Littoral spawning habitats of three southern Arctic charr (Salvelinus alpinus L.) populations

Abstract Arctic charr populations in southern latitudes are nonmigratory, with all life‐stages limited to freshwater lakes and in‐ or out‐flowing tributaries. Although many of these populations are reported to also spawn in lake littorals, little is known about the physical characteristics of putative spawning grounds. A total of 23 discrete spawning sites within three Irish lakes were located by fyke netting of spawning adults and snorkelling in littoral habitats. Spawning sites were found to be long, narrow strips running parallel to the shore at a maximum depth of 124 cm. Spawning sites were limited to areas of coarse mineral substrate with an adequate (c. 8 cm) depth of clean interstitial spaces. In individual lakes, combined areas of spawning sites made up 0.4–0.7% of available littoral. Egg densities varied considerably between sites (33–900·eggs m^?2) and were significantly correlated with gradient and width of spawning sites. No evidence of redd digging was found. The shallow, localised and restricted nature of spawning grounds makes such populations vulnerable to anthropogenically induced postoviposition changes in surface water level, eutrophication processes such as increased lake sedimentation and elevated nutrient status.     

Stream spawning of Arctic charr in Scotland

Abstract –  Scotland is a stronghold for Arctic charr, with about 200 freshwater, wholly loch-resident populations, most of which have yet to be studied. To date, no anadromous populations or individual sea-run charr, have been reported. In Scotland, most Arctic charr spawn in stillwater, during autumn and early winter (September to January), with only one population spawning in spring. Spawning in running water does occur, but has been regarded as rare. This paper examines the status of stream spawning of charr in Scotland, describing instances from 10 populations, although two of these are now extinct. Most stream-spawning charr migrate relatively short distances to running water and construct redds in gently flowing water. The longest recorded migration occurs in Loch Insh, Speyside, with fish travelling 15 km within the River Spey. Information on the extent of stream spawning is essential to help conserve the rich phenotypic and genetic diversity of our remaining charr populations.     

Assessing the food web impacts of an anadromous Arctic charr introduction to a sub‐Arctic watershed using stable isotopes

Anadromous Arctic charr, Salvelinus alpinus (L.), was introduced to a sub‐Arctic river–lake system near the village of Kujjuuaq, Nunavik, and the stable isotope values and diets of key resident fish species were used to assess changes in feeding patterns. Stable isotope values for most species did not differ significantly between the pre‐ and post‐introduction periods, with observed shifts being within the bounds of expected natural variation. Lake chub, Couesius plumbeus (Agassiz), were the single species to show a difference between study periods, with a small but significant increase in δ¹⁵N. No significant post‐introduction changes were seen in lake trout, Salvelinus namaycush (Walbaum), omnivory or in any of the assessed quantitative food web metrics. Gut contents of major fish species similarly showed significant temporal overlap between the pre‐ and post‐introduction periods, and there was no significant change in species' weight–length relationships. The minor ecological impact was interpreted in relation to the availability of open niches exploitable by ecological generalists such as Arctic charr. The explanation accords with the known habitat and feeding flexibility of Arctic charr and the ecological immaturity of sub‐Arctic lakes known to have driven adaptive variation among Arctic charr. Findings suggest that anadromous Arctic charr may be introduced at moderate densities to other sub‐Arctic watersheds without major negative food web consequences for other resident fish species.     

Seasonal variability in the abundance of Arctic charr (Salvelinus alpinus (L.)) recorded using hydroacoustics in Windermere, UK and its implications for survey design

Abstract –  Hydroacoustic surveys are an important tool for assessing the abundance of Arctic charr ( Salvelinus alpinus (L.)) in lakes, the requirement for which is increasing because of new environmental legislation. Consequently, survey demand is now such that assessments must be undertaken with the minimum of effort compatible with scientific validity. One major complication is seasonal variability in abundance, with which surveyors have attempted to deal by defining somewhat arbitrary seasonal sampling windows. The definition of more objective windows was pursued by analysing variations in the seasonal abundance of Arctic charr in Windermere, UK, recorded using day and night hydroacoustic surveys at monthly intervals from 1991 to 2001. Two alternative sampling windows for night surveys free of significant internal seasonal effects were revealed, i.e., December to September and September to December. It was also found that day surveys gave a useful, although lower, estimate of abundance as compared with night surveys.     

