Water bodies located in the agricultural landscape may face a substantial contamination by agrochemicals due to modern agricultural practice. Many insect species inhabiting these water bodies have a complex life-cycle with an aquatic phase as larvae and a terrestrial phase as winged adults when they serve as prey for many aerial insectivores, such as bats and birds. Thus, they may act as vectors, transferring pesticides from the water bodies into the terrestrial food webs. This transfer of pesticides from water to land via emerging insects is still poorly studied. This study investigated the contamination of feces from various insectivorous organisms with different foraging mode and aquatic habitat use, quantified by movement analysis. In total, we detected 16 current-use pesticide residues, two legacy compounds and six metabolites in the feces of three species which had a high selection strength towards aquatic habitats: barn swallow (Hirundo rustica), Western house martin (Delichon urbicum), and the common noctule bat (Nyctalus noctula). In contrast, no substances were detected in feces of European starling (Sturnus vulgaris), a species that uses rather terrestrial habitats and is known to feed on terrestrial insects. The fungicide prochloraz, used to combat fungal grain diseases, was the substance with the highest detection rate of 69% of all samples. Five of the substances detected (i.e., bixafen, diflufenican, dinoterb, prochloraz, simazine) are classified as critical in terms of their potential for bioconcentration and ten out of 18 substances are classified to be of concern for mammal short term dietary uptake. While the direct ecotoxicological effects on the organisms are unclear, our study is one of the first ones to infer the contamination pathway via emerging insects and to highlight that species-specific use of aquatic habitats increases pesticide exposure risk in insect-eating birds and bats in farmland.

The browning of freshwater ecosystems is increasingly evident in temperate and northern regions, with widespread ramifications for lake physics, chemistry, and biology. Contrasting results on how freshwater browning may impact fish have been reported, but there has been no comprehensive examination of how browning may cause cascading effects on individual- to population- to community-level traits of freshwater fishes. We addressed this knowledge gap by summarizing the existing literature and conducting a series of original analyses to: (i) explore the effects of a brown water gradient on populations of eight economically important species of fish across 871 lakes; and (ii) examine how a brown water gradient may influence community trait compositions across 303 lakes. From our literature synthesis, we found that fish growth is often negatively associated with browner waters, despite browning generally showing no effect on fish foraging. We also demonstrated that browner waters had greater abundances of northern pike (Esox lucius) and walleye (Sander vitreus), but lower numbers of lake trout (Salvelinus namaycush), yellow perch (Perca flavescens), largemouth bass (Micropterus salmoides), smallmouth bass (M. dolomieu), and lake whitefish (Coregonus clupeaformis). Moreover, we showed that fish communities were significantly more likely to contain species with larger eyes in browner lakes. Lastly, we examined relationships between various metrics of browning (i.e. dissolved organic carbon, Secchi transparency, water colour) and present a framework for how the effects of freshwater browning on fish may scale from individuals to populations to communities.

Freshwater ecosystems are affected by fluctuations in terrestrially derived dissolved organic carbon (DOC). Increased DOC runoff from watersheds can contribute to the “browning” of freshwater systems, altering water properties and disrupting food web dynamics, with potential impacts on fish. This study examined how carbon sources and the trophic ecology of temperate freshwater fishes varied with DOC concentrations. We used stable isotope ratios of carbon (δ13C) and nitrogen (δ15N) to trace carbon sources in organisms at low trophic levels and to assess the trophic ecology of six fish species from 70 lakes in Sweden, Germany, Canada, and the United States. We found that both pelagic and benthic δ13C decreased as DOC increased, suggesting higher reliance on terrestrial carbon by lower trophic level organisms. Fish responses to elevated DOC were limited to certain species. Mid-trophic level European perch (Perca fluviatilis) increased pelagic diet proportions, while top predators like northern pike (Esox Lucius) and walleye (Sander vitreus) showed nonlinear shifts. Except for walleye, no effect of DOC was detected on fish trophic position. Our findings suggest that DOC alters energy use and diet but not trophic position in some freshwater fish.

