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.     

Niche use and resource partitioning of Arctic charr,European whitefish and grayling in a subarctic lake

Arctic charr and European whitefish are common species in northern Europe, but have a parapatric distribution pattern and rarely coexist in substantial densities within the same lake. Whitefish is considered to be the strongest competitor, usually causing competitive relegation or exclusion of co-occurring charr. Some exceptions do however occur, and the present study addresses niche utilization and resource partitioning of an Arctic charr population coexisting with whitefish and grayling in a subarctic lake. There was a distinct resource partitioning with respect to habitat and diet utilization of the three species. Grayling were exclusively caught in shallow near-shore areas, feeding chiefly on surface insects and Trichoptera larvae. Whitefish dominated in all lake habitats, feeding predominantly on small-sized planktonic and benthic crustaceans. Arctic charr were mainly found along the benthic profile, feeding predominantly on insects and snails and showing a low dietary overlap with whitefish which appeared to have monopolised the zooplankton resources. The highly restricted trophic niche of Arctic charr suggests a severe competitive impact due to the presence of whitefish, but the coexistence of the two species may be facilitated by the presence of grayling.     

Habitat utilization by sympatric arctic charr Salvelinus alpinus L. and brown trout Salmo trutta L. in Lake Atnsjø, south-east Norway

SUMMARY. 1. Habitat utilization, as well as inter- and intraspecific relations of different size groups of arctic charr (Salvelinus alpinus (L.)) and brown trout (Salmo trutta L.) in Lake Atnsjø, south-east Norway, were investigated by analysing food and spatial niches from monthly benthic and pelagic gillnet catches during June-October 1985.
2. Small individuals (150–230 mm) of both arctic charr and brown trout occurred in shallow benthic habitats. However, they were spatially segregated as arctic charr dominated at depths of 5–15 m and brown trout at depths of 0–5 m.
3. Larger (>230 mm) arctic charr and brown trout coexisted in the pelagic zone. Both species occurred mainly in the uppermost 2-3 m of the pelagic, except in August, when arctic charr occurred at high densities throughout the 0–12 m depth interval. On this occasion, arctic charr were segregated in depth according to size, with significantly larger fish in the top 6 m. This was probably due to increased intraspecific competition for food.
4. The two species differed in food choice in both habitats, Arctic charr fed almost exclusively on zooplankton, whereas brown trout had a more variable diet, consisting of surface insects, zooplankton. aquatic insects and fish.
5. The data suggest that the uppermost pelagic was the more favourable habitat for both species. Large individuals having high social position occupied this habitat, whereas small individuals lived in benthic habitat where they were less vulnerable to agonistic behaviour from larger individuals and less exposed to predators. The more aggressive and dominant brown trout occupied the more rewarding part of the benthic habitat.     

Habitat preferences of European grayling in a medium size stream, the Ain river, France

The three grayling Thymallus thymallus age-classes had strong preferences for their local habitat in the Ain river. All age-classes preferred high water velocities between 70 and 110 cm s^-1, confirming that grayling is a typical rheophilic species. An important intraspecific segregation regarding depth occurred, as large individuals preferred deeper water than small ones (optimal ranges=50–60, 80–120 and 100–140 cm for 0+, 1+ and adults respectively). The three age-classes had similar preferences for small substratum articles, with optimal values between 0·5 and 16·0 mm. Preference curves were generally comparable between the two sites, despite important differences in habitat availability. It is emphasized that grayling needs various habitat conditions to achieve its entire life cycle. The increasing scarcity of this species underlines the urgency of protecting lotic habitat integrity and diversity.     

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.     

