A hermaphroditic Arctic charr from Loch Rannoch, Scotland

An individual Arctic charr from Loch Rannoch was found to contain a gonad the anterior portion of which consisted of ovarian tissue. This is an extremely rare condition in salmonid fish and in this instance is possibly the result of environmental contamination.     

How fish-helminth associations arise: an example from Arctic charr in Loch Rannoch

Three sympatric morphs of Arctic charr Salvelinus alpinus occur in Loch Rannoch, Scotland, and are identified by their differing head morphology and diet. These are small-headed benthic, large-headed benthic and pelagic morphs. Six species of endoparasitic helminth were found in the fish, but the morphs had different patterns of infection. Overall infections in pelagic charr were heavier than in large-headed benthics, which were in turn heavier than in small-headed benthics, even though benthic charr live longer than pelagics. Pelagic fish had high prevalences and intensities of pseudophyllidean tapeworms, the intermediate hosts of which are copepods. The prevalence and intensity of metacercariae of Diplostomum sp. (the intermediate hosts of which are snails) were high in the benthic morphs. The results are discussed in terms of the effects of ecological factors on transmission of helminth parasites to their hosts and the evolution of host-parasite associations.     

The genetics of sympatric Arctic charr [Salvelinus alpinus (L.)] populations from Loch Rannoch, Scotland

Two morphologically and ecologically distinct forms of Arctic charr, Salvelinus alpinus , are found in Loch Rannoch, Scotland. The differences in morphology are adaptations to different modes of life, one being pelagic, the other benthic. Both forms have been the subjects of extensive genetic studies including cytogenetics, nuclear and mitochondrial DNA analysis, and protein electrophoresis. Significant differences between the two forms are revealed by some techniques but not others and provide evidence for the reproductive isolation of these two morphs. The findings are discussed in relation to the derivation of sympatry and the phylogenetics of Arctic charr.     

Morphometric analysis of two ecologically distinct forms of Arctic charr, Salvelinus alpinus (L.), in Loch Rannoch, Scotland

Gill–netted samples of Arctic charr from Loch Rannoch, Scotland were bimodal when tested by univariate and multivariate morphometric analyses. The separation into two morphs corresponded very closely (95–98%) with fish classified subjectively in the field as benthic or pelagic, based largely on colour differences and ecological observations. The benthic charr had relatively longer heads, larger eyes and more powerful jaws than the pelagic charr. Unlike sympatric morphotypes described from Scandinavia and Greenland, neither Rannoch morph is dwarfed.     

Trophic polymorphism in introduced bluegill in Japan

We demonstrate that bluegill (Lepomis macrochirus) introduced from their native North American source to Lake Biwa in Japan showed considerable trophic and morphological variations (morphs). Three morphs were found for larger bluegills, specialized in three distinct resources (i.e. plankton, benthic invertebrates and aquatic plant), and two morphs were found for smaller bluegills, specialized in two slightly different prey taxa (i.e. calanoid and cyclopoid plankton). These morphs, especially for the large size class, can be distinguished primarily by differences in midbody length and depth. The dietary specialization and morphological differentiation among individuals were considered to be a result of different requirements for efficient resource utilization, which is a unique example of trophic polymorphism occurring in an introduced fish species in a species-rich fish community.     

Segregation in spawning and early life history among polymorphic Arctic charr, Salvelinus alpinus, in Thingvallavatn, Iceland

The relationships among time of spawning, incubation temperature, timing of first feeding and early growth were examined in four sympatric morphs of Arctic charr in Thingvallavatn, Iceland. Large benthivorous charr spawn in July-August at sites with cold ground-water flow. Planktivorous and piscivorous charr spawn in September-November and are not confined to ground-water sites. The spawning of small benthivorous charr overlaps with that of other morphs. Progeny of large benthivorous charr start feeding 2-3 months earlier than the progeny of autumn spawners. This results in differential size distribution and growth rates of young in the spring.     