Bleak Alburnus alburnus is a highly abundant but understudied fish species, and we know little about the trophic ecology of populations inhabiting rivers in central Europe. From an ecosystem perspective, this fish species is interesting as it is known to feed on surface insects, thereby linking the terrestrial with the aquatic habitat. In a previous study, we demonstrated that this flux is intensified, and dietary contribution of terrestrial insects is higher in fish inhabiting sections of the Spree River, Germany, that are polluted from iron oxides occurring from former lignite mining activities, and thus are characterized by lower abundances of aquatic insects. As terrestrial insects can be considered as food of lower quality (measured as long-chained polyunsaturated fatty acids, n-3 LC-PUFAs) compared to aquatic prey, it is reasonable to assume that the higher contribution of terrestrial insects is related to a lower body condition in fish. In this study, we explore the trophic ecology of riverine A. alburnus and their fitness consequences of feeding on terrestrial insects. We therefore modeled a terrestrial index from stable isotopes of hydrogen (δ2H) measured in the A. alburnus muscle tissue and compared individuals caught in locations upstream of a dam that were greatly influenced by iron oxides, with individuals caught in sections located downstream of a dam where passive remediation technologies are applied. The terrestrial index was significantly higher in A. alburnus caught in locations at high-iron concentrations, characterized by low abundances of aquatic prey, compared to A. alburnus caught in unpolluted habitats at low-iron concentrations. In contradiction to our hypothesis, the terrestrial index had no significant effect on the body condition of A. alburnus (measured as Fulton's condition factor K) in the sections downstream of the dam (i.e., at low-iron concentrations) and a significant positive, albeit weak, effect in sections upstream of the dam (i.e., at high-iron concentrations). However, the condition factor was generally lower in the high-iron section, potentially related to more direct effects of the iron oxide. We conclude that in A. alburnus, terrestrial insects can be considered as the less-favored food, unless the fish occur in environments where the aquatic food is of limited availability. Further research is needed to evaluate the direct and indirect effects, including the internal n-3 LC-PUFA synthesis as an adaption toward low-quality terrestrial prey on the fitness consequences of A. alburnus.

1. Variations in the size and shape of the eye have been observed in many species of fish. As eye size is positively related to visual acuity, larger eyes should favour foraging and detection of predators. 2. However, few studies have examined the variation in eye morphology in relation to the complexity of lake conditions, including environmental perturbation and spatial variation in predation and competition. Such tests are especially important as the degrading of the visual climate is expected due to climate change, where browning, turbidity and variations in structural complexity should set different demands for visual acuity of foraging fish under predation risk. 3. In this study, we tested the variation of the eye size among 667 individuals of an aquatic predator perch, Perca fluviatilis, from littoral and pelagic habitats of 14 lakes. We used Secchi depth to assess the effects of the visual climate of our lake systems, as fish foraging is highly related to visual conditions, and studied eye size variation in relation to the contribution of the pelagic resources to an individual's diet and the risk of predation. 4. Secchi depth, the pelagic contribution to the diet and the percentage of piscivores had significant effects on eye size. 5. These variable outcomes suggest that the lake environment in terms of visual climate, predation landscape and diet are major factors of eye size variation in this generalist predator. As many fish species trade off foraging against predation risk, future studies will show whether the complexity of intra- and interspecific interactions contribute to the variation in eye size in freshwater fish.

