Three sympatric forms of Arctic charr Salvelinus alpinus complex (Salmoniformes,Salmonidae) from Lake Kamkanda,Northern Transbaikalia

We studied morphology, size and age structure, growth, feeding, and variation at microsatellite loci of three forms of Arctic charr Salvelinus alpinus complex (dwarf, small, large) from mountain Lake Kamkanda in River Olekma basin, northern Transbaikalia. The forms differ in meristic and morphometric characters, external appearance and size. The small form distinctly differs from the dwarf and large forms in higher number and length of gill rakers. The forms differ in growth rate; however, differences in growth between the dwarf and the small forms are not as large as between sympatric dwarf and small charr from other Transbaikalian lakes. The large form is heterogeneous in growth rate. The small form matures one year earlier than the dwarf form and has a shorter life span. The dwarf form is a benthophage, the small form is a planktophage, and the large form is a predator. The dwarf form spawns in September, while the small form spawns in November-December, and there is no overlap in their spawning time. The three forms have clear genetic differences, which support their reproductive isolation. It is assumed that the three forms of Arctic charr originated within Lake Kamkanda on the basis of trophic polymorphism and spawning time displacement and attained a high degree of morphological and genetic divergence.     

Ecological polymorphism in Arctic charr

Arctic charr, Salvelinus alpinus (L.), commonly exhibits two coexisting morphotypes, dwarf and normal charr, which are characterized by differences in adult body size and colouration. We tested whether or not the morphotypic differences were genetically determined in rearing experiments with offspring of the two morphs and of their crosses. The experiments suggest that this ecological polymorphism in Arctic charr is largely environmentally determined. When reared under similar conditions, offspring of each of the two morphs differed little in size at the same age, and they had the same early developmental rate and maturation pattern. Moreover, the presence of parr marks along the flanks of the fish, one characteristic of dwarf charr, depended on body size and not on parental morph. Genetic differences between the morphs were suggested for growth rate during the second year of life, and for jaw morphology. Comparisons between charr life histories in captivity and in the wild suggest that ecological polymorphism in Arctic charr is chiefly a result of variation in growth conditions between different habitats.     

Extant and extinct forms of arctic charr Salvelinus alpinus (L.) complex from the Leprindo lake system (Transbaikalia): Differentiation in life history, morphology, and genetics

Interconnected lakes Bol’shoe Leprindo and Maloe Leprindo in Transbaikalia hosted large (extinct) and dwarf charr forms. Rarely “small” individuals intermediate in size between these forms are caught. In order to assess morphological, ecological, and genetic differentiation of sympatric charr forms and parapatric charr populations we studied their meristic and morphometric characters, feeding, breeding, and growth; we also investigated variation at 8 microsatellite loci using DNA isolated both from contemporary and historic samples. Profound differences were found between large and dwarf charr in growth rate, feeding (piscivores and highly specialized zooplanktivores, respectively), spawning time, and morphology. Dwarf charr from the two lakes demonstrate minor differences in morphology and growth rate. “Small” individuals are morphologically similar with dwarf charr and spawn together with them, they are recruited from dwarf form in late ontogeny as the result of transition to piscivorous feeding and growth acceleration. Microsatellite analysis showed that: (1) large and dwarf charr forms display high degree of genetic differentiation and reproductive isolation; (2) dwarf charr from interconnected lakes belong to different isolted populations; (3) “small charr” are genetically identical with dwarfs. The degree of ecomorphological and genetic differentiation between large and dwarf forms places charr from Leprindo Lakes among the most strongly differentiated Arctic charr forms’ flocks known at the vast range of S. alpinus complex.     

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.     

