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.     

Changes in use of lake habitat by experimentally segregated populations of cutthroat trout and Dolly Varden char

The role of competition in a lacustrine community of two salmonid species, cutthroat trout Oncorhynchus clarki and Dolly Varden char Salvelinus malma , was studied in three coastal British Columbia lakes Habitat use by the species alone (allopatric) and in coexistence with each other (sympatric) was investigated by gill netting at 0-40 m depth contours (surface to bottom) so that several habitats (littoral, epipelagic. pelagic, epibenthic) were sampled From June to October, trout used mainly littoral and epipelagic habitats in sympatry and allopatry Char used all habitats in allopatry. and deep habitats (pelagic, epibenthic) not frequented by trout in sympatry The two species were thus spatially segregated with depth in sympatry Diel (day, night) and seasonal changes (spring, summer, autumn) in habitat use were not pronounced The shift in habitat use by experimentally allopatric char but not trout suggests that the effects of competition between sympatric trout and char for habitat resources are greater on the char     

Diel and seasonal habitat and food segregation of three sympatric Coregonus lavaretus forms in a subarctic lake

Habitat use of three sympatric whitefish Coregonus lavaretus forms was determined using hydroacoustics, pelagic trawling and epibenthic gillnetting in the subarctic Lake Muddusjärvi during the day and night in June, August and September. Whitefish constituted 97% of the numerical catches and whitefish with high number of gill rakers (DR) were the most abundant whitefish form. Forms with low numbers of gill rakers used only epibenthic habitats during both the day and night in all study periods: large whitefish with low numbers of gill rakers (LSR) dwelled mainly at depths 0–10 m, whereas small whitefish with low numbers of gill rakers (SSR) used deeper (>10 m) habitats. LSR and SSR whitefish consumed mainly benthic macroinvertebrates during all study occasions. The planktivorous DR whitefish used both epibenthic and pelagic habitats, but vertical habitat selection varied both over time of day and season. In June, when light intensity was continuously high, DR whitefish did not perform diel vertical migrations. In August and September, when dark nights were distinguishable, DR whitefish ascended from the bottom to the pelagic at dusk to feed on zooplankton, and descended at dawn. DR whitefish used pelagic habitats only at the lowest light intensities during the night, which was probably related to the high predation risk from brown trout Salmo trutta .     

Foraging behaviour of non-breeding Western Sandpipers Calidris mauri as a function of sex, habitat and flocking

Individuals within a population may vary considerably in the way they exploit available food resources. If the sexes differ in the size of their feeding apparatus, there can be differences in foraging behaviour and habitat use, hence one sex may be more susceptible to competition. We examined relationships between sexual dimorphism in bill size and foraging behaviour, and habitat and microhabitat use of non-breeding Western Sandpipers Calidris mauri at Bahía Santa María, northwestern Mexico. Western Sandpipers are sexually dimorphic, with females about 15% longer-billed than males. Males used a higher proportion of pecks, had a lower probing–pecking rate, walked at higher rates, foraged at sites with lower water content, and had greater variation in foraging technique than females. Moreover, males decreased their proportion of pecks and foraged at a higher rate than females when they changed from feeding alone to feeding in flocks, suggesting a greater safety advantage or susceptibility to conspecific interference when flock feeding. We compared behaviour and microhabitat usage in three habitats: brackish flats, mangroves, and cattail marshes. Sex-related differences in foraging behaviour and microhabitat use were consistent among habitats. Birds in brackish flats and mangroves used a higher proportion of pecks, foraged at lower rates and walked at higher rates, and foraged at deeper sites, with a lower proportion of water cover, than those in cattail marshes. Sex-related differences in foraging behaviour and microhabitat should reduce the level of competition between sexes, and may account for sex differences in Western Sandpiper distribution observed between habitats in Bahía Santa María.     

Food- and habitat-segregation in sympatric grayling and brown trout

Grayling in the large reservoir Aursjøen, Norway, did not use the pelagic habitat and only trout larger than 187 mm and older than 4+ were caught there. Both species preferred the upper 8 m in the benthic habitat, but grayling were caught deeper than trout in August and September. Grayling were more closely associated with soft substratum than trout. Association with soft substratum decreased with increasing size in grayling, whereas this relationship was reversed in trout. A low number of empty stomachs indicated a low degree of food competition, but despite this the two species segregated in different key prey items—probably due to differences in mouth shapes. The key prey item for grayling was the benthic Eurycercus lamellatus , with zooplankton dominating the diet of the smallest individuals. Large trout (>300 mm) preferred the benthic Lepidurus arcticus , whereas medium trout (160–300 mm) ate predominantly the limnic-pelagic Bythotrephes longimanus , irrespective of habitat. The distribution of key prey items seemed to be the major factor controlling the habitat use, except in small trout which showed a generalist feeding pattern throughout the period. A generalist feeding pattern, a close association to coarse substratum and absence from the pelagic habitat indicated that small trout were negatively influenced by large individuals. Small grayling seemed less affected by aggressive encounters as they were less associated with coarse substratum and ate predominantly high-risk food such as zooplankton. These individuals probably did shoal in order to reduce predation risk.     

