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Boreal lakes experience pronounced seasonal variation in abiotic factors, especially light, temperature, and oxygen. A deep boreal humic lake was sampled year-round to test putative changes in total fish catch, species composition, catch-per-unit-effort (CPUE), habitat use, fish size, and condition. Monthly sampling was conducted in Lake Pääjärvi, southern Finland, during one full year in 2020-2021 as well as in March and August 2021 and 2022. The fish community was dominated by cyprinid species in all months, but the percentage of percid fish caught increased during the warm summer period. Most fish were caught in littoral habitats and the highest catches occurred in summer, but some species (e.g., ruffe, Gymnocephalus cernua, and pikeperch, Sander lucioperca) remained abundant in the winter catch. The body size of fish was larger in the winter catch, while condition factor was higher in summer for most species. Fish species proportions in total catch, CPUE, and average size of fish were closest to the annual mean values in September, which may be used as the optimal period to monitor fish communities of similar deep boreal lakes. Our findings highlight the need for year-round research to reveal the impacts of rising temperatures and diminishing ice-covered periods in fish communities and lake food webs., (© 2024 The Author(s). Journal of Fish Biology published by John Wiley & Sons Ltd on behalf of Fisheries Society of the British Isles.)

Global change may introduce fundamental alterations in phytoplankton biomass and community structure that can alter the productivity of northern lakes. In this study, we utilized Swedish and Finnish monitoring data from lakes that are spatially (135 lakes) and temporally (1995-2019, 110 lakes) extensive to assess how phytoplankton biomass (PB) of dominant phytoplankton groups related to changes in water temperature, pH and key nutrients [total phosphorus (TP), total nitrogen (TN), total organic carbon (TOC), iron (Fe)] along spatial (Fennoscandia) and temporal (25 years) gradients. Using a machine learning approach, we found that TP was the most important determinant of total PB and biomass of a specific species of Raphidophyceae - Gonyostomum semen - and Cyanobacteria (both typically with adverse impacts on food-webs and water quality) in spatial analyses, while Fe and pH were second in importance for G. semen and TN and pH were second and third in importance for Cyanobacteria. However, in temporal analyses, decreasing Fe and increasing pH and TOC were associated with a decrease in G. semen and an increase in Cyanobacteria. In addition, in many lakes increasing TOC seemed to have generated browning to an extent that significantly reduced PB. The identified discrepancy between the spatial and temporal results suggests that substitutions of data for space-for-time may not be adequate to characterize long-term effects of global change on phytoplankton. Further, we found that total PB exhibited contrasting temporal trends (increasing in northern- and decreasing in southern Fennoscandia), with the decline in total PB being more pronounced than the increase. Among phytoplankton, G. semen biomass showed the strongest decline, while cyanobacterial biomass showed the strongest increase over 25 years. Our findings suggest that progressing browning and changes in Fe and pH promote significant temporal changes in PB and shifts in phytoplankton community structures in northern lakes., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

Zooplankton community composition of northern lakes is changing due to the interactive effects of climate change and recovery from acidification, yet limited data are available to assess these changes combined. Here, we built a database using archives of temperature, water chemistry and zooplankton data from 60 Scandinavian lakes that represent broad spatial and temporal gradients in key parameters: temperature, calcium (Ca), total phosphorus (TP), total organic carbon (TOC), and pH. Using machine learning techniques, we found that Ca was the most important determinant of the relative abundance of all zooplankton groups studied, while pH was second, and TOC third in importance. Further, we found that Ca is declining in almost all lakes, and we detected a critical Ca threshold in lake water of 1.3 mg L -1 , below which the relative abundance of zooplankton shifts toward dominance of Holopedium gibberum and small cladocerans at the expense of Daphnia and copepods. Our findings suggest that low Ca concentrations may shape zooplankton communities, and that current trajectories of Ca decline could promote widespread changes in pelagic food webs as zooplankton are important trophic links from phytoplankton to fish and different zooplankton species play different roles in this context., (© 2024 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Eutrophication, i.e. increasing level of nutrients and primary production, is a central environmental change of lakes globally with wide effects on food webs. However, how eutrophication affects the synthesis of physiologically essential biomolecules (omega-3 fatty acids) and their transfer to higher trophic levels at the whole food web level is not well understood. We assessed food web (phytoplankton, zooplankton, and fish) biomass, community structure and fatty acid content (eicosapentaenoic acid [EPA], and docosahexaenoic acid [DHA]), together with fatty acid specific primary production in 12 Finnish boreal lakes covering the total nutrient gradient from oligotrophic to highly eutrophic lakes (4-140 μg TP l -1 ; 413-1814 μg TN l -1 ). Production was measured as the incorporation of 13 C-NaHCO 3 into phytoplankton fatty acids and differentiated into volumetric production (production per litre of water) and productivity (production per phytoplankton biomass). Increases in nutrients led to higher biomass of phytoplankton, zooplankton and fish communities while also affecting community composition. Eutrophication negatively influenced the contribution of phytoplankton biomass preferentially grazed by zooplankton (<35 μm). Total volumetric production saturated at high phytoplankton biomass while EPA volumetric production presented a logarithmic relationship with nutrient increase. Meanwhile, total and EPA productivity had unimodal responses to this change in nutrients. DHA volumetric production and productivity presented large variation with increases in total phosphorus, but a unimodal model best described DHA changes with eutrophication. Results showed that eutrophication impaired the transfer of EPA and DHA into zooplankton and fish, showing a clear negative impact in some species (e.g. perch) while having no effect in other species (e.g. roach, ruffe). Results show non-linear trends in fatty acid production and productivity peaking at nutrient concentrations 22-35 μg l -1 TP followed by a gradual decrease., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. At all stages of this study, national guidelines (FI-564/2013 & FI-487/2013) and the European Union directive (2010/63/EU) on the protection of animals used for scientific purposes were applied., (Copyright © 2023. Published by Elsevier B.V.)

Long-chain polyunsaturated fatty acids (PUFA) are critical for reproduction and thermal adaptation. Year-round variability in the expression of fads2 (fatty acid desaturase 2) in the liver of European perch (Perca fluviatilis) in a boreal lake was tested in relation to individual variation in size, sex, and maturity, together with stable isotopes values as well as fatty acids (FA) content in different tissues and prey items. ARA and DHA primary production was restricted to the summer months, however, perch required larger amounts of these PUFA during winter, as their ARA and DHA muscle content was higher compared to summer. The expression of fads2 in perch liver increased during winter and was higher in mature females. Mature females stored DHA in their gonads already in late summer and autumn, long before the upcoming spring spawning period in May. Lower δ 13 C DHA values in the gonads in September suggest that these females actively synthesized DHA as part of this reproductive investment. Lower δ 13 C ARA values in the liver of all individuals during winter suggest that perch were synthesizing essential FA to help cope with over-wintering conditions. Perch seem able to modulate its biosynthesis of physiologically required PUFA in situations of stress (fasting or cold temperatures) or in situations of high energetic demand (gonadal development). Biosynthesis of physiologically required PUFA may be an important part of survival and reproduction in aquatic food webs with long cold periods., (© 2023. The Author(s).)

Predation is a major evolutionary force determining life-history traits in prey by direct and indirect mechanisms. This study focuses on life-history trait variation in crucian carp (Carassius carassius), a species well known for developing a deep body as an inducible morphological defence against predation risk. Here, the authors tested variation in growth and reproductive traits in 15 crucian carp populations in lakes along a predation risk gradient represented by increasingly efficient predator communities. Lakes were located in south-eastern Norway and were sampled in summer 2018 and 2019. The authors expected crucian carp to attain higher growth rate, larger size, and later age at maturity with increasing predation risk. In the absence of predators, they expected high adult mortality, early maturity and increased reproductive effort caused by strong intraspecific competition. They found that the life-history traits of crucian carp were clearly related to the presence of piscivores: with increasing predation risk, fish grew in body length and depth and attained larger asymptotic length and size at maturity. This growth was evident at young age, especially in productive lakes with pike, and it suggests that fish quickly outgrew the predation window by reaching a size refuge. Contrary to the authors' predictions, populations had similar age at maturity. High-predation lakes also presented low density of crucian carp. This suggests that fish from predator lakes may experience high levels of resource availability due to reduced intraspecific competition. Predation regulated life-history traits in crucian carp populations, where larger size, higher longevity and size at maturity were observed in lakes with large gaped predators., (© 2023 The Authors. Journal of Fish Biology published by John Wiley & Sons Ltd on behalf of Fisheries Society of the British Isles.)

