Effects of Partially Anadromous Arctic Charr (Salvelinus alpinus) Populations on Ecology of Coastal Arctic Lakes

Little research has been conducted on effects of iteroparous anadromous fishes on Arctic lakes. We investigated trophic ecology, fish growth, and food web structure in six lakes located in Nunavut, Canada; three lakes contained anadromous Arctic charr (Salvelinus alpinus) whereas three lakes did not contain Arctic charr. All lakes contained forage fishes and lake trout (Salvelinus namaycush; top predator). Isotope ratios (δ¹³C, δ¹⁵N) of fishes and invertebrates did not differ between lakes with and without anadromous Arctic charr; if anadromous Arctic charr deliver marine-derived nutrients and/or organic matter to freshwater lakes, these inputs could not be detected with δ¹³C and/or δ¹⁵N. Lake trout carbon (C):nitrogen (N) and condition were significantly higher in lakes with Arctic charr (C:N = 3.42, K = 1.1) than in lakes without Arctic charr (C:N = 3.17, K = 0.99), however, and ninespine stickleback (Pungitius pungitius) condition was significantly lower in lakes with Arctic charr (K = 0.58) than in lakes without Arctic charr (K = 0.64). Isotope data indicated that pre-smolt and resident Arctic charr may be prey for lake trout and compete with ninespine stickleback. Linear distance metrics applied to isotope data showed that food webs were more compact and isotopically redundant in lakes where Arctic charr were present. Despite this, lake trout populations in lakes with Arctic charr occupied a larger isotope space and showed greater inter-individual isotope differences. Anadromous Arctic charr appear to affect ecology and feeding of sympatric freshwater species, but effects are more subtle than those seen for semelparous anadromous species.     

Isolation and cross‐familial amplification of 41 microsatellites for the brook charr (Salvelinus fontinalis)

The brook charr (Salvelinus fontinalis; Osteichthyes: Salmonidae) is a phenotypically diverse fish species inhabiting much of North America. But relatively few genetic diagnostic resources are available for this fish species. We isolated 41 microsatellites from S. fontinalis polymorphic in one or more species of salmonid fish. Thirty‐seven were polymorphic in brook charr, 15 in the congener Arctic charr (Salvelinus alpinus) and 14 in the lake charr (Salvelinus namaycush). Polymorphism was also relatively high in Oncorhynchus, where 21 loci were polymorphic in rainbow trout (Oncorhynchus mykiss) and 16 in cutthroat trout (Oncorhynchus clarkii) but only seven and four microsatellite loci were polymorphic in the more distantly related lake whitefish (Coregonus clupeaformis) and Atlantic salmon (Salmo salar), respectively. One duplicated locus (Sfo228Lav) was polymorphic at both duplicates in S. fontinalis.     

Seawater acclimation of diploid and triploid brook charr (Salvelinus fontinalis), diploid Arctic charr (Salvelinus alpinus), and their diploid and triploid hybrids

Seawater acclimation of diploid (FF) and triploid (F2F) brook charr Salvelinus fontinalis , diploid (AA) Arctic charr Salvelinus alpinus , and diploid (FA) and triploid (F2A) hybrids between female brook charr and male Arctic charr was investigated. Triploidization of brook charr and the hybrid did not have any effect on the acclimation. Seawater acclimation of the hybrid was achieved during the experimental period and was comparable to that observed in brook charr. Acclimation could not be ascertained in Arctic charr since the level of cortisol, a stress indicator, was still high at the end of the experiment. No relationship between either length or condition factors and plasma osmolality was observed. Elevated plasma cortisol concentrations in Arctic charr and in diploid or triploid hybrids, both in fresh water and sea water, indicate more favourable rearing conditions for brook charr.     

An atypical brook charr (Salvelinus fontinalis) spawning area

Synopsis Brook charr (Salvelinus fontinalis) spawned successfully on a 0.3 m thick aggregation of waterlogged sticks, woodchips and debris overlying the soft ooze bottom of a small Precambrian Shield lake. Brook charr were apparently attracted to groundwater seeping up through the aggregation and utilized the tangle of various sized sticks as a spawning substrate. Eggs were deposited in late October and alevins emerged from the area in late March, periods that coincide with the conventional spawning of native brook charr on gravel areas in nearby lakes. Our observations support the contention that upwellin, water is more important than bottom type in stimulating brook charr to select a spawning site.     