Habitat use and life history of brown trout (Salmo trutta) and Arctic charr (Salvelinus alpinus) in some low acidity lakes in central Norway

Habitat utilization and the life history of browntrout Salmo trutta and Arctic charr Salvelinus alpinus were investigated in fivesympatric populations and five allopatric brown troutpopulations in Høylandet catchment, a atmosphaericlow deposition area in Mid Norway. There was asignificant inverse correlation in abundance ofepibenthic Arctic charr and brown trout in theselakes, indicating that the latter species is dominant.The largest numbers of sympatric brown trout andArctic charr were caught in epibenthic habitat. In twolakes, brown trout to some extent also occurredpelagically, while pelagic individuals of Arctic charrwere found in all five lakes. The main food items forboth epibenthic and pelagic brown trout wereterrestrial surface insects and chironomid pupae.Zooplankton was the primary food item for Arctic charrin both habitats. Although the age distribution wasvery different in the populations, neither speciesseem to suffer from recruitment failure. There was nosignificant difference in survival rates betweensympatric populations of brown trout and Arctic charr.We found a significant inverse correlation betweenepibenthic catches of brown trout and the mean weightof 4+ fish, the most abundant age group. However, ifusing weight data for three-year-old fish, no suchrelationship was found for Arctic charr. Brown troutand Arctic charr reached asymptotic lengths of197–364 mm and 259–321 mm, respectively. Both speciestypically reached sexual maturity at age 2–3, and nomaturation-induced mortality was evident. We concludethat fish populations in Høylandet lakes areregulated throughout their lifes by inter- andintraspecific competition.     

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.     

Within-population heterogeneity of habitat use by lake trout Salvelinus namaycush

To learn more about the degree of individual variation in resource use by lake trout Salvelinus namaycush , ultrasonic telemetry was used to study their habitat use in a lake without pelagic schooling fish prey. Individuals spent most of their time within the metalimnion in favourable water temperatures. They also made frequent excursions, however, into lake temperatures exceeding their optimum for physiological performance at all temporal scales considered. Their frequent use of nearshore habitats suggested that feeding in littoral areas may be common. Habitat use was highly variable among individuals, but spatial habitat use by individuals showed remarkable consistency between years. In particular, some lake trout exhibited high site fidelity to shallow, nearshore areas, whereas others used deep areas extensively. This level of between-individual variation indicated niche partitioning by depth and the possibility of alternative foraging strategies.     

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.     

Sea trout Salmo trutta in the subarctic: home-bound but large variation in migratory behaviour between and within populations

Anadromous brown trout (sea trout), Salmo trutta, is currently in decline throughout its range, largely due to anthropogenic stressors in freshwater and marine habitats. Acoustic telmetry was utilized to study the marine migration of sea trout post-smolts from three populations in a relatively pristine subarctic fjord system. While at sea, the sea trout spent a substantial part of their time close to their natal river, preferred near shore over pelagic habitats and were strongly surface oriented. Despite a fidelity towards local areas, the sea trout utilized various parts of the fjord system, with maximum dispersion >30 km and total migration distance >300 km. Almost half of the sea trout (44%) migrated between river outlets, indicating that a metapopulation approach may be appropriate when managing neighbouring sea trout populations at high latitudes. Furthermore, the different populations displayed different migratory behaviours in terms of distance migrated, dispersion from origin and the likelihood of leaving their home area. This variation in migratory behaviour is likely influenced by spatiotemporal differences in habitat quality between sites, indicating that local habitat variations may promote population-specific behavioural responses even in relatively confined fjord systems.     

Trophic divergence of Lake Kivu cichlid fishes along a pelagic versus littoral habitat axis

Local adaptation to the littoral and pelagic zones in two cichlid haplochromine fish species from Lake Kivu was investigated using morphometrics. Cranial variation and inferred jaw mechanics in both sexes of the two species across the two habitat types were quantified and compared. Comparisons of littoral versus pelagic populations revealed habitat‐specific differences in the shape of the feeding apparatus. Also, kinematic transmission of the anterior jaw four‐bar linkage that promotes greater jaw protrusion was higher in the pelagic zone than in the littoral zone for both species. Inferred bite force was likewise higher in pelagic zone fish. There were also sex‐specific differences in craniofacial morphology as males exhibited longer heads than females in both habitats. As has been described for other cichlids in the East African Great Lakes, local adaptation to trophic resources in the littoral and pelagic habitats characterizes these two Lake Kivu cichlids. Similar studies involving other types of the Lake Kivu fishes are recommended to test the evidence of the observed trophic patterns and their genetic basis of divergences.     

