Seasonal and ontogenetic shifts in the diet of Arctic charr Salvelinus alpinus in a subarctic lake

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 (δ¹³C) and nitrogen (δ¹⁵N) 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.     

Maternal effects in vulnerability to eye‐parasites and correlations between behavior and parasitism in juvenile Arctic charr

Hatchery‐reared fish show high mortalities after release to the wild environment. Explanations for this include potentially predetermined genetics, behavioral, and physiological acclimation to fish farm environments, and increased vulnerability to predation and parasitism in the wild. We studied vulnerability to Diplostomum spp. parasites (load of eye flukes in the lenses), immune defense (relative spleen size) and antipredator behaviors (approaches toward predator odor, freezing, and swimming activity) in hatchery‐reared juvenile Arctic charr (Salvelinus alpinus) using a nested mating design. Fish were exposed to eye‐fluke larvae via the incoming water at the hatchery. Fish size was positively associated with parasite load, but we did not find any relationship between relative spleen size and parasitism. The offspring of different females showed significant variation in their parasite load within sires, implying a dam effect in the vulnerability to parasites. However, the family background did not have any effect on spleen size. In the mean sire level over dams, the fish from the bolder (actively swimming) families in the predator trials suffered higher loads of eye flukes than those from more cautiously behaving families. Thus, the results indicate potentially maternally inherited differences in vulnerability to eye‐fluke parasites, and that the vulnerability to parasites and behavioral activity are positively associated with each other at the sire level. This could lead to artificial and unintentional selection for increased vulnerability to both parasitism and predation if these traits are favored in fish farm environments.     

Within‐host interactions shape virulence‐related traits of trematode genotypes

Within‐host interactions between co‐infecting parasites can significantly influence the evolution of key parasite traits, such as virulence (pathogenicity of infection). The type of interaction is expected to predict the direction of selection, with antagonistic interactions favouring more virulent genotypes and synergistic interactions less virulent genotypes. Recently, it has been suggested that virulence can further be affected by the genetic identity of co‐infecting partners (G × G interactions), complicating predictions on disease dynamics. Here, we used a natural host–parasite system including a fish host and a trematode parasite to study the effects of G × G interactions on infection virulence. We exposed rainbow trout (Oncorhynchus mykiss) either to single genotypes or to mixtures of two genotypes of the eye fluke Diplostomum pseudospathaceum and estimated parasite infectivity (linearly related to pathogenicity of infection, measured as coverage of eye cataracts) and relative cataract coverage (controlled for infectivity). We found that both traits were associated with complex G × G interactions, including both increases and decreases from single infection to co‐infection, depending on the genotype combination. In particular, combinations where both genotypes had low average infectivity and relative cataract coverage in single infections benefited from co‐infection, while the pattern was opposite for genotypes with higher performance. Together, our results show that infection outcomes vary considerably between single and co‐infections and with the genetic identity of the co‐infecting parasites. This can result in variation in parasite fitness and consequently impact evolutionary dynamics of host–parasite interactions.     

Veligers of an introduced bivalve,Limnoperna fortunei,are a new food resource that enhances growth of larval fish in the Paraná River (South America)

1. Larvae of ‘sábalo’, Prochilodus lineatus, whose adults represent over 60% of overall fish biomass in the Río de la Plata Catchment, have been observed to feed intensively on veligers of the exotic bivalve Limnoperna fortunei. 2. To assess the effects of this dietary shift on the growth of P. lineatus, 28‐day laboratory experiments were carried out feeding newly hatched P. lineatus larvae with three diets: zooplankton artificially enriched with L. fortunei veligers; natural zooplankton; and zooplankton artificially enriched with cladocerans and copepods. The average length, weight and gut contents of the fish larvae were assessed weekly and metabolic rates of fish larvae were measured. 3. Proportions of veligers in gut contents were always higher than those in the experimental diet: 100, 76 and 21% for veliger‐enriched, natural and low‐veliger diets, respectively. Larvae fed a veliger‐enriched diet grew to a significantly larger size than larvae fed the other two diets. In energetic balance comparisons using metabolic rates and prey energy content, all three diets were sufficient to support metabolism and growth. The greatest values of excess energy at the end of each week were in the veliger‐enriched experiments. 4. Feeding on veligers of L. fortunei significantly enhances the growth of P. lineatus larvae and supports the idea that this new and abundant resource is selectively preyed upon by P. lineatus during its larval stage. Higher growth rates may stem from the higher energy contents of veligers compared to crustaceans and/or from the lower energy costs of capturing slower prey.     

