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.     

Temperature–growth patterns of individually tagged anadromous Arctic charr Salvelinus alpinus in Ungava and Labrador,Canada

Individual measurements of annual, or within‐season growth were determined from tag‐recaptured Arctic charr and examined in relation to summer sea surface temperatures and within‐season capture timing in the Ungava and Labrador regions of Eastern Canada. Differences between two years of growth (2010–2011) were significant for Ungava Bay Arctic charr, with growth being higher in the warmer year. Growth of Labrador Arctic charr did not vary significantly among years (1982–1985). Regional comparisons demonstrated that Ungava Arctic charr had significantly higher annual growth rates and experienced warmer temperatures than Labrador Arctic charr. The higher annual growth of Ungava Bay Arctic charr was attributed to the high sea surface temperatures experienced in 2010–2011 and the localised differences in nearshore productivity as compared to Labrador. Within‐season growth rates of Labrador Arctic charr peaked in June, declined towards August and were negatively correlated with the length of time spent at sea and mean experienced sea surface temperatures. A quadratic model relating growth rate to temperature best explained the pattern of within‐season growth. Collectively, results suggest that increases in water temperature may have profound consequences for Arctic charr growth in the Canadian sub‐Arctic, depending on the responses of local marine productivity to those same temperature increases.     

Latitudinal variation in growth and otolith‐inferred field metabolic rates of Canadian young‐of‐the‐year Arctic charr

Countergradient variation (CGV) is defined as genetic variation that counteracts the negative influences of the physical environment, minimising phenotypic variability along an environmental gradient. CGV is thought to have relevance in predicting the response of organisms to climate variability and change. To test the hypothesis that growth rate increased with latitude, consistent with CGV, young‐of‐the‐year (YOY) Arctic charr, Salvelinus alpinus, were examined along a ~27° latitudinal gradient in central and eastern Canada. Growth rates were estimated from fork lengths standardised by the thermal opportunity for growth based on experienced water temperatures derived using otolith oxygen stable isotopes. Results demonstrated patterns consistent with CGV, where northern populations demonstrated faster growth rates. A secondary aim was to test for similar geographical patterns in otolith‐inferred metabolic rates, which reflect the energetic costs of standard metabolic rate (SMR) and other processes such as feeding, locomotion, thermoregulation, reproduction and growth. Results demonstrated a significant, positive relationship between otolith‐inferred metabolic rate and latitude, which may reflect an increase in one, or a combination, of the above‐noted physiological processes. The similar latitudinal pattern in growth and otolith‐inferred metabolic rates suggests greater intake of food per unit of time by northern fish. The phenotypic variation in physiological traits observed here demonstrates the significant adaptability of Arctic charr to different thermal regimes with different growing season lengths. Determining the relative contributions of phenotypic plasticity and genetic variation to the observed latitudinal variation will be critical to predicting the responses of Arctic charr to climate change more accurately.     

Is the winter period a severe bottleneck of anadromous riverine Arctic charr parr?

Abstract –  Winter growth and survival of wild individually tagged juvenile Arctic charr (1+ age) from a riverine anadromous stock, were studied in a small ice-covered (∼6 months) ground water brook (temperature ∼1 °C) connected to Skibotnelva in subarctic northern Norway. The overall winter survival was estimated to be 68% from late October 2005 to early May 2006. The recaptured charr were not significantly different in initial length or weight compared with the nonrecaptured fish suggesting low size-dependent mortality. The majority (98%) of the recaptured charr showed significant increase in size, with a mean increment of 62% from their initial bodyweight and 12% of the charr parr had more than doubled their weight. The mean specific growth rate was calculated to be slightly lower (0.27) than estimated values from a growth model (∼0.35). In addition, the condition factor increased significantly during the field experiment. These results are the first individual growth data on riverine anadromous Arctic charr parr under natural winter conditions, and indicate that charr can grow relatively fast during periods with low temperature and also that ground water brooks can be good over-wintering habitat for juvenile Arctic charr. These results suggest that the winter period is perhaps a less severe bottleneck than previously recognised for the cold-adapted Arctic charr.     

