Arctic Charr Populations Research Articles

Spatiotemporal genetic structure of anadromous Arctic char (Salvelinus alpinus) populations in a region experiencing pronounced climate change

We examined spatio-temporal genetic variation at 53 single nucleotide polymorphisms in anadromous Arctic char populations from Western Greenland, a region experiencing pronounced climate change. The study was based on contemporary and historical samples, the latter represented by DNA extracted from otoliths and scales from the 1950s–1960s. We investigated whether genetic population structure was temporarily stable or unstable, the latter due to relatively small spawning and nursery areas combined with a harsh Arctic environment. Furthermore, in order to evaluate the potential for adaptive responses and local adaptation we estimated effective population size (Ne) and migration rate (m). Temporal stability of genetic population structure was suggested, based on a hierarchical analysis of genetic differentiation showing much higher differentiation among samples from different populations (FCT = 0.091) than among temporal samples from the same populations (FSC = 0.01). This was further supported by a neighbor-joining tree and assignment of individuals that showed high contingency between historical and contemporary samples. Estimates of Ne were high (> 500) in three out of four populations, with a lower estimate in one population potentially reflecting fishing pressure or suboptimal environmental conditions. Estimates of m were in most cases low, ≤ 0.01. Ne and m estimates suggest a potential for adaptive responses and local adaptation. However, long generation time may also cause adaptive responses by microevolution to be unable to track climate change, especially considering the low migration rates that reduce potential evolutionary rescue by gene flow from populations better adapted to the altered environments.

Variation in the magnitude of morphological and dietary differences between individuals among populations of small benthic Arctic charr in relation to ecological factors.

The early stages of intraspecific diversity are important for the evolution of diversification and speciation. Early stages of diversification can be seen in individual specialization, where individuals consume only a portion of the diet of the population as a whole, and how such specialization is related to phenotypic diversity within populations. Here, we study the strength of the relationship between morphological and dietary distances among individuals in eighteen populations of Icelandic small benthic charr. We furthermore studied if the strength of the relationship could be related to variation in local ecological factors these populations inhabit. In all the populations studied, there was a clear relationship between morphological and dietary distances, indicating that fish that had similar morphology were at the same time‐consuming similar food items. Our findings show a systematic variation in the relationship between morphology and diet at early stages of diversification in a highly specialized small benthic charr morph. The results show the importance of fine scale comparisons within populations and furthermore the value that systematic comparisons among populations under parallel evolution can contribute toward our increased understanding of evolutionary and ecological processes.

Smelt (Osmerus eperlanus): Glacial relict, planktivore, predator, competitor, and key prey for the endangered Arctic char in Lake Vättern, southern Sweden

Abstract Smelts are crucial forage prey for many piscivorous fish species and have, as such, been frequently introduced outside their indigenous ranges, a management measure causing controversy, as smelts are also efficient predators and competitors. In Lake Vattern, relict Arctic char, fourhorn sculpin, smelt and seven species of crustaceans form a deep-water food-web, and interact closely as competitors, predators and prey. Here, we analyzed new and archived data back to the late 1960s in order to understand and relate observed life-history shifts in the smelt population, to concurrent adverse life-history changes observed in the endangered Arctic char population. Smelt growth was sigmoidal and the size frequency distribution dominated by fish up to 110 mm and age 0–4, although smelt occasionally reached 250 mm. Smelt diet shifts as the fish grow, from zooplankton, to Mysis relicta, to piscivory for individuals larger than 150 mm. Recruitment commonly demonstrates regular inter-annual pulses of YOY abundance although strong year classes produced during years with higher spring temperature cause post-annual irregularity of year class strength. Young smelt reside in surface water and older fish in deeper water. This ontogenetic shift in distribution depends on temperature occupied, prey availability and risk of inter- and intra-specific predation. Interestingly, even though very small Arctic char feed on smelt, most small Arctic char are found in deep water together with large smelt, and most large Arctic char are found in shallow water with small smelt.

