropower-induced WLR impacts on reservoir fishes and ecosystems, and that there are no one-size-fits-all solutions to mitigate environmental impacts. This knowledge is vital for sustainable planning, management and mitigation of hydropower operations that need to meet the increasing worldwide demand for both renewable energy and ecosystem services delivered by freshwaters.