Mitochondrial DNA variation of Salvelinus species found in Finland

Mitochondrial DNA (mtDNA) was purified from the Arctic charr, Salvelinus alpinus , the brook charr, Salvelinus fontinalis , and the lake charr, Salvelinus namaycush , and digested with restriction enzymes Ava II, Hinf I, Eco R V, Pst I and Xba I. Two Arctic charr samples were from natural populations and they represented two different morphotypes of Arctic charr. All other studied populations were hatchery maintained. Eight additional restriction enzymes and double digestions were employed to study morphotypes of Arctic charr. We distinguished two morphotypes with restriction enzyme Nci I. Sequence divergence among mtDNA types was 2.9–3.8% between S. alpinus and S. fontinalis , 3.4–4.6% between S. alpinus and S. namaycush , and 4.7–5.3% between S. fontinalis and S. namaycush . lntraspecific variation was lowest in Arctic charr, the average of nucleon diversity for three populations being 0.179, while for brook charr and for lake charr nucleon diversity was 0.334 and 0.550, respectively. According to the number of mtDNA types, it is obvious that introduction to Finland and hatchery propagation have not greatly affected the mtDNA variation of brook charr or lake charr.     

Pyloric caecal morphology of brook charr,Salvelinus fontinalis,in relation to diet

Synopsis Brook charr, Salvelinus fontinalis, shifts its diet from zoobenthos to pelagic prey when living sympatrically with white sucker, Catostomus commersoni, in lakes of the Laurentian Shield. We tested the hypothesis that this diet difference would have a significant impact on their pyloric caecal morphology in 5 lakes containing allopatric brook charr populations and 6 other lakes containing both brook charr and white sucker. We observed that the mean length of the most posterior caecum of charr was significantly greater in sympatry than in allopatry (X ± 1 SD: 9.91 ± 1.12 mm versus 8.44 ± 0.67 mm). This is equivalent to an increase of 18% of total pyloric caecal mass (dry weight) in sympatric brook charr. These results indicate that this response to differences in diet, well known in birds, also occurs in fish.     

Molecular genetic examination of the polymorphic Arctic charr Salvelinus alpinus of Thingvallavatn, Iceland

Thingvallavatn, Iceland contains two sympatric morphotypes (benthic and limnetic) of Arctic charr Salvelinus alpinus. Each morphotype is composed of two morphs and these differ markedly in ecology, behaviour and life history. We used molecular genetic approaches to test whether (i) genetic heterogeneity exists among morphs and (ii) if morphs arose in allopatry and came into secondary contact or arose sympatrically within the lake through genetic segregation and/or phenotypic plasticity. Direct sequencing of 275 bp of the mitochondrial DNA (mtDNA) control region, mtDNA restriction fragment length polymorphisms and single locus minisatellite analyses detected insufficient variation to test our hypotheses. Analysis of multilocus minisatellite band sharing detected no significant differences between morphs within the same morphotype. However, significant differences among morphs belonging to different morphotypes suggest some genetic heterogeneity in Thingvallavatn charr. Limnetic charr from Thingvallavatn were more similar to sympatric benthic charr than to allopatric limnetics from two other Icelandic lakes. This suggests that the Thingvallavatn morphs arose sympatrically within the lake rather than in allopatry followed by secondary contact.     

Differentiation of deep-water lake charr <Emphasis Type="Italic">Salvelinus namaycush</Emphasis> in North American lakes

Deep-water morphs of lake charr, Salvelinus namaycush, are found, with one exception, in four of the largest lakes in the world: lakes Superior and Mistassini (QC) and Great Bear and Slave lakes. This paper advances a hypothesis for resource polymorphisms involving two types of deep-water morph, one of which is characteristic of the humper and the other of the siscowet charrs of Lake Superior. My hypothesis states that, first, the humper, or a humper-like morph, diverged postglacially in sympatry from the ancestral common (shallow-water) lake charr and became a feeding specialist on Mysis relicta. Second, in at least two of the four lakes the siscowet, or a siscowet-like charr, diverged as a feeding specialist on postglacially derived forms of deep-water ciscoes. In Lake Superior a successional process may have resulted in dominance of the siscowet at the expense of the humper charr. I concur with a previous inference that the one occurrence of a deep-water charr in a small lake (the above exception) represents emigration from Lake Superior. I further infer that this event involved an early humper charr, which implies that this morphotype had differentiated in Lake Superior in less than 1,900 year. I suggest that innate differences in plasticity, breeding behavior and assortive mating, and philopatry account for why Arctic charr isolate readily in small lakes whereas lake charr do not. My hypothesis assumes divergence of deep-water morphs occurred postglacially, an idea consistent with genetic and biogeographical evidence.     