Mercury is a highly toxic element for consumers, but its relation to amino acids and physiology of wild fish is not well known. The main aim of this study was to evaluate how total mercury content (THg) of northern pike (Esox lucius) is related to amino acids and potentially important environmental and biological factors along a climate-productivity gradient of ten subarctic lakes. Linear regression between THg and sixteen amino acids content [nmol mg -1 dry weight] from white dorsal muscle of pike from these lakes were tested. Lastly, a general linear model (GLM) for age-corrected THg was used to test which factors are significantly related to mercury content of pike. There was a positive relationship between THg and proline. Seven out of sixteen analysed amino acids (histidine, threonine, arginine, serine, glutamic acid, glycine, and aspartic acid) were significantly negatively related to warmer and more productive lakes, while THg showed a positive relationship. GLM model indicated higher THg was found in higher trophic level pike with lower cysteine content and inhabiting warmer and more productive lakes with larger catchment containing substantial proportion of peatland area. In general, THg was not only related to the biological and environmental variables but also to amino acid content., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Authors. Published by Elsevier Inc. All rights reserved.)

Boreal lakes demonstrate pronounced seasonality, where the warm open-water season and subsequent cold and ice-covered season dominate natural cycles. While fish muscle total mercury concentration (mg/kg) [THg] is well documented in open-water summer months, there is limited knowledge on the ice-covered winter and spring mercury dynamics in fish from various foraging and thermal guilds. This year-round study tested how seasonality influences [THg] and its bioaccumulation in three percids, perch (Perca fluviatilis), pikeperch (Sander lucioperca), ruffe (Gymnocephalus cernua), and three cyprinids, roach (Rutilus rutilus), bleak (Alburnus alburnus), and bream (Abramis brama) in deep boreal mesotrophic Lake Pääjärvi, southern Finland. Fish were sampled and [THg] was quantified in the dorsal muscle during four seasons in this humic lake. Bioaccumulation regression slopes (mean ± STD, 0.039 ± 0.030, range 0.013-0.114) between [THg] and fish length were steepest during and after spawning and shallowest during autumn and winter for all species. Fish [THg] was significantly higher in the winter-spring than summer-autumn in all percids, however, not in cyprinids. The lowest [THg] was observed in summer and autumn, likely due to recovery from spring spawning, somatic growth and lipid accumulation. Fish [THg] was best described by multiple regression models (R 2 adj : 52-76%) which included total length and varying combinations of seasonally changing environmental (water temperature, total carbon, total nitrogen, and oxygen saturation) and biotic factors (gonadosomatic index, and sex) in all species. The seasonal variation in [THg] and bioaccumulation slopes across multiple species suggests a need for standardized sampling seasons in long-term monitoring to avoid any seasonality bias. From the fisheries and fish consumption perspective in seasonally ice-covered lakes, monitoring of both winter-spring and summer-autumn would improve knowledge of [THg] variation in fish muscle., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Authors. Published by Elsevier Inc. All rights reserved.)

High latitude ecosystems are experiencing the most rapid warming on earth, expected to trigger a diverse array of ecological responses. Climate warming affects the ecophysiology of fish, and fish close to the cold end of their thermal distribution are expected to increase somatic growth from increased temperatures and a prolonged growth season, which in turn affects maturation schedules, reproduction, and survival, boosting population growth. Accordingly, fish species living in ecosystems close to their northern range edge should increase in relative abundance and importance, and possibly displace cold-water adapted species. We aim to document whether and how population-level effects of warming are mediated by individual-level responses to increased temperatures, shift community structure, and composition in high latitude ecosystems. We studied 11 cool-water adapted perch populations in communities dominated by cold-water adapted species (whitefish, burbot, and charr) to investigate changes in the relative importance of the cool-water perch during the last 30 years of rapid warming in high latitude lakes. In addition, we studied the individual-level responses to warming to clarify the potential mechanisms underlying the population effects. Our long-term series (1991-2020) reveal a marked increase in numerical importance of the cool-water fish species, perch, in ten out of eleven populations, and in most fish communities perch is now dominant. Moreover, we show that climate warming affects population-level processes via direct and indirect temperature effects on individuals. Specifically, the increase in abundance arises from increased recruitment, faster juvenile growth, and ensuing earlier maturation, all boosted by climate warming. The speed and magnitude of the response to warming in these high latitude fish communities strongly suggest that cold-water fish will be displaced by fish adapted to warmer water. Consequently, management should focus on climate adaptation limiting future introductions and invasions of cool-water fish and mitigating harvesting pressure on cold-water fish., (© 2023 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Environmental change, including joint effects of increasing dissolved organic carbon (DOC) and total phosphorus (TP) in boreal northern lakes may affect food web energy sources and the biochemical composition of organisms. These environmental stressors are enhanced by anthropogenic land-use and can decrease the quality of polyunsaturated fatty acids (PUFAs) in seston and zooplankton, and therefore, possibly cascading up to fish. In contrast, the content of mercury in fish increases with lake browning potentially amplified by intensive forestry practises. However, there is little evidence on how these environmental stressors simultaneously impact beneficial omega-3 fatty acid (n3-FA) and total mercury (THg) content of fish muscle for human consumption. A space-for-time substitution study was conducted to assess whether environmental stressors affect Eurasian perch (Perca fluviatilis) allochthony and muscle nutritional quality [PUFA, THg, and their derivative, the hazard quotient (HQ)]. Perch samples were collected from 31 Finnish lakes along pronounced lake size (0.03-107.5 km 2 ), DOC (5.0-24.3 mg L -1 ), TP (5-118 μg L -1 ) and land-use gradients (forest: 50.7-96.4%, agriculture: 0-32.6%). These environmental gradients were combined using principal component analysis (PCA). Allochthony for individual perch was modelled using source and consumer δ 2 H values. Perch allochthony increased with decreasing lake pH and increasing forest coverage (PC1), but no correlation between lake DOC and perch allochthony was found. Perch muscle THg and omega-6 fatty acid (n6-FA) content increased with PC1 parallel with allochthony. Perch muscle DHA (22:6n3) content decreased, and ALA (18:3n3) increased towards shallower murkier lakes (PC2). Perch allochthony was positively correlated with muscle THg and n6-FA content, but did not correlate with n3-FA content. Hence, the quality of perch muscle for human consumption decreases (increase in HQ) with increasing forest coverage and decreasing pH, potentially mediated by increasing fish allochthony., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 The Authors. Published by Elsevier B.V. All rights reserved.)

Global climate change has led to profound alterations of the Arctic environment and ecosystems, with potential secondary effects on mercury (Hg) within Arctic biota. This review presents the current scientific evidence for impacts of direct physical climate change and indirect ecosystem change on Hg exposure and accumulation in Arctic terrestrial, freshwater, and marine organisms. As the marine environment is elevated in Hg compared to the terrestrial environment, terrestrial herbivores that now exploit coastal/marine foods when terrestrial plants are iced over may be exposed to higher Hg concentrations. Conversely, certain populations of predators, including Arctic foxes and polar bears, have shown lower Hg concentrations related to reduced sea ice-based foraging and increased land-based foraging. How climate change influences Hg in Arctic freshwater fishes is not clear, but for lacustrine populations it may depend on lake-specific conditions, including interrelated alterations in lake ice duration, turbidity, food web length and energy sources (benthic to pelagic), and growth dilution. In several marine mammal and seabird species, tissue Hg concentrations have shown correlations with climate and weather variables, including climate oscillation indices and sea ice trends; these findings suggest that wind, precipitation, and cryosphere changes that alter Hg transport and deposition are impacting Hg concentrations in Arctic marine organisms. Ecological changes, including northward range shifts of sub-Arctic species and altered body condition, have also been shown to affect Hg levels in some populations of Arctic marine species. Given the limited number of populations and species studied to date, especially within Arctic terrestrial and freshwater systems, further research is needed on climate-driven processes influencing Hg concentrations in Arctic ecosystems and their net effects. Long-term pan-Arctic monitoring programs should consider ancillary datasets on climate, weather, organism ecology and physiology to improve interpretation of spatial variation and time trends of Hg in Arctic biota., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier B.V. All rights reserved.)

