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Resource polymorphism has been suggested to be a platform for speciation. In some cases resource polymorphism depends on phenotypic plasticity but in other cases on genetic differences between morphotypes, which in turn has been suggested to be the ongoing development of a species pair. Here we study environmentally induced morphological differences in two age classes of Arctic char (Salvelinus alpinus) influencing char performance and diet in relation to resource availability. We found that structurally complex habitats with relatively lower zooplankton densities gave rise to individuals with a deeper body, and a downward positioned tip of the snout compared with individuals from structurally simple habitats with relatively higher zooplankton densities for both age classes. Environment also had an effect on foraging efficiency on zooplankton, with fish from structurally simple habitats had a higher foraging rate than fish from structurally complex habitats. Diet analyses showed that resource use in char mainly depends on the relative abundance of different resources. Therefore, to gain further understanding of resource polymorphism we suggest that future studies must include population dynamic feedbacks by the resources on the consumers.

Question: Does cannibalism lead to resource polymorphism in young Arctic charr (Salvelinusalpinus, Pisces)?

Hypothesis: Cannibals should evoke a low-activity morph that is well adapted to benthivorybut not planktivory, and which differs in morphology compared with a planktivorous morph.

Methods: We reared young-of-the-year charr in laboratory aquaria with and without largercannibalistic charr present. Thereafter, we measured foraging efficiency on pelagic and benthicresources, swimming speed when foraging, and morphology of the young charr.

Conclusions: Living among cannibals did not affect the morphology of the young charr. Italso did not affect the foraging efficiency of the young charr on the benthic resource. However,individuals from cannibal treatments swam closer and had lower foraging efficiency on thepelagic resource.

We investigated productivity at the basal trophic level in 15 unproductive lakes in a gradient ranging from clear-water to brown-water (humic) lakes in northern Sweden. Primary production and bacterial production in benthic and pelagic habitats were measured to estimate the variation in energy mobilization from external energy sources (primary production plus bacterial production on allochthonous organic carbon) along the gradient. Clear-water lakes were dominated by autotrophic energy mobilization in the benthic habitat, whereas humic lakes were dominated by heterotrophic energy mobilization in the pelagic habitat. Whole-lake (benthic + pelagic) energy mobilization was negatively correlated to the light-extinction coefficient, which was determined by colored terrestrial organic matter in the lake water. Thus, variation in the concentration of terrestrial organic matter and its light-absorbing characteristics exerts strong control on the magnitude, as well as on the processes and pathways, of energy mobilization in unproductive lakes. We suggest that unproductive lakes in general are sensitive to input of terrestrial organic matter because of its effects on basal energy mobilization in both benthic and pelagic habitats.

This study quantified new biomass production of algae and bacteria in both benthic and pelagic habitats of clear-water lakes to contrast how carbon from the atmosphere and terrestrial sources regulates whole-lake metabolism. We studied four small unproductive lakes in subarctic northern Sweden during one summer season. The production of new biomass in both benthic and pelagic habitats was calculated as the sum of autotrophic production by algae and heterotrophic production by bacteria using allochthonous organic carbon (OC). Whole-lake production of new biomass was dominated by the benthic habitat (86% +/- 4% [mean +/- SD]) and by primary production (77% +/- 9%). Still, heterotrophic bacteria fueled by allochthonous OC constituted a significant portion of the new biomass production in both benthic (19% +/- 11%) and pelagic habitats (51% +/- 24%). In addition, overall net production (primary production minus respiration) was close to zero in the benthic habitats but highly negative (-163 +/- 81 mg C.m(-2).d(-1)) in pelagic regions of all lakes. We conclude (1) that allochthonous OC supported a significant part of total production of new biomass in both pelagic and benthic habitats, (2) that benthic habitats dominated the whole-lake production of new biomass, and (3) that respiration and net CO2 production dominated the carbon flux of the pelagic habitats and biomass production dominated the benthic carbon flux. Taken together, these findings suggest that previous investigations have greatly underestimated the productivity of clear-water lakes when benthic autotrophic production and metabolism of allochthonous OC have not been measured.

Climate change results in changes in the geographical distribution of species. Species invasion success into a new area is dependent both on the dispersal ability of species as well as the strength and identity of biotic interactions between resident and invading species. Coexistence in intraguild predation (IGP) systems depends on the relative strength of predation and competition interactions which in turn are temperature dependent. We investigated the effects of introducing an intermediate consumer, ninespine stickleback (Pungitius pungitius), into allopatric populations of the omnivorous top predator Arctic char (Salvelinus alpinus). Introductions were performed in lakes with different climate regimes, two tundra lakes and two forest lakes that differed in average summer temperatures with 1.4 ± 0.5 ºC (average ± 1SD). We found that sticklebacks were able to invade and increase in density in both tundra and forest lakes. Sticklebacks had strong negative effects on resource densities which also was reflected in a decreased growth of small char. Increasing stickleback density had a positive effect on growth of large adults and on the maximum size of char. We conclude that stickleback presence is not limited by biological interactions in these systems but rather by dispersion ability. We suggest that the size dependency in the response of char to the invasion of sticklebacks is fundamental for the successful invasion of sticklebacks, and that size dependent interactions including cannibalism play important roles for coexistence in natural IGP-systems.

