Water temperature, light intensity and zooplankton density and the feeding activity of juvenile brook charr (Salvelinus fontinalis)

SUMMARY 1. The objective of this study was to evaluate the effects of zooplankton biomass (as a measure of density), fish biomass, light intensity and water temperature on the attack rate and swimming characteristics (i.e. swimming speed and angle of turn) of juvenile (1+) brook charr (Salvelinus fontinalis) in field enclosures. We used a portable underwater camera system in a series of pelagic enclosures to quantify the feeding behaviour of brook charr over a gradient of natural conditions. 2. In simple linear or non‐linear regression models we found (i) that attack rate and angle of turn were positively related to water temperature, (ii) that attack rate and swimming speed were positively related to zooplankton biomass and light intensity and (iii) that attack rate was positively related to swimming speed. In multiple regression models, fish biomass, light intensity and variance of the angle of turn accounted for 87% of the variation in attack rate. Light intensity and water temperature accounted for 86% of the variation in swimming speed. Fish gut fullness and attack rate accounted for 83% of the variation in the variance of the angle of turn executed by fish. 3. The increase in the number of attacks as zooplankton biomass increases conforms to the general positive functional response observed in other fish species. Our results also support the hypothesis that swimming speed increases with prey biomass. We did not observe a plateau in attack rate as zooplankton biomass increased. As our experiments were performed under natural biotic and abiotic conditions, factors other than zooplankton biomass might affect or limit this response, such as water temperature and light intensity. 4. Because zooplankton biomass was correlated with water temperature and light intensity, it was not possible to evaluate the independent contribution of these factors on the attack rate and swimming characteristics (swimming speeds and angle of turn) of brook charr. However, this study highlighted the impact of these factors on the feeding behaviour of juvenile brook charr when feeding in the pelagic habitat under natural conditions, and their importance in future models of optimal foraging and fish habitat quality.     

Individual variations in habitat use and morphology in brook charr

The specific objectives of this study were to determine if there is individual specialization in habitat use by lacustrine brook charr Salvelinus fontinalis and if so, if specialization is related to fish morphology. Localizations of 28 brook charr equipped with thermosensitive radiotransmitters were recorded during three summers (1991, 1992, and 1993) in two lakes of the Mastigouche Reserve (Québec, Canada). Fifty per cent of the fish were found mainly in the benthic zone (hereafter benthic individuals), 18% in the pelagic zone (pelagic individuals), and 32% travelled regularly between the two zones (generalist individuals). The observed interindividual differences in habitat preference were related to differences in body morphology and coloration: (i) the pectoral fins of benthic and generalist individuals were significantly longer than those of pelagic ones; and (ii) the coloration of the lower flank of benthic and generalist individuals was silver-grey while that of pelagic individuals was red. The results of this study suggest that brook charr inhabiting oligotrophic lakes of the Canadian Shield exhibit trophic polymorphisms, where some individuals are specialists better adapted to feeding in the littoral zone whereas others are specialists better adapted to feeding in the pelagic zone. The potential for reproductive isolation between the two morphs is discussed.     

Trophic polymorphism in brook charr revealed by diet, parasites and morphometrics

Stomach contents, parasite assemblages and morphometrics were compared in brook charr Salvelinus fontinalis from the littoral and pelagic zone of two adjacent lakes on the Canadian Shield. In lac Baie des Onze Îles, fish from the littoral zone had greater abundance of benthic prey in their stomach and were more heavily infected by parasites that use intermediate hosts associated with the littoral zone than fish captured in the pelagic zone. Littoral and pelagic brook charr from this lake also differed in regard to body shape and fin length, with each group being anatomically adapted to exploit their respective habitats. The highly significant correlation between morphometric and parasite canonical scores supports the hypothesis of functional diversification of individuals within lac Baie des Onze Îles. While fish from littoral and pelagic zones of lac Caribou did not differ in terms of diet, parasite assemblages or morphometrics, they were different to fish from lac Baie des Onze Îles in that they were less frequently infected with parasites that use gastropods as intermediate hosts, and had shorter pectoral fins. The inter-lake comparisons suggested that parasite assemblages and morphometrics of brook charr reflected the dominance of the limnetic and littoral habitats in lacs Caribou and lac Baie des Onze Îles, respectively.     

