Food composition, habitat use and growth of stocked and native Arctic charr, Salvelinus alpinus, in Lake Muddusjärvi, Finland

Habitat use, growth and food composition of native and stocked Arctic charr, Salvelinus alpinus (L.), were studied in the subarctic Lake Muddusjärvi, northern Finland, to investigate reasons for poor stocking success. Samples were collected with pelagic and epibenthic gill nets. Stocked and native charr occurred in similar epibethic habitats, whereas pelagic habitat was avoided. Native charr grew fast after shifting to piscivory. Growth rate of stocked charr was slow because only a small proportion of stocked fish became piscivorous during the first year after stocking. During the first lake year, stocked charr divided into slow-growing planktivores and fast-growing piscivores. Piscivorous stocked and native charr consumed only whitefish, Coregonus lavaretus (L.), as their prey. Small-sized (<10 cm) whitefish were preferred when shifting to piscivory.     

Hydroacoustic assessment of mono‐ and polymorphic Coregonus density and biomass in subarctic lakes

Density and biomass estimates of pelagic fish are essential to understand food web interactions and ecosystem functioning. We conducted surveys of six subarctic lakes for assessing both mono‐ and polymorphic whitefish Coregonus lavaretus (L.) populations. Monomorphic whitefish lakes were inhabited by a habitat and diet generalist, large sparsely rakered (LSR) morph, whereas polymorphic whitefish lakes had a littoral benthivorous LSR morph, a pelagic planktivorous densely rakered (DR) morph and in two cases a benthivorous small sparsely rakered (SSR) morph inhabiting the profundal zone. In addition, an introduced specialist zooplanktivore, vendace Coregonus albula (L.), inhabited one of the monomorphic lakes. Hydroacoustics was found to be an appropriate method for estimating coregonid densities and biomasses in large and deep polymorphic lakes occupied by the planktivorous DR morph or vendace, but only during dark nights in autumn. The suitability of hydroacoustic assessment for benthivorous LSR and SSR morphs was low, especially in polymorphic whitefish lakes due to their preference for near‐bottom habitat or shallow areas not sampled with hydroacoustics. The pelagic density of DR morph varied from 330 to 1780 fish·ha^?1 and biomass from 1.4 to 13.3 kg·ha^?1 in polymorphic whitefish lakes, whereas corresponding estimates for LSR morph were 10–320 fish·ha^?1 and 0.5–8.4 kg·ha^?1 in monomorphic whitefish lakes. In general, polymorphism tended to increase the density and biomass of whitefish in the pelagic area compared with monomorphic systems.     

Planktivory and diet-overlap of densely rakered whitefish (Coregonus lavaretus (L.)) in a subarctic lake

Abstract –  Three sympatric whitefish ( Coregonus lavaretus (L.)) forms, one being pelagic and two benthic, segregate available habitat and food resources in subarctic Lake Muddusjärvi, northern Finland. Zooplankton availability in the lake, food composition, diet-overlap and growth of densely rakered (DR) whitefish were examined during June to September to explore the reasons for the small individual size of the pelagic form. DR whitefish used zooplankton as main food item and prey selection followed zooplankton species density proportions in the lake. Zooplankton density and water temperature was highest in July–August. The average lengths of Bosmina spp., Daphnia spp., Calanoida and Cyclopoida in DR whitefish stomach were higher than in zooplankton sample during June–September, except Calanoida in June. Diet-overlap between DR whitefish age groups was high at all months indicating intercohort resource competition. DR whitefish reached sexual maturity at 3 years of age and at the length of 12 cm, after which somatic growth almost ceased. Reason for the small average size and slow growth of DR whitefish were connected to high diet-overlap between age groups and early sexual maturation.     