The functional significance of inherited differences in feeding morphology in a sympatric polymorphic population of Arctic charr

Artificially fertilised eggs from wild-caught Arctic charr parents of two sympatric morphs (benthivorous and planktivorous) from Loch Rannoch, Scotland were reared in the laboratory under identical conditions. During the subsequent 2 years, aspects of their trophic anatomy and feeding behaviour were compared. As previously described for wild-caught fish, charr derived from the benthivorous morph had an increasingly wider mouth gape for a given body length than those derived from the planktivorous morph. The functional significance of these differences in gape was tested by comparing the maximum size of prey that could be handled by each of the two morphs. In both forms, a larger gape enabled larger food particles to be eaten, but the elevation of the regression of maximum prey size on gape was higher in the benthivorous form, indicating the existence of additional morphological and/or behavioural differences influencing the size of prey consumed. When offered a choice between a typical benthic prey item and a typical pelagic food item, charr of benthivorous origin were more likely to feed on the former, whereas those of planktivorous origin were more likely to feed on the latter. Thus inherited differences in gape place constraints on foraging ability and are associated with inherited differences in dietary preference. We conclude that the functional significance of the foraging specialisations indicate a strong selection pressure for the evolution of the divergence and propose that heterochronic growth is the mechanism resulting in the divergence of tropic anatomy.     

Genetic analysis of four sympatric morphs of Arctic charr,Salvelinus alpinus,from Thingvallavatn,Iceland

Synopsis The degree of genetic differentiation among four morphs of Arctic charr (small benthivorous, large benthivorous, piscivorous, and planktivorous) from Thingvallavatn, Iceland, was determined electrophoretically. Five of 36 enzyme loci were found to be polymorphic (Est2, Gpi3, Ldh4, Mdh4, 5 and Pgm2). However, only Est2 and Mdh4,5 showed enough variability to permit statistical analysis of divergence among morphs. All four morphs are very closely related; the values of Nei's (D) range from 0.00004 to 0.00126. These morphs are conspecific and do not represent different evolutionary lineages. There is significant genetic differentiation between the small benthivorous charr and the other three morphs. The relative relatedness of morphs based on gene frequency data is only partially concordant with that based on morphology and ecological specialization. The biological significance of this result is unclear because of the limited number of polymorphic loci upon which the genetic analysis is based and the high degree of relatedness among morphs.     

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.     

Population structure, ecological segregation and reproduction in non-anadromous Arctic charr, Salvelinus alpinus (L), in four unexploited lakes in the Canadian high Arctic

Land-locked populations of Arctic charr in four lakes on Northern Ellesmere Island (80° N) were found to consist of two distinct sizes: 'dwarf' and 'normal' charr. Both groups attained sexual maturity but differed in appearance and habitat. The smaller fish, occupying the more marginal habitats, retained their parr-markings; the larger group had the general characteristics of smolts, being more silvery and without parr-marks. In their juvenile stages, the charr destined to attain the larger group were indistinguishable from members of the smaller group. Although fish in the larger group were capable of cannibalism, this was rarely observed. In general, the fish in the larger group were older than the smaller ones but great variation in size at a given age resulted in certain age classes containing representatives of both groups. The population structure varied considerably between lakes; a high proportion of 'normal' charr correlated well with a high growth rate in the first few years of development. It is postulated that the two groups live in dynamic equilibrium where the advantages of progenesis (retention of juvenile characters by adults) in the smaller type are traded against the larger proportion of the energy resources available to the larger type. The concept of heterochrony in an ecological setting is introduced.     

Genetic differentiation among local populations and morphotypes of Arctic charr, Salvelinus alpinus

The systematica of coexisting morphotypes of Arctic charr, Salvelinus alpinus , has been a matter of dispute ever since the days of Linnaeus. Widespread allelic variation at an esterase locus has led some investigators to propose that the morphotypes reflect a complex of at least three sibling species. We tested this hypothesis by examining 42 electrophoretically detectable loci in natural and transplanted charr populations from 15 localities in S Norway. The absolute values of Nei's genetic distance between morphotypes and populations are small (typically in the order of 0.001), and morphotype changes may occur without accompanying changes in frequencies of esterase alleles. Differentiation among localities explains far more of the total gene diversity than differences between morphotypes in the four cases of naturally occurring sympatric morphotypes examined. The data are consistent with an intraspecific population structuring based on locality, and the multiple species hypothesis is rejected.     

Smolting characters in anadromous and resident Arctic charr, Salvelinus alpinus (L.)

