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.     

Dietary niche expansion and niche shift in native marble trout (Salmo marmoratus) living in sympatry with introduced rainbow trout (Oncorhynchus mykiss)

The non‐native rainbow trout (Oncorhynchus mykiss) has been introduced worldwide for angling purposes and has established self‐reproducing populations in many parts of the world. Introduced rainbow trout often have negative effects on the native salmonid species, ranging from decrease abundance, growth and survival, to their local extinction. Assessing the effects of introduced rainbow trout on the native species is thus crucial to better set up conservation programmes. In this study, we investigated the effects of non‐native rainbow trout on the diet of native marble trout (Salmo marmoratus) living in the Idrijca River (Slovenia). An impassable waterfall separates the stream in two sectors only a few hundred metres apart: a downstream sector (treatment) in which marble trout live in sympatry (MTs) with rainbow trout (RTs) and an upstream sector (control) in which marble trout live in allopatry (MTa). Specifically, we investigated using stable isotopes the effects of rainbow trout on dietary niche, diet composition, body condition, and lipid content of marble trout. We found dietary niche expansion and niche shift in marble trout living in sympatry with rainbow trout. Compared to MTa, MTs had higher piscivory rate and showed higher body condition and prereproduction lipid content. Our results indicate that the presence of rainbow trout did not have negative effects on marble trout diet and condition and that changes in dietary niche of marble trout are likely to be an adaptive response to the presence of rainbow trout, and further research is needed to better understand.     

Dietary plasticity in Arctic charr (Salvelinus alpinus) in response to long‐term environmental change

Corrigan LJ, Winfield IJ, Hoelzel AR, Lucas MC. Dietary plasticity in Arctic charr (Salvelinus alpinus) in response to long‐term environmental change.
Ecology of Freshwater Fish 2011: 20: 5–13. © 2010 John Wiley & Sons A/S Abstract –  In the face of widespread environmental change impacts, there is a need to better understand mechanisms promoting flexibility and resilience of ecosystem components to such change to inform strategies for conservation. Glacial relict species are especially vulnerable to such changes. We investigated the behavioural responses of a native, glacial relict species, Arctic charr (Salvelinus alpinus) to long‐term environmental changes. It was hypothesised that changes in feeding behaviour would occur as a key intermediary to reduction in habitat availability (through climate change and eutrophication) or competitive interactions [with introduced roach (Rutilus rutilus)]. Stomach content analysis was used to assess the diet of 199 charr caught from Windermere, United Kingdom, in the months of March, June, September and November 2003–2007. The results were compared to data from 1940 to 1951 prior to the environmental changes and revealed a marked increase in the contribution of benthic invertebrates in the present diet. Stable isotope analysis confirms the results of the stomach analysis, suggesting that the charr have switched their diet from zooplanktivory towards benthivory. We discuss the possibility that habitat modification and roach population expansion in Windermere have contributed to changes in charr diet. Complementary studies suggest that this diet shift is more likely to be a response to the increasing roach population than to habitat modification; however, further study in Windermere would be required to elucidate the exact mechanism. Long‐term data sets such as these provide information that is useful for determining the directivity of ecological change and the capability of species to respond to change.     

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.     

Stable isotopes and gut content show diet overlap among native and introduced piscivores in a large oligotrophic lake

