Stream spawning of Arctic charr in Scotland

Abstract –  Scotland is a stronghold for Arctic charr, with about 200 freshwater, wholly loch-resident populations, most of which have yet to be studied. To date, no anadromous populations or individual sea-run charr, have been reported. In Scotland, most Arctic charr spawn in stillwater, during autumn and early winter (September to January), with only one population spawning in spring. Spawning in running water does occur, but has been regarded as rare. This paper examines the status of stream spawning of charr in Scotland, describing instances from 10 populations, although two of these are now extinct. Most stream-spawning charr migrate relatively short distances to running water and construct redds in gently flowing water. The longest recorded migration occurs in Loch Insh, Speyside, with fish travelling 15 km within the River Spey. Information on the extent of stream spawning is essential to help conserve the rich phenotypic and genetic diversity of our remaining charr populations.     

Divergent, genetically based feeding behaviour of two sympatric Arctic charr, Salvelinus alpinus (L.), morphs

Abstract –  Arctic charr, Salvelinus alpinus (L.), is one of the several northern fishes that show resource polymorphisms in postglacial lakes. Two reproductively isolated morphs of Arctic charr coexist in distinct ecological niches in the subarctic lake Fjellfrøsvatn, North Norway. Offspring of the two morphs (termed littoral charr and profundal charr) were reared separately but under identical conditions. Their feeding behaviour was compared experimentally using different kinds of live prey. The fishes had no experience with natural prey before the experiments. The littoral morph were more effective in eating live plankton ( Daphnia ) and littoral benthos ( Gammarus ), and had a higher attack rate against pleuston (surface prey, Gerris ) compared with the profundal morph. The two morphs behaved in accordance with expectations from their in situ niche utilisation towards the three prey types. This indicates a case of incipient ecological speciation where divergence in resource utilisation in contrasting niches has evolved adaptations in feeding behaviour by natural selection.     

Latitudinal variation in growth and otolith‐inferred field metabolic rates of Canadian young‐of‐the‐year Arctic charr

Countergradient variation (CGV) is defined as genetic variation that counteracts the negative influences of the physical environment, minimising phenotypic variability along an environmental gradient. CGV is thought to have relevance in predicting the response of organisms to climate variability and change. To test the hypothesis that growth rate increased with latitude, consistent with CGV, young‐of‐the‐year (YOY) Arctic charr, Salvelinus alpinus, were examined along a ~27° latitudinal gradient in central and eastern Canada. Growth rates were estimated from fork lengths standardised by the thermal opportunity for growth based on experienced water temperatures derived using otolith oxygen stable isotopes. Results demonstrated patterns consistent with CGV, where northern populations demonstrated faster growth rates. A secondary aim was to test for similar geographical patterns in otolith‐inferred metabolic rates, which reflect the energetic costs of standard metabolic rate (SMR) and other processes such as feeding, locomotion, thermoregulation, reproduction and growth. Results demonstrated a significant, positive relationship between otolith‐inferred metabolic rate and latitude, which may reflect an increase in one, or a combination, of the above‐noted physiological processes. The similar latitudinal pattern in growth and otolith‐inferred metabolic rates suggests greater intake of food per unit of time by northern fish. The phenotypic variation in physiological traits observed here demonstrates the significant adaptability of Arctic charr to different thermal regimes with different growing season lengths. Determining the relative contributions of phenotypic plasticity and genetic variation to the observed latitudinal variation will be critical to predicting the responses of Arctic charr to climate change more accurately.     

Atlantic salmon Salmo salar L., brown trout Salmo trutta L. and Arctic charr Salvelinus alpinus (L.): a review of aspects of their life histories