Phenotypic plasticity in the morphology of small benthic Icelandic Arctic charr (Salvelinus alpinus)

AbstractIntraspecific phenotypic diversity is the raw material for evolution, so understanding its origin and maintenance is critically important for conservation of biodiversity. Intraspecific diversity in a trait or a suite of traits can result from genetic diversity and/or phenotypic plasticity. The two are, however, not independent as plasticity has been shown to evolve. In this study, we evaluated the importance of phenotypic plasticity in generating morphological diversity in populations of small benthic Arctic charr in Iceland, using a rearing experiment with contrasting modes of feeding. We also examined the association between phenotypic plasticity in offspring groups generated by the contrasting feeding modes and important ecological variables characterising the natural habitats of the respective populations. Although the level of plasticity could not be related to any of the ecological measurements, clear differences in morphological reaction norms among populations suggest that plasticity is an important aspect of morphological diversity of the charr. It is not clear whether that plasticity is adaptive, but it is notable that reaction norms in all populations have similar reaction to the treatments.

Egg Size Versus Number of Offspring Trade-Off: Female Age Rather Than Size Matters in a Domesticated Arctic Charr Population

Understanding how organisms adjust reproductive allocation trade-offs between offspring size versus number (OSN trade-off) is a central question in evolutionary biology. In organisms with indeterminate growth, changes in OSN according to maternal size or age have been reported in numerous taxa. The relative contribution of age and size remains largely unclear, as they are often highly correlated. In this study, we investigated how females adjust the offspring size versus number trade-off and analyzed the relative contribution of female age and size in a domesticated population of Arctic charr Salvelinus alpinus (Linnaeus, 1758) that exhibit large variation in size within five age classes. Our results show that the reproductive output (i.e. as measured by the clutch mass), was strongly correlated to female mass and age suggesting that the proportion of resources allocated to reproduction do not vary along lifetime. Egg mass and fecundity (egg number) increased with female mass overall. However, within an age class, larger females had higher fecundity but egg mass was poorly related to female mass. At the population level, a positive relationship was observed between fecundity and egg mass but within each class age the relation was negative revealing a OSN trade-off. Overall, our results show that, in our model Arctic charr population, allocation trade-off to reproduction and the way females allocate to egg mass and fecundity is largely determined by their age rather than mass.

Stocking activities for the Arctic charr in Lake Geneva: Genetic effects in space and time.

Artificial stocking practices are widely used by resource managers worldwide, in order to sustain fish populations exploited by both recreational and commercial activities, but their benefits are controversial. Former practices involved exotic strains, although current programs rather consider artificial breeding of local fishes (supportive breeding). Understanding the complex genetic effects of these management strategies is an important challenge with economic and conservation implications, especially in the context of population declines. In this study, we focus on the declining Arctic charr (Salvelinus alpinus) population from Lake Geneva (Switzerland and France), which has initially been restocked with allochtonous fishes in the early eighties, followed by supportive breeding. In this context, we conducted a genetic survey to document the evolution of the genetic diversity and structure throughout the last 50 years, before and after the initiation of hatchery supplementation, using contemporary and historical samples. We show that the introduction of exotic fishes was associated with a genetic bottleneck in the 1980–1990s, a break of Hardy–Weinberg Equilibrium (HWE), a reduction in genetic diversity, an increase in genetic structure among spawning sites, and a change in their genetic composition. Together with better environmental conditions, three decades of subsequent supportive breeding using local fishes allowed to re‐establish HWE and the initial levels of genetic variation. However, current spawning sites have not fully recovered their original genetic composition and were extensively homogenized across the lake. Our study demonstrates the drastic genetic consequences of different restocking tactics in a comprehensive spatiotemporal framework and suggests that genetic alteration by nonlocal stocking may be partly reversible through supportive breeding. We recommend that conservation‐based programs consider local diversity and implement adequate protocols to limit the genetic homogenization of this Arctic charr population.

Biomarker response and hypothalamus-pituitary-interrenal axis functioning in Arctic charr from Bjørnøya (74°30′ N), Norway, with high levels of organohalogenated compounds

