Charr Populations Research Articles

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.

First report of the mitochondrial DNA sequences of Chukchi Charr (Salvelinus andriashevi, Salmonidae)

Three mitochondrial DNA regions were sequenced in two individuals of the Chukchi charr Salvelinus andriashevi from Lake Istihed (Chukotka): the entire gene sequences of cytochrome b and cytochrome c oxidase-1, and control region. The low level of sequence divergence detected between the sequences suggests that S. andriashevi is probably an isolated population of the Taranetz charr S. taranetzi. The genealogy of mtDNA haplotypes confirms the phylogenetic closeness of S. andriashevi, S. taranetzi, Salvelinus sp. 4 (Lake Nachikinskoe), S. krogiusae (Lake Dal’nee), S. boganidae and S. elgyticus (Lake Elgygytgyn), and S. a. erythrinus (NWT).

Quantifying long-term population growth rates of threatened bull trout: challenges, lessons learned, and opportunities

Temporal symmetry models (TSM) represent advances in the analytical application of mark–recapture data to population status assessments. For a population of char, we employed 10 years of active and passive mark–recapture data to quantify population growth rates using different data sources and analytical approaches. Estimates of adult population growth rate were 1.01 (95% confidence interval = 0.84–1.20) using a temporal symmetry model (λTSM), 0.96 (0.68–1.34) based on logistic regressions of annual snorkel data (λA), and 0.92 (0.77–1.11) from redd counts (λR). Top-performing TSMs included an increasing time trend in recruitment (f) and changes in capture probability (p). There was only a 1% chance the population decreased ≥50%, and a 10% chance it decreased ≥30% (λMCMC; based on Bayesian Markov chain Monte Carlo procedure). Size structure was stable; however, the adult population was dominated by small adults, and over the study period there was a decline in the contribution of large adults to total biomass. Juvenile condition decreased with increasing adult densities. Utilization of these different information sources provided a robust weight-of-evidence approach to identifying population status and potential mechanisms driving changes in population growth rates.

At the forefront: evidence of the applicability of using environmental DNA to quantify the abundance of fish populations in natural lentic waters with additional sampling considerations

Environmental DNA (eDNA) sampling has proven to be a valuable tool for detecting species in aquatic ecosystems. Within this rapidly evolving field, a promising application is the ability to obtain quantitative estimates of relative species abundance based on eDNA concentration rather than traditionally labor-intensive methods. We investigated the relationship between eDNA concentration and Arctic char (Salvelinus alpinus) abundance in five well-studied natural lakes; additionally, we examined the effects of different temporal (e.g., season) and spatial (e.g., depth) scales on eDNA concentration. Concentrations of eDNA were linearly correlated with char population estimates ([Formula: see text] = 0.78) and exponentially correlated with char densities ([Formula: see text] = 0.96 by area; 0.82 by volume). Across lakes, eDNA concentrations were greater and more homogeneous in the water column during mixis; however, when stratified, eDNA concentrations were greater in the hypolimnion. Overall, our findings demonstrate that eDNA techniques can produce effective estimates of relative fish abundance in natural lakes. These findings can guide future studies to improve and expand eDNA methods while informing research and management using rapid and minimally invasive sampling.

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 (>10°C) and high temperature extremes (>21°C). The downstream migration of adults (>55 cm) began in mid‐June and continued into early July, while the downstream migration of smolts (<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

On the origin and phylogenetic position of Arctic charr (Salvelinus alpinus complex, Salmonidae) from Lake Cherechen’ (middle Kolyma River basin): controversial genetic data

Within the Arctic charr complex (Salvelinus alpinus complex), many different populations and forms with disputable origin and systematic status have been described. Some of them, such as the charr from Lake Cherechen’ (middle Kolyma River basin), combine characters of different phylogenetic groups, representing a possible consequence of former hybridization. The data on 32 allozyme loci and on nucleotide sequences of 501-bp fragments of the mtDNA control region as well as of 899-bp fragments of exon 2 of the RAG1 gene were used for the analysis of the origin of Lake Cherechen’ charr and their phylogenetic relationships with other representatives of the Arctic charr lineage. As was shown previously, the dwarf and large forms of charr from this lake are morphologically similar to other charr populations from the upper Kolyma River, but bear the mtDNA haplotype of northern Dolly Varden Salvelinus malma malma, not of Taranets charr Salvelinus alpinus taranetzi. The analysis of the allozymes and RAG1 gene confirms the affinity of the Lake Cherechen’ charr to the Arctic charr lineage, but it is insufficient to unambiguously attribute them to the Eurasian or Taranets group. The presence of mtDNA of northern Dolly Varden in Lake Cherechen’ Arctic charr and the replacement of their native mtDNA are the result of the introgressive hybridization with S. m. malma. An alternative explanation connected with incomplete lineage sorting seems highly improbable. Our study confirms a postglacial secondary contact of the representatives of different phylogenetic groups of the S. alpinus-S. malma species complex in the Kolyma basin and in the area from Taimyr to Chukotka. It also indicates the need for more thorough analysis of the morphological and genetic diversity of charr from this region as well as caution in taxonomic decisions.

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.

