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.     

Inherited differences in head allometry in polymorphic Arctic charr from Loch Rannoch, Scotland

Planktivorous and benthivorous morphs of wild Arctic charr Salelinus alpinus from Loch Rannoch, Perthshire, Scotland, reared in the laboratory from artificially fertilized eggs under identical conditions, were morphologically distinct (based on a series of head measurements) from an early age. For some morphometric characters, these differences became more marked with increasing body size, reflecting differences in the allometric growth patterns of the two forms. These data show that the observed phenotypic differences between these two trophic variants were, to some extent at least, inherited. Thus it is concluded that the mechanisms regulating expression of phenotype in the sympatric morphs of Loch Rannoch are significantly different from the environmentally determined body size polymorphisms reported from Norway but are closer to the more completely divergent forms from Thingvalavatn, Iceland.     

Trophic polymorphism amongst Arctic charr from Loch Rannoch, Scotland

This paper describes the results of a multivariate and univariate morphometric analysis of three groups of Arctic charr Salvelinus alpinus from Loch Rannoch, Perthshire, Scotland, distinguished a priori on the basis of coloration (bright v. cryptic) and site of capture (from two locations 12 km apart). The analysis produced a clear and highly significant distinction, not only between brightly and cryptically coloured charr, but also between the two cryptically coloured groups, one of which was more robust in terms of several size-independent head measurements. Stomach analysis showed that the brightly coloured charr fed entirely on zooplankton, the less robust, cryptically coloured fish, fed on benthic macroinvertebrates, whereas a significant proportion of the diet of the more robust cryptically-coloured form consisted of other fish. Differences in length at age also distinguished the three forms, with the robust, piscivorous charr, which live longer and potentially reach a larger final size, attaining smaller sizes at a given age. These data clearly confirm the previous identification of a distinct planktivorous morph of Arctic charr in Loch Rannoch and extend the morph analysis by distinguishing two additional morphs, a benthivorous morph and a piscivorous morph that are morphologically and ecologically distinct. The results are discussed in the context of other systems in which sympatric morphs of Arctic charr have been described.     

Behavioural responses of benthic and pelagic Arctic charr to substratum heterogeneity

Responses to environmental heterogeneity were studied in laboratory-reared offspring of two morphs of Arctic charr Salvelinus alpinus from Loch Rannoch, Scotland, one occupying the pelagic zone and feeding predominantly on zooplankton and the other being benthic in habit and feeding mainly on macroinvertebrates. When housed in groups in tanks with a black-and white striped base, benthic charr demonstrated a clear preference for dark areas, whereas pelagic fish positioned themselves at random with respect to substratum colour. In general, pelagic charr were much less aggressive than benthic charr. In pelagic fish, neither spacing nor aggression was affected by the visual heterogeneity of the substrate. In contrast, benthic charr swam closer together and fought more when housed over a uniform as opposed to a non-uniform substratum. The results are discussed in the context of habitat-specific visual requirements and of an interaction between visual complexity and territoriality previously described for Atlantic salmon Salmo salar .     

A morphometric method for estimation of total lipid level in live Arctic charr: a case study of its application on wild fish

Linear body measurements were made on wild Arctic charr Salvelinus alpinus parr (100–200 mm L_F) from two populations in northern Norway during spring (May), summer (June and July) and autumn (October). To reduce handling stress, a method was developed were dorsal and ventral body measurements could subsequently be taken easily from a single picture of the anaesthetized fish. Multiple regression analyses investigated the possible correlation between the body measurements and both total and percentage lipid content of the fish. For both populations and during all seasons, the regression equations gave better estimates for total lipid than for percentage lipid. The regression equations for total lipid accounted for between 67·7 and 89·5% of the variance in lipid content for the different seasons within the lakes. The pooled data within each lake accounted for 62·9 and 81·0% of the variance in total lipid, while the pooled data for both lakes accounted for 67·7% of the variance. In general, the condition factor alone did not give a satisfactory estimate of lipid content of the Arctic charr (r² = 0·003–0·521). Shrinkage on validation values was high (0·20–0·52). Employing a ridge regression method resulted in models with lower r² values and lower shrinkage values (indicating more stable models). Published equations used for hatchery‐reared Arctic charr could not be used on wild fish from the same population. The close correlations between actual and predicted lipid levels found in this study show that morphometric measurements can predict total lipid levels in wild Arctic charr with reasonable accuracy. The most accurate results were obtained when equations were derived from each data set. Therefore, the method has the potential within a single study to estimate lipid levels in live fish as long as some fish can be sacrificed to develop a unique regression equation for each population or experiment.     