Genetic and morphological variation between two forms of lacustrine brook charr

Genetic data suggest that the littoral and pelagic forms of brook charr Salvelinus fontinalis in Lake Bondi are two populations with partial reproductive isolation and non-random mating. Genetic differentiation between the two groups was supported by differences in allele frequencies and by deviation from Hardy-Weinberg equilibrium when the two groups were pooled; no such deviation was observed when fish were divided into littoral and pelagic groups. In contrast to Lake Bondi, no clear evidence of genetic differentiation was observed in Lake Ledoux. Discriminant function analyses of morphological characters support the existence of littoral and pelagic groups in Bondi and Ledoux Lakes. In Lake Bondi, the two groups differed significantly in two shape variables (pelagic fish had shorter dorsal fins, and longer body length posterior to the dorsal fin than littoral ones) whereas in Lake Ledoux, the groups differed in four shape variables (pelagic fish had shorter pectoral fins, shorter dorsal fins, and a shorter and higher caudal peduncle than littoral ones). Discriminant analyses of these characters were effective in reclassifying fish into their appropriate groups in both populations, with an efficiency of 78% for juveniles in Lake Bondi and 69% for adults in Lake Ledoux. Differences in morphology between the two forms are consistent with adaptations required to forage in each zone, i.e. benthic form in the littoral zone and planktivorous form in the pelagic zone.     

Niche segregation of coexisting Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) constrains food web coupling in subarctic lakes

1. Generalist fish species are recognised as important couplers of benthic and pelagic food‐web compartments in lakes. However, interspecific niche segregation and individual specialisation may limit the potential for generalistic feeding behaviour. 2. We studied summer habitat use, stomach contents and stable isotopic compositions of the generalist feeder Arctic charr coexisting with its common resource competitor brown trout in five subarctic lakes in northern Norway to reveal population‐level and individual‐level niche plasticity. 3. Charr and trout showed partial niche segregation in all five lakes. Charr used all habitat types and a wide variety of invertebrate prey including zooplankton, whereas trout fed mainly on insects in the littoral zone. Hence, charr showed a higher potential to promote habitat and food‐web coupling compared to littoral‐dwelling trout. 4. The level of niche segregation between charr and trout and between pelagic‐caught and littoral‐caught charr depended on the prevailing patterns of interspecific and intraspecific resource competition. The two fish species had partially overlapping trophic niches in one lake where charr numerically dominated the fish community, whereas the most segregated niches occurred in lakes where trout were more abundant. 5. In general, pelagic‐caught charr had substantially narrower dietary and isotopic niches and relied less on littoral carbon sources compared to littoral‐caught conspecifics that included generalist as well as specialised benthivorous and planktivorous individuals. Despite the partially specialised planktivorous niche and thus reduced potential of pelagic‐dwelling charr to promote benthic–pelagic coupling, the isotopic compositions of both charr subpopulations suggested a significant reliance on both littoral and pelagic carbon sources in all five study lakes. 6. Our study demonstrates that both interspecific niche segregation between and individual trophic specialisation within generalist fish species can constrain food‐web coupling and alter energy mobilisation to top consumers in subarctic lakes. Nevertheless, pelagic and littoral habitats and food‐web compartments may still be highly integrated due to the potentially plastic foraging behaviour of top consumers.     