Phenotype-specific feeding behaviour of two arctic charr Salvelinus alpinus morphs

The feeding performance of two morphs of a trophically polymorphic fish, the arctic charr Salvelinus alpinus, feeding on zooplankton, was studied in the laboratory. Limnetic morphs, with a fusiform body, terminal mouth, short pectoral fins, and relatively long and dense gill rakers, fed at significantly higher rates on Daphnia and Cyclops than did benthic morphs. The benthic morphs were characterized by a chunky body form, subterminal mouth, long pectoral fins, and relatively short and spaced-out gill rakers. The limnetic morphs attacked the prey at significantly higher rates and were more successful in capture. Benthic morphs were more reluctant to start feeding on zooplankton and reaction time was longer than in limnetic morphs. In the absence as well as presence of zooplankton food, the position of the two morphs in the aquaria differed markedly. The results agree with dietary differences found in field-collected fish. The limnetic morphs consume zooplankton, whereas benthic morphs specialize on Lymnaea gastropods. The results suggest that benthic morphs are phenotypically inferior at using the pelagic food niche. This implies that the current role of competitive or predatory interactions in segregating the morphs may be minor.     

From autochthonous to allochthonous resources: seasonal shifts in food use by stream‐dwelling YOY Arctic charr Salvelinus alpinus through the ice‐free season

Substantial seasonal changes in resource use associated with enhanced water‐column use were revealed in stream‐living YOY Arctic charr Salvelinus alpinus during the ice‐free season. In July, YOY individuals showed a diet dominated by aquatic invertebrates (mainly Chironomidae larvae), but despite the small size of the fish, the abundance of terrestrial insects in their diet increased markedly from July to September (from 1·9 to 62·8%). Similarly, the frequency of surface drifting foragers, i.e. individuals feeding on allochthonous resources, increased from July to September (from 20·6 to 80%); allochthonous resources thus constituting an important energy subsidy for YOY S. alpinus during the late sub‐Arctic summer.     

Experimental evidence for parasite‐induced over‐winter mortality in juvenile Rhodeus amarus

In this study, the effects of the eye fluke Diplostomum pseudospathaceum (Trematoda) infection on over‐winter survival of young‐of‐the‐year (YOY) European bitterling Rhodeus amarus (Cyprinidae) were examined between September 2010 and April 2011. The fish were reared in semi‐natural conditions to ensure that results were not confounded by other parasite infections. The cumulative mortality of R. amarus from November until April was significantly higher in D. pseudospathaceum‐infected fish (57·3%) compared to controls (42·1%). Infection of the parental generation did not have any effect on the mortality of juveniles. The results indicate that D. pseudospathaceum infection increases over‐winter mortality of YOY R. amarus. The possible mechanisms causing mortality are discussed.     

Feeding behavior of the cichlid,Sarotherodon galilaeum: selective predation on Lake Kinneret zooplankton

In laboratory feeding trials, we analyzed the feeding behavior and selectivity of the cichlid, Sarotherodon galilaeum, for zooplankton prey from Lake Kinneret, Israel. The feeding behavior was dependent on fish size. Fish less than 20 mm SL fed on zooplankton as obligate particulate feeders. Fish from 20 to 42 mm SL fed either as particulate feeders or as filter feeders. Fish larger than 62 mm SL fed as obligate filter feeders. Particulate-feeding fish were size selective and had highest feeding electivities for large-sized zooplankton species. Filter-feeding fish had highest feeding electivities for zooplankton species with poor escape ability. In general, S. galilaeum predation pressure would be greatest on Ceriodaphnia reticulata, a large-bodied and easily captured species which is selected by both particulate-feeding and filter-feeding fish.     

An energy-based analysis of particulate-feeding and filter-feeding by blue tilapia,Tilapia aurea

Synopsis Adult blue tilapia, Tilapia aurea, employ filter-feeding as their primary feeding method, but feed as visual, particulate-feeding zooplanktivores as juveniles. We used measurements of oxygen utilization in enclosed chambers to assess filter-feeding energy costs, and videotaped observations of particulate-feeding to compute energy costs for this behavior. Weight-specific costs of filter-feeding are highest for smaller individuals, and decrease exponentially with fish size. Particulate-feeding costs increase with fish size and with distance travelled during attacks on zooplankton prey. These data were used in combination with published information to examine the energetics of the switch in feeding behavior in relation to the profitability of each feeding method. We develop a model which can be used to predict feeding behavior in relation to fish size and plankton array.     

