Phenotypic Differences in Buoyancy and Energetics of Lean and Siscowet Lake Charr in Lake Superior

At least two phenotypes of lake charr, Salvelinus namaycush, coexist in Lake Superior. A lean morph frequents the shallow inshore waters (< 50m) and the fat morph (siscowet) occupies the deeper offshore waters (50–250 m). The objective of this study was to determine if the elevated lipid concentration of siscowets reduces the costs of swimming in deep water. First, we modelled the effects of body composition (lipids) on the costs of swimming by lake charr, and then compared these theoretical results with empirical evidence obtained from Cesium 137-based estimates of food consumption, gross energy conversion, and swimming costs (activity multiplier). The attributes of growth, energy content (kJg^-1), lipid concentrations, and Cesium 137 concentration (Bqg^-1) were obtained from multimesh gillnet catches in eastern Lake Superior (1998 and 1999). The model showed that siscowet (fat) lake charr expended less energy than lean lake charr moving through the water column. Empirical evidence derived from Cesium 137 analysis confirmed that the activity multipliers of siscowets (fat) were less than those for lean charr. These findings support the view that the restoration of the fish community of the predominately deep water of the Great Lakes might be facilitated by the introduction of the fat phenotype.     

Genetic signatures of historical bottlenecks in sympatric lake trout (Salvelinus namaycush) morphotypes in Lake Superior

Humans have played a significant role in reducing levels of genetic diversity and differentiation of many teleost fishes, leading to homogenization across biological entities. We compare patterns of historical and contemporary genetic structure for three sympatric Great Lake??s lake trout (Salvelinus namaycush) morphs (lean, siscowet, and humper) that differ in patterns of habitat occupancy, susceptibility to overfishing and predation by invasive sea lamprey (Petromyzon marinus). Differential susceptibilities to overfishing and predation were expected to result in different impacts to levels of genetic diversity loss for each morphotype. Genetic data was collected for samples at three points in time: 1948 (pre-collapse), 1959 (collapse) and 1990s (current), corresponding to periods of intensive fishing, mortality due to lamprey and recovery, respectively. The lean morph preferentially targeted by the fishery and recognized as highly preyed upon by sea lamprey was more highly impacted genetically than other morphs, as evidenced by greater loss of genetic diversity first during the period of overfishing, then during the period of high sea lamprey abundance once the fishery collapsed. The siscowet morph also experienced genetic bottlenecks during the period of overfishing (pre-collapse period). Results indicate significant levels of genetic differentiation among morphs historically prior to declines in abundance and also among contemporary populations, suggesting that periods of population decline and resurgence in abundance and distribution did not result in loss of genetic distinctiveness among morphs.     

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     

Evidence for genetic differences in the offspring of two sympatric morphs of Arctic charr

The sub-arctic Lake Fjellfrøsvatn, northern Norway, has two morphs of Arctic charr that are reproductively isolated because they spawn 5 months apart. The smaller morph (≤14 cm L_F ) is confined to the profundal zone of the lake and the larger morph is mainly littoral. Three hypotheses were tested: (i) the offspring of the profundal Arctic charr grow slower than the offspring of the littoral Arctic charr under identical conditions, thus indicating a genetic basis for the slow growth of the profundal Arctic charr in the wild; (ii) the wild phenotypes of the two morphs are morphometrically different and the differences are persistent in the offspring; (iii) the offspring of the two morphs have different behaviour traits under similar treatments. The first hypothesis was rejected; offspring of the profundal morph grew slightly better than offspring of the littoral morph at 10° C in the laboratory. The second and third hypotheses were supported by the data. Wild-caught fish of the two morphs differed in several morphometric characters and most of the differences persisted in the offspring. In the laboratory, offspring of the littoral morph were more active, more aggressive and more pelagic than offspring of the profundal morph and naive offspring of the profundal morph were more effective in eating live chironomid larvae than were offspring of the littoral morph. The data for morphometry and behaviour, but not growth, provide evidence for genetic differences between the two Arctic charr morphs of Fjellfrøsvatn.     