Abstract – In past dietary studies kokanee Oncorhynchus nerka were prominent in the diet of Pend Oreille Lake's large piscivores: native bull trout Salvelinus confluentus, cutthroat trout O. clarki and northern pikeminnow Ptychocheilus oregonensis, and introduced lake trout S. namaycush and Kamloops rainbow trout O. mykiss gairdneri. However, kokanee have declined to 10–20% of their former abundance. We therefore initiated this study to understand current predation demands on kokanee and diet overlap among piscivores, using gut content samples and analysis of stable nitrogen (δ¹⁵N) and carbon (δ¹³C) isotopes from the lake's fish and invertebrate community. In gut content samples, kokanee were the main prey item of large [i.e., ≥400 mm total length (TL)] bull and lake trout; a conclusion that was affirmed by stable isotope analysis. Rainbow trout >500 mm TL consumed mostly kokanee, thus there was a high degree of diet overlap among large bull, lake and rainbow trout. Small (i.e., <400 mm TL) rainbow and cutthroat trout diets overlapped, and were composed mostly of littoral benthic invertebrates. However, gut content and stable isotope analysis did not accord for 400–500 mm TL rainbow trout, small lake trout, and large cutthroat trout. In these instances, a linear mixing model using stable isotope results predicted kokanee consumption for each species, but no kokanee were identified in rainbow or lake trout gut content. Gut content and stable isotope analysis of native northern pikeminnow indicated a diet of mostly littoral benthic invertebrates at smaller (100–150 mm TL) lengths, with kokanee becoming more prominent in the diet of individuals >300 mm TL. Percent of kokanee in the diet of northern pikeminnow has declined from a prior study; otherwise piscivore diets have apparently remained unchanged. In this study, judgments as to the feeding of some piscvores, based on gut content alone, would be tenuous because of small sample sizes, but stable isotope analysis provided an efficient means for confirming diets.     

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.     

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.     

Stable isotope evaluation of population‐ and individual‐level diet variability in a large,oligotrophic lake with non‐native lake trout

Non‐native piscivores can alter food web dynamics; therefore, evaluating interspecific relationships is vital for conservation and management of ecosystems with introduced fishes. Priest Lake, Idaho, supports a number of introduced species, including lake trout Salvelinus namaycush, brook trout S. fontinalis and opossum shrimp Mysis diluviana. In this study, we used stable isotopes (δ¹³C and δ¹⁵N) to describe the food web structure of Priest Lake and to test hypotheses about apparent patterns in lake trout growth. We found that isotopic niches of species using pelagic‐origin carbon did not overlap with those using more littoral‐origin carbon. Species using more littoral‐origin carbon, such as brook trout and westslope cutthroat trout Oncorhynchus clarki lewisi, exhibited a high degree of isotopic niche overlap and high intrapopulation variability in resource use. Although we hypothesised that lake trout would experience an ontogenetic diet shift, no such patterns were apparent in isotopic signatures. Lake trout growth rates were not associated with patterns in δ¹⁵N, indicating that variation in adult body composition may not be related to adult diet. Understanding trophic relationships at both the individual and species levels provides a more complete understanding of food webs altered by non‐native species.     

Migrations and harvest rates of Arctic charr (Salvelinus alpinus) in a marine protected area

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Lake trout (Salvelinus namaycush) otoliths indicate effects of climate and lake morphology on growth patterns in Arctic lakes

Climate change is occurring rapidly in the Arctic, and an improved understanding of the response of aquatic biota and ecosystems will be important for this data-limited region. Here, we applied biochronology techniques and mixed-effects modelling to assess relationships among growth increments found on lake trout (Salvelinus namaycush) otoliths (N = 49) captured from 13 lakes on the Arctic Coastal Plain of northern Alaska, observed and modelled climate patterns, and individual-level fish and lake characteristics. We found that annual growth varied by year, fish growth slowed significantly as individuals aged, and females grew faster than males. Lake trout had higher growth in flow-through lakes relative to lakes that were perennially or seasonally connected. Annual growth was positively correlated with observed air temperature measurements from a local weather station for the period 1998–2014, but no clear warming trend was evident for this period. Modelled August air temperatures from 1978–2014 predicted lake trout annual growth (root mean squared error = 0.045 mm) and indicated increasing temperatures and annual lake trout growth over the period 1950–2014. This study demonstrated that biochronology techniques can reconstruct recent climate patterns and provide a better understanding of trends in Arctic lake ecosystems under a changing climate.     