Morphological differences, haematocrit value and chloride cells were examined in downstream migrating Arctic charr, Salvelinus alpinus (L.) from the Hals River, North Norway, and resident charr from Lake Storvann. Fish were classified as visual parr, silvery parr or smolt based on the degree of silvering and the Occurrence of lateral parr marks. On average, 47% of downstream migrating Arctic charr were classified as visual molts, but only 14% of the resident charr. Charr longer than 20 cm fork length were mainly classified as visual molts. Morphometrical analyses of body size and shape revealed that most of the variations could be explained in terms of variations in fork length. Length adjusted ratios of post-anal distances were significantly higher in migrating charr than in resident charr. The number of developed chloride cells, the cell nucleus diameter in the basin of secondary gill lamellae and blood haematocrit values were significantly higher among downstream migrating charr than in resident charr. However, morphological differences between anadromous and resident Arctic charr were not necessarily synchronized with the development of chloride cells.     

Heterochrony in skeletal development and body size in progeny of two morphs of Arctic charr from Thingvallavatn, Iceland

Planktivorus Arctic charr had larger eggs than small benthivorous charr and the progeny of the former were longer (total length) at days 125, 145 and 159 after fertilization. Size differences remained significant after the removal of egg size effect on embryo size. Size of hybrid progeny tended to be similar to their maternal pure progeny group, suggesting maternal effects not directly related to yolk volume. In general, fin ray number increased faster in small benthivorous charr progeny than in planktivorous charr progeny, hybrid progeny tending to have intermediate fin ray numbers. The results indicate that morph differences in embryonic growth and skeletal development have a genetic and maternal component. Results support the hypothesis that in the period from hatching until just after first external feeding small benthivorous charr allocate more energy towards bone development, e.g. formation of fin rays, while planktivorous charr allocate more energy to body growth. The different developmental trajectories may reflect adaptations to discrete differences in habitats between the morphs.     

Evidence for Two Morphotypes of Lake Charr, Salvelinus Namaycush, from Great Bear Lake, Northwest Territories, Canada

The lake charr, Salvelinus namaycush, is generally characterized as a morphologically stable specialized deepwater piscivore. Empirical observations suggested the existence of two or more morphotypes of lake charr in Great Bear Lake, NWT. I gathered morphological data over two seasons from a total of 72 individuals and used principal components analysis to summarize variation in the dataset. UPGMA classification showed that there were two groups in the dataset. I classified individuals into one of two morphotypes using discriminant function analysis. The two forms differed primarily in head proportions. I developed a function, usable in the field, that classifies 80% of all specimens correctly. This study represents one of the first documentations of sympatric morphotypes of lake charr outside the Laurentian Great Lakes and provides the basis for further investigation into origins of and genetic, morphological and ecological differences between the forms.     

Trophic Structure of One Deep-sea Benthic Fish Community in the Eastern Canadian Arctic: Application of Food, Parasites and Multivariate Analysis

Synopsis As new arctic marine fisheries develop there is need for a comprehensive ecosystem approach to long-term management. This approach recognizes the importance of community interactions such as food web structure and trophic patterns. We determined whether hierarchical clustering (guild formation) is an effective method of trophic evaluation in deep-sea Artic fish communities using stomach content and parasite data with size class, and evaluated the application of endohelminth communities (parasite species transmitted in the food) as indicators of trophic status. Cluster analysis using food group abundance with size class of fish revealed the presence of 11 guilds within the community, however the same analysis using parasite data showed little correlation between food and parasites. Redundancy analysis (RDA) within the 11 guilds also revealed no significant correlations between food group and parasite abundance suggesting that this type of ordination is not suited for environments containing mainly generalist feeders. RDA of individual taxa without a priori guild designation found that taxa in benthic deep-sea communities are defined by their ability to exploit prey species in more than one habitat zone. Benthic fish species were significantly correlated with benthic food groups and parasites that utilize benthic intermediate hosts whereas benthopelagic–pelagic species fed on a higher diversity of prey species and were infected by a larger number of non-host specific parasites. Eigenanalysis and Monte Carlo results showed that parasites and food groups are highly correlated, indicating that parasite community analysis is an effective tool for predicting feeding strategies in Arctic marine environments. It also suggests that in most cases endoparasite infections alone could be used for trophic evaluation in the absence of stomach content data.     