Cost of predation avoidance in young-of-year lake trout (Salvelinus namaycush): growth differential in sub-optimal habitats

Selection of habitat to avoid predation may affect the diet of young-of-year (YOY) lake trout (Salvelinus namaycush). YOY lake trout may use inshore habitat to avoid predation; this habitat may be sub-optimal for growth. To test this, YOY lake trout were penned in nearshore and offshore pelagic areas of two arctic lakes. Toolik Lake had a lake trout population, the other lake, S6, did not. YOY lake trout in Toolik Lake lost weight, but those offshore lost less weight. The YOY lake trout in Lake S6 gained weight and those offshore gained more weight. The primary diet item of the YOY lake trout in both lakes during this experiment was the zooplankter Diaptomis probilofensis; it was also one of the most abundant species. However, its density inshore in Lake S6 was similar to inshore and offshore densities in Toolik Lake. The increased availability of alternative zooplankton prey in Lake S6 may account for the growth differential of YOY lake trout in Lake S6 relative to Toolik Lake. Bioenergetic modeling of YOY lake trout suggests that growth similar to that in the offshore of Lake S6 would be necessary for successful recruitment. If the reduced zooplankton availability in Toolik Lake leads to the reduced growth of YOY in the inshore and offshore pelagic areas, then these fish will be more susceptable to winter predation/starvation. For YOY lake trout to survive in Toolik Lake they most likely shift to feeding on benthic prey before the end of their first summer. Dept. of Chemical Engineering     

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 .     

Ecosystem response to earlier ice break‐up date: Climate‐driven changes to water temperature,lake‐habitat‐specific production,and trout habitat and resource use

Climate warming has yielded earlier ice break‐up dates in recent decades for lakes leading to water temperature increases, altered habitat, and both increases and decreases to ecosystem productivity. Within lakes, the effect of climate warming on secondary production in littoral and pelagic habitats remains unclear. The intersection of changing habitat productivity and warming water temperatures on salmonids is important for understanding how climate warming will impact mountain ecosystems. We develop and test a conceptual model that expresses how earlier ice break‐up dates influence within lake habitat production, water temperatures and the habitat utilized by, resources obtained and behavior of salmonids in a mountain lake. We measured zoobenthic and zooplankton production from the littoral and pelagic habitats, thermal conditions, and the habitat use, resource use, and fitness of Brook Trout (Salvelinus fontinalis). We show that earlier ice break‐up conditions created a "resource‐rich" littoral–benthic habitat with increases in zoobenthic production compared to the pelagic habitat which decreased in zooplankton production. Despite the increases in littoral–benthic food resources, trout did not utilize littoral habitat or zoobenthic resources due to longer durations of warm water temperatures in the littoral zone. In addition, 87% of their resources were supported by the pelagic habitat during periods with earlier ice break‐up when pelagic resources were least abundant. The decreased reliance on littoral–benthic resources during earlier ice break‐up caused reduced fitness (mean reduction of 12 g) to trout. Our data show that changes to ice break‐up drive multi‐directional results for resource production within lake habitats and increase the duration of warmer water temperatures in food‐rich littoral habitats. The increased duration of warmer littoral water temperatures reduces the use of energetically efficient habitats culminating in decreased trout fitness.     

Trout induces a shift from preferred habitat types for indigenous species: the example of the indigenous catfish, <Emphasis Type="Italic">Amphilius uranoscopus</Emphasis> (Pfeffer, 1889), on an African montane plateau