The growth of charr, Salvelinus willughbii Günther, in Windermere

The growth of charr ( Salvelinus willughbii Günther ) caught in Windermere from 1941–1952 has been studied. Scales were used for determination of age and back-calculation of length for age. Autumn and spring spawners, males and females, and charr of normal and dwarf growth were treated separately. In fish of normal growth, the spring spawners were significantly smaller than the autumn spawners at ages 1 and 2 years, and significantly larger from age 4 years onwards. There was little difference in growth between males and females within the two spawning populations. Charr of lengths of less than 200 mm at age 4 years were considered to be dwarfs. Mean lengths at capture of male charr were: autumn spawners normal growth 272 mm, dwarf 218 mm; spring spawners normal growth 327 mm, dwarf 194 mm. The oldest recorded age was 8 years.     

Observations on interbreeding between dwarf and normal arctic charr,Salvelinus alpinus,from Stora Rösjön,central Sweden

Synopsis The spawning behaviour of dwarf and normal arctic charr, Salvelinus alpinus, was studied through trough observations in October 1988. The experimental fish consisted of reared, three-summer old progeny of artificially bred dwarf and normal charr from an isolated body of water in central Sweden. The spawning behaviour of three normal female charrs was studied and video-recorded. The study showed that the arctic charr is potentially polyandric both within and between forms. The normal female was courted and mated both by normal and dwarf males. It was also noted that antagonistic behaviour on behalf of the female was much more common towards dwarf males than normal males. Guarding tactics were only observed in normal males, whereas sneaking tactics were employed both by dwarf and normal males. The dwarf as well as normal males mated the female charr either by courting the female or by joining the spawning fish at oviposition. Only dwarf males were observed resting silently on the bottom close to the redd.     

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.     

Late ontogeny growth acceleration and size form transformations in Transbaikalian Arctic charr, <Emphasis Type="Italic">Salvelinus alpinus</Emphasis> complex: evidence from fin ray cross section growth layers

In some polymorphic populations of Arctic charr in Transbaikalia, an individual can transform from a smaller to larger size form during their lifetime as a result of accelerated growth that follows a period of slow growth and reproduction as a small size form. Alternating periods of slow and fast growth are reflected in growth layer patterns visible in fin ray cross sections. Stained microtome fin ray cross sections were used to reveal the incidence of transformations from one form to another. Data were collected from 14 northern Transbaikalian lakes containing two or three sympatric Arctic charr forms (‘dwarf’, ‘small’, and ‘large’) exhibiting varying levels of morphological separation. Individuals recruited from the dwarf or small form were found in varying proportions among the small and/or large form in 12 lakes. Small or large form charr that grew without noticeable acceleration to the adult size typical of the form or experienced accelerated growth as juveniles prior to maturation were also observed. There were no transformations between sympatric forms that differed in the length and number of gill rakers and in some other meristic characters. Results indicate that in the region under study, transformations of sympatric Arctic charr size forms are a widespread, but not a ubiquitous, phenomenon. Such transformations reflect the plasticity of the developmental channels of the forms. In the course of intra-lacustrine form divergence and genetic differentiation, the frequency of the observed transformations decreases to zero.     

Maturation,growth and fecundity of Arctic charr, <Emphasis Type="Italic">Salvelinus alpinus</Emphasis> (L.), life-history variants co-existing in lake systems of Southern Baffin Island,Nunavut, Canada

Two life-history variants of Arctic charr (Salvelinus alpinus), anadromous and lake-resident, have been previously identified in lakes of Southern Baffin Island, Nunavut, Canada. In accordance with classical life-history theory, it is hypothesised that anadromous charr will delay maturation in both size and age, and have increased fecundity (per spawning event) in comparison with lake-resident charr. Sagittal otoliths and biological data were collected for both life-history variants within the three studied lakes: Iqalugaarjuit, Qasigiat and Qinngu. Sagittal otoliths were embedded in epoxy resin, cross-sectioned for age determination, and imaged for back-calculation (size and age). Back-calculated data in each lake were fit to von Bertalanffy growth models for each life-history variant and compared via analysis of residual sums of squares. Anadromous charr had greater mean size at maturity and experienced a delay in mean age at maturity in comparison with lake-residents. The relationship between size and fecundity or egg diameter did not differ between the two life-history variants. Growth models indicate that the overall growth coefficients of lake-resident and anadromous charr were different in all three studied lakes. The Brody growth coefficient for all lake-resident charr populations was greater than anadromous individuals indicating that maximum length was reached at a rapid rate, resulting in a smaller asymptotic length. Indirect evidence suggests that anadromous and lake-resident charr belong to one reproductive population. Future genetic analysis is necessary to further determine the degree of divergence between the life-history variants.     