Phenotypic responses of lacustrine brook charr in relation to the intensity of interspecific competition

We investigated the structure of interindividual variations in the diet of brook charr (Salvelinus fontinalis) based on stomach contents data of 3776 charr captured in 69 lakes of the Canadian Shield (Québec, Canada); 29 of these contained allopatric brook charr populations, 24 contained brook charr and creek chub (Semotilus atromaculatus) and 16 contained brook charr and white sucker (Catostomus commersoni). In any given lake, some of the charr fed almost exclusively on benthic organisms (benthic specialists, i.e., mean percent weight of benthic prey >90%), others, almost exclusively on pelagic prey (pelagic specialists, i.e., mean percent weight of benthic prey <10%), and a lesser proportion were generalist feeders (i.e. mean percent weight of benthic prey between 10 and 90%). The proportion of benthic and pelagic specialists were respectively 41.3 and 18% in allopatric brook charr populations. These proportions fit remarkably well with those based on interindividual preferences in spatial distribution, identified through radio-telemetry in another study done in two lakes of the same area. The proportion of benthic specialists was related to competition for benthic organisms with creek chub and white sucker. The effect of white sucker on brook charr diet was more pronounced than that of creek chub: the proportion of benthic specialists among brook charr decreased from 41.3% in allopatry to 19.7% in sympatry with creek chub, and to 9.9% in sympatry with white sucker. Other response variables of brook charr populations also indicate that white sucker is a stronger competitor than creek chub in this system. Because sucker and chub were introduced in these lakes during the last century, phenotypic responses of brook charr to interspecific competition appear to be rapid. Furthermore, in addition to providing a strong field support to the current hypothesis that polymorphism is promoted by arelaxation of interspecific competition, our results also indicate that phenotypic response of brook charr (i.e. the proportion of each form in a given lake) is related to the intensity of this competition.     

Stable isotope-based determinations of the average temperatures experienced by young-of-the-year Svalbard Arctic charr (Salvelinus alpinus (L.))

Mean ambient water temperatures experienced by individual young-of-the-year (YOY) Arctic charr, Salvelinus alpinus (L.), from a Svalbard lake were estimated using measurements of oxygen stable isotopes (δ¹⁸O) derived from fish otoliths. Otolith-derived water temperatures differed significantly from temperatures recorded at the outlet river of the Dieset watercourse but were consistent with temperatures previously recorded in shallow littoral areas of other Svalbard lakes where YOY charr are commonly found. This indicates that fixed-point monitoring does not necessarily represent the temperatures and thermal habitats used by individual fish. Otolith-derived water temperatures were also positively related to fish length-at-capture and otolith size, although much of the variation remained unexplained. Differences among individuals could be related to variability in food availability and food intake as well as variation in the initial fish size at hatching. Implications for subsequent investigations into how YOY charr respond and adapt to future climate change are discussed.     

The Relevance of Individual Size to Management of Arctic Charr, Salvelinus Alpinus, Populations

Arctic charr, Salvelinus alpinus, tend to form stunted populations presumably due to competition for limited resources. In this context a long-term intensive fishery programme aiming at reducing charr density, and thereby increasing growth and harvestable sizes, was initiated in the early 1980s in a Norwegian alpine lake. Here we present long-term data on catch statistics and changes in mean weights of charr caught with gill nets on spawning grounds, as well as changes in mean weight of juvenile charr caught with funnel traps during the 1990s. Furthermore, we present results from shorter-term studies on growth, size- and age-distribution, and size-related habitat and resource utilisation of Arctic charr and brown trout, Salmo trutta, in this lake. Mean weight (± SD) of charr caught on the spawning grounds increased significantly from 129.8g (±11.9) in the years 1982–1990 to 213.1g (±37.8) in the years 1996–1999, whereas catch per unit of effort decreased significantly. Mean weight of juveniles caught in funnel traps increased significantly from 21.2g (±6.9) in 1993 to 41.9g (±14.8) in 1999. Apparently the increase in weight of spawning charr coincided with the onset of trap fishing for juvenile charr. Compared to generally shallow dwelling trout, charr grew rapidly and were generally found in deeper areas of the lake. Charr went through a distinct size-related niche shift from mainly consuming small zooplankton in the pelagic to consuming large benthic prey in shallower waters. Resource and habitat utilisation in different size-groups of charr and trout are discussed with respect to possible competitive and predatory intra- and interspecific interactions, and with regard to management of charr populations.     