Abstract – Among the species in the family Salmonidae, those represented by the genera Salmo, Salvelinus, and Oncorhynchus (subfamily Salmoninae) are the most studied. Here, various aspects of phenotypic and life‐history variation of Atlantic salmon Salmo salar L., brown trout Salmo trutta L., and Arctic charr Salvelinus alpinus (L.) are reviewed. While many strategies and tactics are commonly used by these species, there are also differences in their ecology and population dynamics that result in a variety of interesting and diverse topics that are challenging for future research. Atlantic salmon display considerable phenotypic plasticity and variability in life‐history characters ranging from fully freshwater resident forms, where females can mature at approximately 10 cm in length, to anadromous populations characterised by 3–5 sea‐winter (5SW) salmon. Even within simple 1SW populations, 20 or more spawning life‐history types can be identified. Juveniles in freshwater can use both fluvial and lacustrine habitats for rearing, and while most smolts migrate to sea during the spring, fall migrations occur in some populations. At sea, some salmon undertake extensive oceanic migrations while other populations stay within the geographical confines of areas such as the Baltic Sea. At the other extreme are those that reside in estuaries and return to freshwater to spawn after spending only a few months at sea. The review of information on the diversity of life‐history forms is related to conservation aspects associated with Atlantic salmon populations and current trends in abundance and survival. Brown trout is indigenous to Europe, North Africa and western Asia, but was introduced into at least 24 countries outside Europe and now has a world‐wide distribution. It exploits both fresh and salt waters for feeding and spawning (brackish), and populations are often partially migratory. One part of the population leaves and feeds elsewhere, while another part stays as residents. In large, complex systems, the species is polymorphic with different size morphs in the various parts of the habitat. Brown trout feed close to the surface and near shore, but large individuals may move far offshore. The species exhibits ontogenetic niche shifts partly related to size and partly to developmental rate. They switch when the amount of surplus energy available for growth becomes small with fast growers being younger and smaller fish than slow growers. Brown trout is an opportunistic carnivore, but individuals specialise at least temporarily on particular food items; insect larvae are important for the young in streams, while littoral epibenthos in lakes and fish are most important for large trout. The sexes differ in resource use and size. Females are more inclined than males to become migratory and feed in pelagic waters. Males exploit running water, near‐shore and surface waters more than females. Therefore, females feed more on zooplankton and exhibit a more uniform phenotype than males. The Arctic charr is the northernmost freshwater fish on earth, with a circumpolar distribution in the Holarctic that matches the last glaciation. Recent mtDNA studies indicate that there are five phylogeographic lineages (Atlantic, Arctic, Bering, Siberian and Acadian) that may be of Pleistocene origin. Phenotypic expression and ecology are more variable in charr than in most fish. Weights at maturation range from 3 g to 12 kg. Population differences in morphology and coloration are large and can have some genetic basis. Charr live in streams, at sea and in all habitats of oligotrophic lakes, including very deep areas. Ontogenetic habitat shifts between lacustrine habitats are common. The charr feed on all major prey types of streams, lakes and near‐shore marine habitats, but has high niche flexibility in competition. Cannibalism is expressed in several cases, and can be important for developing and maintaining bimodal size distributions. Anadromy is found in the northern part of its range and involves about 40, but sometimes more days in the sea. All charr overwinter in freshwater. Partial migration is common, but the degree of anadromy varies greatly among populations. The food at sea includes zooplankton and pelagic fish, but also epibenthos. Polymorphism and sympatric morphs are much studied. As a prominent fish of glaciated lakes, charr is an important species for studying ecological speciation by the combination of field studies and experiments, particularly in the fields of morphometric heterochrony and comparative behaviour.     

Patterns of phenotypic and genetic variability show hidden diversity in Scottish Arctic charr

Abstract –  This study examined the degree and pattern of variability in trophic morphology in Arctic charr ( Salvelinus alpinus L. ) at three spatial scales: across 22 populations from Scotland and between and within two adjacent catchments (Laxford and Shin) in northern Scotland. In addition, the variability at six microsatellite loci between and within the Laxford and Shin systems was determined. Habitat use by charr differed significantly between populations. The pattern of variability in trophic morphology, known to influence foraging ability in charr, showed a very high degree of between-population variation with at least 52% of population pairs showing significant differences in head shape. Trophic morphology and genetic variation was also high over small geographical scales; variation being as high between charr from lakes within the same catchment, as between adjacent catchments. The pattern of both phenotypic and genotypic variation suggests a mosaic of variation across populations with geographically close populations often as distinct from each other as populations with much greater separation. Very low levels of effective migrants between populations, even within the same catchment, suggest that this variation is being maintained by very low straying rates between phenotypically and genetically distinct populations, even when there is no apparent barrier to movement. We conclude that the genetic and phenotypic integrity of charr populations across Scotland is high and that this adaptive radiation constitutes a 'hidden' element of diversity in northern freshwater systems. Two consequences of this are that the population (rather than the species) makes a more rational unit for the consideration of conservation strategies and that the habitat requirements and therefore management needs may differ significantly between populations.     