Fish communities of streams and rivers might be substantially subsidized by terrestrial insects that fall into the water. Although such animal-mediated fluxes are increasingly recognized, little is known about how anthropogenic perturbations may influence the strength of such exchanges. Intense land use, such as lignite mining, may impact a river ecosystem due to the flocculation of iron (III) oxides, thus altering food web dynamics. We compared sections of the Spree River in North-East Germany that were greatly influenced by iron oxides with sections located downstream of a dam where passive remediation technologies are applied. Compared to locations downstream of the dam, the abundance of benthic macroinvertebrates at locations of high iron concentrations upstream of the dam was significantly reduced. Similarly, catch per unit effort of all fish was significantly higher in locations downstream of the dam compared to locations upstream of the dam, and the condition of juvenile and adult piscivorous pike Esox lucius was significantly lower in sections of high iron concentrations. Using an estimate of short-term (i.e., metabarcoding of the gut content) as well as longer-term (i.e., hydrogen stable isotopes) resource use, we could demonstrate that the three most abundant fish species, perch Perca fluviatilis, roach Rutilus rutilus, and bleak Alburnus alburnus, received higher contributions of terrestrial insects to their diet at locations of high iron concentration. In summary, lotic food webs upstream and downstream of the dam greatly differed in the overall structure with respect to the energy available for the highest tropic levels and the contribution of terrestrial insects to the diet of omnivorous fish. Therefore, human-induced environmental perturbations, such as river damming and mining activities, represent strong pressures that can alter the flow of energy between aquatic and terrestrial systems, indicating a broad impact on the landscape level.

Stable isotopes, eye size, and body length for each fish

Trophic transfer efficiency (TTE) is usually calculated as the ratio of production rates between two consecutive trophic levels. Although seemingly simple, TTE estimates from lakes are rare. In our review, we explore the processes and structures that must be understood for a proper lake TTE estimate. We briefly discuss measurements of production rates and trophic positions and mention how ecological efficiencies, nutrients (N, P) and other compounds (fatty acids) affect energy transfer between trophic levels and hence TTE. Furthermore, we elucidate how TTE estimates are linked with size-based approaches according to the Metabolic Theory of Ecology, and how food-web models can be applied to study TTE in lakes. Subsequently, we explore temporal and spatial heterogeneity of production and TTE in lakes, with a particular focus on the links between benthic and pelagic habitats and between the lake and the terrestrial environment. We provide an overview of TTE estimates from lakes found in the published literature. Finally, we present two alternative approaches to estimating TTE. First, TTE can be seen as a mechanistic quantity informing about the energy and matter flow between producer and consumer groups. This approach is informative with respect to food-web structure, but requires enormous amounts of data. The greatest uncertainty comes from the proper consideration of basal production to estimate TTE of omnivorous organisms. An alternative approach is estimating food-chain and food-web efficiencies, by comparing the heterotrophic production of single consumer levels or the total sum of all heterotrophic production including that of heterotrophic bacteria to the total sum of primary production. We close the review by pointing to a few research questions that would benefit from more frequent and standardized estimates of TTE in lakes.

Raw data and R scripts of all analyses of the paper.

Abstract Resource polymorphism is common across taxa and can result in alternate ecotypes with specific morphologies, feeding modes, and behaviors that increase performance in a specific habitat. This can result in high intraspecific variation in the expression of specific traits and the extent to which these traits are correlated within a single population. Although metabolic rate influences resource acquisition and the overall pace of life of individuals it is not clear how metabolic rate interacts with the larger suite of traits to ultimately determine individual fitness. We examined the relationship between metabolic rates and the major differences (habitat use, morphology, and resource use) between littoral and pelagic ecotypes of European perch (Perca fluviatilis) from a single lake in Central Sweden. Standard metabolic rate (SMR) was significantly higher in pelagic perch but did not correlate with resource use or morphology. Maximum metabolic rate (MMR) was not correlated with any of our explanatory variables or with SMR. Aerobic scope (AS) showed the same pattern as SMR, differing across habitats, but contrary to expectations, was lower in pelagic perch. This study helps to establish a framework for future experiments further exploring the drivers of intraspecific differences in metabolism. In addition, since metabolic rates scale with temperature and determine predator energy requirements, our observed differences in SMR across habitats will help determine ecotype‐specific vulnerabilities to climate change and differences in top‐down predation pressure across habitats.