Changes in the density and growth of brown trout (Salmo trutta) after intensive removal of sympatric Arctic charr (Salvelinus alpinus) in the sub‐Arctic lake Møkkelandsvatn,Norway

This study tests the basic hypothesis that the removal of charr, Salvelinus alpinus (L.), would cause an increase in both the growth and density of a sympatric trout population, Salmo trutta L. The charr population was characterised by slow‐growing individuals, with a high proportion of mature fish, that is typical for so‐called overpopulated populations. A total of 31,000 charr was removed from the lake in the period 1990–1992, and the density of younger trout (1+, 2+), but not older trout (3+, 4+), increased. The growth of older trout (3+, 4+) increased, but the evidence for similar growth increases of younger trout (1+, 2+) was limited. From 1989 to 1990, the proportion of trout increased from 30 to only 40% of the total catch, but from 1991 to 1994, it was significantly higher (60–80%) than that of charr. Total trout biomass increased to a maximum in 1992 and then decreased so that the biomass of 1994 was nearly similar to that of 1989, that is before the start of the charr removal. Back‐calculated lengths of trout from otoliths showed that 2+ and 3+ trout caught in the pelagic were growing consistently faster over previous years than those caught in the littoral, while this was not the case for the 4+ fish. Therefore, the hypothesis was partially supported; the growth rate of trout increased (age groups 1+ to 4+), while the density of juvenile trout (1+, 2+), but not the older trout (3+, 4+), increased after the removal of charr.     

Family × Environment interactions in the growth and survival of Arctic charr (Salvelinus alpinus) grown in brackish and fresh water

We investigated whether Fraser strain Arctic charr Salvelinus alpinus L. reared in fresh water (FW) differed in survival and growth from full siblings transferred to brackish water (BW) and whether family performance was consistent throughout 1 year of commercial growth. Fish were grown communally (n = 3600) in either FW or BW from May 2008 to May 2009. Mean family survival in BW was moderately correlated with mean family survival in FW (r = 0.342; P = 0.032). Fish which survived until the conclusion of the experiment were significantly heavier in body weight at the baseline assessment than their full‐sibs which died. Family by treatment interactions were detected for body weight, condition factor and specific growth across all measurement dates (May 2008, October 2008 and May 2009) and growth intervals. Despite the varied response of families to treatment, mean family body weight in FW was correlated phenotypically with BW body weight in October 2008 (r = 0.633, P < 0.0001) and marginally correlated in May 2009 (r = 0.289, P = 0.061). These data suggest that body weight in FW and body weight in BW should be analysed as separate, but correlated traits in Arctic charr breeding programs.     

Interspecific competition between restocked 0+ salmon parr (Salmo salar) and native fish species in a large European river

Atlantic salmon populations have experienced strong declines in most large European rivers. In the Loire Basin, France, a major restocking effort has been carried out on the Allier River to reduce the wild population decline. This study aimed at investigating the trophic insertion of hatchery‐reared salmon parr and whether they compete for resources with native fish species. For this purpose, two riffles located in the upstream and downstream parts of the restocked area have been surveyed monthly following the initial 0+ salmon fry release. 0+ parr densities remained high upstream, with autumnal values greater than those recommended to maximise smolt production. In contrast, downstream densities declined to weak values despite a faster increase in size, probably mainly due to summer temperatures rising above the discomfort threshold. By analysing the vulnerability to predation of ingested invertebrate prey, we highlighted the role of habitat constraints in driving most of the different fish species to a likely opportunistic utilisation of available food resources. As a result, species such as stone loaches and barbel juveniles exhibited narrow trophic niches and high overlaps with 0+ parr. Nonetheless, both abundant prey resources and spatial or temporal habitat segregation have limited interspecific competition, enabling competitive coexistence of sympatric species, except for barbel juveniles in certain circumstances. Therefore, our findings suggest that future restocking programmes should carefully consider the thermal and hydraulic conditions regarding further smolt production as well as their general trophic context, according to the potential inconvenience of restocked salmon fry arrival relative to native species.     