Sexual segregation by Masai giraffes at two spatial scales

In this paper alternative explanations for observed patterns of sexual segregation by giraffes are examined at two spatial scales: within-habitats and within-landscape. Habitats are defined as recognizable plant associations and the landscape as the collection of all available habitat types. The study was conducted in Mikumi National Park, Tanzania. At the within-landscape level, all sex and age classes of giraffes exhibited high degrees of preference for riverine habitats. Sex differences in habitat selection were mostly due to females with young, who tended to select open floodplain habitats in which their vigilance time was lowest. Males, and females without young, preferred more heavily-wooded habitat. Habitat preferences were not related to observed habitat-specific forage intake rates for either males or females. Within habitats, male and female giraffes selected different feeding heights, males feeding higher in the canopy than females. Females showed a strong tendency to generalize with respect to feeding height. It is suggested that a sexual dimorphism–body size hypothesis provides a parsimonious explanation for the observed feeding height selection patterns, whereas a reproductive strategy hypothesis can explain sex-differences in habitat selection patterns within the landscape.     

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.     

Coevolution between flight morphology,vertical stratification and sexual dimorphism: what can we learn from tropical butterflies?

Occurrence patterns are partly shaped by the affinity of species with habitat conditions. For winged organisms, flight‐related attributes are vital for ecological performance. However, due to the different reproductive roles of each sex, we expect divergence in flight energy budget, and consequently different selection responses between sexes. We used tropical frugivorous butterflies as models to investigate coevolution between flight morphology, sex dimorphism and vertical stratification. We studied 94 species of Amazonian fruit‐feeding butterflies sampled in seven sites across 3341 ha. We used wing–thorax ratio as a proxy for flight capacity and hierarchical Bayesian modelling to estimate stratum preference. We detected a strong phylogenetic signal in wing–thorax ratio in both sexes. Stouter fast‐flying species preferred the canopy, whereas more slender slow‐flying species preferred the understorey. However, this relationship was stronger in females than in males, suggesting that female phenotype associates more intimately with habitat conditions. Within species, males were stouter than females and sexual dimorphism was sharper in understorey species. Because trait–habitat relationships were independent from phylogeny, the matching between flight morphology and stratum preference is more likely to reflect adaptive radiation than shared ancestry. This study sheds light on the impact of flight and sexual dimorphism on the evolution and ecological adaptation of flying organisms.     

Seasonal habitat selection by lake trout (<Emphasis Type="Italic">Salvelinus namaycush</Emphasis>) in a small Canadian shield lake: constraints imposed by winter conditions

The need for cold, well-oxygenated waters significantly reduces the habitat available for lake trout (Salvelinus namaycush) during stratification of small temperate lakes. We examined the spatial and pelagic distribution of lake trout over two consecutive summers and winters and tested whether winter increased habitat availability and access to littoral regions in a boreal shield lake in which pelagic prey fish are absent. In winter, lake trout had a narrowly defined pelagic distribution that was skewed to the upper 3 m of the water column and spatially situated in the central region of the lake. Individual core areas of use (50% Kernel utilization distributions) in winter were much reduced (75%) and spatially non-overlapping compared to summer areas, but activity levels were similar between seasons. Winter habitat selection is in contrast to observations from the stratified season, when lake trout were consistently located in much deeper waters (>6 m) and widely distributed throughout the lake. Winter distribution of lake trout appeared to be strongly influenced by ambient light levels; snow depth and day length accounted for up to 69% of the variation in daily median fish depth. More restricted habitat use during winter than summer was in contrast to our original prediction and illustrates that a different suite of factors influence lake trout distribution between these seasons.     

Food of anadromous brown trout at sea

The food of migratory Salmo trutta in coastal marine waters along the Norwegian Skagerrak coast varied significantly with age, season and habitat of the fish. The main prey categories in terms of frequency of occurrence were fishes followed by crustaceans, surface insects and polychaetes. Seasonal variation in diet and within habitats was found, supporting the view that brown trout is an opportunistic feeder. An ontogenetic niche shift was observed with post-smolts feeding on inshore and shallow water prey communities, while larger brown trout fed mainly on pelagic fishes.     