Spatial and temporal distributions of salmonids in two ponds in Newfoundland, Canada

Spatial and temporal distributions of salmonids were examined in Junction Pond, Northeast River, Placentia and Conne Pond, Conne River, Newfoundland using Lundgren multiple-mesh experimental gillnets. Both ponds contain populations of Atlantic salmon and brook trout with Junction Pond also possessing brown trout and resident Arctic charr. For salmon parr there was a significant month effect in distribution of catch rates in both ponds, and in Junction Pond, there was a significant diel effect. There was also significant variation in catch rates by lentic zone. For brook trout, there was a significant lentic zone × month interaction in Junction Pond; in Conne Pond, the main effects lentic zone and month were significant. Highest benthic catch rates of Atlantic salmon parr occurred in the littoral zone of both ponds. Most captures of brook trout also occurred benthically in both ponds; similar to salmon parr, littoral zone catch rates were higher than those of the deeper benthic area in Conne Pond but the reverse was true for Junction Pond. For both salmon parr and brook trout, the deeper benthic area and the pelagic area were relatively important rearing habitats in each pond. The distribution of catches for brown trout (few in number relative to the other species) in Junction Pond was similar to that of brook trout while Arctic charr were found mainly pelagically. Within the benthic area, most Arctic charr were caught at depths beyond the littoral zone. There was a tendency for Altantic salmon parr and brook trout found in the deeper benthic area and the pelagic area to be significantly larger and older than those in littoral zone in each pond. Sizes of Arctic charr did not differ significantly among lentic zones.     

Lipid‐rich zooplankton subsidise the winter diet of benthivorous Arctic charr (Salvelinus alpinus) in a subarctic lake

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Lake size and fish diversity determine resource use and trophic position of a top predator in high‐latitude lakes

Prey preference of top predators and energy flow across habitat boundaries are of fundamental importance for structure and function of aquatic and terrestrial ecosystems, as they may have strong effects on production, species diversity, and food‐web stability. In lakes, littoral and pelagic food‐web compartments are typically coupled and controlled by generalist fish top predators. However, the extent and determinants of such coupling remains a topical area of ecological research and is largely unknown in oligotrophic high‐latitude lakes. We analyzed food‐web structure and resource use by a generalist top predator, the Arctic charr Salvelinus alpinus (L.), in 17 oligotrophic subarctic lakes covering a marked gradient in size (0.5–1084 km²) and fish species richness (2–13 species). We expected top predators to shift from littoral to pelagic energy sources with increasing lake size, as the availability of pelagic prey resources and the competition for littoral prey are both likely to be higher in large lakes with multispecies fish communities. We also expected top predators to occupy a higher trophic position in lakes with greater fish species richness due to potential substitution of intermediate consumers (prey fish) and increased piscivory by top predators. Based on stable carbon and nitrogen isotope analyses, the mean reliance of Arctic charr on littoral energy sources showed a significant negative relationship with lake surface area, whereas the mean trophic position of Arctic charr, reflecting the lake food‐chain length, increased with fish species richness. These results were supported by stomach contents data demonstrating a shift of Arctic charr from an invertebrate‐dominated diet to piscivory on pelagic fish. Our study highlights that, because they determine the main energy source (littoral vs. pelagic) and the trophic position of generalist top predators, ecosystem size and fish diversity are particularly important factors influencing function and structure of food webs in high‐latitude lakes.     

Seasonal fluctuations in macrophyte cover and water transparency of four brown-water lakes: implications for crustacean zooplankton in littoral and pelagic habitats