The physiological dependence of animals on dietary intake of vitamins, amino acids, and fatty acids is ubiquitous. Sharp differences in the availability of these vital dietary biomolecules among different resources mean that consumers must adopt a range of strategies to meet their physiological needs. We review the emerging work on omega-3 long-chain polyunsaturated fatty acids, focusing predominantly on predator-prey interactions, to illustrate that trade-off between capacities to consume resources rich in vital biomolecules and internal synthesis capacity drives differences in phenotype and fitness of consumers. This can then feedback to impact ecosystem functioning. We outline how focus on vital dietary biomolecules in eco-eco-devo dynamics can improve our understanding of anthropogenic changes across multiple levels of biological organization.

Bats are protected by national and international legislation in European countries, yet many species, particularly migratory aerial insectivores, collide with wind turbines which counteracts conservation efforts. Within the European Union it is legally required to curtail the operation of wind turbines at periods of high bat activity, yet this is not practiced at old wind turbines. Based on data from the national carcass repository in Germany and from our own carcass searches at a wind park with three turbines west of Berlin, we evaluated the magnitude of bat casualties at old, potentially poor-sited wind turbines operating without curtailment. We report 88 documented bat carcasses collected by various searchers over the 20-year operation period of this wind park from 2001 to 2021. Common noctule bats (Nyctalus noctula) and common pipistrelles (Pipistrellus pipistrellus) were most often found dead at these turbines. Our search campaign in August and September 2021 yielded a total of 18 carcasses. We estimated that at least 209 bats were likely killed during our field survey, yielding more than 70 casualties/wind turbine or 39 casualties/ MW in two months. Since our campaign covered only part of the migration season, we consider this value as an underestimate. The 20-year period of the wind park emphasises the substantial impact old turbines may have on bat individuals and populations when operating without curtailments. We call for reconsidering the operation procedures of old wind turbines to stop the continuous loss of bats in Germany and other countries where turbine curtailments are even less practiced than in Germany.

The nutritional diversity of resources can affect the adaptive evolution of consumer metabolism and consumer diversification. The omega-3 long-chain polyunsaturated fatty acids eicosapentaenoic acid (EPA; 20:5n-3) and docosahexaenoic acid (DHA; 22:6n-3) have a high potential to affect consumer fitness, through their widespread effects on reproduction, growth and survival. However, few studies consider the evolution of fatty acid metabolism within an ecological context. In this review, we first document the extensive diversity in both primary producer and consumer fatty acid distributions amongst major ecosystems, between habitats and amongst species within habitats. We highlight some of the key nutritional contrasts that can shape behavioural and/or metabolic adaptation in consumers, discussing how consumers can evolve in response to the spatial, seasonal and community-level variation of resource quality. We propose a hierarchical trait-based approach for studying the evolution of consumers' metabolic networks and review the evolutionary genetic mechanisms underpinning consumer adaptation to EPA and DHA distributions. In doing so, we consider how the metabolic traits of consumers are hierarchically structured, from cell membrane function to maternal investment, and have strongly environment-dependent expression. Finally, we conclude with an outlook on how studying the metabolic adaptation of consumers within the context of nutritional landscapes can open up new opportunities for understanding evolutionary diversification.

1. Stable isotopes represent a unique approach to provide insights into the ecology of organisms. δ13C and δ15N have specifically be used to obtain information on the trophic ecology and food web interactions. Trophic discrimination factors (TDF, Δ13C and Δ15N) describe the isotopic fractionation occurring from diet to consumer tissue and these factors are critical for obtaining precise estimates within any application of δ13C and δ15N values. It is widely acknowledged that metabolism influences TDF, being responsible for different TDF between tissues of variable metabolic activity (e.g. liver vs. muscle tissue) or species body size (small vs. large). However, the connection between the variation of metabolism occurring within a single species during its ontogeny and TDF has rarely been considered. 2. Here, we conducted a 9-month feeding experiment to report Δ13C and Δ15N of muscle and liver tissue for several weight classes of Eurasian perch (Perca fluviatilis), a widespread teleost often studied using stable isotopes, but without established TDF for feeding on a natural diet. In addition, we assessed the relationship between the standard metabolic rate (SMR) and TDF by measuring their oxygen consumption of the individuals. 3. Our results showed a significant negative relationship of SMR with Δ13C, and a significant positive relationship of SMR with Δ15N of muscle tissue, but not with TDF of liver tissue. SMR varies inversely with size, which translated into a significantly different TDF of muscle tissue between size classes. 4. In summary, our results emphasize the role of metabolism in shaping specific TDF (i.e. Δ13C and Δ15N of muscle tissue), and especially highlight the substantial differences between individuals of different ontogenetic stages within a species. Our findings thus have direct implications for the use of stable isotope data and the applications of stable isotopes in food web studies.

