Influence of habitat structure and fish density on Atlantic salmon Salmo salar L. territorial behaviour

A laboratory stream experiment was performed to test the influence of young‐of‐ the‐year Atlantic salmon Salmo Salar density on habitat selection ( i.e . pools and riffles) and the combined effect of habitat and density on territoriality. Pools were preferred at low and medium fish density. At higher density, the saturation of the riffles led to an equal fish repartition in both habitats. Aggressive interactions increased according to fish density, and the level of aggression was significantly higher in the pools than in the riffles.     

Static habitat partitioning and dynamic selection by sympatric young Atlantic salmon and brown trout in south-west England streams

Direct underwater observation of micro‐habitat use by 1838 young Atlantic salmon Salmo salar [mean L_T 7·9 ± 3.1(s.d.) cm, range 3·19] and 1227 brown trout Salmo trutta (L_T 10·9 ± 5·0 cm, range 3·56) showed both species were selective in habitat use, with differences between species and fish size. Atlantic salmon and brown trout selected relatively narrow ranges for the two micro‐habitat variables snout water velocity and height above bottom, but with differences between size‐classes. The smaller fishes <7 cm held positions in slower water closer to the bottom. On a larger scale, the Atlantic salmon more often used shallower stream areas, compared with brown trout. The larger parr preferred the deeper stream areas. Atlantic salmon used higher and slightly more variable mean water velocities than brown trout. Substrata used by the two species were similar. Finer substrata, although variable, were selected at the snout position, and differences were pronounced between size‐classes. On a meso‐habitat scale, brown trout were more frequently observed in slow pool‐glide habitats, while young Atlantic salmon favoured the faster high‐gradient meso‐habitats. Small juveniles <7 cm of both species were observed most frequently in riffle‐chute habitats. Atlantic salmon and brown trout segregated with respect to use of habitat, but considerable niche overlap between species indicated competitive interactions. In particular, for small fishes <7 cm of the two species, there was almost complete niche overlap for use of water depth, while they segregated with respect to water velocity. Habitat suitability indices developed for both species for mean water velocity and water depth, tended to have their optimum at lower values compared with previous studies in larger streams, with Atlantic salmon parr in the small streams occupying the same habitat as favoured by brown trout in larger streams. The data indicate both species may be flexible in their habitat selection depending on habitat availability. Species‐specific habitat overlap between streams may be complete. However, between‐species habitat partitioning remains similar.     

Effects of predation, trout density and discharge on habitat use by brown trout, Salmo trutta, in artificial streams

1. The effects of predation risk, fish density and discharge on habitat use by juvenile brown trout, Salmo trutta, in four artificial streams were studied. Each stream contained three habitats, riffles, runs and pools, the latter two each being further divided into shallow margins and deeper mid-regions. 2. The presence of northern pike, Esox Indus, caused trout to decrease use of pool midregions, where pike also occurred, and to increase use of other habitats. Increasing the number of trout caused trout to increase use of pools and the shallow margins of runs. Decreasing discharge reduced the area of the run and pool margins covered by water, thereby reducing use of these areas by trout. 3. Habitat selection indices for the different treatments were calculated. The data indicated that riffles and the mid-regions of runs were preferred habitats, whereas run margins and pools were inferior habitats used when intraspecific fish densities were high. 4. Despite density- and discharge-dependent habitat use by trout, the number of trout consumed by pike was independent of trout density and discharge. 5. The results reveal the flexibility of habitat use by trout and illustrate the potential danger of applying data on habitat use in one stream to others where habitat availability and bioric interactions may differ.     

Linking microhabitat availability and local density of rainbow trout in low-gradient Japanese streams

Quantification of reach-based microhabitat availability for rainbow trout (Oncorhynchus mykiss), considering their microhabitat requirements in two low-gradient streams, northern Japan, was attempted to test for habitat space limitation of local trout density. Underwater observations revealed that fish selected microhabitats of moderate current velocity, relatively greater depth and shorter distance to overhead cover in both streams, although habitat features used and available differed slightly between the streams. Habitat space for fish potentially available in the channel environment was evaluated using principal component analysis (PCA) of both available and used microhabitat. A close relationship was evident in both streams between reach-based microhabitat availability and fish density, which was assessed by a three-pass removal method. Direct estimates of fish microhabitat availability using PCA can contribute to accurate predictions of local fish density and provide insight into the mechanisms responsible for fish–habitat relationships in streams.     