Determining the consistency of thermal habitat segregation within and among Arctic charr morphotypes in Gander Lake,Newfoundland

Abstract – Otolith carbon and oxygen isotope data obtained from distinct genetic and ecological groups of lacustrine Arctic charr, Salvelinus alpinus L., from Gander Lake, Newfoundland, were used to examine hypotheses regarding the consistency of differential habitat use among the groups. Results indicated thermal habitat separation by group, with small ‘pale’ individuals consistently remaining in cooler profundal habitats and larger ‘dark’ individuals more frequently occupying warmer upper water column habitats. Theoretical measures of resource separation and competition indicated lower thermal habitat overlap among the forms and greater within‐form competition. Depth at capture data indicated more varied short‐term use of available lake habitats by ‘dark’ form Arctic charr, possibly as a result of cannibalistic foraging on profundal ‘pale’ form fish. Nominal capture depth data only partially explained observed variation in the mean temperature of occupied thermal habitat, suggesting that capture depth can only be used as a rough index of thermal habitat use. Provided that sufficient thermal gradients exist in the environments being studied, otolith oxygen isotopes provide a useful means of establishing the significance of niche differentiation among individuals.     

Seasonal changes in otolith increment width trajectories and the effect of temperature on the daily growth rate of young sardines

We studied the otolith microstructure and growth of sardine, Sardina pilchardus, in the North Aegean Sea (eastern Mediterranean Sea), using samples of larvae and juveniles that had hatched in winter (November–January) and winter–spring (February–May), respectively. The juveniles had developed during an extended period coinciding with marked pelagic ecosystem changes (from winter, mixed conditions to summer, stratified waters). To examine the relationship between environmental changes and the observed variability in their otolith increment–width trajectories (width‐at‐age), we summarized the shape of trajectories with a four‐parameter set estimated from a growth model fit to each width trajectory. The individual parameter sets were then related to the potential oceanographic conditions that fish experienced during their development, derived from a hydrodynamic–biogeochemical model (POM‐ERSEM), implemented in the sampling area. Substantial seasonal effects were demonstrated on the otolith microstructure (platykurtic versus leptokurtic trajectories in winter‐mixed versus summer‐stratified conditions), which were related to the progressive sea surface warming. In a subsequent step, in order to study the effect of oceanographic conditions on larval and juvenile daily growth rates, a GAM (Generalized Additive Model) analysis of otolith increment widths was carried out, using model‐derived oceanographic parameters and taking into account the ‘inherent otolith growth’, expressed by the explanatory variables ‘previous increment width’ and ‘Age’. Results showed a strong and positive, linear effect of temperature on the growth rate of winter‐caught larvae, whereas in juveniles, which had developed within a wide range of temperatures, an optimum temperature for growth was observed at around 24°C.     

Validation of daily increment formation in otoliths of immature and adult Japanese anchovy <Emphasis Type="Italic">Engraulis japonicus</Emphasis>

In order to validate daily increment formation in otoliths of immature and adult Japanese anchovy Engraulis japonicus, three rearing experiments using chemical marking of otoliths were conducted on adult anchovy in summer 2004 and immature anchovy in summer 2005 and in winter 2006. In the two experiments conducted in summer, the number of otolith microincrements between alizarin complexone (ALC) marks showed that microincrements were formed daily. In the summer 2005 experiment, immature anchovy under conditions of reduced daily food rations also showed daily microincrement formation. Average increment width was 0.9 μm in adults and 1.8–3.1 μm in immature anchovy. In contrast, no clear increments were observed between ALC marks on the otoliths from the experiment in winter 2006, and scanning electron microscope (SEM) observations failed to confirm clear increment formation. We consider that low water temperatures (<13–14°C) restricted otolith growth and lowered the contrast between the discontinuous and the incremental zones of the otolith increments. For age estimation of Japanese anchovy, clear increments wider than about 1 μm in the otolith can be regarded as daily increments. However, daily age estimation of immature and adult anchovy that experience low water temperatures in winter may be difficult due to the obscurity of the increments.     

Using otolith increment widths to infer spatial,temporal and gender variation in the growth of sand whiting Sillago ciliata

Abstract Two methods of otolith increment analysis were used to describe spatial, temporal and gender variation in growth of sand whiting, Sillago ciliata (Cuvier), in four south‐east Australian estuaries. Mean annual standardised otolith increment widths were used as indices of individual lifetime growth rates, while raw otolith increment widths were used to describe variation in growth throughout the life of S. ciliata. Temporal variation in growth was observed at an annual scale, while spatial variation in growth was observed between estuaries. Growth rates increased significantly with decreasing latitude and greater mean sea surface temperatures. A divergence in growth rates between sexes was detected, with females growing faster than males after sexual maturity. This study highlights how otolith increment analyses can: (1) be used to analyse temporal trends in growth from a single sample and (2) provide insight into juvenile growth when samples have an absence of undersized fish.     

