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Subarctic lakes are sentinels of climate change, showing responses in their physical, chemical, and biological properties. However, climate-induced changes in invertebrate diversity and their underlying mechanisms are not fully understood. We explored the relationship between past climate change and taxonomic composition of subfossil cladocerans in a subarctic lake during the last ca. 5700 years. The Cladocera community shifted from specialist to generalist species at ca. 3500 cal years BP, corresponding to the long-term cooling period between the Holocene Thermal Maximum and the Late Holocene. Taxonomic diversity declined driven by the collapse of the keystone herbivorous Daphnia longispina group, pelagic and littoral predators, and phytophilous benthic species, therefore resulting in a simplification of the food web and a reduction of trophic levels. Furthermore, the shift in cladoceran composition was associated with the decline of aquatic primary producers and the development of birch forest, suggesting a potential causal link between dissolved organic carbon dynamics and cladoceran community composition. This study provides empirical evidence of the response of cladocerans to climatic fluctuations and their underlying mechanisms through catchment-mediated processes and direct temperature-induced changes., Competing Interests: Competing interests: The authors declare no competing interests., (© 2024. The Author(s).)

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.)

Reconstructions of past food web dynamics are necessary for better understanding long-term impacts of climate change on subarctic lakes. We studied elemental and stable isotopic composition of sedimentary organic matter, photosynthetic pigments and carbon stable isotopic composition of Daphnia (Cladocera; Crustacea) resting eggs (δ 13 C Clado ) in a sediment record from a small subarctic lake. We examined how regional climate and landscape changes over the last 5800 years affected the relative importance of allochthonous and autochthonous carbon transfer to zooplankton. Overall, δ 13 C Clado values were well in line with the range of theoretical values of aquatic primary producers, confirming that zooplankton consumers in subarctic lakes, even in the long-term perspective, are mainly fuelled by autochthonous primary production. Results also revealed greater incorporations of benthic algae into zooplankton biomass in periods that had a warmer and drier climate and clearer water, whereas a colder and wetter climate and lower water transparency induced higher contributions of planktonic algae to Daphnia biomass. This study thus emphasizes long-term influence of terrestrial-aquatic linkages and in-lake processes on the functioning of subarctic lake food webs.

Dystrophic lakes are widespread in temperate regions and intimately interact with surrounding terrestrial ecosystems in energy and nutrient dynamics, yet the relative importance of autochthonous and allochthonous resources to consumer production in dystrophic lakes remains controversial. We argue that allochthonous organic matter quantitatively dominates over photosynthetic autotrophs in dystrophic lakes, but that autotrophs are higher in diet quality and more important for consumers as they contain essential polyunsaturated fatty acids (PUFA). In a field study, we tested the hypotheses that (1) autochthonous primary production is the main driver for consumer production, despite being limited by light availability and low nutrient supplies, and greater supply of allochthonous carbon, (2) the relative contribution of autotrophs to consumers is directly related to their tissue PUFA concentrations, and (3) methane-oxidizing bacteria (MOB) provide an energy alternative for consumers. Pelagic and benthic consumer taxa representing different trophic levels were sampled from five dystrophic lakes: isopod Asellus aquaticus, megalopteran Sialis lutaria, dipteran Chaoborus flavicans, and perch Perca fluviatilis. Based on carbon and nitrogen stable isotopes, the relative contributions of autochthonous (biofilms and seston) and allochthonous (coarse particulate and dissolved organic matter) resources and MOB to these taxa were 47-79%, 9-44% and 7-12% respectively. Results from fatty acid (FA) analyses show that the relative omega3-FA and PUFA concentrations increased with trophic level (Asellus < Sialis and Chaoborus < Perca). Also, eicosapentaenoic-acid (EPA), omega3-FA and PUFA concentrations increased with the autochthonous contribution in consumers, i.e., a 47-79% biofilm and/or seston diet resulted in tissue EPA of 4.2-18.4, omega3 FAs of 11.6-37.0 and PUFA of 21.6-61.0 mg/g dry mass. The results indicate that consumers in dystrophic lakes predominantly rely on energy from autotrophs and that their PUFA concentrations are dependent on the relative contribution of these autochthonous resources. The limited energy support from MOB suggests they are not negligible and are potentially an integral part of the food webs. Our findings show that autochthonous resources are the main driver of secondary production even in dystrophic lakes and offer new insights into the functioning of these ecosystems.