Phenotypic plasticity, heterochrony and ontogenetic repatterning during juvenile development of divergent Arctic charr (Salvelinus alpinus)

Phenotypic plasticity is a developmental process that plays a role as a source of variation for evolution. Models of adaptive divergence make the prediction that increasing ecological specialization should be associated with lower levels of plasticity. We tested for differences in the magnitude, rate and trajectory of morphological plasticity in two lake populations of Arctic charr (Salvelinus alpinus) that exhibited variation in the degree of resource polymorphism. We reared offspring on diet treatments that mimicked benthic and pelagic prey. Offspring from the more divergent population had lower levels of morphological plasticity. Allometry influenced the rate of shape change over ontogeny, with differences in rate among ecomorphs being minimal when allometric variation was removed. However, plasticity in the spatial trajectory of development was extensive across ecomorphs, both with and without the inclusion of allometric variation, suggesting that different aspects of shape development can evolve independently.     

Reproductive isolation and genetic differentiation of ferox trout from sympatric brown trout in Loch Awe and Loch Laggan, Scotland

Molecular marker studies reported here, involving allozymes, mitochondrial DNA and microsatellites, demonstrate that ferox brown trout Salmo trutta in Lochs Awe and Laggan, Scotland, are reproductively isolated and genetically distinct from co-occurring brown trout. Ferox were shown to spawn primarily, and possibly solely, in a single large river in each lake system making them particularly vulnerable to environmental changes. Although a low level of introgression seems to have occurred with sympatric brown trout, possibly as a result of human-induced habitat alterations and stocking, ferox trout in these two lakes meet the requirements for classification as a distinct biological, phylogenetic and morphological species. It is proposed that the scientific name Salmo ferox Jardine, 1835 , as already applied to Lough Melvin (Ireland) ferox, should be extended to Awe and Laggan ferox.     

Growth, maturation and reproductive investment in Arctic charr

Size and rates of growth in a cohort of 1 + Arctic charr housed in standard conditions were tracked over 12 months (December to December) and mature and immature males and females compared retrospectively. In both sexes, maturing fish were larger than non-maturing ones. In males, this size differential was the result of differences in growth in winter and early spring, but not in the remainder of the study period. In females, size differentials resulted mainly from growth rate differences immediately prior to breeding. In females but not in males, gonadosomatic index was predicted by growth rates in the months leading up to maturation, and among the females that matured, faster growing fish produced more eggs. Lipid reserves in July were correlated negatively with growth during the previous 7 months and, in females only, lipid reserves were significantly lower in maturing fish than in non-maturing fish, indicating that mobilization of lipid energy reserves in maturing fish had commenced by this time. Variation in investment in gonadal tissue, measured as gonadosomatic index, was not explained by variation in July lipid reserves for either males or females. However, July lipid reserves were negatively correlated with egg number, so females investing more in ova exhibited greater depletion of lipid reserves. These results are discussed in the context of the relationship between body condition and the onset of maturation in salmonids, relative investment in reproduction and sexual differences in the cost of reproduction.     

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.     

Thermal adaptation of Arctic charr: experimental studies of growth in eleven charr populations from Sweden, Norway and Britain

1. Experimental growth data for Arctic charr (Salvelinus alpinus L.), all fed on excess rations, from 11 European watercourses between 54 and 70°N were analysed and fitted to a new general growth model for fish. The model was validated by comparing its predictions with the growth rate of charr in the wild. 2. Growth performance varied among populations, mainly because of variation in the maximum growth potential, whereas the thermal response curves were similar. The estimated lower and upper temperatures for growth varied between ?1.7 to 5.3 and 20.8–23.2 °C, respectively, while maximum growth occurred between 14.4 and 17.2 °C. 3. There was no geographical or climatic trend in growth performance among populations and therefore no indication of thermal adaptation. The growth potential of charr from different populations correlated positively with fish body length at maturity and maximum weight in the wild. Charr from populations including large piscivorous fish had higher growth rates under standardised conditions than those from populations feeding on zoobenthos or zooplankton. Therefore, the adaptive variation in growth potential was related to life‐history characteristics and diet, rather than to thermal conditions.