Convergent evolutionary processes driven by foraging opportunity in two sympatric morph pairs of Arctic charr with contrasting post‐glacial origins

The expression of two or more discrete phenotypes amongst individuals within a species (morphs) provides multiple modes upon which selection can act semi‐independently, and thus may be an important stage in speciation. In the present study, we compared two sympatric morph systems aiming to address hypotheses related to their evolutionary origin. Arctic charr in sympatry in Loch Tay, Scotland, exhibit one of two discrete, alternative body size phenotypes at maturity (large or small body size). Arctic charr in Loch Awe segregate into two temporally segregated spawning groups (breeding in either spring or autumn). Mitochondrial DNA restriction fragment length polymorphism analysis showed that the morph pairs in both lakes comprise separate gene pools, although segregation of the Loch Awe morphs is more subtle than that of Loch Tay. We conclude that the Loch Awe morphs diverged in situ (within the lake), whereas Loch Tay morphs most likely arose through multiple invasions by different ancestral groups that segregated before post‐glacial invasion (i.e. in allopatry). Both morph pairs showed clear trophic segregation between planktonic and benthic resources (measured by stable isotope analysis) but this was significantly less distinct in Loch Tay than in Loch Awe. By contrast, both inter‐morph morphological and life‐history differences were more subtle in Loch Awe than in Loch Tay. The strong ecological but relatively weak morphological and life‐history divergence of the in situ derived morphs compared to morphs with allopatric origins indicates a strong link between early ecological and subsequent genetic divergence of sympatric origin emerging species pairs. The emergence of parallel specialisms despite distinct genetic origins of these morph pairs suggests that the effect of available foraging opportunities may be at least as important as genetic origin in structuring sympatric divergence in post‐glacial fishes with high levels of phenotypic plasticity. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Resource polymorphism and diversity of Arctic charr Salvelinus alpinus in a series of isolated lakes

Morphological, dietary and life‐history variation in Arctic charr Salvelinus alpinus were characterized from three geographically proximate, but isolated lakes and one large lake into which they drain in south‐western Alaska. Polymorphism was predicted to occur in the first three lakes because S. alpinus tend to become polymorphic in deep, isolated lakes with few co‐occurring species. Only one morph was evident in the large lake and two of the three isolated lakes. In the third isolated lake, Lower Tazimina Lake, small and large morphs were found, the latter including two forms differing in growth rate. The small morph additionally differed from the two large forms by having more gill rakers and a deeper body than same‐sized individuals of the large morph, consuming more limnetic and fewer benthic resources, having a greater gonado‐somatic index and maturing at a smaller size. The two large forms consumed only slightly different foods (more terrestrial insects were consumed by the medium‐growth form; more snails by the high‐growth form). Trends in consumption of resources with body shape also differed between lakes. Variability in life history of S. alpinus in these Alaskan lakes was as broad as that found elsewhere. This variability is important for understanding lake ecosystems of remote regions where this species is commonly dominant.     

Parallel divergence of sympatric genetic and body size forms of Arctic charr,Salvelinus alpinus,from two Scottish lakes

F _ST and R_ST estimates for Arctic charr from six microsatelite markers collected from two neighbouring Scottish lakes, Loch Maree and Loch Stack, confirm the presence of two distinct genetic groupings representing separate populations within each lake. In both lakes, there was also a clear body size dimorphism, with large and small body size forms that segregated according to genetic grouping. There was evidence of only subtle foraging ecology differences between morphs, with the small body size morph in both lakes being more generalist in its foraging in the summer (consuming mostly plankton but also some macrobenthos) than the large body size morph, which specialized on planktonic prey. Trophic morphology (head and mouth shape) did not differ significantly between morphs (although the small sample size for Maree makes this a preliminary finding). Cluster analysis of the microsatelite data and the presence of private alleles showed that morphologically similar forms in different lakes were not genetically similar, as would be expected under a multiple invasion hypothesis. Thus, the data do not support a hypothesis of a dual invasion of both lakes by two common ancestors but instead suggest an independent origin of the two forms in each lake. Thus parallel sympatric divergence as a result of common selection pressures in both lakes is the most parsimonious explanation of the evolutionary origin of these polymorphisms. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 748–757.     