Resource partitioning and spatial segregation in native and stocked brown trout, Salmo trutta L., and Arctic charr, Salvelinus alpinus (L.), in a hydroelectric reservoir

Abstract. Habitat use, food and spatial segregation in native and stocked brown trout, Salmo trutta L., and Arctic charr, Salvelinus alpinus (L.), were studied during summer 1989 and 1990 in the hydroelectric reservoir Lake Tunhovdfjorden. There was no difference in habitat use and feeding habits between wild and stocked brown trout. In epibenthic areas brown trout lived chiefly down to 2 Secchi disc units, whereas Arctic charr were most abundant between 1 and 4 Secchi disc units. In pelagic areas the catches were low for both species, and they were chiefly confined to surface waters down to 1 Secchi disc unit. The food segregation between brown trout and Arctic charr was almost complete. Both pelagic and epibenthic Arctic charr fed mainly on cladocerans ( Bosmina longispina and Daphnia galeata ), whereas surface insects of terrestrial origin and Arctic charr were the dominant food items for brown trout. Pelagic Arctic charr were significantly older, larger and more homogeneous in size than epibenthic charr. During calm weather schools of Arctic charr were observed cruising with the dorsal fin above the surface.     

Trophic overlap between non‐native brown bullhead (Ameiurus nebulosus) and native shortfin eel (Anguilla australis) in shallow lakes

We quantified trophic overlap between the invasive, non‐native catfish brown bullhead (Ameiurus nebulosus) and the New Zealand native shortfin eel (Anguilla australis) in four peat and riverine lakes using stable isotope (δ¹³C and δ¹⁵N) and gut content analyses. Across all lakes and fish sizes over the austral spring–summer period, shortfin eel guts were dominated numerically by fish prey (57% occurrence cf 42% in brown bullhead), while Diptera larvae were most commonly encountered in guts of brown bullhead (45% cf 14% in eels). Significant differences in % composition of animal contents in guts were detected between fish species and sampling occasions (n = 4) but not between lakes. In contrast, stable isotope signatures of brown bullhead and shortfin eel did differ significantly between lakes but not between sampling occasions, indicating enduring sources of nutrition despite apparently differing ingestion patterns over time. The R mixing model MixSIAR indicated that shortfins likely assimilated higher proportions of fish prey carbon compared to brown bullheads, which appeared to show greater assimilation of invertebrates, consistent with the results of gut content analyses. Isotopic niche regions, calculated in nicheROVER using probabilistic ellipses, indicated that shortfin eels occupied at least c.60% of brown bullhead trophic niche, which occupied less than 30% of eel trophic niche in all but one lake. These estimates suggest that brown bullhead has higher potential to influence shortfin eel nutrition than vice versa, or that a broad trophic niche occupied by eels provides resilience to the effects of overlapping consumption patterns with invasive omnivores.     

Trophic relationships between a native and a nonnative predator in a system of natural lakes

Abstract – Bull trout, a species of char listed as threatened under the US Endangered Species Act, have been displaced from portions of their historic range following the introduction of nonnative lake trout. It has been suggested that competitive exclusion as a result of trophic overlap between bull trout and lake trout may be the causal mechanism associated with displacement of bull trout. This study used stable isotope data to evaluate trophic relationships among native bull trout, nonnative lake trout and other fishes in seven lakes in Glacier National Park (GNP), Montana. Bull trout and lake trout had greater δ¹⁵N values relative to other fishes among lakes (δ¹⁵N ≥ 3.0‰). Lake trout had greater δ¹⁵N values relative to bull trout (δ¹⁵N = +1.0‰). Bull trout had greater δ¹³C values relative to lake trout in six of the seven lakes examined. Although both bull trout and lake trout had greater δ¹⁵N values relative to other fishes within lakes in GNP, differences in δ¹⁵N and δ¹³C between bull trout and lake trout suggest that they are consuming different prey species or similar prey species in different proportions. Therefore, displacement of bull trout as a direct result of complete overlap in food resource use is not anticipated unless diet shifts occur or food resources become limiting. Additionally, future studies should evaluate food habits to identify important prey species and sources of partial dietary overlap between bull trout and lake trout.     