The European whitefish (Coregonus lavaretus) species complex is a classic example of recent adaptive radiation. Here, we examine a whitefish population introduced to northern Finnish Lake Tsahkal in the late 1960s, where three divergent morphs (viz. littoral, pelagic, and profundal feeders) were found 10 generations after. Using demographic modeling based on genomic data, we show that whitefish morphs evolved during a phase of strict isolation, refuting a rapid sympatric divergence scenario. The lake is now an artificial hybrid zone between morphs originated in allopatry. Despite their current syntopy, clear genetic differentiation remains between two of the three morphs. Using admixture mapping, we identify five SNPs associated with gonad weight variation, a proxy for sexual maturity and spawning time. We suggest that ecological adaptations in spawning time evolved in allopatry are currently maintaining partial reproductive isolation in the absence of other barriers to gene flow., (© 2022 The Authors. Evolution published by Wiley Periodicals LLC on behalf of The Society for the Study of Evolution.)

European whitefish is a model species for adaptive radiation of fishes in temperate and subarctic lakes. In northern Europe the most commonly observed morphotypes are a generalist (LSR) morph and a pelagic specialist (DR) morph. The evolution of a pelagic specialist morph is something of an enigma, however, as this region is characterized by long, dark winters with pelagic primary production limited to a brief window in late summer. We conducted the first winter-based study of polymorphic whitefish populations to determine the winter ecology of both morphs, and we combined seasonal diet and stable isotope analysis with several proxies of condition in three polymorphic whitefish populations. The generalist LSR morph fed on benthic and pelagic prey in summer but was solely reliant on benthic prey in winter. This was associated with a noticeable but moderate reduction in condition, lipid content and stomach fullness in winter relative to summer. In contrast, the DR whitefish occupied a strict pelagic niche in both seasons. A significant reduction in pelagic prey during winter resulted in severe decrease in condition, lipid content and stomach fullness in DR whitefish in winter relative to summer, with the pelagic morph apparently approaching starvation in winter. We suggest that this divergent approach to seasonal foraging is associated with the divergent life-history traits of both morphs., (© 2022 Fisheries Society of the British Isles.)

Aim: Understanding the variation in community composition and species abundances (i.e., β-diversity) is at the heart of community ecology. A common approach to examine β-diversity is to evaluate directional variation in community composition by measuring the decay in the similarity among pairs of communities along spatial or environmental distance. We provide the first global synthesis of taxonomic and functional distance decay along spatial and environmental distance by analysing 148 datasets comprising different types of organisms and environments., Location: Global., Time Period: 1990 to present., Major Taxa Studied: From diatoms to mammals., Method: We measured the strength of the decay using ranked Mantel tests (Mantel r ) and the rate of distance decay as the slope of an exponential fit using generalized linear models. We used null models to test whether functional similarity decays faster or slower than expected given the taxonomic decay along the spatial and environmental distance. We also unveiled the factors driving the rate of decay across the datasets, including latitude, spatial extent, realm and organismal features., Results: Taxonomic distance decay was stronger than functional distance decay along both spatial and environmental distance. Functional distance decay was random given the taxonomic distance decay. The rate of taxonomic and functional spatial distance decay was fastest in the datasets from mid-latitudes. Overall, datasets covering larger spatial extents showed a lower rate of decay along spatial distance but a higher rate of decay along environmental distance. Marine ecosystems had the slowest rate of decay along environmental distances., Main Conclusions: In general, taxonomic distance decay is a useful tool for biogeographical research because it reflects dispersal-related factors in addition to species responses to climatic and environmental variables. Moreover, functional distance decay might be a cost-effective option for investigating community changes in heterogeneous environments., Competing Interests: The authors declare no conflicts of interest., (© 2022 The Authors. Global Ecology and Biogeography published by John Wiley & Sons Ltd.)

Dramatic environmental shifts are occuring throughout the Arctic from climate change, with consequences for the cycling of mercury (Hg). This review summarizes the latest science on how climate change is influencing Hg transport and biogeochemical cycling in Arctic terrestrial, freshwater and marine ecosystems. As environmental changes in the Arctic continue to accelerate, a clearer picture is emerging of the profound shifts in the climate and cryosphere, and their connections to Hg cycling. Modeling results suggest climate influences seasonal and interannual variability of atmospheric Hg deposition. The clearest evidence of current climate change effects is for Hg transport from terrestrial catchments, where widespread permafrost thaw, glacier melt and coastal erosion are increasing the export of Hg to downstream environments. Recent estimates suggest Arctic permafrost is a large global reservoir of Hg, which is vulnerable to degradation with climate warming, although the fate of permafrost soil Hg is unclear. The increasing development of thermokarst features, the formation and expansion of thaw lakes, and increased soil erosion in terrestrial landscapes are increasing river transport of particulate-bound Hg and altering conditions for aquatic Hg transformations. Greater organic matter transport may also be influencing the downstream transport and fate of Hg. More severe and frequent wildfires within the Arctic and across boreal regions may be contributing to the atmospheric pool of Hg. Climate change influences on Hg biogeochemical cycling remain poorly understood. Seasonal evasion and retention of inorganic Hg may be altered by reduced sea-ice cover and higher chloride content in snow. Experimental evidence indicates warmer temperatures enhance methylmercury production in ocean and lake sediments as well as in tundra soils. Improved geographic coverage of measurements and modeling approaches are needed to better evaluate net effects of climate change and long-term implications for Hg contamination in the Arctic., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 The Authors. Published by Elsevier B.V. All rights reserved.)

Water browning or brownification refers to increasing water color, often related to increasing dissolved organic matter (DOM) and carbon (DOC) content in freshwaters. Browning has been recognized as a significant physicochemical phenomenon altering boreal lakes, but our understanding of its ecological consequences in different freshwater habitats and regions is limited. Here, we review the consequences of browning on different freshwater habitats, food webs and aquatic-terrestrial habitat coupling. We examine global trends of browning and DOM/DOC, and the use of remote sensing as a tool to investigate browning from local to global scales. Studies have focused on lakes and rivers while seldom addressing effects at the catchment scale. Other freshwater habitats such as small and temporary waterbodies have been overlooked, making the study of the entire network of the catchment incomplete. While past research investigated the response of primary producers, aquatic invertebrates and fishes, the effects of browning on macrophytes, invasive species, and food webs have been understudied. Research has focused on freshwater habitats without considering the fluxes between aquatic and terrestrial habitats. We highlight the importance of understanding how the changes in one habitat may cascade to another. Browning is a broader phenomenon than the heretofore concentration on the boreal region. Overall, we propose that future studies improve the ecological understanding of browning through the following research actions: 1) increasing our knowledge of ecological processes of browning in other wetland types than lakes and rivers, 2) assessing the impact of browning on aquatic food webs at multiple scales, 3) examining the effects of browning on aquatic-terrestrial habitat coupling, 4) expanding our knowledge of browning from the local to global scale, and 5) using remote sensing to examine browning and its ecological consequences., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2021 The Authors. Published by Elsevier B.V. All rights reserved.)