Trophic interactions between introduced lake trout (Salvelinus namaycush) and native Arctic charr (S. alpinus) in a large Fennoscandian subarctic lake

Introduced fishes may have major impacts on community structure and ecosystem function due to competitive and predatory interactions with native species. For example, introduced lake trout (Salvelinus namaycush) has been shown to replace native salmonids and induce major trophic cascades in some North American lakes, but few studies have investigated trophic interactions between lake trout and closely related native Arctic charr (S. alpinus) outside the natural distribution of the former species. We used stomach content and stable isotope analyses to investigate trophic interactions between introduced lake trout and native Arctic charr in large subarctic Lake Inarijärvi in northern Finland. Both salmonids had predominantly piscivorous diets at >280 mm total length and were mainly caught from the deep profundal zone. However, lake trout had a more generalist diet and showed higher reliance on littoral prey fish than Arctic charr, whose diet consisted mainly of pelagic planktivorous coregonids. According to length at age and condition data, lake trout showed slightly faster growth but lower condition than Arctic charr. The results indicate that introduced lake trout may to some extent compete with and prey upon native Arctic charr, but currently have only a minor if any impact on native fishes and food web structure in Inarijärvi. Future monitoring is essential to observe potential changes in trophic interactions between lake trout and Arctic charr in Inarijärvi, as well as in other European lakes where the two salmonids currently coexist.     

Fine‐scale acoustic telemetry reveals unexpected lake trout,Salvelinus namaycush,spawning habitats in northern Lake Huron,North America

Previous studies of lake trout, Salvelinus namaycush, spawning habitat in the Laurentian Great Lakes have used time‐ and labour‐intensive survey methods and have focused on areas with historic observations of spawning aggregations and on habitats prejudged by researchers to be suitable for spawning. As an alternative, we used fine‐scale acoustic telemetry to locate, describe and compare lake trout spawning habitats. Adult lake trout were implanted with acoustic transmitters and tracked during five consecutive spawning seasons in a 19–27 km² region of the Drummond Island Refuge, Lake Huron, using the VEMCO Positioning System. Acoustic telemetry revealed discrete areas of aggregation on at least five reefs in the study area, subsequently confirmed by divers to contain deposited eggs. Notably, several identified spawning sites would likely not have been discovered using traditional methods because either they were too small and obscure to stand out on a bathymetric map or because they did not conform to the conceptual model of spawning habitat held by many biologists. Our most unique observation was egg deposition in gravel and rubble substrates located at the base of and beneath overhanging edges of large boulders. Spawning sites typically comprised <10% of the reef area and were used consistently over the 5‐year study. Evaluation of habitat selection from the perspective of fish behaviour through use of acoustic transmitters offers potential to expand current conceptual models of critical spawning habitat.     

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.     

Recent changes in the spawning grounds of Black Sea anchovy,Engraulis encrasicolus

Towards the end of the 1980s, when the spawning grounds in the northwestern shelf (NWS) of the Black Sea were lingering with the effects of eutrophication and of an exotic invasive ctenophore, a series of basin‐wide international ichthyoplankton surveys pointed out an increase in the anchovies spawning in the southern half of the Black Sea. Later, with the help of international conservation efforts, several key littoral ecosystem components within the anchovy's historical spawning grounds showed signs of recovery. However, the fate of the spawning stock anchovy in the south remained unanswered. In order to present the current situation in the southern Black Sea after two decades, an ichthyoplankton survey adopting the same methodology as previously used was undertaken during the peak spawning season of the Black Sea anchovy (BSa). The survey showed that the density of eggs was by far greater than for any of the surveys conducted previously. A wider geographical distribution of the eggs indicated an increase in the number of vagrants which had drifted away from the known spawning grounds. In contrast, the increased reproductive activity in the south signifies existence of a growing, non‐migrating southern BS stock. This stock seems to utilize the coastal hydrographic features associated with the rim current facilitating escape (loophole) from gelatinous predators such as Mnemiopsis leidyi and Aurelia aurata.     

