Peter Eklöv

The contemporary distribution of genetic diversity in widespread freshwater species reflects a complex interplay between historical processes and recent demographic events. We investigated the postglacial recolonization history of the Eurasian perch ( Perca fluviatilis L.) across its native range spanning Europe and Western Siberia, aiming to understand how historical and recent demographic processes have shaped contemporary genetic diversity in a widespread freshwater species. Using an integrative genomic approach, we combined whole mitochondrial genome resequencing with nuclear SNP‐array genotyping (3660 SNPs) for 382 individuals from 188 locations to reconstruct patterns of lineage divergence, population structure, and admixture. We identified five highly divergent mitochondrial lineages, consistent with the existence of multiple glacial refugia across Southwestern, Southeastern, and Central Europe, as well as Siberia. Nuclear data (3660 SNPs) revealed three major genetic clusters corresponding to Western, Northern, and Southeastern Europe, along with strong regional admixture. The Baltic Sea region emerged as a contemporary hotspot of genetic diversity, likely resulting from the admixture and convergence of distinct maternal lineages during the postglacial recolonization of Northern Europe. Signals of isolation by distance were evident both within and across lineages, highlighting the role of limited dispersal in shaping current genetic patterns. The integration of mitochondrial and nuclear genomic data provided a comprehensive view of the evolutionary history of P. fluviatilis , revealing both deep historical divergence and recent admixture events. The existence of multiple glacial refugia and subsequent secondary contact underscores the complexity of postglacial recolonization processes in freshwater fauna. These findings advance our understanding of how historical and contemporary factors interact to shape biodiversity across Europe.

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. 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. 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. Secchi depth, the pelagic contribution to the diet and the percentage of piscivores had significant effects on eye size. 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. Read the free Plain Language Summary for this article on the Journal blog.

Dissolved organic matter (DOM), often measured as dissolved organic carbon (DOC), plays a fundamental role in influencing the structure and function of lake ecosystems. Due to the myriad ecosystem effects of DOM, widespread observations of long‐term increasing DOM concentrations have received much attention from ecologists. DOM positively influences primary production and consumer production at low concentrations due to the fertilising influence of bound nutrients. However, beyond a unimodal peak in production, a reduced light environment may result in a negative effect on production. This unimodal model has been largely developed and tested in lakes with low to moderate DOM concentrations (i.e., typically ≤10 mg/L DOC). To understand ecological responses in lakes across a larger range in DOM concentrations, we examined the response of benthic invertebrate communities in 148 Swedish lakes with DOM concentrations ranging between 0.67 and 32.77 mg/L DOC. We found that increasing DOM concentrations had a strong effect on invertebrate community composition below c. 10 mg/L. Across this range, abundances of individual taxa both increased and decreased, probably in response to environmental change induced by DOM. However, in lakes above this concentration, increasing DOM had minimal influence on community composition. As DOM concentrations continue to increase, faunal communities in lakes below this 10 mg/L DOC threshold are likely to undergo substantial change whereas those above this threshold are likely to be minimally impacted.

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.

Ecological stability encompasses multiple dimensions of functional and compositional responses to environmental change. Though no single stability dimension used in isolation can fully reflect the overall response to environmental change, a common vulnerability assessment that integrates simultaneously across multiple stability components is highly desirable for ecological risk assessment. We develop both functional and compositional counterparts of a novel, integrative metric of overall ecological vulnerability (OEV). We test the framework with data from a modularized experiment replicated in five lakes over two seasons, examining functional and compositional responses to both pulse and press disturbances across three trophic groups. OEV is measured as the area under the curve integrated over the entire observation period, with the curve delimiting the difference between the disturbance treatment and undisturbed parallel controls, expressed either as the log response ratio of biomass (functional OEV) or community dissimilarity index (compositional OEV). Both, functional and compositional OEV correlated negatively with functional and compositional ‘resistance’, ‘temporal stability’ and ‘final/extent of recovery’ following both pulse and press disturbances, though less so with ‘resilience’ following a pulse disturbance. We also found a positive correlation between functional and compositional OEV, which reveals the potential to also evaluate the intricate linkage between biodiversity and functional change. Our findings demonstrate that OEV comprises a robust framework to: (a) capture simultaneously multiple functional and compositional stability components, and (b) quantify the functional consequences of biodiversity change. Our results provide the basis for an overarching framework for quantifying the overall vulnerability of ecosystems to environmental change, opening new possibilities for ecological risk assessment and management. Synthesis. Ecological stability comprises multiple dimensions that together encapsulate how ecosystems respond to environmental change. Considering these multiple aspects of stability simultaneously often poses a problem in environmental assessments, which frequently require overarching indicators of risk or vulnerability. While an analysis of multiple dimensions allows for deeper exploration of mechanisms, here we develop and test a new univariate indicator that integrates stability aspects under a broad range of disturbance regimes. Using a modularized experiment in Swedish lakes, we show that this integrative measure captures multiple stability dimensions reflecting compositional and functional vulnerability and their relationships between them.