Subarctic lakes are getting warmer and more productive due to the joint effects of climate change and intensive land-use practices (e.g. forest clear-cutting and peatland ditching), processes that potentially increase leaching of peat- and soil-stored mercury into lake ecosystems. We sampled biotic communities from primary producers (algae) to top consumers (piscivorous fish), in 19 subarctic lakes situated on a latitudinal (69.0-66.5° N), climatic (+3.2 °C temperature and +30% precipitation from north to south) and catchment land-use (pristine to intensive forestry areas) gradient. We first tested how the joint effects of climate and productivity influence mercury biomagnification in food webs focusing on the trophic magnification slope (TMS) and mercury baseline (THg baseline) level, both derived from linear regression between total mercury (log 10 THg) and organism trophic level (TL). We examined a suite of environmental and biotic variables thought to explain THg baseline and TMS with stepwise generalized multiple regression models. Finally, we assessed how climate and lake productivity affect the THg content of top predators in subarctic lakes. We found biomagnification of mercury in all studied lakes, but with variable TMS and THg baseline values. In stepwise multiple regression models, TMS was best explained by negative relationships with food chain length, climate-productivity gradient, catchment properties, and elemental C:N ratio of the top predator (full model R 2 = 0.90, p < 0.001). The model examining variation in THg baseline values included the same variables with positive relationships (R 2 = 0.69, p = 0.014). Mass-standardized THg content of a common top predator (1 kg northern pike, Esox lucius) increased towards warmer and more productive lakes. Results indicate that increasing eutrophication via forestry-related land-use activities increase the THg levels at the base of the food web and in top predators, suggesting that the sources of nutrients and mercury should be considered in future bioaccumulation and biomagnification studies., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2021 The Authors. Published by Elsevier B.V. All rights reserved.)

Phenotypic plasticity can be expressed as changes in body shape in response to environmental variability. Crucian carp ( Carassius carassius ), a widespread cyprinid, displays remarkable plasticity in body morphology and increases body depth when exposed to cues from predators, suggesting the triggering of an antipredator defense mechanism. However, these morphological changes could also be related to resource use and foraging behavior, as an indirect effect of predator presence. In order to determine whether phenotypic plasticity in crucian carp is driven by a direct or indirect response to predation threat, we compared twelve fish communities inhabiting small lakes in southeast Norway grouped by four categories of predation regimes: no predator fish, or brown trout ( Salmo trutta ), perch ( Perca fluviatilis ), or pike ( Esox lucius ) as main piscivores. We predicted the body shape of crucian carp to be associated with the species composition of predator communities and that the presence of efficient piscivores would result in a deeper body shape. We use stable isotope analyses to test whether this variation in body shape was related to a shift in individual resource use-that is, littoral rather than pelagic resource use would favor the development of a specific body shape-or other environmental characteristics. The results showed that increasingly efficient predator communities induced progressively deeper body shape, larger body size, and lower population densities. Predator maximum gape size and individual trophic position were the best variables explaining crucian carp variation in body depth among predation categories, while littoral resource use did not have a clear effect. The gradient in predation pressure also corresponded to a shift in lake productivity. These results indicate that crucian carp have a fine-tuned morphological defense mechanism against predation risk, triggered by the combined effect of predator presence and resource availability., Competing Interests: Authors have no conflict of interest to declare., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Climate change in the Arctic is outpacing the global average and land-use is intensifying due to exploitation of previously inaccessible or unprofitable natural resources. A comprehensive understanding of how the joint effects of changing climate and productivity modify lake food web structure, biomass, trophic pyramid shape and abundance of physiologically essential biomolecules (omega-3 fatty acids) in the biotic community is lacking. We conducted a space-for-time study in 20 subarctic lakes spanning a climatic (+3.2°C and precipitation: +30%) and chemical (dissolved organic carbon: +10 mg/L, total phosphorus: +45 µg/L and total nitrogen: +1,000 µg/L) gradient to test how temperature and productivity jointly affect the structure, biomass and community fatty acid content (eicosapentaenoic acid [EPA] and docosahexaenoic acid [DHA]) of whole food webs. Increasing temperature and productivity shifted lake communities towards dominance of warmer, murky-water-adapted taxa, with a general increase in the biomass of primary producers, and secondary and tertiary consumers, while primary invertebrate consumers did not show equally clear trends. This process altered various trophic pyramid structures towards an hour glass shape in the warmest and most productive lakes. Increasing temperature and productivity had negative fatty acid content trends (mg EPA + DHA/g dry weight) in primary producers and primary consumers, but not in secondary nor tertiary fish consumers. The massive biomass increment of fish led to increasing areal fatty acid content (kg EPA + DHA/ha) towards increasingly warmer, more productive lakes, but there were no significant trends in other trophic levels. Increasing temperature and productivity are shifting subarctic lake communities towards systems characterized by increasing dominance of cyanobacteria and cyprinid fish, although decreasing quality in terms of EPA + DHA content was observed only in phytoplankton, zooplankton and profundal benthos., (© 2020 John Wiley & Sons Ltd.)

Adaptive radiation is the diversification of species to different ecological niches and has repeatedly occurred in different salmonid fish of postglacial lakes. In Lake Tinnsjøen, one of the largest and deepest lakes in Norway, the salmonid fish, Arctic charr ( Salvelinus alpinus (L.)), has likely radiated within 9,700 years after deglaciation into ecologically and genetically segregated Piscivore, Planktivore, Dwarf, and Abyssal morphs in the pelagial, littoral, shallow-moderate profundal, and deep-profundal habitats. We compared trait variation in the size of the head, the eye and olfactory organs, as well as the volumes of five brain regions of these four Arctic charr morphs. We hypothesised that specific habitat characteristics have promoted divergent body, head, and brain sizes related to utilized depth differing in environmental constraints (e.g., light, oxygen, pressure, temperature, and food quality). The most important ecomorphological variables differentiating morphs were eye area, habitat, and number of lamellae. The Abyssal morph living in the deepest areas of the lake had the smallest brain region volumes, head, and eye size. Comparing the olfactory bulb with the optic tectum in size, it was larger in the Abyssal morph than in the Piscivore morph. The Piscivore and Planktivore morphs that use more illuminated habitats have the largest optic tectum volume, followed by the Dwarf. The observed differences in body size and sensory capacities in terms of vision and olfaction in shallow and deepwater morphs likely relates to foraging and mating habitats in Lake Tinnsjøen. Further seasonal and experimental studies of brain volume in polymorphic species are needed to test the role of plasticity and adaptive evolution behind the observed differences., Competing Interests: The authors declare no conflicts of interest., (© 2020 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Modern speciation theory has greatly benefited from a variety of simple mathematical models focusing on the conditions and patterns of speciation and diversification in the presence of gene flow. Unfortunately the application of general theoretical concepts and tools to specific ecological systems remains a challenge. Here we apply modeling tools to better understand adaptive divergence of whitefish during the postglacial period in lakes of northern Fennoscandia. These lakes harbor up to three different morphs associated with the three major lake habitats: littoral, pelagic, and profundal. Using large-scale individual-based simulations, we aim to identify factors required for in situ emergence of the pelagic and profundal morphs in lakes initially colonized by the littoral morph. The importance of some of the factors we identify and study - sufficiently large levels of initial genetic variation, size- and habitat-specific mating, sufficiently large carrying capacity of the new niche - is already well recognized. In addition, our model also points to two other factors that have been largely disregarded in theoretical studies: fitness-dependent dispersal and strong predation in the ancestral niche coupled with the lack of it in the new niche(s). We use our theoretical results to speculate about the process of diversification of whitefish in Fennoscandia and to identify potentially profitable directions for future empirical research.