An experimental test of the genetic component of the ontogenetic habitat shift in Arctic charr (Salvelinus alpinus)

Abstract— Fry of the Arctic charr, Salvelinus alpinus , were experimentally stocked into a small fish-free lake to test the hypothesis that the size-dependent habitat shift from the epibenthic to the pelagic habitat is genetically determined. The charr originated from a nearby lake inhabiting predatory brown trout Salmo trutta. The cohort of stocked charr was investigated for three years. The Arctic charr started to exploit the pelagic habitat in their first summer at a size of 7–9 cm in contrast to about 15 cm in the donor lake. In the next two summers, the pelagic fraction of the cohort increased. The main fraction lived in epibenthic areas, utilizing the same prey as pelagic charr. Water temperature moderated the habitat use of juveniles such that they avoided warm (>16°C) waters and resided in cool, deep areas. The result was consistent with the hypothesis of a tradeoff between feeding benefit and the predation risk producing spatial segregation of Arctic charr and demonstrated that the fish can facultatively respond to predation risk and adjust the size at which they migrate to the pelagic zone to feed on zooplankton.     

Survival, growth, sexual maturation and reproduction of brook charr, Salvelinus fontinalis (Mitchill), Arctic charr, Salvelinus alpinus L., and their hybrids

Survival of hybrids (FA, AF) between brook charr, Salvelinus fontinalis (Mitchill), (FF) and Arctic charr, Salvelinus alpinus L., (AA) were compared at three fish farms. Survival of hybrids and Arctic charr was impaired in warm water. In cold water, brook charr and hybrids showed similar survival percentages until the spawning period. Each spawning period was followed by an increase of mortality, but to a much higher degree in brook charr. Growth was studied in the best (cold water) fish farm. During the first 2 years of rearing, brook charr had the highest relative weight. The weight of the two hybrids approached that of brook charr in the third year because of the drastic decrease of growth rate in brook charr after the spawning period. No differences between genotypes of either females or males could be detected in the eviscerated weights, the dressing percentages, or the gonado-somatic indexes. The two hybrids matured sexually, but their sperm quality was inferior to that of the pure species. Crosses between the 4-year-old breeders from the F1 population (FF, FA, AF and AA) were achieved. Survival from the eyed stage to 6 weeks post-hatching was significantly lower in F2 progeny than in back-crosses or pure species crosses. Even though the survival following spawning periods and the total biomass produced were better in hybrids than in brook charr raised in cold water, we believe the farming of this hybrid should not be carried out. The full development of sexually mature hybrids does not resolve problems linked with precocious sexual maturation in brook charr farming carried out in eastern Canada.     

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.     

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.     

Solar and lunar influences on the spawning activity of Pacific bluefin tuna (Thunnus orientalis) in the south‐western North Pacific spawning ground

Pacific bluefin tuna (Thunnus orientalis) migrate to the south‐western North Pacific spawning grounds (area off Taiwan and the Ryukyu Islands) from mid‐April to early July. Reproductive traits of the species in this area have been studied for some time; however, temporal changes in spawning activity during this season are not well understood. In this study, the spawning fraction (proportion of females with post‐ovulatory follicles to total mature females) in relation to solar calendar date and lunar day was investigated using a generalized linear model. Sampling was conducted almost every day throughout the fishing season at Ishigaki Fishing Port (24°21′N, 124°09′E), southern Japan between 2012 and 2014, and a total of 212 ovaries were histologically observed. Spawning fraction continued to increase from the beginning to the end of the season. This indicates that the peak season of occurrence in the area and the peak of spawning activity did not co‐occur. Spawning fraction also changed with the lunar day and significantly increased around the new moon. Eggs and early larvae hatched around the new moon may have reduced the predation risk during night‐time. These findings improve the current knowledge of reproductive biology and resource evaluation of Pacific bluefin tuna and indicate that spawning activity is not constant, and has two or three peaks in each season.     

Habitat use, size and age structure in sympatric brown trout (Salmo trutta) and Arctic charr (Salvelinus alpinus) stocks: resistance of populations to change following harvest

Abstract– Habitat use and population dynamics in brown trout Salmo trutta and Arctic charr Salvelinus alpinus were studied in an oligotrophic lake over a period of 10 years. Previous studies showed that the species segregated by habitat during summer. While brown trout occupied the surface water down to a depth of 10 m, Arctic charr were found deeper with a maximum occurrence at depth 10–15 m. Following the removal of a large number of intermediate sized fish in 1988–89, habitat segregation between the species broke down and Arctic charr were found in upper waters, while brown trout descended to deeper waters. The following year, both species were most frequently found in surface waters at depths of 0–5 m. During the last four years, the species reestablished their original habitat segregation despite another removal experiment of intermediate-sized fish in 1992–1994. The removal of fish resulted in an increased proportion of large (≥ 25 cm) fish in both species. Furthermore, the charr stock responded by reduced abundance and increased size-at-age. The results revealed plasticity and strong resistance to harvest populations of brown trout and Arctic charr. This is probably due to internal mechanisms of intraspecific competition within each population, which result in differential mortality among size classes.     