This study investigated the impact of introduced rainbow trout, Oncorhynchus mykiss, on the distribution and feeding of mountain catfish, Amphilius uranoscopus, on the Nyika Plateau, Malawi. Twenty-four sites were sampled over three different periods in three rivers. Fish habitat units were identified as separate riffle or pool, about 100 m in length, at each site. Each habitat unit was sampled for fish, invertebrates and physical habitat characteristics. Twenty four and 20 habitat units were sampled, respectively, from sites with catfish and sites with trout and catfish. In the absence of trout, the mountain catfish was associated with all depth ranges, with strong preference to shallow and moderate depth, and moderate to fast flow on coarse substratum type (gravel, pebble and boulder). In the presence of trout, the catfish was frequently associated with very shallow depth and slow flow. In its natural habitat, the catfish fed randomly, but preferred the most abundant invertebrate taxa, especially black fly larvae (Simuliidae). In the presence of trout, the catfish preferred mostly the chironomids. The preference by catfish for Simuliidae, also preferred by trout, was less in the trout streams than in its natural habitat. The prey taxa in the catfish stream were diverse, and consisted of large invertebrate predators. Trout streams were dominated by few prey taxa, especially black flies and chironomids. The catfish of the Nyika Plateau may represent genetically unique populations in southern Africa. Introductions of trout into rivers where they currently do not occur on the Nyika should be prevented in order to maintain the genetic diversity of the Amphilius uranoscopus species complex. Handling editor: J. A. Cambray     

Foraging habitat determines predator–prey size relationships in marine fishes

Predator–prey size (PPS) relationships are determined by predator behaviour, with the likelihood of prey being eaten dependent on their size relative to that of the consumer. Published PPS relationships for 30 pelagic or benthic marine fish species were analysed using quantile regression to determine how median, lower and upper prey sizes varied with predator size and habitat. Habitat effects on predator foraging activity/mode, morphology, growth and natural mortality are quantified and the effects on PPS relationships explored. Pelagic species are more active, more likely to move by caudal fin propulsion and grow more rapidly but have higher mortality rates than benthic species, where the need for greater manoeuvrability when foraging in more physically complex habitats favours ambush predators using pectoral fin propulsion. Prey size increased with predator size in most species, but pelagic species ate relatively smaller prey than benthic predators. As pelagic predators grew, lower prey size limits changed little, and prey size range increased but median relative prey size declined, whereas the lower limit increased and median relative prey size was constant or increased in benthic species.     

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

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Empirical evaluation of phenotype–environment correlation and trait utility with allopatric and sympatric whitefish, Coregonus lavaretus (L.), populations in subarctic lakes

Adaptive phenotypic divergence of sympatric morphs in a single species may have significant evolutionary consequences. In the present study, phenotypic impacts of predator on zooplankton prey populations were compared in two northern Finnish lakes; one with an allopatric whitefish, Coregonus lavaretus (L.), population and the other with three sympatric whitefish populations. First, we examined whether there were phenotypic associations with specific niches in allopatric and sympatric whitefish. Second, trait utility (i.e. number of gillrakers) of allopatric and sympatric whitefish in utilizing a pelagic resource was explored by comparing predator avoidance of prey, prey size in environment, and prey size in predator diet. The allopatric living large sparsely rakered (LSR) whitefish morph, was a generalist using both pelagic and benthic niches. In contrast, sympatric living whitefish morphs were specialized: LSR whitefish was a littoral benthivore, small sparsely rakered whitefish was a profundal benthivore and densely rakered (DR) whitefish was a pelagic planktivore. In the lake with allopatric whitefish, zooplankton prey did not migrate vertically to avoid predation whereas, in the lake with sympatric whitefish, all important prey taxa migrated significantly. Trait utility was observed as significantly smaller size of prey in environment and predator diet in the lake with DR whitefish than in the lake with only LSR whitefish.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 561–572.     

Unidirectional prey-predator facilitation: apparent prey enhance predators' foraging success on cryptic prey

Food availability can strongly affect predator-prey dynamics. When change in habitat condition reduces the availability of one prey type, predators often search for other prey, perhaps in a different habitat. Interactions between behavioural and morphological traits of different prey may influence foraging success of visual predators through trait-mediated indirect interactions (TMIIs), such as prey activity and body coloration. We tested the hypothesis that foraging success of stream-dwelling cutthroat trout (Onchorhyncus clarki) on cryptically coloured, less-active benthic prey (larval mayfly; Paraleptophebia sp.) can be enhanced by the presence of distinctly coloured, active prey (larval stonefly shredder; Despaxia augusta). Cutthroat trout preyed on benthic insects when drifting invertebrates were unavailable. When stonefly larvae were present, the trout ate most of the stoneflies and also consumed a higher proportion of mayflies than under mayfly only treatment. The putative mechanism is that active stonefly larvae supplied visual cues to the predator that alerted trout to the mayfly larvae. Foraging success of visual predators on cryptic prey can be enhanced by distinctly coloured, active benthic taxa through unidirectional facilitation to the predators, which is a functional change of interspecific interaction caused by a third species. This study suggests that prey-predator facilitation through TMIIs can modify species interactions, affecting community dynamics.     