Strategies of zooplanktivory shape the dynamics and diversity of littoral plankton communities: a mesocosm approach

Planktivorous fish can exert strong top‐down control on zooplankton communities. By incorporating different feeding strategies, from selective particulate feeding to cruising filter feeding, fish species target distinct prey. In this study, we investigated the effects of two species with different feeding strategies, the three‐spined stickleback (Gasterosteus aculeatus (L.)) and roach (Rutilus rutilus (L.)), on a low‐diversity brackish water zooplankton community using a 16‐day mesocosm experiment. The experiment was conducted on a small‐bodied spring zooplankton community in high‐nutrient conditions, as well as a large‐bodied summer community in low‐nutrient conditions. Effects were highly dependent on the initial zooplankton community structure and hence seasonal variation. In a small‐bodied community with high predation pressure and no dispersal or migration, the selective particulate‐feeding stickleback depleted the zooplankton community and decreased its diversity more radically than the cruising filter‐feeding roach. Cladocerans rather than copepods were efficiently removed by predation, and their removal caused altered patterns in rotifer abundance. In a large‐bodied summer community with initial high taxonomic and functional diversity, predation pressure was lower and resource availability was high for omnivorous crustaceans preying on other zooplankton. In this community, predation maintained diversity, regardless of predator species. During both experimental periods, predation influenced the competitive relationship between the dominant calanoid copepods, and altered species composition and size structure of the zooplankton community. Changes also occurred to an extent at the level of nontarget prey, such as microzooplankton and rotifers, emphasizing the importance of subtle predation effects. We discuss our results in the context of the adaptive foraging mechanism and relate them to the natural littoral community.     

Spatiotemporal and gender‐specific parasitism in two species of gobiid fish

Parasitism is considered a major selective force in natural host populations. Infections can decrease host condition and vigour, and potentially influence, for example, host population dynamics and behavior such as mate choice. We studied parasite infections of two common marine fish species, the sand goby (Pomatoschistus minutus) and the common goby (Pomatoschistus microps), in the brackish water Northern Baltic Sea. We were particularly interested in the occurrence of parasite taxa located in central sensory organs, such as eyes, potentially affecting fish behavior and mate choice. We found that both fish species harbored parasite communities dominated by taxa transmitted to fish through aquatic invertebrates. Infections also showed significant spatiotemporal variation. Trematodes in the eyes were very few in some locations, but infection levels were higher among females than males, suggesting differences in exposure or resistance between the sexes. To test between these hypotheses, we experimentally exposed male and female sand gobies to infection with the eye fluke Diplostomum pseudospathaceum. These trials showed that the fish became readily infected and females had higher parasite numbers, supporting higher susceptibility of females. Eye fluke infections also caused high cataract intensities among the fish in the wild. Our results demonstrate the potential of these parasites to influence host condition and visual abilities, which may have significant implications for survival and mate choice in goby populations.     

Effects of the sea louse Lepeophtheirus salmonis on temporal changes in cortisol,sex steroids,growth and reproductive investment in Arctic charr Salvelinus alpinus

Groups of mature (5+ year old) Arctic charr Salvelinus alpinus held in sea water were exposed for 34 days to either a high (mean ±s.e . 0·15 ± 0·01 sea lice Lepeophtheirus salmonis g^?1 fish mass) (HI), medium (0·07 ± 0·00 sea lice g^?1 fish mass) (MI) or no [control (C)] sea‐lice infection during early stages of gonad development (June to July). Infection with sea lice resulted in increased plasma cortisol concentrations and this was related to intensity of infection; females tended to have higher cortisol concentrations than males at high infection intensities (HI group: female c. 130 ng ml^?1; male c. 80 ng ml^?1). Plasma osmolality (C c. 330, MI c. 350 and HI c. 415 mOsm) and chloride concentrations (C c. 135, MI c. 155 and HI c. 190 mM) increased significantly with infection intensity, indicating osmoregulatory problems in infected fish. A strong positive relationship between plasma osmolality and cortisol concentration was recorded. Plasma sex‐steroid concentrations were influenced negatively by sea‐lice infection, particularly in the HI group, and were inversely related to plasma cortisol concentrations. The most heavily infected fish postponed the initiation of reproductive development until exposed to fresh water and timing of ovulation tended to be delayed in these fish. Growth rate and condition were negatively influenced by sea‐lice infection and growth rate was inversely related to plasma cortisol concentrations. Sea‐lice infection resulted in mortality among females in the HI group, and the proportion of maturing females was lower in the MI group (46%) than in the controls (85%). Egg production in the MI and HI groups was c. 50 and 30% of the C group. Egg size, embryonic survival and fry mass did not differ across groups. Sea lice influence reproductive development and egg production in S. alpinus, and consequently these parasites may influence populations via sublethal effects on broodfish, affecting growth and condition, and their reproductive output.     