Morphological divergence between three Arctic charr morphs – the significance of the deep‐water environment

Morphological divergence was evident among three sympatric morphs of Arctic charr (Salvelinus alpinus (L.)) that are ecologically diverged along the shallow‐, deep‐water resource axis in a subarctic postglacial lake (Norway). The two deep‐water (profundal) spawning morphs, a benthivore (PB‐morph) and a piscivore (PP‐morph), have evolved under identical abiotic conditions with constant low light and temperature levels in their deep‐water habitat, and were morphologically most similar. However, they differed in important head traits (e.g., eye and mouth size) related to their different diet specializations. The small‐sized PB‐morph had a paedomorphic appearance with a blunt head shape, large eyes, and a deep body shape adapted to their profundal lifestyle feeding on submerged benthos from soft, deep‐water sediments. The PP‐morph had a robust head, large mouth with numerous teeth, and an elongated body shape strongly related to their piscivorous behavior. The littoral spawning omnivore morph (LO‐morph) predominantly utilizes the shallow benthic–pelagic habitat and food resources. Compared to the deep‐water morphs, the LO‐morph had smaller head relative to body size. The LO‐morph exhibited traits typical for both shallow‐water benthic feeding (e.g., large body depths and small eyes) and planktivorous feeding in the pelagic habitat (e.g., streamlined body shape and small mouth). The development of morphological differences within the same deep‐water habitat for the PB‐ and PP‐morphs highlights the potential of biotic factors and ecological interactions to promote further divergence in the evolution of polymorphism in a tentative incipient speciation process. The diversity of deep‐water charr in this study represents a novelty in the Arctic charr polymorphism as a truly deep‐water piscivore morph has to our knowledge not been described elsewhere.     

Parallel phenotypic evolution of skull-bone structures and head measurements of Arctic charr morphs in two subarctic lakes

Interspecific morph variations in trophic morphology related to skull-bones and head traits is associated to ecological segregation of Arctic charr morphs (genus Salvelinus) in two sub-arctic lakes (Fjellfrøsvatn and Skogsfjordvatn, Norway). The replicated morph pair, the profundal spawning benthivorous PB-morph and the littoral spawning omnivorous LO-morph of Arctic charr, diverge along the shallow-deep-water resource axis. In Skogsfjordvatn there is also a profundal spawning piscivorous PP-morph. The PB-morphs from both lakes have similar skull-bone traits and head morphology such as elongated jaw-bones, small opercular bones and relatively longer heads. The PP-morph also has an elongated head, relatively small opercular bones as well as larger jaw-bones. In contrast, the LO-morphs in both lakes have shorter jaw-bones, larger opercular bones in addition to relatively small heads. However, some small non-parallel differences exist among the morphs from the two lakes. Overall, all profundal morphs (PB and PP) have relatively similar skull-bone structures, suggesting adaptations to the deep-water environment but also to their separated dietary niches. There is strong evidence for parallel evolution with some local adaptations in skull-bones and head morphology of the PB-morph and the LO-morph from separate lakes.     

Omega-3 Fatty Acids in Fish from the Laurentian Great Lakes Tribal Fisheries

Dietary fish must be assessed for benefits and risks to formulate risk management strategies. This article demonstrates that Laurentian Great Lakes (GL) freshwater species are good sources of omega-3 fatty acids using new data from a small sample (n = 7) of Lake Superior siscowet lake trout (Salvelinus namaycush siscowet) and five other GL fish species’ data. For Lake Superior (LS) siscowets, the saturates, mono-unsaturates, and poly-unsaturates composed 20.1, 40.7, and 39.1% of total lipid weight, respectively. Omega-3 poly-unsaturates (PUFAs) in these fish were more than twice the omega-6 (omega 3/6 ratio = 2.4). The LS lake trout data were combined with earlier LS data collected during the 1980s for eight other species and from five species of Lake Erie fish. All the GL freshwater species were compared with seven other published marine and freshwater fish studies from other global regions. PUFAs were compared based on latitude and marine versus freshwater origin. Differences between marine and freshwater species in omega-3 fatty acid were less at higher latitudes. GL freshwater fish species can be a good source of beneficial fats like marine fish and must be accounted in effective risk communications involving persistent bioaccumulative toxicants in dietary fish.     

Atypical swimbladders of lake charr, Salvelinus namaycush, from Great Slave Lake, Northwest Territories, Canada

Swimbladder walls of lake charr, Salvelinus namaycush, from Great Slave Lake (GSL), Northwest Territories, Canada, were unusually thick for the species. The thinnest sections of the GSL bladders (mean = 2.44mm, range = 1.1–4.4mm) were significantly thicker (P = 0.001) than lake charr swimbladders collected from two small Province of Ontario lakes (means = 0.65 and 0.92mm), whose populations were assumed to be representative of the species. Variance in wall thickness was also greater in GSL lake charr than in charr from two small lakes (P < 0.02). Within individuals, some of the GSL bladder walls were markedly irregular in thickness, but whether these anomalies exist in situ or were artifacts of preservation remains uncertain. The bulk of the tissue in the thickest sections of the GSL swimbladders was in the tunica serosa (outer layer). The extent of the modification of the GSL swimbladders is extraordinary for northern fishes in postglacial lakes.     