A major goal of evolutionary science is to understand how biological diversity is generated and altered. Despite considerable advances, we still have limited insight into how phenotypic variation arises and is sorted by natural selection. Here we argue that an integrated view, which merges ecology, evolution and developmental biology (eco evo devo) on an equal footing, is needed to understand the multifaceted role of the environment in simultaneously determining the development of the phenotype and the nature of the selective environment, and how organisms in turn affect the environment through eco evo and eco devo feedbacks. To illustrate the usefulness of an integrated eco evo devo perspective, we connect it with the theory of resource polymorphism (i.e. the phenotypic and genetic diversification that occurs in response to variation in available resources). In so doing, we highlight fishes from recently glaciated freshwater systems as exceptionally well-suited model systems for testing predictions of an eco evo devo framework in studies of diversification. Studies on these fishes show that intraspecific diversity can evolve rapidly, and that this process is jointly facilitated by (i) the availability of diverse environments promoting divergent natural selection; (ii) dynamic developmental processes sensitive to environmental and genetic signals; and (iii) eco evo and eco devo feedbacks influencing the selective and developmental environments of the phenotype. We highlight empirical examples and present a conceptual model for the generation of resource polymorphism - emphasizing eco evo devo, and identify current gaps in knowledge., (© 2019 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

Resource polymorphism-whereby ancestral generalist populations give rise to several specialised morphs along a resource gradient-is common where species colonise newly formed ecosystems. This phenomenon is particularly well documented in freshwater fish populations inhabiting postglacial lakes formed at the end of the last ice age. However, knowledge on how such differential exploitation of resources across contrasting habitats might be reflected in the biochemical compositions of diverging populations is still limited, though such patterns might be expected. Here, we aimed to assess how fatty acids (FA)-an important biochemical component of animal tissues-diverged across a polymorphic complex of European whitefish (Coregonus lavaretus) and their closely related monomorphic specialist congener vendace (Coregonus albula) inhabiting a series of six subarctic lakes in northern Fennoscandia. We also explored patterns of FA composition in whitefish's predators and invertebrate prey to assess how divergence in trophic ecology between whitefish morphs would relate to biochemical profiles of their key food web associates. Lastly, we assessed how information on trophic divergence provided by differential FA composition compared to evidence of resource polymorphism retrieved from more classical stomach content and stable isotopic (δ13C, δ15N) information. Examination of stomach contents provided high-resolution information on recently consumed prey, whereas stable isotopes indicated broad-scale patterns of benthic-pelagic resource use differentiation at different trophic levels. Linear discriminant analysis based on FA composition was substantially more successful in identifying whitefish morphs and their congener vendace as distinct groupings when compared to the other two methods. Three major FA (myristic acid, stearic acid, and eicosadienoic acid) proved particularly informative, both in delineating coregonid groups, and identifying patterns of pelagic-benthic feeding throughout the wider food web. Myristic acid (14:0) content and δ13C ratios in muscle tissue were positively correlated across fish taxa, and together provided the clearest segregation of fishes exploiting contrasting pelagic and benthic niches. In general, our findings highlight the potential of FA analysis for identifying resource polymorphism in animal populations where this phenomenon occurs, and suggest that this technique may provide greater resolution than more traditional methods typically used for this purpose., Competing Interests: The authors have declared that no competing interests exist.

Temporally (1965-2015) and spatially (55°-70°N) extensive records of total mercury (Hg) in freshwater fish showed consistent declines in boreal and subarctic Fennoscandia. The database contains 54 560 fish entries ( n: pike > perch ≫ brown trout > roach ≈ Arctic charr) from 3132 lakes across Sweden, Finland, Norway, and Russian Murmansk area. 74% of the lakes did not meet the 0.5 ppm limit to protect human health. However, after 2000 only 25% of the lakes exceeded this level, indicating improved environmental status. In lakes where local pollution sources were identified, pike and perch Hg concentrations were significantly higher between 1965 and 1990 compared to values after 1995, likely an effect of implemented reduction measures. In lakes where Hg originated from long-range transboundary air pollution (LRTAP), consistent Hg declines (3-7‰ per year) were found for perch and pike in both boreal and subarctic Fennoscandia, suggesting common environmental controls. Hg in perch and pike in LRTAP lakes showed minimal declines with latitude, suggesting that drivers affected by temperature, such as growth dilution, counteracted Hg loading and food web exposure. We recommend that future fish Hg monitoring sampling design should include repeated sampling and collection of pollution history, water chemistry, fish age, and stable isotopes to enable evaluation of emission reduction policies.

Climate change is resulting in increased temperatures and precipitation in subarctic regions of Europe. These changes are extending tree lines to higher altitudes and latitudes, and enhancing tree growth enabling intensification of forestry into previously inhospitable subarctic regions. The combined effects of climate change and land-use intensification extend the warm, open-water season in subarctic lakes and increase lake productivity and may also increase leaching and methylation activity of mercury within the lakes. To assess the joint effects of climate and productivity on total mercury (THg) bioaccumulation in fish, we conducted a space-for-time substitution study in 18 tributary lakes of a subarctic watercourse forming a gradient from cold pristine oligotrophic lakes in the northern headwaters to warmer and increasingly human-altered mesotrophic and eutrophic systems in the southern lower reaches. Increasing temperature, precipitation, and lake productivity were predicted to elevate length- and age-adjusted THg concentrations, as well as THg bioaccumulation rate (the rate of THg bioaccumulation relative to length or age) in muscle tissue of European whitefish (Coregonus lavaretus), vendace (Coregonus albula), perch (Perca fluviatilis), pike (Esox lucius), roach (Rutilus rutilus) and ruffe (Gymnocephalus cernua). A significant positive relationship was observed between age-adjusted THg concentration and lake climate-productivity in vendace (r 2  = 0.50), perch (r 2  = 0.51), pike (r 2  = 0.55) and roach (r 2  = 0.61). Higher climate-productivity values of the lakes also had a positive linear (pike; r 2  = 0.40 and whitefish; r 2  = 0.72) or u-shaped (perch; r 2  = 0.64 and ruffe; r 2  = 0.50) relationship with THg bioaccumulation rate. Our findings of increased adjusted THg concentrations in planktivores and piscivores reveal adverse effects of warming climate and increasing productivity on these subarctic fishes, whereas less distinct trends in THg bioaccumulation rate suggest more complex underlying processes. Joint environmental stressors such as climate and productivity should be considered in ongoing and future monitoring of mercury concentrations., (Copyright © 2018 Elsevier B.V. All rights reserved.)

Populations of fishes provide valuable services for billions of people, but face diverse and interacting threats that jeopardize their sustainability. Human population growth and intensifying resource use for food, water, energy and goods are compromising fish populations through a variety of mechanisms, including overfishing, habitat degradation and declines in water quality. The important challenges raised by these issues have been recognized and have led to considerable advances over past decades in managing and mitigating threats to fishes worldwide. In this review, we identify the major threats faced by fish populations alongside recent advances that are helping to address these issues. There are very significant efforts worldwide directed towards ensuring a sustainable future for the world's fishes and fisheries and those who rely on them. Although considerable challenges remain, by drawing attention to successful mitigation of threats to fish and fisheries we hope to provide the encouragement and direction that will allow these challenges to be overcome in the future., (© 2018 The Authors. Journal of Fish Biology published by John Wiley & Sons Ltd on behalf of The Fisheries Society of the British Isles.)

The first few months of life is the most vulnerable period for fish and their optimal hatching time with zooplankton prey is favored by natural selection. Traditionally, however, prey abundance (i.e., zooplankton density) has been considered important, whereas prey nutritional composition has been largely neglected in natural settings. High-quality zooplankton, rich in both essential amino acids (EAAs) and fatty acids (FAs), are required as starting prey to initiate development and fast juvenile growth. Prey quality is dependent on environmental conditions, and, for example, eutrophication and browning are two major factors defining primary producer community structures that will directly determine the nutritional quality of the basal food sources (algae, bacteria, terrestrial matter) for zooplankton. We experimentally tested how eutrophication and browning affect the growth and survival of juvenile rainbow trout ( Oncorhynchus mykiss ) by changing the quality of basal resources. We fed the fish on herbivorous zooplankton ( Daphnia ) grown with foods of different nutritional quality (algae, bacteria, terrestrial matter), and used GC-MS, stable isotope labeling as well as bulk and compound-specific stable isotope analyses for detecting the effects of different diets on the nutritional status of fish. The content of EAAs and omega-3 (ω-3) polyunsaturated FAs (PUFAs) in basal foods and zooplankton decreased in both eutrophication and browning treatments. The decrease in ω-3 PUFA and especially docosahexaenoic acid (DHA) was reflected to fish juveniles, but they were able to compensate for low availability of EAAs in their food. Therefore, the reduced growth and survival of the juvenile fish was linked to the low availability of DHA. Fish showed very low ability to convert alpha-linolenic acid (ALA) to DHA. We conclude that eutrophication and browning decrease the availability of the originally phytoplankton-derived DHA for zooplankton and juvenile fish, suggesting bottom-up regulation of food web quality.