Reliance on lakes by salmon,trout and charr (Oncorhynchus,Salmo and Salvelinus): An evaluation of spawning habitats,rearing strategies and trophic polymorphisms

Salmonid fishes may reside within or migrate between stream and lake habitats, or undergo anadromous migrations between freshwater and the ocean. While the degree of anadromy of salmonids has been thoroughly compared, no analogous review has examined the degree of lake use. To assess the extent of reliance on lake habitat in this family, we considered 16 species of salmon, trout and charr from the genera Oncorhynchus, Salmo and Salvelinus, comparing their (a) use of lakes as spawning habitat, (b) rearing strategies in lakes, and (c) occurrence and diversity of lacustrine trophic polymorphism. In identifying the primary life‐history patterns of each species and exploring the lesser‐known lacustrine behaviours, we found that the extent of reliance on lakes exhibits a negative association with the degree of anadromy. Oncorhynchus rely least on lakes, Salmo to an intermediate level and Salvelinus the most, opposite of the general prevalence of anadromy among these genera. Lakes are critical to adfluvial and lake‐resident salmonids, but they also support anadromous and fluvial life histories by providing spawning, rearing, overwintering and/or summer refuge habitat. Adfluviality, although a non‐anadromous life history, consists of similar migration‐related traits and behaviours as anadromy, including the parr–smolt transformation, sex‐biased patterns of migration and residency, and the presence of precocious males. Lakes support life‐history variants, reproductive ecotypes and trophic morphs unique to lacustrine habitat. Therefore, conservation of salmonids is dependent on maintaining the diversity and quality of their habitats, including lakes.     

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.     

The effect of lake morphometry on thermal habitat use and growth in Arctic charr populations: implications for understanding climate‐change impacts

Oxygen stable isotope temperature reconstruction methods were used to estimate mean experienced summer temperatures from growth zones within individual Arctic charr otoliths sampled from lakes with contrasting morphologies but proximate locations. For either lake, otolith‐estimated temperatures were not significantly related to back‐calculated growth. Fish in the smaller lake evidenced an increase in growth with age related to increasing use of cooler thermal habitats, with the use of thermal habitat possibly governed by predation risks. No relationships between age, growth or temperature were observed in the larger lake. Significant negative effects on back‐calculated growth were observed due to increasing air temperatures in the smaller and shallower lake, possibly owing to warmer surface and littoral waters and a limited amount of preferred cool‐water habitat. A similar relationship was not observed in the larger and deeper lake and indicated that resident Arctic charr were not as vulnerable to the impacts of temperature warming, possibly because of better behavioural thermoregulation opportunities in the cooler, deeper lake. Results provide evidence for differing climate‐influenced growth outcomes among proximately located populations, with outcomes likely to depend on the differences among habitats, including lake size and morphometry which may act to influence fish densities in available preferred thermal habitats.     

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

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Evidence for the recent extinction of two Arctic charr Salvelinus alpinus (L.) populations in the West of Ireland

1. This paper highlights the recent extinctions of two native Arctic charr populations in the West of Ireland, one population from the Lough Conn catchment, and a second from the Lough Corrib catchment, respectively. 2. The distribution of charr populations in the two catchments is reviewed. The current distribution information is drawn from recent fish stock surveys (Lough Corrib in 1996, and Lough Conn in 1997 and 1998). Previous records of charr in the literature and unpublished data are described to place the current distribution of charr within these catchments into a historical perspective. 3. The importance of the charr population in Lough Mask (Lough Corrib catchment) is emphasized, as it is one of the last extant examples of charr from a large Irish alkaline lake. 4. Impacts on charr populations in both catchments are discussed and conservation measures proposed. Copyright © 2001 John Wiley & Sons, Ltd.     

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