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.     

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.     

Chromosomes and constitutive heterochromatin distribution in Arctic charr,Salvelinus alpinus (L.) (Pisces: Salmonidae)

The chromosomes, together with their C and Q banding patterns, of Arctic charr from Loch Rannoch, Scotland are described. Q banding revealed fewer bands than C banding and variation between individuals for the number of C bands was found. These results suggest that subsets of heterochromatin exist and that there may be an individual variation in DNA content.     

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.     

Body composition in baboons: Evaluating a morphometric method

The objective of this study was to determine whether noninvasive morphometric measurements of olive baboons (Papio cynocephalus anubis) can reliably predict lean body mass and fat mass in this species. Crown-rump length, triceps circumference, and skinfold measures at the neck, subscapular, suprailiac, and triceps sites were obtained prior to necropsy from 21 clinically normal infant baboons at 18 weeks of age and from 22 clinically normal adolescent baboons at 5 years of age. At necropsy, the lean body mass and body fat mass were measured using gravimetric methods. Pearson's correlations and regression analysis were used to test the relationship between direct measures of lean and fat mass obtained at necropsy with calculated estimates based on morphometric measures obtained prior to death. Our null hypothesis was that the morphometric measures (individually or in combination) were not related to direct measures of fat mass or lean mass. Non-linear and multiple regression estimators, by age and gender, were derived and provided increased predictability. Our results indicate that (1) morphometric measures can accurately predict lean body mass in male and female baboons; (2) morphometric measures used to predict lean body mass change with age; (3) morphometric measures are strongly associated with body fat mass at 18 weeks of age but are not as strongly associated with body fat mass in 5-year-old baboons; (4) triceps circumference provides the best single indicator of lean body mass for both genders and age periods; (5) baboons are like humans in that adolescent females tend to accumulate body fat while males of the same age tend to develop lean mass; and (6) combinations of these morphometric measurements explain between 70% and 100% of the variability and can be used to estimate lean and fat mass in baboons.     

Effect of diets enriched in Delta6 desaturated fatty acids (18:3n-6 and 18:4n-3), on growth, fatty acid composition and highly unsaturated fatty acid synthesis in two populations of Arctic charr (Salvelinus alpinus L.)

This study aimed to test the hypothesis that diets containing relatively high amounts of the Delta6 desaturated fatty acids stearidonic acid (STA, 18:4n-3) and gamma-linolenic acid (GLA, 18:3n-6), may be beneficial in salmonid culture. The rationale being that STA and GLA would be better substrates for highly unsaturated fatty acid (HUFA) synthesis as their conversion does not require the activity of the reputed rate-limiting enzyme, fatty acid Delta6 desaturase. Duplicate groups of two Arctic charr (Salvelinus alpinus L.) populations with different feeding habits, that had been reported previously to show differences in HUFA biosynthetic capacity, were fed for 16 weeks on two fish meal based diets containing 47% protein and 21% lipid differing only in the added lipid component, which was either fish oil (FO) or echium oil (EO). Dietary EO had no detrimental effect on growth performance and feed efficiency, mortalities, or liver and flesh lipid contents in either population. The proportions of 18:2n-6, 18:3n-3, 18:3n-6, 18:4n-3, 20:3n-6 and 20:4n-3 in total lipid in both liver and flesh were increased by dietary EO in both populations. However, the percentages of 20:5n-3 and 22:6n-3 were reduced by EO in both liver and flesh in both strains, whereas 20:4n-6 was only significantly reduced in flesh. In fish fed FO, HUFA synthesis from both [1-(14)C]18:3n-3 and [1-(14)C]20:5n-3 was significantly higher in the planktonivorous Coulin charr compared to the demersal, piscivorous Rannoch charr morph. However, HUFA synthesis was increased by EO in Rannoch charr, but not in Coulin charr. In conclusion, dietary EO had differential effects in the two populations of charr, with HUFA synthesis only stimulated by EO in the piscivorous Rannoch morph, which showed lower activities in fish fed FO. However, the hypothesis was not proved as, irrespective of the activity of the HUFA synthesis pathway in either population, feeding EO resulted in decreased tissue levels of n-3HUFA and 20:4n-6. This has been observed previously in salmonids fed vegetable oils, and thus the increased levels of Delta6 desaturated fatty acids in EO did not effectively compensate for the lack of dietary HUFA.     