Trophic specialization in Arctic charr Salvelinus alpinus (Pisces; Salmonidae): morphological divergence and ontogenetic niche shifts

Lake Thingvallavatn supports four trophic morphs of Arctic charr, Salvelinus alpinus (L.); two of the morphs are benthic (small and large benthivorous charr) one exploits pelagic waters (planktivorous charr) and the fourth is found in both habitats (piscivorous charr). The morphological variation among these morphs was analysed by use of principal component analysis and canonical discriminant analysis. The benihic morphs have a short lower jaw and long pectoral fins. The benthic fish also have fewer gillrakers than the other morphs. Small and large benthivorous charrs attain sexual maturity from 2 and 6 years of age, and at fork lengths from 7 and 22 cm, respectively. Small benthivorous charr retain their juvenile parr marks as adults, have beige ventral colours, and are frequently melanized under the lower jaw. Planktivorous and piscivorous charr attain sexual maturity from 4 and 6 years of age, from fork lengths of 15 and 23 cm, respectively. This phenotypic polymorphism is associated with habitat utilization and diet of the fish, and has probably arisen within the lake system through diversification and niche specialization. The pelagic morphs apparently stem from a single population, and are possibly diversified through conditional niche shifts which affect ontogeny. Juveniles reaching a body length of 23 cm may change from zooplankton to fish feeding. Asymptotic length increases thereby from 20.5 cm in planktivorous charr to 30.2 cm in piscivorous charr. The benthic morphs appear to represent separate populations, although both feed chiefly on the gastropod Lymnaea peregra. Their co-existence seems to be facilitated by size dependent constraints on habitat use. The small morph (asymptotic length 13.3 cm) exploit the interstitial crevices in the lava block substratum, whereas the large morph (asymptotic length 55.4 cm) live epibenthically.     

Biomass and Production of Lake Charr During the Acidification and PH Recovery of A Small Ontario Lake

The biomass and production of lake charr, Salvelinus namaycush, was studied in Lake 223, a lake that was intentionally acidified to pH 5.0 and then allowed to gradually recover, and in Lake 224, a natural lake of neutral pH. During the first 6 years pH decreased, biomass of Lake 223 lake charr increased, primarily due to high recruitment during the initial years of acidification. Biomass then decreased the final year of acidification. Biomass of Lake 223 lake charr remained low during the subsequent 10 years of pH recovery and 5 additional years after lake pH had returned to neutral pH. Production of lake charr decreased during acidification and increased during pH recovery. Production was still at least an order of magnitude lower at the end of this study than 22 years earlier before lake pH was lowered. The continued low production was caused primarily by low abundance during the years of pH recovery and following years. Production per individual fish in Lake 223 decreased during acidification and quickly increased during pH recovery to values similar to that prior to acidification. Production, biomass, and production per individual varied little between years for lake charr in Lake 224.     

Cost of predation avoidance in young-of-year lake trout (Salvelinus namaycush): growth differential in sub-optimal habitats

Selection of habitat to avoid predation may affect the diet of young-of-year (YOY) lake trout (Salvelinus namaycush). YOY lake trout may use inshore habitat to avoid predation; this habitat may be sub-optimal for growth. To test this, YOY lake trout were penned in nearshore and offshore pelagic areas of two arctic lakes. Toolik Lake had a lake trout population, the other lake, S6, did not. YOY lake trout in Toolik Lake lost weight, but those offshore lost less weight. The YOY lake trout in Lake S6 gained weight and those offshore gained more weight. The primary diet item of the YOY lake trout in both lakes during this experiment was the zooplankter Diaptomis probilofensis; it was also one of the most abundant species. However, its density inshore in Lake S6 was similar to inshore and offshore densities in Toolik Lake. The increased availability of alternative zooplankton prey in Lake S6 may account for the growth differential of YOY lake trout in Lake S6 relative to Toolik Lake. Bioenergetic modeling of YOY lake trout suggests that growth similar to that in the offshore of Lake S6 would be necessary for successful recruitment. If the reduced zooplankton availability in Toolik Lake leads to the reduced growth of YOY in the inshore and offshore pelagic areas, then these fish will be more susceptable to winter predation/starvation. For YOY lake trout to survive in Toolik Lake they most likely shift to feeding on benthic prey before the end of their first summer. Dept. of Chemical Engineering     