Brown trout (Salmo trutta L.) and Arctic charr (Salvelinus alpinus (L.)) as predators on three sympatric whitefish (Coregonus lavaretus (L.)) forms in the subarctic Lake Muddusjärvi

Abstract  – Brown trout ( Salmo trutta L.) and Arctic charr ( Salvelinus alpinus (L.)) use whitefish ( Coregonus lavaretus (L.)) as their main prey in the subarctic Lake Muddusjärvi. Brown trout dwelled in littoral and pelagic habitat, whereas Arctic charr lived only in epibenthic habitat. Both species shifted to whitefish predation at a length of 20–30 cm. At this size, brown trout fed on larger whitefish than Arctic charr. Whitefish occur in three sympatric forms, differing in their habitat, ecology and morphology. Both the predators preyed primarily upon the small-sized, densely rakered whitefish form (DR), which was the most numerous whitefish form in the lake. DR used both epibenthic and pelagic habitat, whereas two sparsely rakered whitefish forms dwelled (LSR and SSR) only in epibenthic habitat: LSR in littoral and SSR in profundal areas. Sparsely rakered whitefish forms had minor importance in predator diet.     

Recruitment of sympatric vendace (Coregonus albula) and whitefish (C. lavaretus) is affected by different environmental factors

Vendace and whitefish in Lake Osensjøen (boreal south‐east Norway) were studied by means of gillnet test fishing (1976–2013) and hydro acoustic acquisition (1986–2011). Vendace increased in number between 1998 and 2009 while growth and size at maturity decreased. The relative density of whitefish decreased in the pelagic habitat, whereas growth and size remained the same. Both species exhibited varying year‐class strength. Whereas strong year‐classes of both species became less frequent after 1980 than before this, this seemed to change after 2000, especially for vendace. Generalised additive models suggested a strong positive relationship between July/August mean air temperature and year‐class strength of both vendace and whitefish. Whitefish recruitment was also negatively affected by the new regulation regime implemented since 1981, and positively correlated with water level after hatching and by late ice off. The reason for the diverging impacts of environmental factors on the two species may be that vendace spawn in deeper waters that whitefish do. The results indicate that increasing summer temperatures benefit recruitment of both species, whereas low water level and early ice off will harm whitefish recruitment. Both trends are caused by climate warming. It may be speculated that increased density of the specialised plankton feeding vendace may affect the algal community through increased predation on herbivorous zooplankton and potentially affect the trophic state of the lake.     

Trophic ecology of brown trout (Salmo trutta L.) in subarctic lakes

In subarctic lake systems, fish species like brown trout are often important predators, and their niche performance is a key characteristic for understanding trophic interactions and food web functioning at upper trophic levels. Here, we studied summer habitat use and stomach contents of brown trout under both allopatric and sympatric conditions in six subarctic lakes to reveal its trophic role, and population‐ and individual‐level niche plasticity. In allopatry, brown trout mainly used the littoral habitat, but also less commonly used the pelagic zone. In sympatry with stickleback, there was always a considerable habitat overlap between the two species. In contrast, sympatric populations of brown trout and Arctic charr generally revealed a distinct habitat segregation. In the sympatric systems, in general, there was a distinct resource partitioning between the trout and charr, whereas the observed diet overlap between trout and stickleback was much larger. Trout modified their individual dietary specialisation between the littoral and pelagic zone, always being lower in the pelagic. Piscivorous behaviour of trout was only found in sympatric systems, possibly contributing to a competitive advantage of trout over charr and stickleback. Hence, the trophic level of trout was strongly related to the fish community composition, with a higher trophic level in sympatric systems where piscivorous behaviour was frequent. These changes in the trophic level of trout linked with the observed food resource partitioning might be an important mechanism in the ecosystem functioning of subarctic lakes to allow coexistence among sympatric‐living fish species.     