State-dependent feed acquisition among two strains of hatchery-reared Arctic charr

The effects of genetic and environmental factors on aggression and feeding hierarchies were studied using X-radiography to measure food intake by hatchery-reared individuals of two strains (Hammerfest and Svalbard) of Arctic charr Salvelinus alpinus . A reduction in food rations and/or water current speed increased intraspecific aggression, and both factors led to increased interindividual variability in food intake, increasing the coefficient of variation (CV). Following a return to pre-manipulation conditions, CVs decreased to their original level. In control groups, CVs and share of group meals were stable throughout the experiment. The increase in CVs following manipulation was the result of a small number of dominant individuals obtaining a high share of the meal. Restriction in food ration affected share of meals, specific growth rates and the frequencies of non-feeding fish, while reductions in water current speed affected only share of meals. Feeding hierarchies were size-dependent in the control groups. In contrast, no relationships between body weight and feeding rank were evident in groups in which food ration or water current speed were reduced. A small, but consistent, difference was revealed in feeding hierarchy responses between the two strains.     

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.     

Seasonal variation and sex differences in the nutritional status in two local populations of wild Japanese macaques

Seasonal variations and sex differences in the nutritional status in two local populations of wild Japanese macaques (Macaca fuscata fuscata) were examined. It was hypothesized that the ecological condition and/or reproductive strategies of each sex determine the nutritional condition and its seasonal fluctuation in each sex. Morphometric measures such as body mass, thoracic and femoris circumferences, skinfold thickness in four places (triceps, biceps femoris, subscapular, and abdomen), and wet mass of mesenteric and omental fat were used for comparisons between sexes, seasons, and populations. Animals of the Shimane population were larger than those of the Boso population in most morphometric measures, abdominal skinfold, and mesenteric and omental fat mass, suggesting environmental and/or genetic differences in the two populations. Females of both populations had larger skinfolds and mesenteric and omental fat mass than males, indicating that females had more fat than males. Females showed seasonality in most measures, having two peaks of body mass, thoracic and femoris circumferences, abdominal skinfold, and mesenteric and omental fat masses in early spring and late fall. In contrast, males exhibited no clear seasonal variations for most measurements, except for biceps femoris and subscapular skinfolds, which showed peaks in summer. Most morphometric measurements significantly correlated to each other, particularly in females, but most skinfolds had no correlation with other measurements. These findings suggest that sexual dimorphism in body composition and its fluctuation may be affected by the different reproductive strategies of males and females.     

Variation in scale shape among alternative sympatric phenotypes of Arctic charr Salvelinus alpinus from two lakes in Scotland

Landmark‐based geometric morphometric analysis was used to detect differences in scale shape between ecologically distinct phenotypes of Arctic charr Salvelinus alpinus coexisting in the same lake. Relative warp analysis and standard multivariate analyses of the partial warps, obtained after a Procrustes superimposition, showed that scale landmarks were efficient in discriminating among two closely related alternative phenotypes within each of the two lakes. In Loch Tay, S. alpinus exhibited a bimodal body size‐frequency distribution among sexually mature fish, whereas in Loch Awe, S. alpinus are unimodal in body size but segregated into two distinct spawning phenotypes. In both lakes, alternative phenotypes showed significant differences in foraging ecology, habitat use and life history. It is probable that differences in scale shape reflect differences in ecology of these forms.     

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.     

