Dual fuels: intra-annual variation in the relative importance of benthic and pelagic resources to maintenance, growth and reproduction in a generalist salmonid fish.
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 d(13)C and d(15)N isotopes)- and long-term (season: muscle d(13)C and d(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.
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.
Department of Forestry and Wildlife Management, Inland Norway University of Applied Sciences, Campus Evenstad, NO2418, Elverum, Norway. kjartan.ostbye@ibv.uio.no.
Marine threespine sticklebacks colonized and adapted to brackish and freshwater environments since the last Pleistocene glacial. Throughout the Holarctic, three lateral plate morphs are observed; the low, partial and completely plated morph. We test if the three plate morphs in the brackish water Lake Engervann, Norway, differ in body size, trophic morphology (gill raker number and length), niche (stable isotopes; d15N, d13C, and parasites (Theristina gasterostei, Trematoda spp.)), genetic structure (microsatellites) and the lateral-plate encoding Stn382 (Ectodysplasin) gene. We examine differences temporally (autumn 2006/spring 2007) and spatially (upper/lower sections of the lake - reflecting low versus high salinity).
All morphs belonged to one gene pool. The complete morph was larger than the low plated, with the partial morph intermediate. The number of lateral plates ranged 8-71, with means of 64.2 for complete, 40.3 for partial, and 14.9 for low plated morph. Stickleback d15N was higher in the lower lake section, while d13C was higher in the upper section. Stickleback isotopic values were greater in autumn. The low plated morph had larger variances in d15N and d13C than the other morphs. Sticklebacks in the upper section had more T. gasterostei than in the lower section which had more Trematoda spp. Sticklebacks had less T. gasterostei, but more Trematoda spp. in autumn than spring. Sticklebacks with few and short rakers had more T. gasterostei, while sticklebacks with longer rakers had more Trematoda. spp. Stickleback with higher d15N values had more T. gasterostei, while sticklebacks with higher d15N and d13C values had more Trematoda spp. The low plated morph had fewer Trematoda spp. than other morphs.
Trait-ecology associations may imply that the three lateral plate morphs in the brackish water lagoon of Lake Engervann are experiencing ongoing divergent selection for niche and migratory life history strategies under high gene flow. As such, the brackish water zone may generally act as a generator of genomic diversity to be selected upon in the different environments where threespine sticklebacks can live.
The temporal window of ecological adaptation in postglacial lakes: a comparison of head morphology, trophic position and habitat use in Norwegian threespine stickleback populations.
Studying how trophic traits and niche use are related in natural populations is important in order to understand adaptation and specialization. Here, we describe trophic trait diversity in twenty-five Norwegian freshwater threespine stickleback populations and their putative marine ancestor, and relate trait differences to postglacial lake age. By studying lakes of different ages, depths and distance to the sea we examine key environmental variables that may predict adaptation in trophic position and habitat use. We measured trophic traits including geometric landmarks that integrated variation in head shape as well as gillraker length and number. Trophic position (Tpos) and niche use (a) were estimated from stable isotopes (d(13)C, d(15)N). A comparison of head shape was also made with two North American benthic-limnetic species pairs.
We found that head shape differed between marine and freshwater sticklebacks, with marine sticklebacks having more upturned mouths, smaller eyes, larger opercula and deeper heads. Size-adjusted gillraker lengths were larger in marine than in freshwater stickleback. Norwegian sticklebacks were compared on the same head shape axis as the one differentiating the benthic-limnetic North American threespine stickleback species pairs. Here, Norwegian freshwater sticklebacks with a more "limnetic head shape" had more and longer gillrakers than sticklebacks with "benthic head shape". The "limnetic morph" was positively associated with deeper lakes. Populations differed in a (mean?±?sd: 0.76?±?0.29) and Tpos (3.47?±?0.27), where a increased with gillraker length. Larger fish had a higher Tpos than smaller fish. Compared to the ecologically divergent stickleback species pairs and solitary lake populations in North America, Norwegian freshwater sticklebacks had similar range in Tpos and a values, but much less trait divergences.
Our results showed trait divergences between threespine stickleback in marine and freshwater environments. Freshwater populations diverged in trophic ecology and trophic traits, but trophic ecology was not related to the elapsed time in freshwater. Norwegian sticklebacks used the same niches as the ecologically divergent North American stickleback species pairs. However, as trophic trait divergences were smaller, and not strongly associated with the ecological niche, ecological adaptations along the benthic-limnetic axis were less developed in Norwegian sticklebacks.
Notes
Cites: J Fish Biol. 2009 Nov;75(8):2000-3620738669
Cites: Proc Biol Sci. 2000 Dec 7;267(1460):2375-8411133026
Cites: Evolution. 2009 Jul;63(7):1740-5319228184
Cites: Am Nat. 1997 Jun;149(6):1009-2918811261
Cites: Evolution. 2004 Jun;58(6):1274-8115266976
Cites: Mol Ecol. 2011 Jun;20(12):2483-9321443674
Cites: Proc Biol Sci. 2011 Jan 22;278(1703):233-820685715
Total mercury concentrations in liver and muscle of European whitefish (Coregonus lavaretus (L.)) in a subarctic lake - Assessing the factors driving year-round variation.
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, d13C, 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.