The Welsh Torgoch (Salvelinus alpinus): a short review of its distribution and ecology

Abstract –  The Arctic charr Salvelinus alpinus (L.), or Torgoch ('red belly') in Welsh, is found in seven lakes in North Wales comprising three native populations (Bodlyn, Cwellyn and Padarn) and four translocated populations (Cowlyd, Diwaunedd, Dulyn and Ffynnon Llugwy). The Welsh charr populations have not been studied much, but the available data are summarised in this short review. Analysis of stomach contents of the native charr populations shows that they are planktivores feeding mainly on cladocerans and copepods. Adult fish attain a size of 14–24 cm (fork length) and fecundity is low (100–800 eggs per female). The available growth data reveal that Welsh charr grow at a similar rate and attain a size similar to most other planktivorous charr populations that have been studied in the UK. Genetic studies using allozymes indicate that the three native populations are distinct from each other.     

Assessing the food web impacts of an anadromous Arctic charr introduction to a sub‐Arctic watershed using stable isotopes

Anadromous Arctic charr, Salvelinus alpinus (L.), was introduced to a sub‐Arctic river–lake system near the village of Kujjuuaq, Nunavik, and the stable isotope values and diets of key resident fish species were used to assess changes in feeding patterns. Stable isotope values for most species did not differ significantly between the pre‐ and post‐introduction periods, with observed shifts being within the bounds of expected natural variation. Lake chub, Couesius plumbeus (Agassiz), were the single species to show a difference between study periods, with a small but significant increase in δ¹⁵N. No significant post‐introduction changes were seen in lake trout, Salvelinus namaycush (Walbaum), omnivory or in any of the assessed quantitative food web metrics. Gut contents of major fish species similarly showed significant temporal overlap between the pre‐ and post‐introduction periods, and there was no significant change in species' weight–length relationships. The minor ecological impact was interpreted in relation to the availability of open niches exploitable by ecological generalists such as Arctic charr. The explanation accords with the known habitat and feeding flexibility of Arctic charr and the ecological immaturity of sub‐Arctic lakes known to have driven adaptive variation among Arctic charr. Findings suggest that anadromous Arctic charr may be introduced at moderate densities to other sub‐Arctic watersheds without major negative food web consequences for other resident fish species.     

An experimental comparison of cannibalistic response in different Arctic charr (Salvelinus alpinus (L.)) stocks

Abstract– The cannibalistic response of four domesticated Arctic charr stocks were compared experimentally using a radiographic technique. Charr from Arctic Spitsbergen (80°N) consistently exhibited a much higher frequency of cannibalistic response than fish from three charr strains from mainland Norway (70°N). The results indicate that the extent of cannibalistic feeding in Arctic charr is population specific and support the hypothesis that charr from Arctic populations have stronger cannibalistic tendencies than do fish from more temperate regions.     

Conservation and management of the Arctic charr: a forward view

Abstract –  In this study, we synthesised the views of 34 participants in a workshop to consider the status and future conservation and management of the Arctic charr ( Salvelinus alpinus L.). These are expressed as a series of resolutions deriving from a conference on Arctic charr conservation held in Perth, Scotland in June 2004. Agreed resolutions from the conference were as follows. (i) The unique diversity of Arctic charr should be recognised for the contribution it makes to biodiversity of northern aquatic communities. (ii) The recognition by the public, nongovernmental organisations and national governments of the importance of Arctic charr in northern ecosystems is an important step to the management and protection that should be pursued. (iii) The taxonomic status of highly variable charr populations requires to be reviewed. (iv) There should be a separate system, complementary to the taxonomic one, which systematically catalogues the biological diversity of S. alpinus . (v) The Arctic charr provides a unique natural resource to study the process of evolution and this requires the highest level of protection from anthropogenic effects. (vi) The status of conservation policy for Arctic charr needs to be urgently reviewed to take account of its unique position in the fauna.     