The dynamics of crustacean zooplankton in the littoral and pelagic zones of four forest lakes having variable water qualities (colour range 130–340 mg Pt l⁻¹, Secchi depth 70–160 cm) were studied. The biomass of zooplankton was higher in the littoral zone than in the pelagic zone only in the lake having the highest transparency. In the three other lakes, biomass was significantly higher in the pelagic zone than in the littoral zone. In the two lakes with highest transparency, the littoral biomass of cladocerans significantly followed the development of macrophyte vegetation, and cladoceran biomass reached the maximum value at the time of highest macrophyte coverage. In lakes with lowest transparency, littoral zooplankton biomass developed independently of macrophyte density and decreased when macrophyte beds were densest. The seasonal development of the littoral copepod biomass did not follow the development of macrophytes in any of the lakes. The mean size of cladocerans in the pelagic zone decreased with increasing Secchi depth of the lake, whereas in the littoral zone no such phenomenon was detected. Seasonally, when water transparency increased temporarily in two of the lakes, the mean size of cladocerans in the pelagic zone decreased steeply. For copepods, no relationship between water transparency and body size was observed. The results suggested that in humic lakes the importance of the littoral zone as a refuge decreases with decreasing transparency of the water and that low water transparency protects cladocerans from fish predation. All the observed between-lake differences could not be explained by fish predation, but were probably attributed to the presence of chaoborid larvae with variable densities. Feeding efficiency of chaoborids is not affected by visibility and thus they can obscure the relationship between water quality, fish density, and the structure of crustacean zooplankton assemblages. Handling editor: S. I. Dodson     

Density,length distribution,and diet of age-0 arctic charrSalvelinus alpinus in the surf zone of Thingvallavatn,Iceland

Synopsis Population densities of age-0 arctic chaff in the surf zone averaged 1.83 and 4.70 fish m^-2 in August 1984 and June 1985, respectively. Length variation of the littoral fish was low in early summer, increasing in late summer and autumn. Newly emerged charr, ∼ 20 mm long, appeared in the stony shallow water zone during both May and June. From length variation and variation in mouth position of the young charr, it is concluded that at least two of the four chaff morphs in the lake are present in the surf zone during spring and summer. In August, some of the larger age-0 charr had moved out from the surf zone, into the pelagic and the deeper epibenthic waters. The food of young littoral charr was dominated by large chironomid larvae (instar 3 and 4) and pupae Contribution from the Thingvallavatn project     

The biology and feeding ecology of Arctic charr in the Kerguelen Islands

Subsequent to their introduction in the 1950s, Arctic charr Salvelinus alpinus have been able to establish a self-sustaining population that has adapted to the unique conditions of the sub-Antarctic Kerguelen Islands. Here, 48 individuals (198–415 mm) were caught with gillnets and their basic biology and feeding ecology were examined using stable isotope analysis. The Lac des Fougères population split use of littoral and pelagic resources evenly, although larger fish relied more heavily on littoral production and appear to follow the size-dependent life history habitat template seen in many Scandinavian lakes where smaller sized individuals occupy the pelagic zone and larger individuals dominate the littoral habitat. In Kerguelen, Arctic charr mature at the same ages (5.6 years) as Arctic charr in both sub-Arctic and Arctic lakes. Although mortality was average in comparison to comparator sub-Arctic lakes, it was high in comparison to Arctic lakes. Maximal age (>7+) was at the lower end of the range typically seen in sub-Arctic lakes. Although they inhabit a resource-poor environment, Kerguelen Arctic charr showed no evidence of cannibalism. Thus, while Arctic charr can survive and reproduce in the relatively unproductive Kerguelen lake environments, survival and growth nevertheless appear to be traded off against survival and longevity. The uniqueness of the population location and the recency of its introduction suggest that further monitoring of the population has the potential to yield valuable insights into both the adaptability of the species and its likely responses to ongoing large-scale environmental change as represented by climate change.     

Temporal stability of individual feeding specialization may promote speciation

1.  Inter-individual differences in trophic behaviour are considered important in the disruptive selection process for resource specialization and may represent an early phase in the evolution of polymorphism and adaptive radiation. Here, we provide evidence of high stability of individual trophic niches of a fish predator from a 15-year study.
2.  Individual resource specialization was investigated by combining data from analyses of stomach contents (recent trophic niche), trophically transmitted parasites (long-term niche) and trophic morphology (niche adaptations) from single specimens of a postglacial fish (Arctic charr) population sampled from contrasting pelagic and littoral habitats.
3.  Based on the relationships between morphology, parasites and diet, high inter-individual temporal consistency of narrow niches (zooplanktivorous vs . benthivorous) was evident through the ontogeny of the charr, indicating low degree of switching both in habitat utilization and feeding strategy of individual fish. Co-occurrence of differently specialized behavioural phenotypes was sustained over multiple generations.
4.  The stable long-term habitat and feeding specializations may represent an important initial step in an adaptive radiation process, and our findings suggest a case of sympatric speciation into two incipient forms diverging along the littoral–pelagic resource axis.     