Foraging mode variation in three stream‐dwelling salmonid fishes

Despite long‐standing interest in foraging modes as an important element of animal space use, few studies document and compare individual foraging mode differences among species and ecological conditions in the wild. We observed and compared foraging modes of 61 wild Arctic charr, Salvelinus alpinus, 42 brown trout, Salmo trutta, and 50 Atlantic salmon, Salmo salar, in their first growing season over a range of habitats in 10 Icelandic streams. We found that although stream salmonids typically sit‐and‐wait to ambush prey from short distances, Arctic charr were more mobile during prey search and prior to prey attack than Atlantic salmon, whereas brown trout were intermediate. In all three species, individuals that were mobile during search were more likely to be moving when initiating attacks on prey, although the strength and the slope of this relationship differed among species. Arctic charr also differed from salmon and trout as more mobile individuals travelled longer distances during prey pursuits. Finally, coupled with published data from the literature, salmonid foraging mobility (both during search and prior to attack) clearly decreased from still water habitats (e.g., brook charr), to slow‐running waters (e.g., Arctic charr) to fast‐running waters (e.g., Atlantic salmon). Hence, our study suggests that foraging mode of young salmonids can vary distinctly among related species and furthers our understanding of the behavioural mechanisms shaping the geographical distribution of wild salmonids.     

Winter movement activity patterns of anadromous Arctic charr in two Labrador lakes

Anadromous Arctic charr, Salvelinus alpinus, feed in the marine environment for several months during the summer and migrate back to fresh water in late summer to spawn and/or overwinter. While overwintering, anadromous Arctic charr are generally believed to reduce or cease feeding, and they are poorly described in their winter movement activity. This study used telemetry data collected from two locations to describe overwintering movement activity, including interindividual variation. Movement activity declined markedly during the ice‐covered period, suggesting opportunistic maintenance feeding was used as an energy conservation strategy. Fall and spring movement was correlated with daylight hours, and ice break‐up had a significant effect on the timing of outmigration. Movement activity was negatively correlated with body length, with smaller individuals being more active than larger fish. Although general activity patterns were evident, there were significant differences among individuals, particularly during spring immediately prior to lake departure. Lake size and individual differences in metabolic rate may account for some of this variation.     

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 influence of physical watercourse parameters on the degree of anadromy in different lake populations of Arctic charr (Salvelinus alpinus (L.)) in northern Norway

The influence of physical watercourse parameters on the degree of anadromy was studied in 15 lake populations of Arctic charr (Salvelinus alpinus (L.)) in northern Norway. The anadromy of the different populations was scored based on the following criteria; marine parasites. fish length and length at maturity. There were significant negative correlations between the anadromy scores and both the length of the outlet rivers of the different lakes and a migration barrier index for the different watercourses (combining migration distance and water velocity). It is hypothsized that these correlations were due to differences in energy expenditure for charr during upstream migration and/or differences in predation pressure on charr during river migration. Short rivers with relatively high water velocity seem to be the most preferable conditions for the anadromous life-history strategy. There were no correlations between the degree of anadromy and mean river slope or minimum summer water discharge.     

The marine temperature and depth preferences of Arctic charr (Salvelinus alpinus) and sea trout (Salmo trutta), as recorded by data storage tags

Eleven Arctic charr (Salvelinus alpinus) (370–512 mm) and eight sea trout (Salmo trutta) (370–585 mm in length) were tagged externally or internally with depth‐ and temperature‐measuring data‐storage tags (DST) before they were released into the sea in the Alta Fjord in north Norway in June 2002. All sea trout were recaptured after they spent 1–40 days at sea, while all Arctic charr were recaptured after 0.5–33 days at sea. On average, trout preferred water about 0.6 m deeper and 1.3°C warmer than Arctic charr. Arctic charr spent >50% of their time between 0 and 1 m depth, while trout spent >50% of their time between 1 and 2 m depth. Both species spent >90% of their time in water no deeper than 3 m from the water surface. However, sea trout dove more frequently and to greater depths (max. 28 m) than Arctic charr (max. 16 m), and these deep dives were most frequently performed at the end of the sea migration. Arctic charr demonstrated a diel diving pattern, staying on average about 0.5 m deeper between 08:00 hours and about 15:00 hours than during the rest of the 24 h, even though there was continuous daylight during the experiments. When comparing data obtained from the DSTs with temperature measurements within the fjord system, the two species were observed to select different feeding areas during their sea migration, the sea trout choosing the inner and warmer parts of the fjord, in contrast to the Arctic charr that preferred the outer, colder parts of the fjord. The observed differences in migration behaviour between the two species are discussed in relation to species preferences for prey and habitat selection, and their optimal temperatures for growth.     

