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  • Rolls, Robert J.; Hayden, Brian; Kahilainen, Kimmo K. (2017)
    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.
  • Ohlund, Gunnar; Bodin, Mats; Nilsson, Karin A.; Ohlund, Sven-Ola; Mobley, Kenyon B.; Hudson, Alan G.; Peedu, Mikael; Brännström, Åke; Bartels, Pia; Prabel, Kim; Hein, Catherine L.; Johansson, Petter; Englund, Göran (2020)
    Lake-dwelling fish that form species pairs/flocks characterized by body size divergence are important model systems for speciation research. Although several sources of divergent selection have been identified in these systems, their importance for driving the speciation process remains elusive. A major problem is that in retrospect, we cannot distinguish selection pressures that initiated divergence from those acting later in the process. To address this issue, we studied the initial stages of speciation in European whitefish (Coregonus lavaretus) using data from 358 populations of varying age (26-10,000 years). We find that whitefish speciation is driven by a large-growing predator, the northern pike (Esox lucius). Pike initiates divergence by causing a largely plastic differentiation into benthic giants and pelagic dwarfs: ecotypes that will subsequently develop partial reproductive isolation and heritable differences in gill raker number. Using an eco-evolutionary model, we demonstrate how pike's habitat specificity and large gape size are critical for imposing a between-habitat trade-off, causing prey to mature in a safer place or at a safer size. Thereby, we propose a novel mechanism for how predators may cause dwarf/giant speciation in lake-dwelling fish species.
  • Kozak, Natalia; Ahonen, Salla A.; Keva, Ossi; ostbye, Kjartan; Taipale, Sami J.; Hayden, Brian; Kahilainen, Kimmo K. (2021)
    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 degrees N), climatic (+3.2 degrees 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 (log10THg) 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 R2 = 0.90, p < 0.001). The model examining variation in THg baseline values included the same variables with positive relationships (R2 = 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. (c) 2021 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY license (http://
  • Eloranta, Antti P.; Kahilainen, Kimmo K.; Amundsen, Per-Arne; Knudsen, Rune; Harrod, Chris; Jones, Roger I. (2015)
    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.
  • Vesterinen, Jussi; Keva, Ossi; Kahilainen, Kimmo K.; Strandberg, Ursula; Hiltunen, Minna; Kankaala, Paula; Taipale, Sami J. (2021)
    Littoral benthic primary production is considered the most important energy source of consumers in subarctic lakes. We analyzed essential fatty acid (EFA) and amino acid (EAA) content of 23 littoral benthic macroinvertebrate taxa as well as cladocerans and copepods from pelagic and littoral habitats of 8-9 subarctic lakes to compare their nutritional quality. Pelagic crustacean zooplankton had significantly higher EFA and total FA content (on average 2.6-fold and 1.6-fold, respectively) than littoral macroinvertebrates in all our study lakes. Specifically, docosahexaenoic acid (DHA), one of the most important EFA for juvenile fish, was almost exclusively found in pelagic copepods. In littoral macroinvertebrates, onlyLymnaea(Gastropoda),Eurycercus(Cladocera), andGammarus(Amphipoda) contained a low amount of DHA, whereas most littoral invertebrate taxa contained moderate amounts of eicosapentaenoic acid (EPA). The difference in DHA content may explain why so many generalist fish shift their diet to pelagic zooplankton at their peak abundance in mid/late-summer. Meanwhile, the differences in EAA content between pelagic zooplankton and littoral invertebrates were much lower than for EFA suggesting a wider availability of EAA in subarctic lakes, except for methionine. In the studied subarctic lakes, EFA and EAA variation in consumers was more related to taxon-specific than lake-specific characteristics. This indicates that climate-induced changes in the abundance and community structure of zooplankton vs. littoral macroinvertebrates will be important parameters in determining the availability of EFA and EAA to juvenile fish, and potentially fish production.
  • Keva, Ossi; Hayden, Brian; Harrod, Chris; Kahilainen, Kimmo K. (2017)
    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 mu g/g, muscle: 0.24 +/- 0.05 mu g/g) and lowest in summer (liver: 0.87 +/- 0.72 mu g/g, muscle: 0.19 +/- 0.04 mu g/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 bio-accumulated 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, delta C-13, 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. (C) 2017 Elsevier Ltd. All rights reserved.