Northern lakes are facing rapid environmental alterations—including warming, browning, and/or changes in nutrient concentrations—driven by climate change. These environmental changes can have profound impacts on the synthesis and trophic transfer of polyunsaturated fatty acids (PUFA), which are important biochemical molecules for consumer growth and reproduction. Zooplankton are a key trophic link between phytoplankton and fish, but their biochemical responses to environmental change are not well understood. In this study, we assess the trends in fatty acid (FA) composition of zooplankton taxa among 32 subarctic and temperate lakes across broad climate‐productivity and ecosystem size gradients. We found that genus‐level taxonomy explained most FA variability in zooplankton (54%), suggesting that environmental changes that alter the taxonomic composition also affect the FA composition of zooplankton communities. Furthermore, the FA responses and their underlying environmental drivers differed between cladocerans and copepods. Cladocerans, including widespread Bosmina spp. and Daphnia spp., showed pronounced responses across the climate‐productivity gradient, with abrupt declines in PUFA, particularly eicosapentaenoic acid and arachidonic acid in warmer, browner, and more eutrophic lakes. Conversely, calanoid copepods had high and relatively stable PUFA levels across the gradient. In addition, all zooplankton taxa increased in stearidonic acid levels in larger lakes where PUFA‐rich cryptophytes were more abundant. Overall, our results suggest that climate‐driven environmental alterations pose heterogeneous impacts on PUFA levels among zooplankton taxa, and that the negative impacts of climate warming are stronger for cladocerans, especially so in small lakes. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

We combined paleolimnological reconstructions and space-for-time substitutions to unravel chironomid biodiversity responses to climate change in subarctic mountains across temporal and spatial scales. Using sediment records, we found that long-term temporal changes in chironomid taxonomic diversity were mainly induced by the temperature tolerance/optimum of species, while little changes in functional diversity were found due to the replacement of similar functional-type taxa within the community. Overall, paleolimnological reconstructions suggested the selection of larger chironomid species by long-term climate cooling and little changes in trophic guilds. Space-for-time substitutions showed, however that low-elevation lakes with forested have more sediment-feeding taxa and larger larvae than high-elevation lakes, thus, suggesting the selection of large chironomid morphotypes with a sediment-feeding mode under warmer climate. Space-for-time substitutions and paleolimnological reconstructions, therefore, gave contrasting results for the link between climate and functional diversity of chironomid communities, likely because space-for-time substitutions failed to match the extent of both spatial and temporal climatic gradients. We suggest that future studies must address biodiversity issues across both temporal and spatial scales as an improved understanding of biodiversity responses to climate change may help us to understand how biodiversity will be affected by ongoing and future change. [ABSTRACT FROM AUTHOR]