Adaptive radiation is the evolution of ecological and phenotypical diversity. It arises via ecological opportunity that promotes the exploration of underutilized or novel niches mediating specialization and reproductive isolation. The assumed precondition for rapid local adaptation is diversifying natural selection, but random genetic drift could also be a major driver of this process. We used 27 populations of European whitefish ( Coregonus lavaretus ) from nine lakes distributed in three neighboring subarctic watercourses in northern Fennoscandia as a model to test the importance of random drift versus diversifying natural selection for parallel evolution of adaptive phenotypic traits. We contrasted variation for two key adaptive phenotypic traits correlated with resource utilization of polymorphic fish; the number of gill rakers and the total length of fish, with the posterior distribution of neutral genetic differentiation from 13 microsatellite loci, to test whether the observed phenotypic divergence could be achieved by random genetic drift alone. Our results show that both traits have been under diversifying selection and that the evolution of these morphs has been driven by isolation through habitat adaptations. We conclude that diversifying selection acting on gill raker number and body size has played a significant role in the ongoing adaptive radiation of European whitefish morphs in this region.

Resource polymorphism, whereby ancestral trophic generalists undergo divergence into multiple specialist morphs, is common in salmonid fish populations inhabiting subarctic lakes. However, the extent to which such resource specialization into the three principal lake habitats (littoral, profundal, and pelagic) affects patterns of contaminant bioaccumulation remains largely unexplored. We assessed total mercury concentrations (THg) of European whitefish (Coregonus lavaretus (L.)) and their invertebrate prey in relation to potential explanatory variables across 6 subarctic lakes, of which three are inhabited by polymorphic (comprised of four morphs) and three by monomorphic populations. Among invertebrate prey, the highest THg concentrations were observed in profundal benthic macroinvertebrates, followed by pelagic zooplankton, with concentrations lowest in littoral benthic macroinvertebrates in both lake types. Broadly similar patterns were apparent in whitefish in polymorphic systems, where average age-corrected THg concentrations and bioaccumulation rates were the highest in pelagic morphs, intermediate in the profundal morph, and the lowest in the littoral morph. In monomorphic systems, age-corrected THg concentrations were generally lower, and showed pronounced lake-specific variation. In the polymorphic systems, we found significant relationships between whitefish muscle tissue THg concentration and gill raker count, resource use, lipid content and maximum length, whilst no such relationships were apparent in the monomorphic systems. Across all polymorphic lakes, the major variables explaining THg in whitefish were gill raker count and age, whereas in monomorphic systems, the factors were lake-specific. Whitefish resource polymorphism across the three main lake habitats therefore appears to have profound impacts on THg concentration and bioaccumulation rate. This highlights the importance of recognizing such intraspecific diversity in both future scientific studies and mercury monitoring programs., (Copyright © 2017 Elsevier B.V. All rights reserved.)

Subarctic lakes are characterised by extreme seasonal variation in light and temperature which influences growth, maturation, condition and resource use of fishes. However, our understanding of how seasonal changes affect mercury concentrations of fishes is limited. We conducted a year-round study (3 ice-covered months, 3 open-water months) with open-water inter-annual aspect (3 years: samples from August/September), focusing on total mercury (THg) concentrations and ecological characteristics of a common freshwater fish, European whitefish (Coregonus lavaretus (L.)) from a subarctic lake. We measured THg concentrations from tissues with fast (liver, n = 164) and moderate (muscle, n = 225) turnover rates, providing information on THg dynamics over different temporal scales. In both tissues, lipid-corrected THg concentrations were highest in winter (liver: 1.70 ± 0.88 μg/g, muscle: 0.24 ± 0.05 μg/g) and lowest in summer (liver: 0.87 ± 0.72 μg/g, muscle: 0.19 ± 0.04 μg/g). THg concentrations increased in winter following the summer-autumn dietary shift to pelagic zooplankton and starvation after spawning. Whitefish THg concentrations decreased towards summer, and were associated with consumption of benthic macroinvertebrates and subsequent growth dilution. Mercury bioaccumulated in both tissues with age, both showing the strongest regression slopes in winter and lowest in summer. THg concentrations in liver and muscle tissue were correlated throughout the year, however the correlation was lowest in summer, indicating high metabolism during somatic growing season in summer and growth dilution. Multiple linear regression models explained 50% and 55% of the THg variation in liver and muscle both models dominated by seasonally-variable factors i.e. sexual maturity, δ 13 C, and condition factor. Seasonally varying bioaccumulation slopes and the higher level of intra-annual variation (21%) in whitefish THg concentration in muscle than the inter-annual accumulation (8%) highlight the importance of including seasonal factors in future THg studies., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Ecological speciation - whereby an ancestral founder species diversifies to fill vacant niches - is a phenomenon characteristic of newly formed ecosystems. Despite such ubiquity, ecosystem-level effects of such divergence remain poorly understood. Here, we compared the trophic niche of European whitefish (Coregonus lavaretus) and their predators in a series of contrasting subarctic lakes where this species had either diversified into four ecomorphologically distinct morphs or instead formed monomorphic populations. We found that the trophic niche of whitefish was almost three times larger in the polymorphic than in the monomorphic lakes, due to an increase in intraspecific specialisation. This trophic niche expansion was mirrored in brown trout (Salmo trutta), a major predator of whitefish. This represents amongst the first evidence for ecological speciation directly altering the trophic niche of a predator. We suggest such mechanisms may be a common and important - though presently overlooked - factor regulating trophic interactions in diverse ecosystems globally.

Climate change and species invasions represent key threats to global biodiversity. Subarctic freshwaters are sentinels for understanding both stressors because the effects of climate change are disproportionately strong at high latitudes and invasion of temperate species is prevalent. Here, we summarize the environmental effects of climate change and illustrate the ecological responses of freshwater fishes to these effects, spanning individual, population, community and ecosystem levels. Climate change is modifying hydrological cycles across atmospheric, terrestrial and aquatic components of subarctic ecosystems, causing increases in ambient water temperature and nutrient availability. These changes affect the individual behavior, habitat use, growth and metabolism, alter population spawning and recruitment dynamics, leading to changes in species abundance and distribution, modify food web structure, trophic interactions and energy flow within communities and change the sources, quantity and quality of energy and nutrients in ecosystems. Increases in temperature and its variability in aquatic environments underpin many ecological responses; however, altered hydrological regimes, increasing nutrient inputs and shortened ice cover are also important drivers of climate change effects and likely contribute to context-dependent responses. Species invasions are a complex aspect of the ecology of climate change because the phenomena of invasion are both an effect and a driver of the ecological consequences of climate change. Using subarctic freshwaters as an example, we illustrate how climate change can alter three distinct aspects of species invasions: (1) the vulnerability of ecosystems to be invaded, (2) the potential for species to spread and invade new habitats, and (3) the subsequent ecological effects of invaders. We identify three fundamental knowledge gaps focused on the need to determine (1) how environmental and landscape characteristics influence the ecological impact of climate change, (2) the separate and combined effects of climate and non-native invading species and (3) the underlying ecological processes or mechanisms responsible for changes in patterns of biodiversity.

Winter conditions are rapidly changing in temperate ecosystems, particularly for those that experience periods of snow and ice cover. Relatively little is known of winter ecology in these systems, due to a historical research focus on summer 'growing seasons'. We executed the first global quantitative synthesis on under-ice lake ecology, including 36 abiotic and biotic variables from 42 research groups and 101 lakes, examining seasonal differences and connections as well as how seasonal differences vary with geophysical factors. Plankton were more abundant under ice than expected; mean winter values were 43.2% of summer values for chlorophyll a, 15.8% of summer phytoplankton biovolume and 25.3% of summer zooplankton density. Dissolved nitrogen concentrations were typically higher during winter, and these differences were exaggerated in smaller lakes. Lake size also influenced winter-summer patterns for dissolved organic carbon (DOC), with higher winter DOC in smaller lakes. At coarse levels of taxonomic aggregation, phytoplankton and zooplankton community composition showed few systematic differences between seasons, although literature suggests that seasonal differences are frequently lake-specific, species-specific, or occur at the level of functional group. Within the subset of lakes that had longer time series, winter influenced the subsequent summer for some nutrient variables and zooplankton biomass., (© 2016 The Authors. Ecology Letters published by CNRS and John Wiley & Sons Ltd.)