Feed-growth relationships of Arctic charr transferred from freshwater to saltwater at different seasons

Groups of Arctic charr, Salvelinus alpinus (L.), originating from an anadromous stock, were transferred directly from freshwater to seawater (35 S), from freshwater to brackish water (20 S), or were gradually adapted to seawater over a period of 10 days. A control group was kept in freshwater. Feed intake and growth were then monitored during the following 59 days. Two experiments were carried out, one during winter (December–January) and the other during summer (June–July). Water temperature was maintained at 8 C. All fish had been held under natural photoperiod and temperature conditions from the end of the first feeding period (mid-April) until the experiments were carried out. Rates of growth did not differ between the groups of charr held in fresh- and brackish water, but growth of these groups was better than that of fish reared in seawater. Relationships between individual feed intake (FI) and individual specific growth rates (SGR), within groups were expressed as: ln (SGR+1) = a + ln (FI+1), and regression lines were compared using analysis of covariance. In winter, the regression line for fish transferred directly to seawater had a significantly steeper slope, and a lower elevation, than the lines for fish transferred to brackish water or held in freshwater. Fish transferred gradually to seawater had an intermediate position. In summer, the regressions for all groups were parallel with intercepts in the seawater groups being significantly lower than in the freshwater group. This indicates that the gross feed conversion efficiency (FCE) in Arctic charr was lower in seawater than in freshwater. The results are discussed in relation to digestion, gut function and osmoregulation. yc]Keywords xc]Arctic charr (Salvelinus alpinus (L.)) xc]Feed utilization xc]Salinity xc]Season     

Development and use of an Arctic charr cell line to study antiviral responses at extremely low temperatures

Arctic charr (Salvelinus alpinus) are the northernmost distributed freshwater fish and can grow at water temperatures as low as 0.2 °C. Other teleost species have impaired immune function at temperatures that Arctic charr thrive in, and thus, charr may maintain immune function at these temperatures. In this study, a fibroblastic cell line, named ACBA, derived from the bulbus arteriosus (BA) of Arctic charr was developed for use in immune studies at various temperatures. ACBA has undergone more than forty passages at 18 °C over 3 years, while showing no signs of senescence‐associated β‐galactosidase activity and producing nitric oxide. Remarkably, ACBA cells survived and maintained some mitotic activity even at 1 °C for over 3 months. At these low temperatures, ACBA also continued to produce MH class I proteins. After challenge with poly I:C, only antiviral Mx proteins were induced while MH proteins remained constant. When exposed to live viruses, ACBA was shown to permit viral infection and replication of IPNV, VHSV IVa and CSV at 14 °C. Yet at the preferred temperature of 4 °C, only VHSV IVa was shown to replicate within ACBA. This study provides evidence that Arctic charr cells can maintain immune function while also resisting infection with intracellular pathogens at low temperatures.     

Modelling the life-history variation of Arctic charr

Abstract –  A model based on proximate considerations of life histories of Atlantic salmon, Salmo salar , was examined for its applicability to fit the variation in life-history of wild Arctic charr, Salvelinus alpinus , based on a qualitative assessment of information related to growth and lipid dynamics of Arctic charr. The original salmon model is discussed in context of modifications required to account for added complexities in the life history of Arctic charr in relation to anadromy versus residency. A study from North Norway shows that individual charr that emigrate from the lakes to the sea, maintain a high growth rate in the lake in late summer and early autumn compared with resident fish. Their relatively low lipid level in autumn combined with a high rate of change of lipid during winter was associated with postponement of maturation in the anadromous individuals. Individuals that remain resident in the lake arrested growth in autumn. Their high lipid level in autumn combined with a low rate of change of lipid during winter was associated with maturation the following summer, without emigration from freshwater. Results from this and other related studies show similarities with the model derived from lipid and growth dynamics of Atlantic salmon. The adjusted charr model illustrates possible proximate explanations for the high variation in life-history strategies of Arctic charr. However, the model does not account for the characteristic return migration of immature charr into freshwater several weeks after their entry to the sea. The proximate physiological stimulus for this movement of immature fish is not entirely clear.     