Size-scaling of zooplankton foraging in Arctic charr

Prey capture rate (number of prey s⁻¹) and the mode of feeding of Arctic charr Salvelinus alpinus were studied by performing foraging experiments with two sizes (1·1 and 1·8 mm) of Daphnia longispina prey. Arctic charr were particulate feeders at all densities tested. Adjusted for the effect of prey density, the capture rate showed a hump-shaped relationship with Arctic charr size for both sizes of D. longispina . Estimated attack rates ( a ) also tended to show a hump-shaped relationship with fish size. The estimated size-scaling exponent of the attack rate function, however, was relatively small, implying small changes in attack rate over fish sizes. Simultaneous estimations of a and handling time were used in combination with published data on fish metabolism and dry mass rations of prey to estimate maintenance resource density of prey as a function of Arctic charr mass. Maintenance resource densities increased monotonically with Arctic charr size, and rapidly as optimum fish size relative to attack rate on prey was passed.     

Pyloric caecal morphology of brook charr,Salvelinus fontinalis,in relation to diet

Synopsis Brook charr, Salvelinus fontinalis, shifts its diet from zoobenthos to pelagic prey when living sympatrically with white sucker, Catostomus commersoni, in lakes of the Laurentian Shield. We tested the hypothesis that this diet difference would have a significant impact on their pyloric caecal morphology in 5 lakes containing allopatric brook charr populations and 6 other lakes containing both brook charr and white sucker. We observed that the mean length of the most posterior caecum of charr was significantly greater in sympatry than in allopatry (X ± 1 SD: 9.91 ± 1.12 mm versus 8.44 ± 0.67 mm). This is equivalent to an increase of 18% of total pyloric caecal mass (dry weight) in sympatric brook charr. These results indicate that this response to differences in diet, well known in birds, also occurs in fish.     

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 .     

The impact of intra- and interspecific interactions on young-of-the-year brook charr, in temperate lakes

The abundance, growth, spatial distribution, and feeding habits of five allopatric brook charr, Salvelinus fontinalis, populations (young-of-the-year, 0+ juveniles; YOY) were compared with five other populations living sympatrically with white sucker, Catostomus commersoni. The study was made in oligotrophic lakes of the Laurentian Shield (Québec, Canada) during three sampling periods in 1989 (July, August and September). The abundance of YOY charr was significantly higher in allopatric than in sympatric populations (45·3 ± 3·8 vs 3·4 ± 3·8 fish/lake caught in 1773 m² of gillnets; P<0·005). The mean length of YOY charr did not differ among allopatric and sympatric populations at each sampling period; July: 60·2 ± 3·0 vs 60·0 ± 4·5 mm; August: 61·9 ± 4·5 vs 63·2 ± 4·1 mm; September: 77·9 ± 8·7 vs 77·3 ± 7·8 mm respectively. Horizontal distribution of allopatric YOY charr did not differ from that of sympatric charr, 65% of the fish being captured within the first 2 m depth and the rest between 2 and 7 m depth. In contrast, the vertical distribution of allopatric YOY charr from both communities was significantly different; 81% of allopatric charr were captured within 0·5 m from the substrate compared to 64% for sympatric charr (P<0·001). Differences in vertical distribution of the fish were related to differences in diet; allopatric charr fed mainly on benthic and large planktonic organisms whereas sympatric charr fed less on these organisms and more on terrestrial organisms. In the lake where YOY charr were most abundant, individuals were spatially segregated into two groups; one ‘littoral’, found in 0–2m depth, and one ‘profundal’, found in 3–6 m depth. Growth, condition, and feeding habits of charr from the two groups were different, especially during the last sampling period.     