Patterns of growth and nutrient deposition in lake trout (Salvelinus namaycush), brook trout (Salvelinus fontinalis) and their hybrid, F1 splake (Salvelinus namaycush × Salvelinus fontinalis) as a function of water temperature

Several fish species of the genus Salvelinus are used for stocking freshwaters in North and South America and much of Europe but there is little information about their growth and carcass composition. Lake trout (S. namaycush), brook trout (S. fontinalis) and their hybrid F₁ splake (S. namaycush × S. fontinalis) (initial body weight ca. 2–4 g) were raised at 6.4, 10.6 and 14.9 °C to examine growth and nutrient deposition as a function of water temperature. In all species, weight gain and feed intake increased significantly with water temperature and feed efficiency was significantly lower at 6.4 than at 10.6 and 14.9 °C. In brook trout, Thermal‐unit Growth Coefficient (TGC) growth rate was significantly lower at 6.4 than at 10.6 and 14.9 °C, while in F₁ splake TGC was only lower at 6.4 than 10.6 °C. Expressed in terms of relative composition a significant effect of temperature was observed. In all species, moisture content decreased while crude protein, lipid, ash and energy contents increased with increased temperature. Expressed in absolute terms, however, a significant effect of temperature was not observed. In all species carcass contents increased significantly with increased live body weight and were best described by simple linear equations. Gross energy concentration was significantly affected by both water temperature and body weight. These data indicate that the growth of these species is a function of water temperature and, in absolute terms, carcass composition is mainly a function of body weight and not water temperature. Also, such simple linear equations bode well for modification of existing feed requirement and waste outputs models; improving their applicability to these species.     

Correlation between dietary lipid:protein ratios and plasma growth and thyroid hormone levels in juvenile Arctic charr, Salvelinus alpinus (Linnaeus)

Sexually immature Arctic charr, Salvelinus alpinus (Linnaeus), were fed one of five isoenergetic practical diets of differing lipid:protein ratios (0.98, 0.67, 0.41, 0.26, 0.19) for an 84‐day period to examine the influence of diet composition on growth, and growth hormone (GH) and thyroid hormone physiology. All five diets supported growth at approximately the same rate, but the diet with a lipid:protein ratio of 0.98 had the lowest weight gain and highest food conversion ratios. A GH enzyme‐linked immunosorbent assay (ELISA), developed for use with oncorhynchid fishes, was validated for use with Arctic charr. Plasma GH concentrations were significantly higher in fish fed the diet with a lipid:protein ratio of 0.98, and there were significant direct and inverse correlations between plasma GH levels and dietary lipid and protein content respectively. There were no significant differences in pre‐ and post‐prandial plasma GH concentrations for any group. There were significant post‐prandial elevations of plasma triiodothyronine (T₃) and thyroxine (T₄) for fish fed the lower lipid:protein ratio diets, but there were no differences related to the diets. The results are discussed in terms of GH as a factor in the regulation of lipid and protein homeostasis in fishes.     

Epitheliocystis in lake trout Salvelinus namaycush (Walbaum) is associated with a β‐proteobacteria

Lake trout Salvelinus namaycush (Walbaum) raised for stocking experienced yearly (2011–13) winter epizootics of epitheliocystis. Affected fish were dispersed on the bottom of the tank, had decreased feed and fright response, and mortality often reached 40%. Peak mortality occurred within 3 weeks of the appearance of clinical signs, and outbreaks typically lasted 6 weeks. Affected fish had no gross lesions but histologically had branchial epithelial necrosis and lamellar hyperplasia, with small to large numbers of scattered epithelial cells containing 10‐ to 20‐μm inclusions. A longitudinal study was undertaken of one annual outbreak, and lamellar hyperplasia was most closely associated with mortality. The number of inclusions was statistically greater (P < 0.05) before and during peak mortality, but inclusions were present in low numbers before clinical signs occurred. Results of histochemical staining, immunohistochemistry and transmission electron microscopy supported the presence of a β‐proteobacteria rather than a Chlamydiales bacterium within inclusions. PCR primers to identify Chlamydiales did not give consistent results. However, the use of universal 16S rDNA bacterial primers in conjunction with laser capture microdissection of inclusions demonstrated that a β‐proteobacteria was consistently associated with affected gills and is more likely the cause of the disease in lake trout.