The ecology of ruffe,Gymnocephalus cernuus (Pisces: Percidae) introduced to Mildevatn,western Norway

Synopsis Diet, habitat use, diel and seasonal activity and a number of population parameters were studied on ruffe,Gymnocephalus cernuus, introduced to Mildevatn, western Norway. This lake is sited outside the natural range of the ruffe and has a lower fish diversity and a different fish species composition than within its native range. From June through September the ruffe was planktivorous and mainly caught at 4 to 6 m depth in the benthic zone. At other times of year ruffe was feeding on zoobenthos and caught deeper in the benthic zone. Ruffe was mainly day active. Zooplankton feeding during summer is the clearest difference compared to ruffe populations living within its natural range. Presence of large zooplankton organisms available for ruffe is suggested as the main reason for the difference found in food choice. The availability of large zooplankton is probably due to community structure caused by a predator and lack of interspecific competition for zooplankton in the deeper parts of the lake. Piscivorous brown trout.Salmo trutta, restrict the habitat of threespined stickleback,Gasterosteus aculeatus, to the zone of littoral vegetation, allowing high densities of larger zooplankton species likeBythotrephes longimanus to be present in the lake. Brown trout is present only in the upper light and well oxygenated parts of the lake, leaving a refuge for the ruffe, where they can feed on the rich zooplankton community.     

Competitive habitat displacement of brown trout by Siberian sculpin: the role of size and density

Habitat competition in brown trout Salmo trutta and Siberian sculpin Cottus poecilopus was investigated by varying density, fish size, and species composition in stream channels providing areas of different substratum particle sizes. In allopatry, both small (52 ± 4 mm L _T) and large (86 ± 6 mm L _T) brown trout exhibited strong preference for the intermediate (8–11 cm diameter) and large (17–21 cm) gravel substrata. There was a tendency for more brown trout to occupy finer (2–4 cm) substrata with increasing density, in particular for large brown trout. Also, more small brown trout were observed on finer substrata when tested with large brown trout, suggesting interspecific competition for restricted space. Both small (56 ± 6 mm L _T) and large (88 ± 10 mm L _T) Siberian sculpin preferred the large gravel in all tests, and did not change their substratum preferences much with increasing densities, suggesting higher tolerance for 'crowding'. The large Siberian sculpin preferred the coarser substratum, and the largest individuals were consistently found on it. In sympatry with large Siberian sculpin, habitat displacement of brown trout occurred, indicative of interspecific competition. A higher proportion of small and large brown trout occupied the finer substrata than in allopatry. Habitat selection by large Siberian sculpin appeared to be unaffected by species composition and density. Small Siberian sculpin were displaced to finer substrata when tested with large Siberian sculpin, suggesting intraspecific competition. The results indicate that Siberian sculpin are potential habitat competitors for young brown trout.     