Census and effective population sizes of white‐spotted charr (Salvelinus leucomaenis) in a fragmented landscape

Several habitat models have been proposed to predict population size for stream fishes and to guide habitat assessment and monitoring techniques. However, most models do not incorporate the potential advantage of molecular genetic markers. We conducted a field survey and microsatellite DNA analyses to quantify the relationships among genetic diversity, census/effective population size and habitat variables in fragmented populations of white‐spotted charr (Salvelinus leucomaenis). The census population size significantly increased with the stream length, the number of pools and a pool‐riffle sequence index, a proxy for channel‐unit habitat type complexity within reaches. Population density was correlated with the pool‐riffle sequence index only. Genetic diversity and effective population size were not correlated with the habitat variables or census population size. There was a lack of isolation‐by‐distance population structure in the studied populations. Our results suggest that stream length and the number of pools within reaches associated with habitat complexity are the habitat variables that explain the majority of variation in population size of white‐spotted charr. Our findings provide further evidence that census population size per se is a poor indicator of the inclusive genetic diversity within populations in a fragmented landscape.     

Dietary plasticity of Arctic charr (Salvelinus alpinus) facilitates coexistence with competitively superior European whitefish (Coregonus lavaretus)

Abstract – Habitat use and diet of Arctic charr (Salvelinus alpinus) coexisting with European whitefish (Coregonus lavaretus) and grayling (Thymallus thymallus) were studied in one deep and two relatively shallow subarctic lakes in northern Norway. Stomach content and stable isotope analyses revealed clear and temporally stable resource partitioning between the species in all three lakes. Arctic charr had a wide and flexible trophic niche and was the only piscivorous species. In contrast, whitefish and grayling had remarkably stable planktivorous and benthivorous niches, respectively. In the deepest lake, Arctic charr together with grayling mainly utilised littoral benthos, while piscivory was more prevalent in Arctic charr in the two shallower lakes. In one of the shallow lakes, whitefish was apparently relegated to the inferior profundal niche because of dominance of the littoral by grayling. Our results suggest that Arctic charr may not necessarily need an extensive profundal zone as a refuge, but can coexist with whitefish if a third competing fish species like grayling occurs in the littoral habitat or if profitable small prey fish are available. The study demonstrates that strong dietary plasticity of Arctic charr is instrumental in the observed coexistence with the commonly competitively superior whitefish.     

Dietary plasticity in Arctic charr (Salvelinus alpinus) in response to long‐term environmental change

Corrigan LJ, Winfield IJ, Hoelzel AR, Lucas MC. Dietary plasticity in Arctic charr (Salvelinus alpinus) in response to long‐term environmental change.
Ecology of Freshwater Fish 2011: 20: 5–13. © 2010 John Wiley & Sons A/S Abstract –  In the face of widespread environmental change impacts, there is a need to better understand mechanisms promoting flexibility and resilience of ecosystem components to such change to inform strategies for conservation. Glacial relict species are especially vulnerable to such changes. We investigated the behavioural responses of a native, glacial relict species, Arctic charr (Salvelinus alpinus) to long‐term environmental changes. It was hypothesised that changes in feeding behaviour would occur as a key intermediary to reduction in habitat availability (through climate change and eutrophication) or competitive interactions [with introduced roach (Rutilus rutilus)]. Stomach content analysis was used to assess the diet of 199 charr caught from Windermere, United Kingdom, in the months of March, June, September and November 2003–2007. The results were compared to data from 1940 to 1951 prior to the environmental changes and revealed a marked increase in the contribution of benthic invertebrates in the present diet. Stable isotope analysis confirms the results of the stomach analysis, suggesting that the charr have switched their diet from zooplanktivory towards benthivory. We discuss the possibility that habitat modification and roach population expansion in Windermere have contributed to changes in charr diet. Complementary studies suggest that this diet shift is more likely to be a response to the increasing roach population than to habitat modification; however, further study in Windermere would be required to elucidate the exact mechanism. Long‐term data sets such as these provide information that is useful for determining the directivity of ecological change and the capability of species to respond to change.     