The populations of Arctic charr (Salvelinus alpinus) residing in Lake Ellasjøen at Bjørnøya Island in the Norwegian Arctic (74° 30′N, 19° 00′E) possess substantially higher levels of organohalogenated compounds (strongly dominated by polychlorinated biphenyls, PCBs) than conspecifics residing in other, proximate lakes on the island. In the present study we sampled large (<400g), immature charr from Lake Ellasjøen (high PCB levels) and Lake Laksvatn (reference lake, low PCB levels) by hook and line for an immediate blood sampling, and blood and tissue sampling after a 1h confinement stressor. This was done in order to investigate possible effects of pollutants on an acute stress performance in a high-latitude fish species by comparing muscle PCB levels, hepatic cytochrome P4501A (CYP1A) biomarker activation and functioning of the hypothalamus-pituitary-interrenal (HPI) axis between these two populations of Arctic charr. As expected sum PCB muscle levels were 8-fold higher on a wet weigh basis, and 19-fold higher on a lipid weight basis, in charr from Ellasjøen than in charr from Laksvatn. This was accompanied by a 3.5-fold higher liver cyp1a mRNA abundance in the Ellasjøen charr compared to Laksvatn charr. Brain transcript levels encoding glucocorticoid receptor 1 and 2 (GR2) and corticotropin-releasing factor, and pituitary transcript levels encoding GR2 and proopiomelanocortin A1 and A2 were higher in Ellasjøen charr than in Laksvatn charr, while interrenal transcript levels encoding melanocortin 2 receptor and steroidogenic acute regulatory protein were lower. There were no differences in plasma cortisol concentration between the two charr populations immediately after capture and one hour after confinement. The strong biomarker response to OHCs and altered mRNA abundances of key genes related to HPI axis functioning in the Ellasjøen charr suggest endocrine disruptive effects of OHCs in this charr population. Possible ecological implications are not known, but it cannot be excluded that a slower growth rate in Ellasjøen charr compared to Laksvatn charr due to an increased metabolic demand associated with the activation of xenobiotic defense and detoxification systems may have contributed to the lower body mass of Ellasjøen charr compared to Laksvatn charr.

Single nucleotide polymorphism markers for analysis of historical and contemporary samples of Arctic char (Salvelinus alpinus)

Arctic regions are predicted to experience drastic temperature increases in the future, which may impact species negatively by changing their environments and habitats. One such species is the Arctic Char (Salvelinus alpinus), which is an important target for local fisheries and the top predator in many freshwater systems. Here we present 53 SNPs for population genetic inferences in the species. They are tested in contemporary and historical samples and will provide a useful resource for future studies of genetic structure and for genetic monitoring of Arctic char populations.

The physiological costs of prey switching reinforce foraging specialization.

Sympatric speciation is thought to be strongly linked to resource specialization with alternative resource use acting as a fundamental agent driving divergence. However, sympatric speciation through niche expansion is dependent on foraging specialization being consistent over space and time. Standard metabolic rate is the minimal maintenance metabolic rate of an ectotherm in a post-absorptive and inactive state and can constitute a significant portion of an animal's energy budget; thus, standard metabolic rate and growth rate are two measures frequently used as an indication of the physiological performance of individuals. Physiological adaptations to a specific diet may increase the efficiency with which it is utilized, but may have an increased cost associated with switching diets, which may result in a reduced standard metabolic rate and growth rate. In this study, we use the diet specialization often seen in polymorphic Arctic charr (Salvelinus alpinus) populations to study the effects of different prey on standard metabolic rate and growth rate as well as the effects that early prey specialization may have on the ability to process other prey types efficiently. We found a significant effect of prey type on standard metabolic rate and growth rate. Furthermore, we found evidence of diet specialization with all fish maintaining a standard metabolic rate and growth rate lower than expected when fed on a diet different to which they were raised, possibly due to a maladaptation in digestion of alternative prey items. Our results show that early diet specialization may be reinforced by the elevated costs of prey switching, thus promoting the process of resource specialization during the incipient stages of sympatric divergence.

Water level regulation affects niche use of a lake top predator, Arctic charr (Salvelinus alpinus)

AbstractWater level fluctuations are expected to deteriorate the littoral zone in heavily regulated hydropower reservoirs, but there is limited empirical evidence of how food webs and fish populations are affected. We contrasted the size, growth, condition, niche use (i.e., habitat and diet), and parasite infection of allopatric Arctic charr (Salvelinus alpinus) populations in two neighboring and comparable Norwegian mountain lakes. We hypothesized that the presumed abiotic and biotic deterioration of the littoral zone would lead to reduced abundance and growth as well as to increased pelagic niche use and reduced niche width of the charr in the heavily regulated Govdajavri (maximum regulation amplitude 24 m) as compared with the unregulated Cazajavri. Our stable isotope and parasite data showed that charr had a slightly narrower and more pelagic feeding niche in the regulated than in the unregulated lake. The relative abundance of charr was lower in the regulated lake, but no between‐lake differences were observed in charr condition, and the charr grew slightly faster in the regulated than in the unregulated lake. Our study suggests that impaired littoral production can alter food webs in alpine hydropower reservoirs and induce a pelagic niche shift by top predators. These results argue for further investigations of hydropower impacts on lake food webs along with other factors that influence the abundance and niche use of fish, such as intraspecific interactions and compensatory growth, which may partly mask the potential impacts.