Intra- and interspecific life history differences in sympatric grayling and brown trout in a Norwegian reservoir

Grayling Thymallus thymallus in Lake Aursjøen, Norway, showed a remarkably uniform growth pattern throughout life, whereas brown trout Salmo trutta showed far more variation. In addition, a narrower age-length interval of maturation was found in grayling. The restricted life history variation in grayling is discussed and it is suggested that all grayling of Lake Aursjøen experience similar environmental conditions as juveniles, which induces low phenotypic variation. In contrast the existence of several spawning populations, adapted to as many as 28 different tributaries, may have created large life history variation in Aursjøen trout. Logistic models revealed that both age and length had significant, simultaneous effects on the maturation of both species. Furthermore, the sexes of trout differed in maturation patterns, i.e. males matured earlier and at smaller sizes than female conspecifics, but no difference was found between the sexes of grayling. In addition, larger sex-specific growth differences were found in trout. An absence of early maturing males in grayling and their presence in trout is discussed as a possible explanation of the restricted life history variation found between sexes of grayling. Male grayling experienced a larger mortality rate than did females, whereas no such differences were found in trout. It is suggested that grayling males invest more in reproduction than do females, due primarily to large investments in breeding behaviour. The equal mortality rates found for both sexes of trout, albeit males starting to spawn earlier than females, is probably explained by a female-selective fishing mortality.     

Diet of Arctic charr (Salvelinus alpinus (L.)) and brown trout (Salmo trutta L.) in sympatry in two high altitude alpine lakes

Lake Pisses and Lake Labarre are two oligotrophic high altitude alpine lakes that have sympatric populations of Arctic charr and brown trout. These two lakes have similar morphometric, physical and chemical characteristics. The zooplanktonic and benthic fauna show little diversity. But the density of benthos (Chironomidae) and zooplankton is higher in Lake Pisses. The fish fauna of Lake Pisses is slightly more abundant than that of Lake Labarre, althought in both lakes fish density is low. A study of the diet of the two species revealed differences. In Lake Pisses, where the food supply is better, Arctic charr takes exclusively pelagic and benthic prey, whereas in Lake Labarre it also takes exogenous prey and thus comes into competition with trout. Length and body weight growth rates for Arctic charr are higher in Lake Pisses than in Lake Labarre. For trout, maximum length recorded was in Lake Pisses. The results show that the abundance of Chironomidae favours coexistence of the two species in Lake Pisses and confirm that, in the face of shortage of food, Arctic charr is better adapted than trout. This revised version was published online in July 2006 with corrections to the Cover Date.     

Food of the roughhead grenadier, Macrourus berglax, Lacepede in North Norwegian waters

Over 30 different prey species were found in the stomachs of roughhead grenadiers, Macrourus berglax , caught at depths of approximately 700 m off the coast of northern Norway. Crustaceans and fish were the principal prey categories and large, mature grenadiers are classified as being generalist feeders taking predominantly benthopelagic organisms. There is evidence for an ontogenetic shift in dietary composition, with a change from benthic prey to an increased reliance on benthopelagic prey as the fish increase in size and this results in reduced intraspecific competition for food.     

Diet composition and prey preference of tench, Tinca tinca (L.), common bream, Abramis brama (L.), perch, Perca fluviatilis L. and roach, Rutilus rutilus (L.), in two contrasting gravel pit lakes: potential trophic overlap with wildfowl

The abundance of benthic macroinvertebrate taxa and the relative abundance of zooplankton taxa were compared with the diet of bream, tench. perch and roach from two gravel pit lakes during May-July 1986 and 1987. Significant food preferences were demonstrated between species and between lakes. Chironomid pupae dominated the macroinvertebrate diet of perch, roach, ate predominantly either Spirogyra sp. (St Peters Lake) or Daphniu hyalina (Main Lake).
The Main Lake, bream fed largely upon chironomid larvae and the bivalve Sphuerium but in St Peters Lake they positively selected a variety of less abundant benthic invertebrate taxa. Bream switched from benthos to zooplankton in the Main Lake in 1986. Tench ate large numbers of Aselhs and showed positive selection of various macroinvertebrate prey in St Peters but ate D. hjulim in Main Lake. Tropic overlap for chironomids and other macroinvertebrate prey was demonstrated between perch, bream and tench, and potentially with wildfowl which used the gravel pits for breeding and wintering.