A comparison of growth patterns between a stunted and two large predatory Arctic charr populations under identical hatchery conditions

Arctic charr are characterized by an extensive variability in growth and body size in natural waters. Although growth traits may involve a significant heritable component, most of this intraspecific variation presumably is environmentally induced and thus attributable to phenotypic plasticity. In the present study, size-at-age and length–weight relationship (body condition) were assessed for three Finnish Arctic charr populations of different geographical origins and extreme size forms (a stunted vs. two large-growing, predatory charr) held under standardized rearing conditions for 3 years (up to 37 months after hatching). In particular, our interest was to investigate whether the differences in growth between the large and the stunted charr as observed in the wild populations would diminish when the fish are offered suitable food in abundance. Population-specific mean body size and condition differed significantly in 0+, 1+, 2+ and 3+ fish. However, the identical rearing conditions resulted in the originally stunted charr reaching a comparable final mean size (317 mm/427 g) as the large charr populations (343 mm/510 g and 359 mm/497 g). Some individuals were of the same size as their parents at spawning already at the age of 0+ years. Furthermore, length–weight regression residuals of the stunted charr developed to a notably high level, indicating the largest final condition mean. The increase of size variation (CV of weight) in stunted charr lasted for over two growth seasons, whereas in large charr it remained stable since the end of the first summer. Variations in mortality and sexual maturation at age 2 seemed to be less relevant factors affecting overall growth performance. The study demonstrates an example of the high plasticity involved in the growth of fish: the stunted charr possess a tremendous capacity for growth in a benign environment, virtually corresponding to that observed in the large predatory populations.     

The size structure of lacustrine Arctic charr (Pisces: Salmonidae) populations

The size structure of Arctic charr (Salvelinus alpinus) populations was analysed using published information; 44% of populations were bimodal, with both large (normal) and small (dwarf) morphs occurring within a cohort. The remaining populations were unimodal, consisting of normal-sized or of stunted adults. Bimodal populations increased in frequency with latitude and were characteristic of large, deep lakes with few fish species. The age and size at which bimodality developed, the size difference between the morphs, and the frequency of cannibalism in charr populations increased with latitude. A variety of genotypic and phenotypic explanations are examined. The evidence for specific differences between the morphs is unconvincing. Various competition and predation hypotheses fail to explain the occurrence of bimodality. Cannibalism does not cause bimodality despite being strongly associated with it since bimodality develops before charr become cannibalistic. Much of the variation in charr size structure is suggested to be a consequence of increased seasonality in food supply in more northerly environments, coupled with feeding size thresholds. The latter result in larger members of a cohort being able to maintain growth rates on seasonally abundant prey while smaller individuals which cannot catch these items form a second mode of more slowly growing fish. Bimodality is documented in a number of other, predominantly northern, fish species.     

Patterns of development of the coldwater lacustrine-riverine form of the Drjagin charr (genus <Emphasis Type="Italic">Salvelinus</Emphasis>) from Lake Lama (the Taimyr Peninsula) during the larval period

Reproduction of the lacustrine-riverine form of the Drjagin charr (genus Salvelinus) from Lake Lama (the Noril’sk-Pyasina aquatic system on the Taimyr Peninsula) in a mountain tributary, the Bunisyak River, as well as the formation and development of skeletal elements in embryos and larvae reared under different experimental conditions are described. Patterns of the development and the heterochrony between the rates of somatic growth and osteogenesis are an adaptation to the extremely low water temperature and oligotrophic spawning habitat of the mountain stream. Still, the formation of vertebral centrae is retarded even at rather high rearing temperatures. As compared to larvae of the small normal (nondwarf) planktivorous lacustrine form of Arctic charr from Lake Davatchan (Transbaikalia) reared at 3°C, larvae of Drjagin charr from Lake Lama reared at 1°C are characterized by accelerated growth and heterochronies of osteogenesis. The divergence of the lacustrine-riverine form of Drjagin charr from the generalized form of Salvelinus alpinus in morphology begins during the larval period of the ontogeny.     