Using phenotypic variation to determine conservation value: application of a novel approach to Arctic charr

Abstract –  There is a very high degree of discrete variation in phenotype between populations of Arctic charr. This takes the form of variation not only in morphometric and meristic characters traditionally used to distinguish species, but also in characteristics of life-history, behaviour, coloration and ecology. This variability has a number of consequences, one of these is that there is a strong case for the conservation of populations with extreme phenotypes. However, if variation is discrete between populations but continuous across many populations, this poses difficulties in separating those populations of high conservation value from those of lower conservation value. In this paper we describe a statistical technique which enables populations on the extreme edges of the range of phenotypic variation to be identified and apply this to the morphometric characters of charr from 25 populations from across Scotland and Ireland. The technique enables the identification of any proportion of the most extreme phenotypes. When applied to our data, one population (Loch More) was in the top 2 percentile of the most extreme phenotypes from across the range of all populations included. Three populations were within the top 10% most extreme phenotypes (Lochs More, Uaine and Earn) and a further five within the top 20% (previously mentioned lochs plus Lochs Eck, Merkland, Uaine, Talla and Lough Nalughraman). This technique can potentially be used on any species and on any suite of characteristics as an objective measure of conservation value of a population within a continuous phenotypic range.     

Resource partitioning between lake-dwelling Atlantic salmon (Salmo salar L.) parr, brown trout (Salmo trutta L.) and Arctic charr (Salvelinus alpinus (L.))

Abstract – Resource partitioning between Atlantic salmon parr, brown trout and Arctic charr was studied throughout the ice-free season in a north Norwegian lake. Juvenile salmon and trout (≤160 mm) utilized the littoral zone and juvenile charr the profundal, while adult trout and charr (>160 mm) were found in both. Juvenile salmon and trout had a similar diet, although trichopteran larvae were more important for the trout and chironomid pupae and three-spined sticklebacks for the salmon parr. Small salmon and trout parr (≤120 mm) had a higher diet overlap than larger parr (121–160 mm). The feeding habits of adult trout were similar to that of juvenile trout, but the former took larger prey items. At the population level, both salmon and trout were generalistic feeders with a broad diet, but at the individual level, both species had specialized on a single or a few prey categories. Juvenile charr were segregated from salmon and trout in both habitat and food utilization; they had a narrow diet consisting of chironomids and zooplankton, possibly reflecting their confinement to the profundal habitat which have a low diversity of potential prey. Larger charr also took zoobenthos and sticklebacks in the littoral zone. ^Note     

Arctic charr in Britain and Ireland – 15 species or one?

Abstract –  From both a modern and a historical perspective there is little doubt that the Arctic charr, Salvelinus alpinus , in Britain and Ireland (as well as elsewhere) is a 'difficult' species. Historically 15 separate species have been recognised from populations in Britain and Ireland and there have been recent attempts to reassert these specific names. Here we review the evidence for the status of these 'species'. We conclude that the evidence for these 15 being afforded full species status is poor. However, both historical and contemporary data show that Salvelinus alpinus (Linnaeus 1758) in Britain and Ireland exhibits levels of variability in form that is much greater than in other species. We argue that a comprehensive genotypic and phenotypic survey of charr populations in Britain and Ireland is required to determine the full extent of variability and the status of populations with a view of providing suitable protection.     

Atlantic cod recovery from the Allee effect zone: contrasting ecological and evolutionary rescue

The ability of a population to recover from disturbances is fundamental for its persistence. Impaired population recovery might be associated with a demographic Allee effect. Immigration from adjacent populations could accelerate the recovery not only by promoting population growth beyond the Allee effect threshold but also by bringing in advantageous genotypes. We explore the nature and role of ecological and evolutionary rescue in an Atlantic cod (Gadus morhua Gadidae) population fished below its Allee effect threshold. We utilize an eco‐evolutionary model and simulate scenarios, where the target population evolves in response to selective fishing and sample immigrants from (a) a source population similarly adapted to fishing (post‐fishing genotypes) or (b) an unexploited source population with natural genetic and phenotypic diversity (pre‐fishing genotypes). Immigration of pre‐fishing genotypes enhances the recovery due to the larger body sizes and the flow of associated genes. Post‐fishing immigrants can also promote the population abundance recovery, but they increase uncertainty about recovery times as compared to pre‐fishing immigrants and do not promote evolutionary recovery. Our results stress the importance of maintaining genetic and phenotypic diversity and suggest that marine protected areas can serve as an important source of immigrants to promote both the demographic and evolutionary recovery of exploited populations.     