Plankton dynamics in a tropical floodplain lake: fish, nutrients, and the relative importance of bottom-up and top-down control

1. Two enclosure experiments were carried out in Laguna Bufeos, a neotropical várzea lake located in the floodplain of River Ichilo (Bolivia). The experiments aimed (i) to assess the relative importance of bottom‐up and top‐down control on the plankton community, (ii) to assess the relative impact of direct and indirect effects of planktivorous fish on the zooplankton, and (iii) to attempt to identify the mechanisms responsible for these effects. 2. During the first experiment, bottom‐up control seemed to dominate the planktonic food web. Compared with fishless enclosures, oxygen concentrations, chlorophyll a levels and the population densities of all cladoceran zooplankton taxa increased in enclosures with fish. Birth rates of Moina minuta, the dominant taxon, were substantially higher in the presence than in the absence of fish, whereas death rates did not differ between treatments. These results are the first to suggest that the positive effects of fish on crustacean zooplankton via effects on nutrient cycling and the enhancement of primary production can compensate for losses because of fish‐related mortality. 3. During the second experiment, the direction of control appeared to vary between trophic levels: the phytoplankton appeared to be bottom‐up controlled whereas the zooplankton was mainly top‐down controlled. Chlorophyll a concentrations were enhanced by both fish and nutrient additions. The majority of the zooplankton taxa were reduced by the presence of fish. Birth rates of most cladoceran taxa did not differ between treatments, whereas death rates were higher in the enclosures with fish than in the fishless enclosures. Bosminopsis deitersi reached higher densities in the presence of fish, probably because of a release from predation by Chaoborus. 4. We convincingly showed strong deviations from trophic cascade‐based expectations, supporting the idea that trophic cascades may be weak in tropical lakes.     

The extinction of Arctodiaptomus alpinus (Copepoda) following the introduction of charr into a small alpine lake Dvojno Jezero (NW Slovenia)

In 1991 about twenty-five age 2+ specimens of the Arctic charr (Salvelinus alpinus (L.) (length between 100 and 130 mm) were introduced into the fishless small mountain lake Dvojno Jezero (= Twin Lake) (area 0.5+0.4 ha; max. depth 8 m, altitude 1670 m) in NW Slovenia. The first spawning of the Arctic charr was observed in autumn 1994 and regularly each year thereafter. Fish of different sizes were observed from 1995 onward each year but a fish census was never carried out. Autumn zooplankton samples, collected before the introduction of Arctic charr, contained adults (including ovigerous females) of two copepod species: Cyclops abyssorum tatricus and Arctodiaptomus alpinus. No planktonic Cladocera were found. Seven years after the introduction of fish standing crop of zooplankton declined 100-fold. The zooplankton contained only a few copepodites (IV and V) of C. a. tatricus. Simultaneously, water transparency declined and the concentration of chlorophyll a increased and filamentous green algae and picoplankton became abundant in the littoral zone of both lakes.     

Variation in scale shape among alternative sympatric phenotypes of Arctic charr Salvelinus alpinus from two lakes in Scotland

Landmark‐based geometric morphometric analysis was used to detect differences in scale shape between ecologically distinct phenotypes of Arctic charr Salvelinus alpinus coexisting in the same lake. Relative warp analysis and standard multivariate analyses of the partial warps, obtained after a Procrustes superimposition, showed that scale landmarks were efficient in discriminating among two closely related alternative phenotypes within each of the two lakes. In Loch Tay, S. alpinus exhibited a bimodal body size‐frequency distribution among sexually mature fish, whereas in Loch Awe, S. alpinus are unimodal in body size but segregated into two distinct spawning phenotypes. In both lakes, alternative phenotypes showed significant differences in foraging ecology, habitat use and life history. It is probable that differences in scale shape reflect differences in ecology of these forms.     