Low Annual Fidelity and Early Upstream Migration of Anadromous Arctic Char in a Variable Environment

AbstractThe life histories of anadromous Arctic Char Salvelinus alpinus are complex and vary greatly between populations and environments. Here, we detail key aspects of the physical environment and life history of a population of Arctic Char from Nulahugyuk Creek, Nunavut, Canada, to characterize migration traits in a highly variable environment. Over the course of this migration, creek discharge declined precipitously, forcing Arctic Char to migrate through shallow water with large diel temperature fluctuations (&gt;10°C) and high temperature extremes (&gt;21°C). The downstream migration of adults (&gt;55 cm) began in mid‐June and continued into early July, while the downstream migration of smolts (&lt;30 cm) began in late June and continued until late July. The upstream adult migration began in late June and ended in late July, far earlier than most upstream migrations in the region. There was no appreciable upstream migration of juveniles, and Arctic Char 30 to 55 cm in length were absent from the up‐ and downstream migrations. The average age at first migration was 4 years, and the youngest adult Arctic Char migrating upstream were 8 and 9 years old. The missing size‐ and age‐classes, and the fact that most upstream migrants were near reproductive maturity, indicate that Arctic Char in this system typically leave at a length of 19 cm and an age of 4 years and do not return for 4 to 5 years, when they are ready to reproduce. Anadromous Nulahugyuk Creek Arctic Char appear to employ strategies that limit their exposure to restrictive migratory conditions and facilitate their existence in an otherwise uninhabitable system. Understanding such population‐specific migratory strategies is critical to the management of Arctic Char fisheries, which are comprised of populations with highly diverse life histories, and to our understanding of how these life histories may contribute to the adaptability and persistence of the species as climate change progresses.Received November 24, 2015; accepted March 23, 2016 Published online July 28, 2016

Contrasting levels of strays and contemporary gene flow among anadromous populations of Arctic charr, Salvelinus alpinus (L.), in northern Norway

Arctic charr, Salvelinus alpinus (L.), occur as anadromous, resident, and landlocked populations within their circumpolar distribution. While resident Arctic charr reside in freshwater their entire life, anadromous Arctic charr migrate to sea in the summer due to the increased resource availability, but return to freshwater to overwinter and/or spawn. The migratory behaviour in Arctic charr is generally restricted to the northern range of their distribution area. The objective of this study was to estimate the amount of straying and contemporary gene flow among anadromous Arctic charr from eight river systems in northern Norway to illuminate the role of straying for the contemporary genetic stability of the populations. The results revealed that Arctic charr frequently stray among watercourses, especially among those in close geographical proximity, although strays were found nearly 500 km from their river of origin. While straying was revealed among six river systems, the results also revealed limited or no contemporary gene flow among the nine populations identified in the eight sampled river systems. The study also shows that Arctic charr may be a potential host for spreading diseases and parasites among watercourses.

Change in relative abundance of Atlantic salmon and Arctic charr in Veidnes River, Northern Norway: a possible effect of climate change?

Temperature changes affect salmonids across a hierarchy of spatial and temporal scales, so that shifts in thermal river regimes may influence interspecific interactions in sympatric species. In Northern Norway, anadromous populations of Arctic charr (Salvelinus alpinus) and Atlantic salmon (Salmo salar) coexist, but in recent years landings of Arctic charr have decreased, while those of Atlantic salmon have been stable or even increased. Here we studied relative abundance, habitat use and growth rate of sympatric stream-living juveniles of both species in Veidnes River, where they are the only fish species present. In 1998/2000, juvenile Arctic charr dominated, especially in the upper and colder part of the river. In 2010, however, Atlantic salmon juveniles were now prominent in all habitat types, whereas nearly all Arctic charr were captured in slow-flowing water near the river bank. Summer air temperature has increased in the region during the last decade. Positive correlations between summer temperatures and back-calculated growth rates were documented in both species, but the growth response was significantly higher in Atlantic salmon. Accordingly, we suggest that juvenile Atlantic salmon may benefit from a warmer climate in northernmost Norway, at the expense of the more cold-water-adapted Arctic charr.