Population structure and conservation genetics of anadromous white-spotted char (Salvelinus leucomaenis) on Hokkaido Island: Detection of isolation-by-distance

To increase the quantity of anadromous and freshwater fish resources and conserve natural populations effectively, it is important to identify conservation units and population structures, as well as to ensure sufficient genetic variability within populations. Here, we used 12 microsatellite loci to evaluate the population structure, effective population size, and bottlenecks in seven anadromous white-spotted char (Salvelinus leucomaenis leucomaenis) populations inhabiting the rivers of Hokkaido Island, Japan. Low migration rates were detected among populations, with significant genetic differentiation being observed, suggesting high homing rates. In addition, isolation-by-distance was observed among the evaluated populations, indicating that the populations in this region are at equilibrium between migration and drift. We identified a genetic bottleneck footprint in all seven of the analyzed white-spotted char populations by using the M ratio test. In contrast, heterozygote excess tests showed that all seven populations showed no signatures of population decline. This discrepancy may have been caused by differences in statistical power among tests. Alternatively, this discrepancy may be consistent with a strong founder effect during the late Pleistocene, followed by a subsequent low migration rate among populations. In conclusion, future conservation genetic management strategies should ensure that anadromous white-spotted char populations successfully exhibit homing behavior in the rivers of Hokkaido Island, Japan.

Subcellular distribution of trace elements and liver histology of landlocked Arctic char (Salvelinus alpinus) sampled along a mercury contamination gradient

We sampled landlocked Arctic char (Salvelinus alpinus) from four lakes (Small, 9-Mile, North, Amituk) in the Canadian High Arctic that span a gradient of mercury contamination. Metals (Hg, Se, Tl, and Fe) were measured in char tissues to determine their relationships with health indices (relative condition factor and hepatosomatic index), stable nitrogen isotope ratios, and liver histology. A subcellular partitioning procedure was employed to determine how metals were distributed between potentially sensitive and detoxified compartments of Arctic char livers from a low- and high-mercury lake (Small Lake and Amituk Lake, respectively). Differences in health indices and metal concentrations among char populations were likely related to differences in feeding ecology. Concentrations of Hg, Se, and Tl were highest in the livers of Amituk char, whereas concentrations of Fe were highest in Small and 9-Mile char. At the subcellular level we found that although Amituk char had higher concentrations of Tl in whole liver than Small Lake char, they maintained a greater proportion of this metal in detoxified fractions, suggesting an attempt at detoxification. Mercury was found mainly in potentially sensitive fractions of both Small and Amituk Lake char, indicating that Arctic char are not effectively detoxifying this metal. Histological changes in char livers, mainly in the form of melano-macrophage aggregates and hepatic fibrosis, could be linked to the concentrations and subcellular distributions of essential or non-essential metals.

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.

Socially induced reproductive synchrony in a salmonid: an approximate Bayesian computation approach

Reproductive synchrony is a widespread phenomenon found in many taxa, including plants and corals. However, compared with synchrony caused by environmental cues, knowledge of socially induced reproductive synchrony is limited, partly due to the difficulty of experimentally manipulating and/or making detailed behavioral observations of populations in the wild. In this study, we developed a novel modeling framework combining an individual-based model, a hierarchical Bayesian model, and an approximate Bayesian computation (ABC) to elucidate socially induced reproductive synchrony. This method was applied to time-series redd (i.e., spawning nests) count data in 30 wild populations of stream-dwelling Dolly Varden charr. The model with reproductive synchrony explained all the redd count data, whereas the null model, which did not include the synchrony, failed to reproduce the observed data in several populations. In addition, our models suggest that Dolly Varden should be able to adjust spawning by up to a week following other females to produce synchrony. No significant correlation was observed between reproductive timing and environmental factors, suggesting that the major cue for the synchrony was social rather than environmental. The presence of reproductive synchrony within but not among local populations suggests that predator satiation is not the main driver of the synchrony; rather, other mechanisms must exist in the Dolly Varden, such as induced monogamy or polygamy, or avoidance of nest superimposition. This study has demonstrated the effectiveness of using individual-based and hierarchical modeling together with an ABC parameter estimation method in behavioral ecological studies.

Effects of age and rearing environment on genetic parameters of growth and body weight and heritability of skin pigmentation in Arctic charr (Salvelinus alpinus L.)

The Swedish breeding programme for Arctic charr aims to improve traits that are important for farming. Selection has been ongoing since the 1980s, and the aim of this study was to re-estimate the genetic parameters in the selected population. In addition, the genetics of skin pigmentation was studied, as previous work indicated a relationship between pigmentation and the ability to cope with stress.One hundred twenty-five full sib families of Arctic charr were hatched and reared separately until they were individually tagged (n=12,266). In connection with the tagging, three replicates were formed, which were subsequently reared at different sites. The genetic relationships were known from a seven-generation pedigree. Body weight and length were at one site measured at six occasions until 27months old, and these measurements were used to estimate the DGC (daily growth coefficient) and condition factor. At a second site fish were measured at 21 and 27months age and at a third site at 27months age.Skin pigmentation was measured using image analysis. The final weight at 27months of age varied widely between rearing sites, ranging from 529g to 1443g. Weight heritability differed among the sites, at 0.45, 0.11, and 0.07. At one of the sites, where growth was followed closely, weight heritability varied among growth periods, ranging from 0.01 to 0.25, and similar results were obtained for the DGC. In contrast, condition factor heritability did not show such pronounced variations within and among sites. The magnitude of the differences in heritability of weight and growth among different sites that was observed here was not observed previously with this charr population. In particular, the low heritability among the fastest growing fish is unprecedented.Skin pigmentation showed a high heritability of 0.76, indicating a strong additive genetic component. At most ages, pigmentation showed a weak positive genetic correlation with growth. The genotype environment interaction (GEI) was estimated using genetic correlations among sites. For the final weight, genetic correlations were high, indicating weak GEI, although the presence of GEI was suggested from the DGC data. Statement of relevanceThe ms will impact charr breeding for commercial farming.

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.