Arctic and sub‐Arctic lakes in northern Europe are increasingly threatened by climate change, which can affect their biodiversity directly by shifting thermal and hydrological regimes, and indirectly by altering landscape processes and catchment vegetation. Most previous studies of northern lake biodiversity responses to environmental changes have focused on only a single organismal group. Investigations at whole‐lake scales that integrate different habitats and trophic levels are currently rare, but highly necessary for future lake monitoring and management.We analysed spatial biodiversity patterns of 74 sub‐Arctic lakes in Norway, Sweden, Finland, and the Faroe Islands with monitoring data for at least three biological focal ecosystem components (FECs)—benthic diatoms, macrophytes, phytoplankton, littoral benthic macroinvertebrates, zooplankton, and fish—that covered both pelagic and benthic habitats and multiple trophic levels.We calculated the richnessrelative (i.e. taxon richness of a FEC in the lake divided by the total richness of that FEC in all 74 lakes) and the biodiversity metrics (i.e. taxon richness, inverse Simpson index (diversity), and taxon evenness) of individual FECs using presence–absence and abundance data, respectively. We then investigated whether the FEC richnessrelative and biodiversity metrics were correlated with lake abiotic and geospatial variables. We hypothesised that (1) individual FECs would be more diverse in a warmer and wetter climate (e.g. at lower latitudes and/or elevations), and in hydrobasins with greater forest cover that could enhance the supply of terrestrial organic matter and nutrients that stimulated lake productivity; and (2) patterns in FEC responses would be coupled among trophic levels.Results from redundancy analyses showed that the richnessrelative of phytoplankton, macrophytes, and fish decreased, but those of the intermediate trophic levels (i.e. macroinvertebrates and zooplankton) increased with decreasing latitude and/or elevation. Fish richnessrelative and diversity increased with increasing temporal variation in climate (temperature and/or precipitation), ambient nutrient concentrations (e.g. total nitrogen) in lakes, and woody vegetation (e.g. taiga forest) cover in hydrobasins, whereas taxon richness of macroinvertebrates and zooplankton decreased with increasing temporal variation in climate.The similar patterns detected for richnessrelative of fish, macrophytes, and phytoplankton could be caused by similar responses to the environmental descriptors, and/or the beneficial effects of macrophytes as habitat structure. By creating habitat, macrophytes may increase fish diversity and production, which in turn may promote higher densities and probably more diverse assemblages of phytoplankton through trophic cascades. Lakes with greater fish richnessrelative tended to have greater average richnessrelative among FECs, suggesting that fish are a potential indicator for overall lake biodiversity.Overall, the biodiversity patterns observed along the environmental gradients were trophic‐level specific, indicating that an integrated food‐web perspective may lead to a more holistic understanding of ecosystem biodiversity in future monitoring and management of high‐latitude lakes. In future, monitoring should also focus on collecting more abundance data for fish and lower trophic levels in both benthic and pelagic habitats. This may require more concentrated sampling effort on fewer lakes at smaller spatial scales, while continuing to sample lakes distributed along environmental gradients. [ABSTRACT FROM AUTHOR]

Summary 1. Fatty acids (FAs) have been widely applied as trophic biomarkers in aquatic food web studies. However, current knowledge of inter- and intraspecific variation in consumer FA compositions across spatial and temporal scales is constrained to a few pelagic taxa. 2. We analysed the FAs of 22 taxa of benthic macroinvertebrates, zooplankton and fish collected from the littoral, pelagic and profundal habitats of nine boreal oligotrophic lakes over spring, summer and autumn. We quantified and compared the FA variance partitions contributed by species identity (i.e. an integrative effect of phylogenetic origin, life history and functional feeding guild of individual taxa), site and season using partial redundancy analysis both on all consumers and on benthic arthropods alone. 3. Species identity alone contributed 84.4 and 72.8% of explained FA variation of all consumers and benthic arthropods, respectively. Influences of site, season and all joint effects accounted for 0-11.3% only. Fatty acid profiles of primary consumers differentiated below class level, but those of predators were distinguishable only when they became more taxonomically distinct (i.e. among classes or higher). 4. Pelagic and profundal consumers showed stronger reliance on autochthonous resources than did their littoral counterparts as reflected by their higher ω3 to ω6 FA ratios. Polyunsaturated FAs (PUFAs) were increasingly retained with trophic levels, and saturated FAs (e.g. FA 16 : 0) gradually reduced. Ecologically, this trade-off enhances the trophic transfer efficiency and confirms the importance of PUFA-rich autotrophs in aquatic food webs. 5. Our findings indicate strong interspecific differences in FA requirements and assimilation among aquatic consumers from a wide range of taxonomic levels, habitats and lakes. Consumers were able to maintain homoeostasis in FA compositions across spatial and temporal changes in resource FAs, but consumer homoeostasis did not limit the effectiveness of FAs as trophic biomarkers. [ABSTRACT FROM AUTHOR]