Water temperatures influence the marine area use of Salvelinus alpinus and Salmo trutta

The migratory behaviour and spatial area use of sympatric Arctic charr Salvelinus alpinus and brown trout Salmo trutta were investigated during their marine feeding migration. The likelihood of finding individuals of both species in the inner or outer fjord areas was dependent on water temperature in the inner area (especially for S. alpinus), the temperature difference between the inner and outer areas (especially for S. trutta) and fish fork length (both species). The strongest predictor was the water temperature in the inner area, and particularly S. alpinus left this area and moved to the outer areas with increasing temperatures in the inner area. At 8° C in the inner area, the likelihood of finding S. alpinus in the outer areas was >50%. This predictor had a smaller effect on S. trutta, and the likelihood of finding S. trutta in the outer areas only started to increase at around 14° C. The relationships between temperature and area use did not correspond to the species' optimal growth temperatures, but to their previously documented temperature preferences. Individuals of both species used mainly the littoral fjord areas, and to a lesser extent the pelagic areas. In conclusion, temperature differences between the inner and outer marine areas probably resulted in the segregated area use between the species, because water temperatures or factors influenced by temperature affected their migratory behaviour and habitat use differently. The results indicate that increased marine temperatures with global warming may lead to increased spatial overlap between S. trutta and S. alpinus, which again may lead to increased interspecific competition during their marine phase, and with S. alpinus probably being the more negatively affected.     

Climate and productivity shape fish and invertebrate community structure in subarctic lakes

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Finding the optimal vertical distribution: behavioural responses of Daphnia pulicaria to gradients of environmental factors and the presence of fish

1. The vertical distribution of zooplankton results from active habitat choice aiming to optimise fitness gain in a system of trade‐offs. 2. Using large, controlled indoor mesocosms (Plön Plankton Towers), we monitored the behavioural response of Daphnia pulicaria to vertical gradients of temperature, food, oxygen and light, in the presence and absence of fish predation. 3. In the absence of fish, Daphnia distributed as predicted by an ideal ‘free distribution with costs’. If the food was distributed homogeneously, they stayed in the warm epilimnion, while they balanced their time dwelling in epi‐ and hypolimnion if the food was concentrated in a deep‐water maximum. 4. However, oxygen depletion in the hypolimnion, representing an additional cost, prevented Daphnia from completely exploiting the hypolimnetic food maximum. Consequently, the proportion dwelling in the hypolimnion was larger if oxygen was not limiting. 5. Fish predation had an overwhelming effect, driving Daphnia into the hypolimnion under all experimental conditions. If permitted by oxygen availability, Daphnia used the whole hypolimnion, but oxygen depletion reduced their possible habitat to the upper hypolimnion with oxygen concentrations above c. 0.7 mg L^?1. As fish were less tolerant of low oxygen, the layer below the thermocline formed a predation refuge for Daphnia.     

Clonal integration facilitates invasiveness of the alien aquatic plant Myriophyllum aquaticum L. under heterogeneous water availability

The avoidance of visually feeding fish has long been considered as the primary driver of diel vertical migration of zooplankton. The diurnal vertical distribution of Cyclops gr. abyssorum, Arctodiaptomus alpinus, and Daphnia gr. longispina from 13 alpine lakes with fish (Salvelinus fontinalis) and without, was compared in order to understand whether fish in transparent lakes reduce the presence of large zooplankton from the irradiated zone. We used the light level at each sampling depth and the size of each specimen as proxies of predation risk, and we tested two predictions: (P1) the relative abundance of zooplankton in the well-lit surface waters vs. the darker waters will be greater in fishless lakes; (P2) the size of zooplankton in the well-lit surface waters vs. the deeper, darker waters will be greater in fishless lakes. We did not find any evidence of the validity of P1, but we confirmed P2 for Arctodiaptomus alpinus. These results support with new field data the Transparency Regulator Hypothesis, which argues that in transparent lakes, fish predation is less important for the vertical distribution of zooplankton than ultraviolet radiation, and further suggest that zooplankton size rather than vertical distribution may be more effective against visual predators in transparent lakes.     

Time–place learning and leader–follower relationships in Arctic charr Salvelinus alpinus

Feeding activity from a larger refuge site into two visually separated feeding sites with temporally restricted food availability, one in the morning and one in the evening was studied in duplicate groups of Arctic charr Salvelinus alpinus. A passive integrated transponder (PIT) system enabled continuous monitoring of individual movements between the sites. Both groups synchronized their diel pattern of visit activity to the two feeding sites when food was available. One group showed significant anticipatory visit activity into both feeding sites during the hours before the feed was available, suggesting a time and place learning of resource availability. The anticipatory activity of the other group was, however, less pronounced and only occurred into one of the feeding sites. Individual S. alpinus entered the feeding sites independently and no obvious patterns of leaders and followers were identified. All S. alpinus gained mass and moved between a refuge and the feeding sites. Different strategies of how individual S. alpinus utilized the feeding sites were not correlated with growth.