Taxonomic status of the six-band morph of Cyphotilapia frontosa (Perciformes: Cichlidae) from Lake Tanganyika, Africa

Six- and seven-band morphs have been identified in a cichlid, Cyphotilapia frontosa, that is endemic to Lake Tanganyika. These color morphs have allopatric distributions; the six-band morph is widespread in the northern half of the lake while the seven-band morph is restricted to Kigoma on the east coast of the lake. Because no specimens of the seven-band morph have been available for taxonomic study except for the holotype of C. frontosa, the taxonomic status of these morphs has not been discussed. In a recent survey at the lake, 21 specimens of the seven-band morph were collected. A comparison of these with existing collection specimens of the six-band morph showed significant differences in morphometric and meristic characters; however, because all characters largely overlapped between these morphs, they are regarded as conspecific.     

Incipient speciation through niche expansion: an example from the Arctic charr in a subarctic lake

Two reproductive isolated morphs of Arctic charr (Salvelinus alpinus), termed profundal and littoral charr according to their different spawning habitats, co-occur in the postglacial lake Fjellfr?svatn in North Norway. All profundal charr live in deep water their entire life and have a maximum size of 14cm, while the littoral charr grow to 40cm. Some small and young littoral charr move to the profundal zone in an ontogenetic habitat shift in the ice-free season and the rest of the population remains in epilimnic waters. The two morphs had different diet niches in the profundal zone: the profundal charr ate typical soft-bottom prey (chironomid larvae, pea mussels and benthic copepods), while the young littoral charr mainly consumed crustacean zooplankton. In four other lakes without a profundal morph (i.e. monomorphic populations), young charr also performed ontogenetic habitat shifts to the profundal zone and fed on zooplankton. The profundal morph of Fjellfr?svatn therefore utilize a food resource niche that neither the littoral morph nor comparable monomorphic populations exploit. This suggests that intraspecific resource competition has driven incipient ecological speciation of the profundal charr of Fjellfr?svatn. The exploitation of the soft-bottom resources by the profundal charr supports earlier experimental findings that the profundal morph is genetically different in trophic behaviour and morphology. The sympatric ecological divergence within the profundal habitat is possible because unexploited food resources (soft-bottom profundal prey) are available. Apparently, this represents a case of incipient segregation by expansion to new resource types (niche invasion), and not by subdivision of one broad ancestral niche.     

The trace element contents in tissues of Dolly Varden <Emphasis Type="Italic">Salvelinus malma</Emphasis> Walbaum, 1792 sympatric morphs from Lake Kronotskoe (Kamchatka)

The liver, gonads, gills, and muscles of five trophically distant charr morphs from Lake Kronotskoe, as well as silty sediments of the lake, were tested for trace-element contents. Concentrations of Mn, Fe, Ni, Cu, Zn, Rb, Sr, As, and Se were detectable in the fish tissues, while the xenobiotics Hg, Cd, and Pb were not determined. The current data indicate a strong correlation between trace-element content and the ecology of each morph and no relationship to the fish length. In the majority, the Ni, Cu, and Zn levels were strongly dependent on trophic biology. The contents of Rb, Mn, and Se were dependent on the distribution of the charr morphs along the depth gradient and correlated with the accumulation of these elements in the bottom sediments.     

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.     

Does global warming threaten the dynamics of Arctic charr in Lake Geneva?

In France, the Arctic charr (Salvelinus alpinus) is native in only two lakes (Lakes Bourget and Geneva), in the most southerly part of its distribution area. It is a profundal morph living at depths of between 30 and 100 m in Lake Geneva. Following considerable stocking of Lake Geneva with juvenile Arctic charr and some good results during the 1980s, catches are currently declining. Several hypotheses have been proposed to explain this decrease, but the potential influence of warming of the lake in the 1990s has not been considered. This article studies the relationship between the strength of the cohorts and the temperature in Lake Geneva from 1992 to 2002, and discusses the various potential causes of the population collapse observed. The findings reveal close correlation between water temperature and the strength of the cohorts. It is concluded that the recent warming of Lake Geneva may have a significant direct or indirect impact on the Arctic charr population.