Temperature‐ and light‐dependent ratio of energy gain to metabolic costs explains spatial and temporal habitat use of zooplanktivorous fish

Understanding the forces that drive habitat selection of species in communities is important in both ecology and evolution. In nature, species face variation in competition, predation and physical characters among habitats. Vendace (Coregonus albula (L.)) is a specialised zooplanktivorous fish predominantly using deeper water in lakes during summer, while roach (Rutilus rutilus (L.)) uses mainly the shallow littoral zone as well as the upper layer of the pelagic zone. To understand mechanisms behind habitat use of these species, I first conducted a predation experiment to investigate their sensitivity to predation by perch (Perca fluviatilis L.). Second, I performed a foraging experiment using different temperature and light treatments. I then used metabolic calculations to estimate energetic costs when foraging. I found no difference between species regarding sensitivity to predation. Vendace was the most efficient forager on zooplankton but also swam faster spending more energy compared to roach. Roach had a comparatively high metabolic rate in the lowest temperature, where their foraging efficiency was lowest. The energy gain ratio at 6°C was highest for vendace, while it was lowest for roach. In the highest temperature (18°C) and the lowest light level (1 lux), both species were similar in their energy gain ratio. The relative energy gain ratio provides a mechanism to explain habitat distribution for the two species. An increased understanding of the role of metabolism in combination with biotic interactions and habitat use may help to foresee effects of environmental change for different species.     

Trophic structure of fish assemblages from Mamoré River floodplain lakes (Bolivia)

Abstract –  The fish assemblage of the floodplain of the Mamoré River (Bolivia) was estimated in eight lakes, corresponding to four habitat types, situated on an environmental gradient related to the river distance: lakes situated near the river, in the forested floodplain, at the floodplain edge and lakes isolated in the savanna. This paper documents the diet of 71 fish species (among the 140 recorded) and compares the taxonomic and trophic structure of fish assemblages between four lake types. The diet analysis was conducted to determine five trophic guilds: algivores/iliophages, herbivores, zooplanktivores, invertivores and piscivores. The taxonomic and trophic structures of the fish assemblages were not similar in the different lake types of the Mamoré River. The trophic structure of assemblages showed a coarse pattern of dominance of algivores/iliophages and invertivores, but different situations were observed in relative abundance of the trophic groups in relation to the spatial position of the lakes (except for piscivores). Lakes close to the river appeared more favourable to the microphages (algivores/iliophages, zooplanktivores) although remote lakes appeared more favourable to the macrophages (invertivores, herbivores). These results support the general idea that fish distribution follows a pattern linked to the ecology of the species, and related to environmental characteristics of the lakes.     

Simulated carrying capacities of fish in Norwegian fjords

The influence of oceanographic and meteorological conditions and topography on the carrying capacity of organisms in coastal areas of western Norway is investigated by field studies and dynamic modelling. Published data on Calanus finmarchicus , the dominant species in the zooplankton biomass of west Norwegian coastal waters, demonstrate a strong gradient, with high densities at the outer coastal areas and lower densities at the innermost localities. This gradient spans several orders of magnitude. A similar tendency is found in the data from pelagic trawling and from beach seine catches, and simulation modelling indicates that fish production depends on zooplankton availability. According to the model, the carrying capacity level of fish production is high in the outer areas and decreases with distance from the outer coast according to the decrease in advected biomass of zooplankton. The simulations indicate that while growth prospects for individuals at lower trophic levels vary with the distance from the coast, potential for growth seems largely unaffected by the distance from the coast for the sublittoral piscivores (e.g. cod). This indicates that, for immigration of a new sublittoral piscivore to a habitat, it does not matter whether the habitat is situated close to the outer coast or far from it. A consequence for stock enhancement may be that a small-scale release of sublittoral piscivores will be equally productive in most coastal areas, while the outcome of a large-scale release will be strongly dependent on distances from the coast. The outcome may be high, but variable, in the outer areas and low, but probably more constant in the innermost areas, provided that releases of juvenile, sublittoral, piscivorous fish do not result in a fish stock density in excess of the carrying capacity level.     

Does habitat occupancy by lake trout and lake whitefish in large lakes match published thermal habitat envelopes?