Latitudinal variation in growth among Arctic charr in eastern North America: evidence for countergradient variation?

Biological data from 66 populations of Arctic charr, Salvelinus alpinus, from eastern North America were analysed to test the applicability of the countergradient hypothesis as an explanation of differences in seasonally adjusted growth rates. Samples were obtained along a 37° latitudinal gradient and partitioned among anadromous, normal lacustrine, and dwarf lacustrine Arctic charr morphotypes. Models relating length-at-age or age-specific growth rates to latitude were estimated for each morphotype. Length-at-age declined with latitude for anadromous and lacustrine charr. Age-specific growth rates also varied with latitude, particularly for normal lacustrine charr. Results of analyses provide support for the countergradient hypothesis in growth performance of normal lacustrine morphotypes, where northern populations compensate for the shorter growth season with a greater rate of growth than southern populations. Anadromous charr exhibited equivocal evidence of countergradient variation, while results for dwarf lacustrine Arctic charr populations were inconclusive owing to the limited range of ages, and latitudes for which data were available.     

Size at maturity of fluvial white-spotted charr, <Emphasis Type="Italic">Salvelinus leucomaenis</Emphasis>, around the Lake Biwa water system varies with habitat size

Size at maturity of fluvial white-spotted charr, Salvelinus leucomaenis, was studied in small headwater tributaries of nine rivers around the Lake Biwa water system, Japan. Threshold size at maturity in both sexes showed significant positive relationships with water discharge, indicating that smaller threshold sizes at maturity of fluvial white-spotted charr occurred in smaller habitats. These results provide a link between size at maturity and habitat size and have important implications for the management of both habitats and white-spotted charr populations.     

Correlated contemporary evolution of life history traits in New Zealand Chinook salmon, Oncorhynchus tshawytscha

Size at age and age at maturity are important life history traits, affecting individual fitness and population demography. In salmon and other organisms, size and growth rate are commonly considered cues for maturation and thus age at maturity may or may not evolve independently of these features. Recent concerns surrounding the potential phenotypic and demographic responses of populations facing anthropogenic disturbances, such as climate change and harvest, place a premium on understanding the evolutionary genetic basis for evolution in size at age and age at maturity. In this study, we present the findings from a set of common-garden rearing experiments that empirically assess the heritable basis of phenotypic divergence in size at age and age at maturity in Chinook salmon (Oncorhynchus tshawytscha) populations introduced to New Zealand. We found consistent evidence of heritable differences among populations in both size at age and age at maturity, often corresponding to patterns observed in the wild. Populations diverged in size and growth profiles, even when accounting for eventual age at maturation. By contrast, most, but not all, cases of divergence in age at maturity were driven by the differences in size or growth rate rather than differences in the threshold relationship linking growth rate and probability of maturation. These findings help us understand how life histories may evolve through trait interactions in populations exposed to natural and anthropogenic disturbances, and how we might best detect such evolution.     

Rule of age and size at maturity: individual variation in the maturation history of resident white-spotted charr

Individual growth and maturation histories, age, and size at maturity of resident white-spotted charr Salvelinus leucomaenis were examined in a tag-recapture study in a natural river over 3 years. Slow-growing fish reached sexual maturity not only at an older age, but also at a smaller size than fast-growing fish, although females had a larger threshold size at maturity than males at each age. It is suggested that these patterns result from adaptive phenotypic plasticity that depends on individual growth conditions.     