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.     

Annual variation in otolith increment widths of walleye pollock (Gadus chalcogrammus) larvae in Funka Bay,Hokkaido, Japan

To clarify relationships between year‐class strength and larval growth of walleye pollock (Gadus chalcogrammus), and oceanographic conditions in the Pacific stock off Hokkaido and Tohoku, Japan, we undertook conductivity/temperature/depth (CTD) observations and investigated larval densities, larval otolith increment widths and larval prey densities (of copepod nauplii) of the 2008, 2009, 2010 and 2011 yr classes in Funka Bay. Oyashio Coastal Water (OCW) flowed into the bay in late February in 2008, 2010 and 2011, and the mean water temperatures decreased to 1.9–3.1 °C in March. OCW was not observed in 2009, and it was warm in late February (≥3.4 °C). Increment widths of lapillar otoliths during the yolk‐sac stage were wide in 2009 and 2011, medium in 2010 and narrow in 2008. Increment widths during the first‐feeding stage tended to become wider as the hatch month progressed, and the annual variation during the first‐feeding stage was larger than that of the yolk‐sac stage. The densities of the primary food for the larvae were high in 2008 when larval increment widths were narrowest, so the effect of prey abundance on larval growth appeared to be small. The ranking of the larval abundance in March was nearly coincident with that of the increment width during the larval stage. We, therefore, suggest that the larval growth rate is associated with the mortality rate and that the growth–mortality hypothesis may be applicable to walleye pollock in Funka Bay. Feeding success under warm water conditions may be an important factor that contributes towards high growth rates.     

Parallelism in thermal growth response in otoliths and scales of brown trout (Salmo trutta L.) from alpine lakes independent of genetic background

Low density in natural populations of salmonids has predominantly been managed by stocking of non‐native conspecifics. Due partly to domestication, introduced non‐native fish may be maladapted under natural conditions. Interbreeding between introduced and wild individuals may therefore impair local adaptation and potentially population viability. Brown trout (Salmo trutta L.) from three headwaters (with stocked fish) and three interconnected lakes (with native fish) on the Hardangervidda mountain plateau, southern Norway, were tested for differences in thermal effects on scale and otolith growth. Otolith and scale annuli widths from immature brown trout showed positive correlation with mean annual summer temperature for all six sampled populations. In mature individuals, a similar positive thermal correlation was evident for the otoliths only. Interannuli width measurements from scales indicate a halt in somatic growth for brown trout in this alpine environment when reaching ages between 7 and 9 winters, coinciding with age at maturity. Our study indicates that otolith growth follows summer temperature even when individuals do not respond with somatic growth in these populations and that introduced brown trout and introgressed populations have similar thermal growth responses. Due to the continued otolith growth after stagnation in somatic growth and the impact of fluctuations in summer temperature, the utilisation of otolith annuli widths for back calculation of length at age should be treated with caution.     

Feeding,growth and environmental requirements of Arctic charr: a review of aquaculture potential

Interest in the cultivation of Arctic charr arose during the 1970s, and research into charr farming was instigated in the Nordic countries and in Canada. Most work has been conducted on fish from anadromous populations, although land-locked freshwater populations of Arctic charr have also received attention. Research has also been carried out in the British Isles and in the alpine regions of central Europe, where land-locked populations of charr. Small-scale commercial farming is now carried out in several countries of northern Europe and North America, and charr are reared for restocking purposes in a number of countries.Growth of charr is rapid during the early freshwater rearing stages, and quite good rates of growth can be achieved at low water temperatures. Growth may be submaximal if charr are reared in systems designed for other salmonids, and problems may arise when charr are held at low stocking densities. Growth and food conversion can be improved by exposing the fish to water currents, forcing them to swim at moderate speeds. Growth in seawater has been reported as being highly variable, probably as a result of the use of inappropriate rearing techniques and owing to the seasonal changes in the hypo-osmoregulatory ability of the charr.Prospects for aquaculture development and areas requiring further research effort are briefly discussed.