The effects of temperature on top consumer populations in subarctic lake communities were studied by contrasting two lake pairs in different climate regimes: one pair on the low alpine tundra and one pair in the subalpine birch forest. We measured zooplankton and macroinvertebrate biomasses over the season and estimated population density and size structure of the top consumer Arctic char (Salvelinus alpinus). Furthermore, we modelled char growth using literature data on temperature dependent search rate, handling time and metabolic demands. The forest lakes were warmer than the tundra lakes. Char in the forest lakes were larger and had a higher individual growth compared to char in the tundra lakes, while population density and biomasses of char were not different between the forest and the tundra lakes. There were no differences in macroinvertebrate and zooplankton resource levels available for char between lake pairs. Our modeling of char growth revealed that higher temperature increased growth of char at the observed resource densities, suggesting that the higher temperature in the forest lakes was primarily the cause of the higher growth of char in these lakes. We suggest that cannibalism in char may regulate char recruitment and thereby population density and biomass of char leading to effects of increasing temperature on consumer biomass and consumer individual growth different from what is expected in pure consumer-resource systems. Our results emphasize the importance of feedbacks within ecosystems when addressing effects of climate change and increasing temperature on lake communities.

Cross-ecosystem movements of material and energy are ubiquitous. Aquatic ecosystems typically receive material that also includes organic matter from the surrounding catchment. Terrestrial-derived (allochthonous) organic matter can enter aquatic ecosystems in dissolved or particulate form. Several studies have highlighted the importance of dissolved organic carbon to aquatic consumers, but less is known about allochthonous particulate organic carbon (POC). Similarly, most studies showing the effects of allochthonous organic carbon (OC) on aquatic consumers have investigated pelagic habitats; the effects of allochthonous OC on benthic communities are less well studied. Allochthonous inputs might further decrease primary production through light reduction, thereby potentially affecting autotrophic resource availability to consumers. Here, an enclosure experiment was carried out to test the importance of POC input and light availability on the resource use in a benthic food web of a clear-water lake. Corn starch (a C-4 plant) was used as a POC source due to its insoluble nature and its distinct carbon stable isotope value (delta C-13). The starch carbon was closely dispersed over the bottom of the enclosures to study the fate of a POC source exclusively available to sediment biota. The addition of starch carbon resulted in a clear shift in the isotopic signature of surface-dwelling herbivorous and predatory invertebrates. Although the starch carbon was added solely to the sediment surface, the carbon originating from the starch reached zooplankton. We suggest that allochthonous POC can subsidize benthic food webs directly and can be further transferred to pelagic systems, thereby highlighting the importance of benthic pathways for pelagic habitats.

Background

In recent decades, many attempts have been made to restore eutrophic lakes through biomanipulation. Reducing the populations of planktivorous and benthivorous fish (either directly or through stocking of piscivorous fish) may induce ecosystem changes that increase water transparency and decrease the risk of algal blooms and fish kills, at least in the short term. However, the generality of biomanipulation effects on water quality across lake types and geographical regions is not known. Therefore, we have undertaken a systematic review of such effects in eutrophic lakes in temperate regions throughout the world.

Methods

Searches for literature were made using online publication databases, search engines, specialist websites and bibliographies of literature reviews. Search terms were developed in English, Danish, Dutch and Swedish. Identified articles were screened for relevance using inclusion criteria set out in an a priori protocol. To reduce the risk of bias, we then critically appraised the combined evidence found on each biomanipulation. Data were extracted on outcomes such as Secchi depth and chlorophyll a concentration before, during and/or after manipulation, and on effect modifiers such as lake properties and amounts of fish removed or stocked.

Results

Our searches identified more than 14,500 articles. After screening for relevance, 233 of them remained. After exclusions based on critical appraisal, our evidence base included useful data on 128 biomanipulations in 123 lakes. Of these interventions, 85% had been made in Europe and 15% in North America. Meta-analysis showed that removal of planktivores and benthivores (with or without piscivore stocking) leads to increased Secchi depth and decreased chlorophyll a concentration during intervention and the first three years afterwards. Piscivore stocking alone has no significant effect. The response of chlorophyll a levels to biomanipulation is stronger in lakes where fish removal is intense, and in lakes which are small and/or have high pre-manipulation concentrations of total phosphorus.

Conclusions

Our review improves on previous reviews of biomanipulation in that we identified a large number of case studies from many parts of the world and used a consistent, repeatable process to screen them for relevance and susceptibility to bias. Our results indicate that removal of planktivorous and benthivorous fish is a useful means of improving water quality in eutrophic lakes. Biomanipulation tends to be particularly successful in relatively small lakes with short retention times and high phosphorus levels. More thorough fish removal increases the efficacy of biomanipulation. Nonetheless successes and failures have occurred across a wide range of conditions.