Early microhabitat use by age 0 year brook charr Salvelinus fontinalis in lakes

The early habitat use of age 0 year brook charr Salvelinus fontinalis in three north temperate lakes which differ in terms of shoreline physical habitat is described. In the two lakes, which contained abundant shoreline woody debris and inundated vegetation, brook charr were observed in extremely close proximity with these habitat features, near shore and near the surface. Fish were absent from open areas away from shore unless in close proximity with fallen floating logs near the surface, extending offshore. In a third lake that had no woody debris or inundated shoreline vegetation, brook charr were observed exclusively in close proximity with the shoreline itself, and near the surface. In all three lakes, fish were most closely associated with the shoreline and with woody debris and inundated vegetation (when present) shortly after emergence, and significantly farther from shore and deeper in the water column thereafter.     

Lake enrichment and the status of Windermere charr, Salvelinus alpinus (L.)

All English populations of Arctic charr, Salvelinus alpinus (L.), are found in the Lake District (northwest England). There are at least four races of charr in Windermere, the largest lake in England; the North and South basins of the lake each contain two distinct races that spawn in autumn and spring respectively. The spring spawners in both basins probably represent less than 15% of the total population in the lake.
Changes in the population density of charr in the lake are described briefly and examined in relation to the trophic status of the lake. Other factors that could possibly affect the charr population are reviewed, especially the influence of climate change.     

Diversification, Sympatric Speciation, and Trophic Polymorphism of Arctic Charr, Salvelinus Alpinus Complex, in Transbaikalia

In northern Transbaikalia, independently evolving landlocked populations of Arctic charr are found in mountain lakes. To assess the diversity of charr in this region, speciation modes involved in the evolution of charr forms, and the role of trophic polymorphism in their divergence, we studied the morphology and feeding of dwarf, small, and large forms of Arctic charr from a number of Transbaikalian lakes. Meristic data on charr from five lakes support the earlier conclusion that the three forms do not represent separate lineages but have independently diverged in sympatry in each of the lakes. In 10 lakes, the dwarf form showed varying degrees of differentiation from normal (small and large) charr in meristic characters (up to morphologically distinct and presumably reproductively isolated groupings), which is viewed as various levels of sympatric divergence. Indexes of gill raker length in fish from 20 lakes vary among populations of both dwarf and normal charr, with forms having short and long rakers being sympatric in some of these lakes. However, the index can be used only for comparing charr of different forms up to about 32cm fork length (FL) because it is strongly negatively correlated with size in larger fish. The study of charr diets in 21 lakes indicates that large charr are piscivorous whereas dwarf and small charr feed on a wide range of invertebrates, partitioning these resources in different ways. Planktivores, including very specialized ones, and non-planktivores (benthic feeders, insectivores) can be identified within the small and dwarf forms. The proportion of plankton in the diets of dwarf and small charr is positively correlated with the number and length of gill rakers while the proportion of benthos is negatively correlated. Allopatric planktivorous and non-planktivorous small charr differ in body proportions; parallel emergence of such morphotypes in different parts of the range is a characteristic feature of the Salvelinus alpinus complex.     

Growth and food of young-of-the-year brook charr,Salvelinus fontinalis,in lake and creek environments

Synopsis Growth and diet of brook charr,Salvelinus fontinalis, during their first weeks of exogenous feeding were compared between the nearshore zone of a central Ontario lake and its small, inlet creeks. Food selection was related to size and age of charr and possibly availability of food items. Mean growth was similar between habitats despite differences in consumed food types and caloric values. Differences in social behaviour may have been responsible for different patterns of growth between habitats. The importance of creek habitats to the dynamics and fitness of lake populations of brook charr are suggested.     

Effects of Fish-farm Activity on the Limnetic Community Structure of Brown Trout, Salmo trutta, and Arctic Charr, Salvelinus alpinus

Establishment of four fish-farms during the period 1971 to 1994 in the oligotrophic lake Skogseidvatnet affected Arctic charr, Salvelinus alpinus, but not brown trout, Salmo trutta. From 1971 to 1987, an increase in mean individual size of Arctic charr was recorded, while the mean individual size of brown trout remained stable. Arctic charr were found to use deeper benthic areas than brown trout. Approximately 8% of the Arctic charr population (>26cm), were found to switch to waste food from fish-farms, resulting in a novel feeding habitat for the species. They were, however, found in gillnets distant from the fish farm cages, indicating high mobility. The habitat segregation between the two species can most likely be explained by selective differences and asymmetric competition with brown trout as the dominant species. Based on the present results, changes in the Arctic charr population may be due to increased food availability and due to a new habitat use as a waste food feeder. The reason for the brown trout population to have remained stable with respect to mean size, growth pattern and habitat use, may be due to a different diet choice than Arctic charr in this lake. Brown trout were found to feed mainly on terrestrial insects, while Arctic charr fed mainly on zooplankton and on waste food.     