Divergence and parallelism of parasite infections in Arctic charr morphs from deep and shallow lake habitats

Parasite communities can show large differences in species composition between sympatric host species. Here, we assessed how divergent resource use of polymorphic populations of Arctic charr from two environmentally similar sub-arctic lakes was related to parasite community composition. Large similarities in parasite infections were found for both the reproductively isolated littoral-spawning omnivore LO-morph and the profundal-spawning benthivore PB-morph, reflecting cross-lake similarities in resource use. Furthermore, whereas the PB-morph had the lowest richness and abundance of parasite species in both lakes, a third morph, the profundal-spawning piscivorous PP-morph (one lake only), had the highest parasite richness and accumulated parasites by preying upon fish. These results highlight that host ecology and abiotic factors are important for structuring parasite communities. The dissimilarities in the local parasite species pool between the lakes suggest that stochastic events as well as transmission abilities are involved in structuring the parasite communities. The parallel divergent parasite infections between morphs may form an arena for parasite-mediated selection promoting the incipient speciation process in concert with other factors. The deep-water PB-morphs may avoid parasites by specialising in parasite-poor but low-productive habitats, while the piscivorous PP-morph aggregates infections of detrimental parasites through specialising on energetically rich but generally more heavily parasitised prey.

Contrasting life-history strategies facilitated by cannibalism in a stunted Arctic charr population

Life-history tradeoffs between energy investments in reproduction versus somatic growth may be highly important for fish populations suffering from food limitations. This study addresses life-history tradeoffs in a stunted Arctic charr population from a subarctic lake sampled annually over a 12-year period. The vast majority of charr matured early, grew slowly toward average adult sizes of 13–14 cm, and had a short longevity with few fish reaching ages older than 6 years. Some gender differences in life-history strategies related to sexual maturation were revealed, likely due to energetic constraints from high cost of egg production. Some charr followed a highly different growth trajectory, growing rapidly and attaining lengths >40 cm. These fast-growing individuals matured later and reached higher ages than the stunted fish. Hence, there was a distinct tradeoff between early versus late sexual maturation, the former strategy resulting in short generation time enhancing the survival up to first spawning and the latter being related to a dietary shift to cannibalism resulting in increased growth and body size, and reproduction at a postponed maturation age. This dual pattern was sustained over the 12-year study period, apparently reflecting two contrasting stable strategies.

Morphological, ecological and behavioural differentiation of sympatric profundal and pelagic Arctic charr (Salvelinus alpinus) in Loch Dughaill Scotland

Phenotypic variation in populations of fishes that inhabit postglacial lakes is often associated with trophic specialisations. A common sympatric foraging divergence seen in Arctic charr is into either plankton or littoral-zoobenthos feeding specialisms. In this study, we report a sympatric polymorphic Arctic charr population which is not centred on this divergence but instead manifests as a plankton (pelagic)—profundal zoobenthos foraging specialisms. The head shape of profundal fish was round and robust, the body thick set and pectoral fins long and wide. In contrast, the head of pelagic fish was pointed and slender, the body fusiform in shape and with short, narrow pectoral fins. There was no difference between profundal and pelagic fish in gill raker number. Body lipid content was significantly higher in pelagic fish as were the number or Diphyllobothrium cysts. The carbon isotope ratio was more heavily depleted in profundal fish. There was no dietary overlap in the prey items recovered from stomach contents of profundal and pelagic fish. We suggest the proximate driver behind the sympatric divergence was the successful exploitation of the profundal zone. The consequences of this have led to the development of adaptations in morphology and behaviour to support and maintain this divergence.