An evaluation of condition indices and predictive models for noninvasive estimates of lipid mass of migrating Common Yellowthroats, Ovenbirds, and Swainson's Thrushes

ABSTRACT.   Noninvasive methods of measuring lipid mass in birds are widely used, but not frequently evaluated. I evaluated the ability of three noninvasive indicators of fat content (fat scores, body mass, body mass/wing chord) and regression models to predict lipid mass in two migratory songbirds previously unexamined in this context—Common Yellowthroat ( Geothlypis trichas ) and Ovenbird ( Seiurus aurocapillus ). I also examined the accuracy of these methods for Swainson's Thrushes ( Catharus ustulatus ) for comparison to a previous study. Fat score, body mass, and body mass/wing chord were highly correlated with chemically extracted lipid mass in each species. In all three species, birds with no visible subcutaneous fat possessed considerable quantities of fat, ranging from 9.8 to 19.7% of total dry body mass. Forward-selected regression models explained 69−87% of lipid mass variation, with prediction errors of 14.6−27.5%. An existing predictive model for the Swainson's Thrush overestimated lipid mass by an average of 92%. Fat score, body mass, and the regression models generated here are reliable predictors of lipid mass in two of the three migrating species examined. The accuracy of the methods, in addition to their low cost and simplicity, justifies their continued use in field studies of birds.     

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, ?? , ??–??.     

Differences in susceptibility of anadromous and resident stocks of Arctic charr to infections of Gyrodactylus salaris, under experimental conditions

The ability of Gyrodactylus salaris , an important pathogen of the Atlantic salmon Salmo salar , in Norway, to infect anadromous and resident stocks of the Arctic charr, Salvelinus alpinus , has been examined in the laboratory. Resident charr (Korssjoen stock) exposed to heavily infected salmon, were considered innately resistant as they lost their infections within 21 days when individually isolated. Isolated anadromous harr (Hammerfest stock) remained infected for up to 150 days, although most infections disappeared within 30–50 days. In many cases the parasite population grew initially, but growth was limited after 20–30 days and infections subsequently disappeared. At the same time, shoals of 50 anadromous charr, swimming in the tanks containing the individually isolated fish in floating cages, remained infected for up to 280 days. Charr isolated from these shoals after 115 days and subsequently monitored individually lost their infections within 30 days, although the parasite persisted within the shoals for a further 75–135 days. This suggests that G. salaris , persisted on shoaling charr despite an immune response which led to the elimination of parasites from isolated hosts. The Hammerfest stock of anadromous charr supports G. salaris , in the laboratory, and the extended period of survival on this host suggests that charr may be important in the epidemiology of G. salaris , in northern Norway.     

Relationships among body composition measures in community-dwelling older women

Objective: To examine whether simple anthropometric measures provide a good estimate of total and visceral fat in 146 community‐dwelling, older white women (mean age, 74.0 ± 4.1 years). Research Methods and Procedures: Total body fat and visceral fat were measured using electron beam computed tomography (EBT). Anthropometric parameters (height, weight, BMI, sagittal diameter, and waist circumference) were measured using standard techniques. Total percentage body fat was assessed using DXA. Spearman correlations were used to examine the association between the measures. Linear regression, controlling for age, was used to examine the associations between the anthropometric parameters and total and visceral body fat measured by EBT. Results: Correlations among body composition measures ranged from ρ = 0.46 to 0.93 (p < 0.0001). EBT total fat was strongly correlated with both DXA estimates of total percentage fat (ρ = 0.86) and BMI (ρ = 0.89). Separate linear regression models indicated that BMI, waist circumference, sagittal diameter, and DXA total percentage fat were each independently related to EBT total fat. BMI had the strongest linear relationship, explaining 80% of the model variance (p < 0.0001). Linear regression indicated that BMI, waist circumference, and sagittal diameter were each independently related to EBT visceral fat, with BMI and sagittal diameter explaining ～53% of the model variance (p < 0.0001). Discussion: The use of simple anthropometric measures such as BMI, sagittal diameter, and waist circumference may be an appropriate alternative for more expensive techniques when assessing total fat but should be used with caution when estimating visceral body fat.