Sexual dimorphism is common across the animal kingdom, but the contribution of environmental factors shaping differences between the sexes remains controversial. In ectotherms, life-history traits are known to correlate with latitude, but sex-specific responses are not well understood. We analyzed life-history trait variation between the sexes of European perch ( Perca fluviatilis L.), a common freshwater fish displaying larger female size, by employing a wide latitudinal gradient. We expected to find sex-dependent latitudinal variation in life-history variables: length at age, length increment, and size at maturity, with females showing consistently higher values than males at all latitudes. We further anticipated that this gender difference would progressively decrease with the increasingly harsh environmental conditions toward higher latitude. We hypothesized that growth and length increment would decrease and size/age at maturity would increase at higher latitudes. Our results confirmed female-biased sexual size dimorphism at all latitudes and the magnitude of sexual dimorphism diminished with increase in latitude. Growth of both sexes decreased with increase in latitude, and the female latitudinal clines were steeper than those of males. Hence, we challenge two predominant ecological rules (Rensch's and Bergmann's rules) that describe common large-scale patterns of body size variation. Our data demonstrate that these two rules are not universally applicable in ectotherms or female-biased species. Our study highlights the importance of sex-specific differences in life-history traits along a latitudinal gradient, with evident implications for a wide range of studies from individual to ecosystems level.

Fish are an important source of eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA) for birds, mammals and humans. In aquatic food webs, these highly unsaturated fatty acids (HUFA) are essential for many physiological processes and mainly synthetized by distinct phytoplankton taxa. Consumers at different trophic levels obtain essential fatty acids from their diet because they cannot produce these sufficiently de novo. Here, we evaluated how the increase in phosphorus concentration (eutrophication) or terrestrial organic matter inputs (brownification) change EPA and DHA content in the phytoplankton. Then, we evaluated whether these changes can be seen in the EPA and DHA content of piscivorous European perch (Perca fluviatilis), which is a widely distributed species and commonly consumed by humans. Data from 713 lakes showed statistically significant differences in the abundance of EPA- and DHA-synthesizing phytoplankton as well as in the concentrations and content of these essential fatty acids among oligo-mesotrophic, eutrophic and dystrophic lakes. The EPA and DHA content of phytoplankton biomass (mgHUFAg -1 ) was significantly lower in the eutrophic lakes than in the oligo-mesotrophic or dystrophic lakes. We found a strong significant correlation between the DHA content in the muscle of piscivorous perch and phytoplankton DHA content (r=0.85) as well with the contribution of DHA-synthesizing phytoplankton taxa (r=0.83). Among all DHA-synthesizing phytoplankton this correlation was the strongest with the dinoflagellates (r=0.74) and chrysophytes (r=0.70). Accordingly, the EPA+DHA content of perch muscle decreased with increasing total phosphorus (r 2 =0.80) and dissolved organic carbon concentration (r 2 =0.83) in the lakes. Our results suggest that although eutrophication generally increase biomass production across different trophic levels, the high proportion of low-quality primary producers reduce EPA and DHA content in the food web up to predatory fish. Ultimately, it seems that lake eutrophication and brownification decrease the nutritional quality of fish for human consumers., (Copyright © 2016 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Freshwater food webs can be partly supported by terrestrial primary production, often deriving from plant litter of surrounding catchment vegetation. Although consisting mainly of poorly bioavailable lignin, with low protein and lipid content, the carbohydrates from fallen tree leaves and shoreline vegetation may be utilized by aquatic consumers. Here we show that during phytoplankton deficiency, zooplankton (Daphnia magna) can benefit from terrestrial particulate organic matter by using terrestrial-origin carbohydrates for energy and sparing essential fatty acids and amino acids for somatic growth and reproduction. Assimilated terrestrial-origin fatty acids from shoreline reed particles exceeded available diet, indicating that Daphnia may convert a part of their dietary carbohydrates to saturated fatty acids. This conversion was not observed with birch leaf diets, which had lower carbohydrate content. Subsequent analysis of 21 boreal and subarctic lakes showed that diet of herbivorous zooplankton is mainly based on high-quality phytoplankton rich in essential polyunsaturated fatty acids. The proportion of low-quality diets (bacteria and terrestrial particulate organic matter) was <28% of the assimilated carbon. Taken collectively, the incorporation of terrestrial carbon into zooplankton was not directly related to the concentration of terrestrial organic matter in experiments or lakes, but rather to the low availability of phytoplankton.

Variation in presumably neutral genetic markers can inform us about evolvability, historical effective population sizes and phylogeographic history of contemporary populations. We studied genetic variability in 15 microsatellite loci in six native landlocked Arctic charr (Salvelinus alpinus) populations in northern Fennoscandia, where this species is considered near threatened. We discovered that all populations were genetically highly (mean FST ≈ 0.26) differentiated and isolated from each other. Evidence was found for historical, but not for recent population size bottlenecks. Estimates of contemporary effective population size (Ne) ranged from seven to 228 and were significantly correlated with those of historical Ne but not with lake size. A census size (NC) was estimated to be approximately 300 individuals in a pond (0.14 ha), which exhibited the smallest Ne (i.e. Ne/NC = 0.02). Genetic variability in this pond and a connected lake is severely reduced, and both genetic and empirical estimates of migration rates indicate a lack of gene flow between them. Hence, albeit currently thriving, some northern Fennoscandian populations appear to be vulnerable to further loss of genetic variability and are likely to have limited capacity to adapt if selection pressures change.

Prey preference of top predators and energy flow across habitat boundaries are of fundamental importance for structure and function of aquatic and terrestrial ecosystems, as they may have strong effects on production, species diversity, and food-web stability. In lakes, littoral and pelagic food-web compartments are typically coupled and controlled by generalist fish top predators. However, the extent and determinants of such coupling remains a topical area of ecological research and is largely unknown in oligotrophic high-latitude lakes. We analyzed food-web structure and resource use by a generalist top predator, the Arctic charr Salvelinus alpinus (L.), in 17 oligotrophic subarctic lakes covering a marked gradient in size (0.5-1084 km(2)) and fish species richness (2-13 species). We expected top predators to shift from littoral to pelagic energy sources with increasing lake size, as the availability of pelagic prey resources and the competition for littoral prey are both likely to be higher in large lakes with multispecies fish communities. We also expected top predators to occupy a higher trophic position in lakes with greater fish species richness due to potential substitution of intermediate consumers (prey fish) and increased piscivory by top predators. Based on stable carbon and nitrogen isotope analyses, the mean reliance of Arctic charr on littoral energy sources showed a significant negative relationship with lake surface area, whereas the mean trophic position of Arctic charr, reflecting the lake food-chain length, increased with fish species richness. These results were supported by stomach contents data demonstrating a shift of Arctic charr from an invertebrate-dominated diet to piscivory on pelagic fish. Our study highlights that, because they determine the main energy source (littoral vs. pelagic) and the trophic position of generalist top predators, ecosystem size and fish diversity are particularly important factors influencing function and structure of food webs in high-latitude lakes.