Habitat occupancy patterns of lake trout (Salvelinus namaycush) and lake whitefish (Coregonus clupeaformis) in two large Canadian Shield lakes were modelled based on detections of fish from repeated depth‐stratified surveys over several summers. Lake trout and lake whitefish consistently occupied sites outside traditional thermal envelopes and were not detected at some sites within these ranges. This included the metalimnion and shallow epilimnion for lake trout and lake whitefish in Lake Opeongo. Physical habitat covariates were not important in defining lake trout habitat in both lakes. Physical habitat as represented by the hardness/softness gradient based on acoustic substrate surveys was important for lake whitefish in Lake Opeongo but not in Smoke Lake. In addition, thermal envelopes for lake whitefish differed between the lakes possibly because of differences in substrate slope. The wash zone of lakes, where the thermocline contacts the substrate, appears to be a physical habitat feature for lake whitefish in some lakes. Lake whitefish also exhibited diurnal activity behaviour that was reflected through greater detection rates in the morning versus the afternoon. By accounting for imperfect detection, true estimated overall occupancy of lake trout and lake whitefish increased 0.15–0.30 over naïve occupancy. Thermal habitat envelopes for lake trout and lake whitefish are wider than previously thought. Lake trout occupied a consistent thermal habitat envelope while lake whitefish varied between lakes likely because of lake specific differences in basin morphology and wash zone.     

Impaired littoral energy pathways cause a shift to pelagic resources by fish in recovering lake food webs

Fish communities may increase in biomass and productivity due to energy subsidies from the littoral invertebrate community. In lakes recovering from acidification and metal contamination, such as those in Sudbury, Ontario, Canada, impaired benthic invertebrate communities (i.e., low diversity with higher abundance of small‐bodied taxa) allowed a critical test of the role of these littoral pathways on fish diet. We compared fish abundance, diversity, diet and biomass in eight recovering and eight reference lakes and related availability of the main littoral and pelagic invertebrate groups to fish diet regime using stable isotope analysis. A Bayesian mixing model (MixSIR) was used to estimate diet likelihood, and convex hull analysis was used to estimate trophic niche space of fish communities. Fish biomass did not differ between impaired and reference lakes despite substantial differences in potential diet. Fish depended strongly on littoral benthos in the reference lakes but consumed more pelagic food in the impaired lakes. The trophic niche of the focal, most common fish species (i.e., yellow perch, smallmouth bass, pumpkinseed and brown bullhead) was larger in the impaired lakes. We attributed these differences to low diversity at the highest trophic levels of fish communities in the impaired lakes as well as to depauperate benthic invertebrate communities. In contrast to the food webs of most temperate lakes, fish in impaired lakes preyed less on littoral invertebrates yet still managed to maintain a reference lake level of biomass standing crop by relying more on pelagic resources – macro zooplankton such as Chaoborus.     

Trophic niche segregation among native whitefish and invasive vendace in a north Norwegian lake system

Introductions and invasions of non-native species alter nutrient cycling and trophic dynamics resulting in significant ecological disturbance. Stable isotope data were used to test for evidence of invader-induced trophic niche differences in a north Norwegian lake system differentially dominated by native European whitefish (Coregonus lavaretus) morphotypes and invasive vendace (Coregonus albula). Aspects of both realised trophic niche position and trophic niche width were affected by the invader, with the effects varying by whitefish morphotype. Densely rakered pelagic whitefish demonstrated a relatively lower reliance on pelagic resources in the presence of the invader and the isotopic niche size was relatively larger in conjunction with the broadening of the prey base. Within the benthic-dwelling sparsely rakered whitefish morphotype, the trophic impacts of invading vendace were size-dependent, with larger individuals experiencing niche compression. Smaller sparsely rakered whitefish increased, contrary to our hypothesis, reliance on pelagic-based energy in the face of invasion. Our findings demonstrate that the trophic ecology of invaded systems can differ in multiple and subtle ways that have consequences for community- and ecosystem-level energy flows, which if persisting over time are likely to have implications for the recruitment, growth and reproductive rate of the native fishes.     