Inland waters receive and process large amounts of colored organic matter from the terrestrial surroundings. These inputs dramatically affect the chemical, physical, and biological properties of water bodies, as well as their roles as global carbon sinks and sources. To understand the complex changes associated with allochthonous inputs, experiments are needed. However, manipulative studies, especially at ecosystem scales, require large amounts of dissolved organic matter with optical and chemical properties resembling indigenous organic matter. Here we compared the chemical and biological impacts of two leonardite products (HuminFeed (HF) and SuperHume (SH)) and a freshly derived reverse osmosis concentrate of organic matter (RO) in a set of comprehensive mesocosm- and laboratory-scale experiments and analyses. The chemical properties of RO concentrate and the leonardite products were very different with leonardite products being low and RO being high in carboxylic functional groups. Light had a strong impact on the properties of leonardite products, including loss of color and increased particle formation. Furthermore, HF had drastic impacts on bacteria as light stimulated bacterial production and modified community composition, while dark conditions appeared to inhibit bacterial processes. While none of the browning agents inhibited the growth of the tested phytoplankton, Gonyostomum semen , leonardite products had detrimental effects on zooplankton abundance and Daphnia reproduction. We conclude that the effects of browning agents extracted from leonardite are in sharp contrast to those originating from terrestrially-derived DOM. Hence, they should be used with great caution in experimental studies on the consequences of terrestrial carbon for aquatic systems.

Fatty acids (FAs) are key nutrients for fitness which take part in multiple physiological processes over the ontogeny of organisms. Yet, we lack evidence on how FA nutrition mediates life-history trade-offs and ontogenetic niche shifts in natural populations. In a field study, we analyzed ontogenetic changes in the FAs of Eurasian perch ( Perca fluviatilis L.), a widespread fish that goes through ontogenetic niche shifts and can have high individual niche specialization. Diet explained most of the variation in the FA composition of perch dorsal muscle over early ontogeny (28%), while the total length explained 23%, suggesting that perch significantly regulated FA composition over early ontogeny. Condition explained 1% of the remaining variation. 18:3n-3 (ALA) and 18:4n-3 (SDA) indicated planktivory; 18:1n-7, benthivory; and 22:6n-3 (DHA), piscivory in perch diet. Conversely, perch regulated long-chained polyunsaturated fatty acids (PUFAs), such as 20:5n-3 (EPA), 20:4n-6 (ARA) and 22:6n-3 (DHA) over ontogeny, emphasizing the role of such FAs in early growth and sexual maturation. Adult perch increasingly retained 16:1n-7 and 18:1n-9 suggesting higher energy storage in older perch. Furthermore, differences in DHA availability in diet correlated with intra-cohort differences in perch growth, potentially hindering the overall use of benthic resources and promoting earlier shifts to piscivory in littoral habitats. Overall, this study indicates that in addition to diet, internal regulation may be more important for FA composition than previously thought. Differences between FA needs and FA availability may lead to life-history trade-offs that affect the ecology of consumers, including their niche.