Adaptive radiation is considered an important mechanism for the development of new species, but very little is known about the role of thermal adaptation during this process. Such adaptation should be especially important in poikilothermic animals that are often subjected to pronounced seasonal temperature variation that directly affects metabolic function. We conducted a preliminary study of individual lifetime thermal habitat use and respiration rates of four whitefish (Coregonus lavaretus (L.)) morphs (two pelagic, one littoral and one profundal) using stable carbon and oxygen isotope values of otolith carbonate. These morphs, two of which utilized pelagic habitats, one littoral and one profundal recently diverged via adaptive radiation to exploit different major niches in a deep and thermally stratified subarctic lake. We found evidence that the morphs used different thermal niches. The profundal morph had the most distinct thermal niche and consistently occupied the coldest thermal habitat of the lake, whereas differences were less pronounced among the shallow water pelagic and littoral morphs. Our results indicated ontogenetic shifts in thermal niches: juveniles of all whitefish morphs inhabited warmer ambient temperatures than adults. According to sampling of the otolith nucleus, hatching temperatures were higher for benthic compared to pelagic morphs. Estimated respiration rate was the lowest for benthivorous profundal morph, contrasting with the higher values estimated for the other morphs that inhabited shallower and warmer water. These preliminary results suggest that physiological adaptation to different thermal habitats shown by the sympatric morphs may play a significant role in maintaining or strengthening niche segregation and divergence in life-history traits, potentially contributing to reproductive isolation and incipient speciation.

Ecological systems are often characterized as stable entities. However, basal productivity in most ecosystems varies between seasons, particularly in subarctic and polar areas. How this variability affects higher trophic levels or entire food webs remains largely unknown, especially in these high-latitude regions. We undertook a year-long study of benthic (macroinvertebrate) and pelagic (zooplankton) resource availability, along with short (day/days: stomach content)-, medium (month: liver δ(13)C and δ(15)N isotopes)- and long-term (season: muscle δ(13)C and δ(15)N isotopes) assessments of resource use by a generalist fish, the European whitefish, in a deep, oligotrophic, subarctic lake in northern Europe. Due to the long ice-covered winter period, we expected to find general benthic reliance throughout the year, but also a seasonal importance of zooplankton to the diet, somatic growth and gonadal development of whitefish. Benthic and pelagic resource availability varied between seasons: peak littoral benthic macroinvertebrate density occurred in mid-winter, whereas maximum zooplankton density was observed in summer. Whitefish stomach content revealed a reliance on benthic prey items during winter and pelagic prey in summer. A seasonal shift from benthic to pelagic prey was evident in liver isotope ratios, but muscle isotope ratios indicated a year-round reliance on benthic macroinvertebrates. Whitefish activity levels as well as somatic and gonadal growth all peaked during the summer, coinciding with the zooplankton peak and the warmest water temperature. Stable isotopes of muscle consistently depicted the most important resource, benthic macroinvertebrates, whereas short-term indicators, that is, diet and stable isotopes of liver, revealed the seasonal significance of pelagic zooplankton for somatic growth and gonad development. Seasonal variability in resource availability strongly influences consumer growth and reproduction and may also be important in other ecosystems facing pronounced annual weather fluctuations., (© 2014 The Authors. Journal of Animal Ecology © 2014 British Ecological Society.)

Understanding how a monophyletic lineage of a species diverges into several adaptive forms has received increased attention in recent years, but the underlying mechanisms in this process are still under debate. Postglacial fishes are excellent model organisms for exploring this process, especially the initial stages of ecological speciation, as postglacial lakes represent replicated discrete environments with variation in available niches. Here, we combine data of niche utilization, trophic morphology, and 17 microsatellite loci to investigate the diversification process of three sympatric European whitefish morphs from three northern Fennoscandian lakes. The morphological divergence in the gill raker number among the whitefish morphs was related to the utilization of different trophic niches and was associated with reproductive isolation within and across lakes. The intralacustrine comparison of whitefish morphs showed that these systems represent two levels of adaptive divergence: (1) a consistent littoral-pelagic resource axis; and (2) a more variable littoral-profundal resource axis. The results also indicate that the profundal whitefish morph has diverged repeatedly from the ancestral littoral whitefish morph in sympatry in two different watercourses. In contrast, all the analyses performed revealed clustering of the pelagic whitefish morphs across lakes suggesting parallel postglacial immigration with the littoral whitefish morph into each lake. Finally, the analyses strongly suggested that the trophic adaptive trait, number of gill rakers, was under diversifying selection in the different whitefish morphs. Together, the results support a complex evolutionary scenario where ecological speciation acts, but where both allopatric (colonization history) and sympatric (within watercourse divergence) processes are involved.

Species introductions are considered one of the major drivers of biodiversity loss via ecological interactions and genetic admixture with local fauna. We examined two well-recognized fish species, native whitefish (Coregonus lavaretus) and introduced vendace (Coregonus albula), as well as their morphological hybrids in a single lake to test for selection against hybrids and backcrosses in the wild. A representative random subsample of 693 individuals (27.8%) was taken from the total catch of coregonids. This subsample was examined with the aim to select c. 50 individuals of pure whitefish (n = 52), pure vendace (n = 55) and putative hybrid (n = 19) for genetic analyses. The subsequent microsatellites and mitochondrial (mt) DNA analyses provided compelling evidence of hybridization and introgression. Of the 126 fish examined, four were found to be F(1) , 14 backcrosses to whitefish and seven backcrosses to vendace. The estimates of historical gene flow suggested higher rates from introduced vendace into native whitefish than vice versa, whereas estimates of contemporary gene flow were equal. Mitochondrial introgression was skewed, with 18 backcrosses having vendace mtDNA and only three with whitefish mtDNA. Hybrids and backcrosses had intermediate morphology and niche utilization compared with parental species. No evidence of selection against hybrids or backcrosses was apparent, as both hybrid and backcross growth rates and fecundities were high. Hybrids (F(1) ) were only detected in 2 year-classes, suggesting temporal variability in mating between vendace and whitefish. However, our data show that hybrids reached sexual maturity and reproduced actively, with backcrosses recorded from six consecutive year-classes, whereas no F(2) individuals were found. The results indicate widespread introgression, as 10.8% of coregonids were estimated to be backcrosses., (© 2011 Blackwell Publishing Ltd.)

1. The adaptive radiation of fishes into benthic (littoral) and pelagic (lentic) morphs in post-glacial lakes has become an important model system for speciation. Although these systems are well studied, there is little evidence of the existence of morphs that have diverged to utilize resources in the remaining principal lake habitat, the profundal zone. 2. Here, we tested phenotype-environment correlations of three whitefish (Coregonus lavaretus) morphs that have radiated into littoral, pelagic and profundal niches in northern Scandinavian lakes. We hypothesized that morphs in such trimorphic systems would have a morphology adapted to one of the principal lake habitats (littoral, pelagic or profundal niches). Most whitefish populations in the study area are formed by a single (monomorphic) whitefish morph, and we further hypothesized that these populations should display intermediate morphotypes and niche utilization. We used a combination of traditional (stomach content, habitat use, gill raker counts) and more recently developed (stable isotopes, geometric morphometrics) techniques to evaluate phenotype-environment correlations in two lakes with trimorphic and two lakes with monomorphic whitefish. 3. Distinct phenotype-environment correlations were evident for each principal niche in whitefish morphs inhabiting trimorphic lakes. Monomorphic whitefish exploited multiple habitats, had intermediate morphology, displayed increased variance in gillraker-counts, and relied significantly on zooplankton, most likely due to relaxed resource competition. 4. We suggest that the ecological processes acting in the trimorphic lakes are similar to each other, and are driving the adaptive evolution of whitefish morphs, possibly leading to the formation of new species.

Seasonal and ontogenetic shifts in the diet of Arctic charr Salvelinus alpinus were studied in a deep, ultra-oligotrophic lake in subarctic Finland from both stomach contents and the stable carbon (delta(13)C) and nitrogen (delta(15)N) isotope compositions of muscle and liver tissues. Both diet and isotope results indicated that the S. alpinus population relied mainly on littoral benthic energy sources. The strong littoral reliance appeared largely independent of season or fish size, although the data lacked small (total length, L(T), <130 mm) and young (<3 years) S. alpinus. Liver isotope values of intermediate-sized S. alpinus (200-350 mm), however, suggested exploitation of the increase in the abundance of pelagic zooplankton in the late open-water season. The results suggest that, in general, a strong littoral reliance of fishes can be a feature in subarctic lakes throughout the year. Due to its faster isotopic turnover rate and thus higher resolution for temporal diet changes, liver could be more commonly used in stable-isotope studies of fish trophic niche shifts instead of using only the less responsive muscle tissue.