Dietary plasticity of Arctic charr (Salvelinus alpinus) facilitates coexistence with competitively superior European whitefish (Coregonus lavaretus)

Abstract – Habitat use and diet of Arctic charr (Salvelinus alpinus) coexisting with European whitefish (Coregonus lavaretus) and grayling (Thymallus thymallus) were studied in one deep and two relatively shallow subarctic lakes in northern Norway. Stomach content and stable isotope analyses revealed clear and temporally stable resource partitioning between the species in all three lakes. Arctic charr had a wide and flexible trophic niche and was the only piscivorous species. In contrast, whitefish and grayling had remarkably stable planktivorous and benthivorous niches, respectively. In the deepest lake, Arctic charr together with grayling mainly utilised littoral benthos, while piscivory was more prevalent in Arctic charr in the two shallower lakes. In one of the shallow lakes, whitefish was apparently relegated to the inferior profundal niche because of dominance of the littoral by grayling. Our results suggest that Arctic charr may not necessarily need an extensive profundal zone as a refuge, but can coexist with whitefish if a third competing fish species like grayling occurs in the littoral habitat or if profitable small prey fish are available. The study demonstrates that strong dietary plasticity of Arctic charr is instrumental in the observed coexistence with the commonly competitively superior whitefish.     

Resource use of crucian carp along a lake productivity gradient is related to body size,predation risk,and resource competition

Generalist fish species can feed on a wide resource spectrum and across trophic levels depending on resource availability and trophic interactions. Crucian carp (Carassius carassius) represents a good candidate species to investigate variation in the trophic ecology of generalist fish as it can be found in highly variable fish communities and its resource use is well documented. In this study, we explored the trophic ecology of crucian carp at the individual and population levels using stable isotope and gut content analysis. We tested if trophic resource use varied according to lake productivity, predation risk, intra- and interspecific competition, or individual fish size. We found that crucian carp resource preference was highly variable among and within lakes. In predator-free lakes, small crucian carp occurred in high densities, showed increased interindividual specialisation, and relied mainly on pelagic zooplankton. In presence of predators, large crucian carp occurred in low densities and included greater proportions of benthic macroinvertebrates in their diet. This shift in resource use was further favoured in productive, shallow lakes where littoral prey were probably abundant. Resource partitioning was an important factor determining crucian carp niche use, as fish had higher trophic position in absence of other cyprinids. Crucian carp showed highly dynamic resource use and food preferences in response to variable environmental conditions. Overlooking complex diet preferences of generalist fish may lead to an oversimplification of freshwater community dynamics.     

Multispecies functional responses reveal reduced predation at high prey densities and varied responses among and within trophic groups

Understanding predator–prey interactions is critical for marine fisheries and ecosystem management as they shape community structure, regulate prey populations and present energy demands critical for community sustainability. We examined multispecies functional responses of 17 fishes (48 predator‐size combinations) spanning piscivores, planktivores and benthivores for the northeast US continental shelf. Similar to previous work, linear relationships between predation and prey density (Holling type I response) were not supported, since model estimates of handling time were greater than zero for the prey considered. Instead, a clear majority of the predators sampled were Holling type III feeders (sigmoidal; prey switching or learning). For piscivores, nearly all responses were Holling type III with the exception of one being Holling type II (hyperbolic; satiation). Planktivores and benthivores exhibited a combination of type IV (feeding confusion at high prey density) and Holling type III responses. The relationships were predator‐ and prey‐dependent, which is counter to assumptions that are often made of trophic groups. Decreased predation at high prey densities (type IV response) present among planktivores and benthivores suggests an overestimation of predation can occur if ignored. This contrasts with fish and squid prey which primarily invoked a Holling type III response. Functional responses are key to modelling trophic interactions for multispecies and ecosystem models. By refining these inputs in a multispecies context with empirical data, we can advance our understanding of whole‐shelf ecology and improve decision‐making tools for resource management.     