The second population of Arctic charr Salvelinus alpinus complex (Salmoniformes, Salmonidae) in the Lake Baikal basin, the highest mountain charr population in Russia

We report the finding of the second population of Arctic charr in Lake Baikal basin, in a nameless lake in the Barguzin mountain range in the outlet of the Svetlaya River. The lake is situated at an altitude of 1766 m above sea level; this is the highest lake inhabited by this species in the territory of Russia. Two abundant charr forms, dwarf and small, were recorded in the lake; data on their ecology, morphology, age composition, growth, and microsatellite variation are presented. Both forms feed mainly on zooplankton, though neither has pronounced morphological traits characteristic of specialized plankton-feeders. Dwarf and small charrs are rather close in meristic characters and in the length of gill rakers, but differ substantially in allele frequencies and allele diversity at the microsatellite loci, which is indicative of a high degree of reproductive isolation between them. The discovered population is the only one among Transbaikalian charr populations, that, due to its remoteness, has not been affected by man. For that reason, it can be considered as an etalon of an undisturbed natural charr population. The lake and its surroundings should receive official protected status.     

Polymorphic segregation in Arctic charr Salvelinus alpinus (L.) from Vatnshlidarvatn, a shallow Icelandic lake

We studied the salmonid fish Arctic charr, Salvelinus alpinus , in a small and shallow landlocked lake in NW Iceland. The lake is productive but die only fish present is Arctic charr. Despite the apparent absence of discrete benthic and limnetic habitats for fish, two forms of Arctic charr are found in the lake. They show subde differences in morphology related to swimming performance and manoeuvrability, but differences in life history such as growth, and age and size at sexual maturation are more pronounced. Both forms have benthic feeding habits with one form consuming greater number of species than the other. We suggest that the segregation of these forms is based on the evolution of a specialist from a local generalist and that this has been made possible by the absence of a common fish competitor in similar lakes, the threespined stickleback Gasterosteous aculeatus.     

The population, survival, growth, reproduction and food of arctic charr, Salvelinus alpinus (L.), in four unexploited lakes in Greenland

The material comprised 1205 arctic charr caught by gillnets and electrofishing in the ice-free season 1982 in four interconnected lakes. The lakes were similar in biotic and abiotic factors and the arctic charr populations were therefore treated as one population. Two size groups of fish older than three years, called 'small charr' and 'large charr', were found. The two groups differed in feeding habits, growth rates, age of maturity and spawning frequencies. However, they did not differ in the frequencies of the F- and S- serum esterase alleles, and were in accord with the model proposed by Johnson (1976) which states that the 'small charr' and 'large charr' are two different forms of the same population. 'Small charr' recruit to the 'large charr' by entering a period of fast growth. In the investigated lakes this occurred when the 'small charr' were 3–10 years old. Some indications of rematuration of the 'small charr' when entering the group of 'large charr' were found.     

Reproduction and development in the dwarf form of Arctic charr Salvelinus alpinus from Lake Davatchan, Transbaikalia

Biological characteristics in the deepwater dwarf form of Arctic charr Salvelinus alpinus from Lake Davatchan, Transbaikalia, are described. In nine from fifteen attempts of artificial insemination of eggs from a female by sperm from a male, developing eggs are obtained. Embryonic and larval development is described at the initial temperature 1.6–3.6°C and subsequent (from early gastrulation to the transition of larvae to exogenous feeding) average temperature 4.4 and 8.6°C. To take into account the data obtained previously (Pavlov et al., 1990, 1993b), the comparison of early ontogeny is conducted between the dwarf and small forms of charr from Lake Davatchan and the charr from Lake Leprindokan (two lakes are separated by the ridge 8 km in width). A substantial similarity in early ontogeny between the dwarf and small forms of charr from Lake Davatchan suggests that genes controlling development of the embryos and prelarvae until the transition of the young to exogenous feeding have not been subjected to natural selection determining a considerable differentiation between two forms and their reproductive isolation. Published in Voprosy Ikhtiologii, 2008, Vol. 48, No. 1, pp. 102–119. The article was translated by the author.