Summer use of a small stream by fish and crayfish and exchanges with adjacent lentic macrohabitats

1. Movements between a stream reach and two adjacent lentic macrohabitats, a beaver pond and a lake, were followed for the Appalachian crayfish and two fish species, brook charr and brown bullhead, over an 85‐d period from early June to late August, and were analysed in relation to water level, maximum water temperature, photoperiod length, lunar luminosity, and age, by use of time‐series regressions. 2. Brook charr showed strong net immigration to the stream reach for underyearling (age class 0+) fish but net emigration for 1+ fish. Both immigration and emigration were positively related to water level and temperature; migratory responses to temperature were age‐specific. 3. Brown bullhead used the stream primarily as a corridor for downstream migration from the beaver pond to the lake. As with brook charr, water level and temperature had a positive effect on movement and responses were stronger in younger individuals. 4. Crayfish emigrated from the stream during the summer. Movements were positively related to increases in water level and temperature, with responses of 1+ crayfish much stronger than those of older individuals. 5. The results indicate that the stream tributary played different roles for brook charr (nursery), brown bullhead (dispersal corridor) and Appalachian crayfish (early summer refuge), and suggest that variation in water level or temperature resulting from climatic change or local anthropogenic activities might modify connectivity between macrohabitats, particularly for younger individuals.     

Habitat use of arctic charrSalvelinus alpinus in Thingvallavatn,Iceland

Synopsis Habitat use by four morphs of arctic charr,Salvelinus alpinus, was investigated in Thingvallavatn, Iceland, by sampling with pelagic and benthic gill nets. Sampling was done in May/June and August/September. Greatest abundance of fish was recorded in the littoral and epipelagic zone in early autumn. Catches were low in early summer. The four morphs are partly segregated in habitat. Small (SB-) and large benthivorous (LB-) charr have a more restricted spatial distribution than piscivorous (PI-), and especially planktivorous (PL-) charr. Both benthivorous morphs are mainly found in the littoral zone, and occur in largest numbers in stony shallows at depths between 0 and 10 m. PL-charr usually dominates in numbers in all habitats. PI-charr is most abundant in epibenthic habitats, although numbers are always low. All morphs are caught in higher numbers at night than during the day, but the diurnal activity difference is highest among SB-charr. The habitat use by different morphs is as may be expected from their morphology and diets. Within the population of PL-charr, young and small fish are more abundant on the bottom than in the pelagic zone, and there is a surplus of females in the pelagic zone. Along the benthic profile, young, small and immature PL-charr are more abundant in deep than in shallow waters. The results are discussed in relation to food supply, competition and predation. Possible reasons for the occurrence of four arctic charr morphs are also discussed.Contribution from the Thingvallavatn project.     

The food of Charr,*Salvelinus willughbii (Günther), in Windermere

The stomach contents of 1015 charr, (Salvelinus willughbii Günther)] from Windermere, of which 658 were feeding fish, were examined. The fish were caught over a period of several years and in every month of the year. The data were analysed by two methods, by a points system, which takes into account the abundance and volume of the food organisms, and by the frequency of occurrence of each organism. Charr of all sizes had fed mainly on planktonic Crustacea (particularly Cladocera); larval and pupal chironomids and Chaoborus were also important. From November to April most stomachs of charr netted on the spawning grounds contained charr eggs. The diet of charr is compared with that of seven other fish species in Windermere. There was almost no overlap between the food of charr, which was obtained in the pelagic region, and that of the seven other species, which was mostly bottom fauna, obtained from the littoral region. Thus by occupation of these two different niches the charr and the seven other fish in no way compete for food in Windermere. Some limited data from six other lakes in the English Lake District are presented, in these planktonic Crustacea are also found to be the main food.