Does morphological variation between young‐of‐the‐year perch from two Swedish lakes depend on genetic differences?

Heynen M, Hellström G, Magnhagen C, Borcherding J. Does morphological variation between young‐of‐the‐year perch from two Swedish lakes depend on genetic differences?
Ecology of Freshwater Fish 2010: 19: 163–169. © 2009 John Wiley & Sons A/S Abstract – Different local environmental conditions have often been found to generate phenotypic diversity. In the present study we examined morphological differences between young‐of‐the‐year perch from two lake populations with differences in size‐specific predation risk. A common garden setup was used to examine the genetic and environmental components of the morphological variation. We found differences in head and jaw length and slight differences in body depth between the wild young‐of‐the‐year perch from Lake Ängersjön and Lake Fisksjön. The differences found between the wild fish from the two lakes were not maintained under common garden rearing. The observed morphological divergence between the wild young‐of‐the‐year perch from Lake Ängersjön and Lake Fisksjön seems to stem mainly from a plastic response to different environmental conditions in the two lakes. It is clear that the morphological traits are not influenced by direct reaction to the size‐specific risk of cannibalism, but probably stem from a combination of different environment characteristics, including resource and habitat use, and the density of other piscivores, such as pike.     

Foraging mode variation in three stream‐dwelling salmonid fishes

Despite long‐standing interest in foraging modes as an important element of animal space use, few studies document and compare individual foraging mode differences among species and ecological conditions in the wild. We observed and compared foraging modes of 61 wild Arctic charr, Salvelinus alpinus, 42 brown trout, Salmo trutta, and 50 Atlantic salmon, Salmo salar, in their first growing season over a range of habitats in 10 Icelandic streams. We found that although stream salmonids typically sit‐and‐wait to ambush prey from short distances, Arctic charr were more mobile during prey search and prior to prey attack than Atlantic salmon, whereas brown trout were intermediate. In all three species, individuals that were mobile during search were more likely to be moving when initiating attacks on prey, although the strength and the slope of this relationship differed among species. Arctic charr also differed from salmon and trout as more mobile individuals travelled longer distances during prey pursuits. Finally, coupled with published data from the literature, salmonid foraging mobility (both during search and prior to attack) clearly decreased from still water habitats (e.g., brook charr), to slow‐running waters (e.g., Arctic charr) to fast‐running waters (e.g., Atlantic salmon). Hence, our study suggests that foraging mode of young salmonids can vary distinctly among related species and furthers our understanding of the behavioural mechanisms shaping the geographical distribution of wild salmonids.     

Feeding tactics and body condition of two introduced populations of pumpkinseed Lepomis gibbosus: taking advantages of human disturbances?

Abstract –  Feeding tactics, body condition and size structure of two populations of pumpkinseed Lepomis gibbosus from Cabañeros National Park (Guadiana River basin, central Spain) were compared to provide insight into the ecological requirements favouring levels of success/failure in relation to human intervention. Habitat, benthic macroinvertebrates and pumpkinseed were quantified in Bullaque (regulated flow, affected by agricultural activities) and Estena (natural conditions) rivers, from May to September of 2005 and 2006. Significant differences were found in the limnological characteristics between the two rivers. Spatial and temporal variations in diet composition were likely related to opportunistic feeding and high foraging plasticity. Diet diversity was higher in Bullaque River. Electivity of benthic prey showed variation between sized individuals and populations. The body condition of pumpkinseed was better in Bullaque River. Current results show that this invasive species is favoured by human intervention in Bullaque River.