Fishing down or fishing up in Chinese freshwater lakes

Changes in mean trophic level (MTL) of catches have been widely used to reflect the impact of industrial fisheries on aquatic ecosystems because this measure represents the relative abundance of fished species across the trophic level spectrum. In this study, fisheries data from six important freshwater lakes at the middle‐lower Yangtze River and Huaihe River reach of Southern China from 1949 to 2009 were used to evaluate changes in catch MTL. After fishery markets opened at 1985, fish catches increased significantly in all the lakes. Lakes Poyang and Dongting, which were dominated by omnivores and connected to the Yangtze River, showed no significant change in catch MTL before and after 1985. Catch MTL in lakes Taihu and Hongze increased significantly due to an increase in the proportion of pelagic zooplanktivorous. Catches in Lake Chaohu were dominated by zooplankton‐feeding lake anchovy, Coilia ectenes Temminck & Schlegel and icefish, Neosalanx taihuensis Chen, while Lake Donghu was dominated by phytoplanktivorous carps. Due to low biodiversity, catch MTL of these two lakes showed no significant change before and after 1985. Both fisheries‐based and human activities‐based drivers influenced the structure and catch MTL of fisheries in Chinese freshwater lakes.     

Foraging specialisms, prey size and life-history patterns: a test of predictions using sympatric polymorphic Arctic charr (Salvelinus alpinus)

Abstract –  We use arguments based on optimal foraging theory to predict body size constraints and the consequences of these on a range of life-history traits in three trophic specialist morphs of Arctic charr, Salvelinus alpinus , living in sympatry in Loch Rannoch, Scotland. As predicted, foraging specialists feeding on small prey items with a narrow size range showed evidence of deterministic growth; the ultimate body size of macrobenthos feeders being larger ( L _∞ = 238 mm) than that of planktivores ( L _∞ = 216 mm). In contrast, the piscivorous morph showed no evidence of reaching a maximal body size.
The two size-constrained morphs (benthivores and planktivores) matured earlier and died younger (living for up to 11 and 7 years, respectively, in this study) than did the piscivorous charr which showed continuous growth up to at least 17 years.
The pattern of annual reproductive investment in maturing individuals was complex. Planktivores invested in larger eggs than the other two forms, but benthivores produced a greater number of eggs than planktivores, which in turn produced more than piscivores. Planktivorous males had a greater investment in mean testis weight than the other two forms.
Lifetime reproductive output was the greatest in the benthivorous charr, intermediate in planktivorous and the lowest in the piscivorous charr when measured either as fecundity or as gonadal weight. We conclude that constraints imposed upon foraging specialists by foraging efficiency is a significant driver of body size and ultimately reproductive investment in gape-limited foraging salmonids.     

黄海鱼类功能群及其对浮游动物捕食的季节变化

用2006年9月-2007年8月对黄海中南部进行的5次断面调查,通过胃含物分析、聚类分析、功能群划分以及估算摄食量等方法,研究了黄海鱼类群落功能群组成及其对浮游动物摄食的季节变化。结果表明,黄海鱼类群落包括7个功能群,除初春以外,黄海鱼类群落主要以浮游生物食性功能群为主。春夏季的功能群组成简单,秋冬季的功能群组成变得复杂。黄海鱼类群落摄食的浮游动物种类主要有11种,春季摄食的浮游动物量最大,夏秋季逐渐减少,直至冬季摄食的浮游动物量最少,初春摄食的浮游动物量有所回升。不同季节摄食的浮游动物种类,以及摄食量都有较大差异,仅太平洋磷虾在各个月份的食物中均有出现,其余浮游动物饵料种类均是季节性的。     

Rainbow trout effects on zooplankton in the reproductive area of the critically endangered hooded grebe

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Numerical and behavioural response of cyprinids to the introduction of predatory brook trout in two oligotrophic lakes in northern Ontario

Abstract –  Response of potential prey to the introduction of piscivores in a whole ecosystem is generally inferred from small-scale studies. Using distance sampling, two lakes that were initially without predators and three control lakes where predators were previously present were examined for the response of prey species to the introduction of brook trout ( Salvelinus fontinalis ). Responses monitored included numerical changes in abundance, changes in habitat use patterns, and changes in the mean shoal sizes of potential prey species. No statistical changes in abundance or habitat use patterns were detected, whereas there was a significant increase in the mean shoal sizes of potential prey species in lakes where predators were added. We conclude that, in some natural systems, shoaling may be the first response potential prey initiate to reduce mortality risk.