Stable isotopes represent a unique approach to provide insights into the ecology of organisms. δ¹³C and δ¹⁵N have specifically been 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. Here, we conducted a 9‐month feeding experiment to report Δ¹³C and Δ¹⁵N of muscle and liver tissues 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 the oxygen consumption of the individuals. 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. 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.

Phenotypic plasticity is common among animal taxa. While there are clearly limits and likely costs to plasticity, these costs are unknown for most organisms. Further, as plasticity is partially genetically determined, the potential magnitude of exhibited plasticity may vary among individuals. In addition to phenotypic plasticity, various animal taxa also display sexual size dimorphism, a feature ultimately thought to arise due to differential size-dependent fitness costs and benefits between sexes. We hypothesized that differential selection acting on males and females can indirectly select for unequal genetically defined plasticity potential between the sexes. We evaluate this possibility for Eurasian perch (Perca fluviatilis), a species that displays modest sexual size dimorphism and habitat-related morphological plasticity. Using 500-year simulations of an ecogenetic agent-based model, we demonstrate that genetically determined morphological plasticity potential may evolve differently for males and females, leading to greater realized morphological variation between habitats for one sex over the other. Genetically determined potential for plasticity evolved differently between sexes across (a) various sex-specific life-history differences and (b) a variety of assumed costs of plasticity acting on both growth and survival. Morphological analyses of Eurasian perch collected in situ were consistent with model predictions: realized morphological variation between habitats was greater for females than males. We suggest that due to sex-specific selective pressures, differences in male and female genetically defined potential for plasticity may be a common feature across organisms.

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.

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.

Aquatic and terrestrial ecosystems differ fundamentally in the abundance of long‐chained polyunsaturated fatty acids (PUFAs), such as eicosapentaenoic acid (EPA; 20:5n‐3), which are produced by aquatic algae, but only in low quantities by terrestrial plants. Aquatic insects, such as Chironomidae (non‐biting midges) feed on algae during their larval stage, making them rich in EPA and therefore high‐quality prey for insectivores after emergence. However, the magnitude of EPA subsidies from aquatic insects may be different among water bodies in response to abiotic (e.g. nutrient load) as well as biotic factors (e.g. food web structure). To test the predation effects of crucian carp (Carassius carassius), nutrient concentrations, and Chironomidae community composition on the fatty acid export from aquatic ecosystems, we conducted a 25‐day experiment across 20 1,500‐L mesocosms covering a total phosphorus (TP) gradient of 20–1,000 µg/L. Twice a week, we collected adult emerging Chironomidae and found differences in fatty acid composition in the two most abundant chironomid species emerging from the mesocosms. Two PUFAs, α‐linolenic acid and EPA, contributed to most of the variation in Chironomidae fatty acid content across the nutrient gradient. Whereas the proportions of α‐linolenic acid were positively correlated to the mesocosm TP concentration, we found a negative correlation for the proportions of Chironomidae EPA and mesocosm TP concentration. However, despite lower biomass‐specific EPA content at higher TP, higher biomass of emerging Chironomidae at intermediate TP concentrations resulted in higher total export of PUFAs from water to land. Predation pressure from carp decreased the biomass of emerging Chironomidae on average 8‐fold. Chironomidae biomass showed a hump‐shaped relationship along the TP‐gradient and was strongly influenced by periphyton biomass. Export rates of EPA and fatty acids in general responded in a quadratic manner along the nutrient gradient, reaching a maximum value at a TP of 400 µg/L and decreasing thereafter. These findings highlight that the export of fatty acids from aquatic systems via adult Chironomidae is highly dependent on fish predation pressure, but also the nutrient concentrations of the system.