Hydroelectric reservoirs retain large volumes of water and have a global impact on sea level, elemental cycles, and biodiversity. Using data from a total of 90 historical and recent surveys in nine regulated and eight unregulated alpine and subalpine lakes, we show an additional large effect of reservoirs, i.e., that impoundment causes drastically decreased fish growth and thereby great negative consequences for inland fisheries in Scandinavia. Following a long period (40-65 years) after impoundment, the length and mass of Arctic charr () of the single age class 4+ years was, on average, 35% and 72% lower, respectively, in impounded versus natural lakes in northern Scandinavia. The effect was stronger at higher altitudes and can be mitigated by addition of inorganic nutrients. We suggest that the decreased fish growth is a consequence of lowered ecosystem productivity, oligotrophication, caused by impoundment, resulting in erosion and loss of the littoral ecosystem as well as delayed flooding and leakage of nutrients from the riparian zone until after the growing season. [ABSTRACT FROM AUTHOR]

The transfer of energy and nutrients from plants to animals is a key process in all ecosystems. In lakes, inefficient transfer of primary producer derived energy can result in low animal growth rates, accumulation of nuisance phytoplankton blooms and dissipation of energy from the ecosystem. Most research on carbon transfer efficiency in pelagic food webs has focused on either food quantity or food quality, with the latter considered separately as either elemental stoichiometry or biochemical composition. The natural occurrence and magnitude of these types of growth limitations and their combined effects on Daphnia, a keystone grazer in pelagic freshwater ecosystems, are largely unknown. Our empirical models predict that the strength and nature of food quantity and quality limitation varies greatly with lake trophic state (total phosphorus, TP) and that Daphnia growth rates and thus energy and nutrient transfer efficiency are highest in lakes with intermediate trophic status (TP 10–25 μg l−1). We predict that food availability place the greatest constraint on Daphnia growth in nutrient poor lakes (TP≤4 μg l−1). Phosphorus limitation of Daphnia growth increased with decreasing TP, but the overall effect was never predicted to be the dominant constraining factor. Eicosapentaenoic acid (EPA, 20:5ω3) limitation was predicted to occur in both nutrient poor and nutrient rich lakes and placed the primary constraint on food quality in the most productive lakes. Two contrasting EPA-models gave different results on the magnitude of EPA-limitation, implying that additional food quality factors decrease Daphnia growth at high TP. In conclusion, the model predicts that Daphnia growth should peak in mesotrophic lakes, food quantity will place the greatest constraint on growth in oligotrophic lakes and EPA will primarily limit growth in eutrophic lakes. [ABSTRACT FROM AUTHOR]

Management actions often need to be taken in the absence of ecological information to mitigate the impact of pressing environmental problems. Managers counteracted the detrimental effects of cultural acidification on aquatic ecosystems during the industrial era using liming to salvage biodiversity and ecosystem services. However, historical contingencies, i.e., whether lakes were naturally acidic or degraded because of acidification, were largely unknown and therefore not accounted for in management. It is uncertain whether liming outcomes had a potentially detrimental effect on naturally acidic lakes. Evidence from paleolimnological reconstructions allowed us to analyze community structure in limed acidified and naturally acidic lakes, and acidified and circumneutral references. We analyzed community structure of phytoplankton, zooplankton, macroinvertebrates (littoral, sublittoral, profundal), and fish between 2000 and 2004. Naturally acidic limed lakes formed communities that were not representative of the other lake types. The occurrence of fish species relevant for ecosystem service provisioning (fisheries potential) in naturally acidic limed lakes were confounded by biogeographical factors. In addition, sustained changes in water quality were conducive to harmful algal blooms. This highlights a pathological outcome of liming lakes when their naturally acidic conditions are not accounted for. Because liming is an important socialecological system, sustained ecological change of lakes might incur undesired costs for societies in the long term. [ABSTRACT FROM AUTHOR]