Parallel evolution of profundal Arctic charr morphs in two contrasting fish communities

A population of small-sized Arctic charr living their entire life cycle in deep water was recently found in Lake Skogsfjordvatn, Northern Norway. Their phenotype and life history closely resembles the profundal charr morph in another subarctic Norwegian lake, Fjellfrosvatn. Fjellfrosvatn has one littoral omnivorous and one small profundal benthivorous (PB) charr morph coexisting with brown trout, whereas Skogsfjordvatn has migratory salmon, trout, charr and eel populations in addition to threespined sticklebacks, resident trout and three resident charr morphs of which one is a large piscivorous profundal morph. We predicted that the small profundal charr morph in Skogsfjordvatn would have a similar trophic niche to the PB-morph in Fjellfrosvatn and tested this by using stomach content and stable isotope (δ13C and δ15N) analyses. Profundal benthic invertebrates, mainly chironomid larvae, Pisidium sp. mussels and small crustaceans, dominated the diets, and the δ13C and δ15N values indicated almost identical long-term trophic niches of the PB-morphs in both lakes. The prediction was therefore supported. This seems to indicate that the small profundal Arctic charr morphs in these lakes have evolved in parallel, in nearly identical and stable habitat and dietary niches in spite of likely different ecological interactions within the contrasting fish communities.

How to contain a tapeworm epidemic-testing the efficiency of different catch methods to reduce the translocated final hostEsox luciusin an alpine lake

Half a century after introducing the final host Northern pike (Esox lucius L. 1758) into Lake Grundlsee in Austria, a sudden massive infection with the pike tapeworm Triaenophorus crassus Forel 1868 was noticed in the population of the intermediate host Arctic charr (Salvelinus umbla (L. 1758)) and led to the closure of this famous fishery. In an effort to contain the epidemic, a total of 1163 pike were removed over a period of three years (2008–2010). The efficiencies of gill and fyke netting as well as electro and spear fishing were assessed. All four methods were costly with EUR 37–371 to catch a single fish. During intensive fishing, catch per unit effort and fish lengths of pike declined. Fyke netting coupled with electro fishing turned out to be the most practicable strategy to reduce the pike stock. After another three years, prevalence of T. crassus in Arctic charr could be reduced by 40% in 2013. However, it will be impossible to completely eradicate pike and its parasite in the lake.

Genetic Variability and Structuring of Arctic Charr (Salvelinus alpinus) Populations in Northern Fennoscandia.

Variation in presumably neutral genetic markers can inform us about evolvability, historical effective population sizes and phylogeographic history of contemporary populations. We studied genetic variability in 15 microsatellite loci in six native landlocked Arctic charr (Salvelinus alpinus) populations in northern Fennoscandia, where this species is considered near threatened. We discovered that all populations were genetically highly (mean F ST ≈ 0.26) differentiated and isolated from each other. Evidence was found for historical, but not for recent population size bottlenecks. Estimates of contemporary effective population size (N e) ranged from seven to 228 and were significantly correlated with those of historical N e but not with lake size. A census size (N C) was estimated to be approximately 300 individuals in a pond (0.14 ha), which exhibited the smallest N e (i.e. N e/N C = 0.02). Genetic variability in this pond and a connected lake is severely reduced, and both genetic and empirical estimates of migration rates indicate a lack of gene flow between them. Hence, albeit currently thriving, some northern Fennoscandian populations appear to be vulnerable to further loss of genetic variability and are likely to have limited capacity to adapt if selection pressures change.

Recorded bimodal length frequency distributions of Arctic charr, Salvelinus alpinus (L.), and brown trout, Salmo trutta L.: an effect of both population structure and sampling bias

The length distributions of gillnet catches obtained by fishing on unexploited or slightly exploited Arctic charr, Salvelinus alpinus (L.), and brown trout, Salmo trutta L., populations are frequently bimodal. To test the hypothesis that these length distributions may not only be due to sampling bias, but also an effect of population structure, fleets of gillnets were used in sampling from allopatric populations of Arctic charr and brown trout, where the population numbers had been estimated by mark–recapture experiments. In both populations, the length distributions of the populations and the corresponding gillnet catches appeared as bimodal. The second mode of the length frequency distributions consisted of individuals from many age classes accumulated within a restricted length range, which is in turn a consequence of stagnation or low annual growth in old fish. Our results show that the gillnet catches by the use of fleets of gillnets described the overall bimodal length frequency distributions of the populations, although the fraction of larger fish may be significantly overestimated due to a strong increase in gillnet catchability by increasing fish size.