Comparative ontogeny of photobehavioural responses of charrs (Salvelinus species)

Synopsis The development of photobehavioural responses in brook (Salvelinus fontinalis) and lake (S. namaycush) charr was studied by monitoring the intrasubstrate movements and concurrent photoresponse behaviour of incubated embryos and alevins. Photoresponse behaviour of both F-1 hybrids of the parent species was also recorded. All embryos initially moved downward in the substrate, however brook charr descended farther and faster into the substrate than did lake charr. Photoresponse tests demonstrated a similar pattern of photoresponse transformation from a photonegative to a photopositive state in both species. However, photoresponse reversal was faster, more extensive and occurred later in brook charr than in lake charr. Patterns of photoresponse change in F-1 hybrids were intermediate between those of the parent species. Photoresponse shifts preceded the onset of alevin emergence in both species. occurring when differential development of various morphological characters existed. Developmental states of characters were synchronously maximal towards the end of alevin emergence. Intermediate measures of morphological development were observed for F-1 hybrids. Possible functions and mechanisms of photoresponse transitions are discussed in relation to ecological differences between the species.     

Behaviour-Genetics of Lake Charr (Salvelinus namaycush) and Brook Charr (S. fontinalis): Observation of Backcross and F2 Generations

Brook charr (Salvelinus fontinalis) defend territories and show high levels of agonistic behaviour. In contrast, lake charr (S. namaycush) are non-territorial and rarely show any agonistic behaviour. The genetic determinance of behavioural differences between these species was confirmed by the behavioural similarity of brook charr backcross hybrids and brook charr and a significant maternal effect on hybrid behavioural phenotypes. Backcross and F₂ generations showed either one of two distinct behavioural strategies, one aggressive and territorial and the other non-aggressive and non-territorial. It seems most likely that these strategies are conditional on individual phenotypes, such as size of nearby conspecifics. However, precise measurements of costs and benefits of each strategy are required to distinguish between these alternate hypotheses for either species. Each action pattern was an independent pattern of variation and a distinct behavioural unit. However, patterns were coordinated with all other patterns at a higher hierarchical level (corresponding to the particular behavioural strategy). It seems most likely that these strategies would be controlled by a polygenic system, even though the nature of our data does not allow us to make firm conclusions in this regard.     

Microgeographic population structure of brook charr: a comparison of microsatellite and mark-recapture data

Polymorphism at five microsatellite genetic markers (genotyped n  = 496) and mark-recapture tagging data (tagged n  = 9813) were used to define the population structure of brook charr, Salvelinus fontinalis from the Indian Bay watershed, Newfoundland, Canada. Despite the absence of physical barriers to migration among lakes, both genetic and tagging data suggest that brook charr in each lake represent reproductively isolated populations. Exact tests comparing allele frequencies, θ (global value = 0·063), R st (global value = 0·052), individual assignment tests, and Nei's genetic distance provided congruent estimates of population subdivision in agreement with the tagging data (only 2·2% of recaptures were lake-to-lake). The genetic structure of the brook charr populations corresponded with the geographic structure of the drainage basin on a qualitative level, although linear distance over water was not significantly correlated with the tagging data or the genetic distance measures. The agreement between the tagging and the genetic data suggest that microsatellite markers can be useful tools for defining real biological units. The results also suggest that brook charr exhibit microgeographic population structure at the watershed scale, and that this is the scale at which conservation and management of this salmonid might best be implemented.     

Effects of population,family, and diet on craniofacial morphology of Icelandic Arctic charr (Salvelinus alpinus)

We evaluated hypotheses of intralacustrine diversification and plastic responses to two diet environments in Icelandic Arctic charr (Salvelinus alpinus). Full‐sib families of progeny of wild polymorphic charr from two lakes where morphs vary in their degree of phenotypic and ecological divergence were split, with half of the offspring reared on a benthic and half on a limnetic type of diet to estimate family norms of reaction. We focused on variation in craniofacial traits because they are probably functionally related to diet and complement a previous study of body shape in these charr. A hierarchical analysis of phenotypic variation between lakes, pairs of morphs within each lake, and two families within each morph found that phenotypic variation partitioned between families relative to morphs was reduced in the more ecologically diversified population, which is consistent with adaptive diversification. The effect size of plastic responses between lake populations was similar, suggesting little difference in the degree of canalization in contrast to a previous analysis of body form plasticity. Thus, the role that plastic morphological responses play in the adaptive diversification of morphs and different lake populations of Arctic charr may depend on the trait. © 2013 The Linnean Society of London