Abstract Phenotypic divergence in response to divergent natural selection between environments is a common phenomenon in species of freshwater fishes. Intraspecific differentiation is often pronounced between individuals inhabiting lakes versus stream habitats. The different hydrodynamic regimes in the contrasting habitats may promote a variation of body shape, but this could be intertwined with morphological adaptations to a specific foraging mode. Herein, I studied the divergence pattern of the European minnow (Phoxinus phoxinus), a common freshwater fish that has received little attention despite its large distribution. In many Scandinavian mountain lakes, European minnows are considered as being invasive and were found to pose threats to the native fish populations due to resource competition. Minnows were recently found to show phenotypic adaptations in lake versus stream habitats, but the question remained if this divergence pattern is related to differences in resource use. I therefore studied the patterns of minnow divergence in morphology (i.e., using geometric morphometrics) and trophic niches (i.e., using stomach content analyses) in the lake Ånnsjön and its tributaries to link the changes in body morphology to the feeding on specific resources. Lake minnows showed a strong reliance on benthic Cladocera and a more streamlined body shape with a more upward facing snout, whereas stream minnows fed on macroinvertebrates (larvae and adults) to a higher degree and had a deeper body with a snout that was pointed down. Correlations showed a significant relationship of the proportion of macroinvertebrates in the gut and morphological features present in the stream minnows. The results of this study highlight the habitat‐specific divergence pattern in morphology and resource use in this ubiquitous freshwater fish. Consequently, interspecific interactions of invasive minnows and the native fish population could differ in the respective food webs and resource competition could target different native fish species in the contrasting habitats.

AbstractIndividual specialization is a common phenomenon throughout the animal kingdom. Many studies have identified intraspecific competition as one of the main drivers for individual feeding specialization. These studies have mainly considered the quantity of resources, commonly overlooking qualitative aspects of the diet. For example, highly unsaturated fatty acids of the ω‐3 class (ω‐3 HUFAs) are related to optimal health and growth in consumers. However, little is known on direct fitness consequences for consumers of natural populations that specialize on high‐quality resources, such as those rich in ω‐3 HUFAs. Despite being such an important qualitative aspect of the diet, it is still unknown whether natural populations show among‐individual variation in their choice on prey items that are either rich or poor in HUFAs, and how it affects individual performances. In this study, we investigated whether there is individual feeding specialization and whether it is related to fitness benefits, in a population of perch (Perca fluviatilis) in the Baltic Sea. The contribution of pelagic planktivorous fish to the diet varied from 17% to 61% among perch individuals, as depicted by stable isotope mixing models. This variation in diet was also qualitative, as the ω‐3 HUFA content differed among prey types. Specialization on the high‐quality resource pelagic planktivorous fish was associated with the proportions of ω‐3 HUFA in the individuals’ muscles and individuals among those with the highest proportions of ω‐3 HUFAs had the greatest relative gonad weight (gonadosomatic index, GSI), a proxy for reproductive investment. Thus, our results highlight the function of food quality for individual specialization and its potential to have direct fitness benefits, playing a major role in shaping ecological interactions.

According to apparent competition theory, sharing a predator should cause indirect interactions among prey that can affect the structure and the dynamics of natural communities. Though shifts in prey dominance and predator resource use along environmental gradients are rather common, empirical evidence on the role of indirect prey-prey interactions through shared predation particularly with increasing productivity, is still scarce. In an 8-week lake mesocosm experiment, we manipulated both the addition of inorganic nutrients and the presence of generalist fish predators (crucian carp, Carassius carassius L.), to test for the effects of indirect interactions through shared predation along a productivity gradient. We found that apparent mutualism (indirect positive interaction) between benthic and pelagic prey strongly affected short-term responses of aquatic food webs to increasing productivity in the presence of a generalist fish. Increasing productivity favoured the relative abundance of benthic prey, following trends in natural productive lake systems. This led to a shift in fish selectivity from pelagic to benthic prey driven by changes in fish behaviour, which resulted in apparent mutualism due to the lower and delayed top-down control of pelagic prey at increasing productivity. Our results show empirical evidence that the coupling of multiple production pathways can lead to strong indirect interactions through shared predation, whereby prey dynamics on short time-scales are highly dependent on the foraging behaviour of generalist predators. This mechanism may play an important role in short-term responses of food webs across environmental gradients.