Lake morphometry predicts the degree of habitat coupling by a mobile predator

Habitat coupling is an ecosystem process whereby semi-discontinuous habitats are connected through the movement of energy and nutrients by chemical, physical or biological processes. One oft-cited example is that of littoral–pelagic coupling in lakes. Theory has argued that such habitat coupling may be critical to food web dynamics, yet there have been few empirical studies that have quantified ecological factors that affect the degree of habitat coupling in ecosystems. Specifically, the degree to which habitat coupling occurs across important physical gradients has largely been ignored. To address this, we investigate the degree of littoral habitat coupling (i.e. the degree to which a top predator lake trout, Salvelinus namaycush, derives energy from the littoral zone) along a gradient of lake shape, where lake shape modifies the relative quantity of coupled epilimnetic benthic and pelagic habitats within each lake. Herein we demonstrate that littoral habitat coupling is intensified in simple circular lakes compared to their reticulate counterparts in seven Canadian Shield lakes. Although the more reticulate lakes had larger areas of epilimnetic benthic habitat, littoral food sources comprised 11% compared to 24% of lake trout diet in reticulate and circular lakes, respectively. This heightened interaction in circular lakes also appears to translate into increased omnivory in more circular lakes compared to reticulate lakes such that lake trout of circular lakes have a significantly lower trophic position than lake trout of reticulate lakes (F_1,5=6.71 p=0.05). These results suggest that it is the accessibility of littoral production via thermal refugia, and not the amount of littoral production, that determines the degree to which lake trout couple littoral and pelagic habitats in lakes.     

Habitat use and mating system of the houbara bustard (Chlamydotis undulata undulata) in a semi-desertic area of North Africa: implications for conservation

Studies of the movements and home-ranges of houbara bustards (Chlamydotis undulata undulata) showed sexual and seasonal differences in the use of space, with a polygynous mating system similar to an ‘exploded-lek’ or a ‘resource-defence-polygyny’, that remains undefined. We used the arthropod biomass as an index of the trophic quality of six defined habitats and we radio-tracked 7 females and 13 males to test whether sexual and seasonal variations in habitat use were related to resource availability, and to verify if critical resources for breeding females were monopolised by males. We analysed habitat selection in both sexes separately. We used the habitat type composition of buffer zones around radio-locations to study annual and seasonal habitat selection and to identify preferred habitats, using the chi-square goodness-of-fit test. Habitat use between sexes and between seasons were compared using MANOVA based on log-ratios of habitat proportions. During the year, and in each season, both sexes appeared to be significantly selective for habitats in comparison to their availability. But males avoided esparto grass, while females used all habitats. Habitat use differed between sexes in the breeding season, but not in the non-breeding season. In spring, when food resources were abundant and uniformly distributed in space, males preferred ‘temporarily flooded areas’ and females preferred ‘reg with tall perennials’ that offered both food and cover for brooding. Critical resources were not monopolised by males and the mating system fulfilled the definition of the ‘exploded-lek’. Leks are key sites for reproduction and should be considered as priority areas in further conservation plans.     

Seasonal and spatial microhabitat selection and segregation in young Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L., in a Norwegian river

Seasonal microhabitat selection by sympatric young Atlantic salmon and brown trout was studied by diving. Both species, especially Atlantic salmon, showed seasonal variation with respect to surface and mean water velocities and depth. This variation is partly attributed to varying water flows and water temperatures. In winter the fish sought shelter in the substratum. A spatial variation in habitat use along the river due to different habitat availabilities was observed. Both species occupied habitats within the ranges of the microhabitat variables, rather than selecting narrow optima. It is hypothesized that the genetic basis allows a certain range to the behavioural response. Microhabitat segregation between the two species was pronounced, with brown trout inhabiting the more slow-flowing and partly more shallow stream areas. Atlantic salmon tolerated a wider range of water velocities and depths. Habitat suitability curves were produced from both species. It is suggested that habitat suitability curves that are based on observations of fish occupancy of habitat at median or base flow may not be suitable in habitat simulation models, where available habitat is projected at substantially greater water flows.     

The distribution of salmonids in upland streams in relation to depth and gradient

The distribution of various age classes of salmon and trout was assessed in upland streams by electrofishing. Water depths and site gradients were measured and correlated to fish densities. The fry of both species were significantly more abundant in shallow water; up to 75·3% of salmon fry and 72·2% of trout fry were captured in sites of mean depth < 20 cm. Older trout were found mainly in the deeper areas, with a maximum of 7·4% captured in sites < 20 cm mean depth. Yearling fish were found in all the depth-ranges sampled, but with a tendency for higher numbers in mid-range depths. There were similar correlations in the abundance of each age class with the actual areas of shallow, mid-range and deep water habitat available within sites. Correlations of fish density with gradient indicated that trout were limited in their distribution to areas of lower flow, whereas salmon were not. Since depth and gradient were significantly negatively correlated, there was an apparent preference of trout for slightly deeper habitats than the equivalent year classes of salmon. The observed habitat segregation is discussed in terms of competition and selection.