The importance of spatial scale in habitat selection by European beaver

We evaluated habitat selection by European beaver Castor fiber L. across a spatial gradient from local (within the family territory) to a broad, ecoregional scale. Based on aerial photography, we assessed the habitat composition of 150 beaver territories along the main water bodies of the Vistula River delta (northern Poland) and compared these data with 183 randomly selected sites not occupied by the species. The beavers preferred habitats with high availability of woody plants, including shrubs, and avoided anthropogenically modified habitats, such as arable lands. Within a single family territory, we observed decreasing woody plant cover with increasing distance from a colony centre, which suggests that beaver habitat preferences depend on the assessment of both the abundance and spatial distribution of preferred habitat elements. We tested the importance of spatial scale in beaver habitat selection with principal coordinates of neighbour matrices analysis, which showed that the geographical scale explained 46.7% of the variation in habitat composition, while the local beaver density explained only 10.3% of this variability. We found two main spatial gradients that were related to the broad spatial scale: first, the most important gradient was related to the largest distances between beaver sites and was independent of woody plant cover and the local beaver site density. The second most important gradient appeared more locally and was associated with these variables. Our results indicate that European beaver habitat selection was affected by different scale‐related phenomena related 1) to central place foraging behaviour, which resulted in the clumped distribution of woody plants within the territory, and 2) local population density and woody plant cover. Finally, 3) habitat selection occurs independently across the largest spatial scale studied (e.g. between watersheds), which was probably due to the limited natal dispersal range of the animals.     

‘Prudent habitat choice’: a novel mechanism of size‐assortative mating

Assortative mating, an ubiquitous form of nonrandom mating, strongly impacts Darwinian fitness and can drive biological diversification. Despite its ecological and evolutionary importance, the behavioural processes underlying assortative mating are often unknown, and in particular, mechanisms not involving mate choice have been largely ignored so far. Here, we propose that assortative mating can arise from ‘prudent habitat choice’, a general mechanism that acts under natural selection, and that it can occur despite a complete mixing of phenotypes. We show that in the cichlid Eretmodus cyanostictus size‐assortative mating ensues, because individuals of weaker competitive ability ignore high‐quality but strongly competed habitat patches. Previous studies showed that in E. cyanostictus, size‐based mate preferences are absent. By field and laboratory experiments, here we showed that (i) habitat quality and body size are correlated in this species; (ii) territories with more stone cover are preferred by both sexes in the absence of competition; and (iii) smaller fish prudently occupy vacant territories of worse quality than do larger fish. Prudent habitat choice is likely to be a widespread mechanism of assortative mating, as both preferences for and dominance‐based access to high‐quality habitats are generic phenomena in animals.     

Habitat quality influences population distribution, individual space use and functional responses in habitat selection by a large herbivore

Identifying factors shaping variation in resource selection is central for our understanding of the behaviour and distribution of animals. We examined summer habitat selection and space use by 108 Global Positioning System (GPS)-collared moose in Norway in relation to sex, reproductive status, habitat quality, and availability. Moose selected habitat types based on a combination of forage quality and availability of suitable habitat types. Selection of protective cover was strongest for reproducing females, likely reflecting the need to protect young. Males showed strong selection for habitat types with high quality forage, possibly due to higher energy requirements. Selection for preferred habitat types providing food and cover was a positive function of their availability within home ranges (i.e. not proportional use) indicating functional response in habitat selection. This relationship was not found for unproductive habitat types. Moreover, home ranges with high cover of unproductive habitat types were larger, and smaller home ranges contained higher proportions of the most preferred habitat type. The distribution of moose within the study area was partly related to the distribution of different habitat types. Our study shows how distribution and availability of habitat types providing cover and high-quality food shape ungulate habitat selection and space use.     

Diel Habitat Partitioning by Bull Charr and Cutthroat Trout During Fall and Winter in Rocky Mountain Streams

We used underwater observation to determine diel habitat partitioning between bull charr, Salvelinus confluentus, and cutthroat trout, Oncorhynchus clarki, during fall and winter (0.1–8.3°C) in two Rocky Mountain streams that differed in habitat availability. The majority (>70%) of both species emerged from concealment cover at night, though bull charr exhibited a greater tendency for nocturnal behavior than cutthroat trout. Differences in day and night counts were most pronounced at temperatures <3°C, when very few fish of either species were observed in the water column during the day, but both species were common at night. Both species used concealment cover of large woody debris and boulder substrate crevices in deep pools during the day. At night, fish emerged from cover and habitat use shifted to shallow water with low cover. Microhabitat partitioning among species and size classes occurred at night, cutthroat trout moving into shallower, faster water that was farther from cover compared to bull charr. Smaller fish of both species occupied focal positions in slower, shallower water closer to the substrate than larger fish. Large, mixed-species aggregations also were common in beaver ponds both day and night. High variation in diel and site-specific winter habitat use suggests the need for caution in developing habitat suitability criteria for salmonids based solely on daytime observations or on observations from a few sites. Our results support the need to incorporate nocturnal habitat use and partitioning in studies of salmonid ecology.     

Importance of rare habitats and riparian zones in a tropical forest fragment: preferential use by Tayassu pecari, a wide-ranging frugivore

In forest fragments, rare habitats contribute to heterogeneity and may provide unique resources for frugivorous species like peccaries with spatially and temporally complex patterns of range use. This study examined seasonal habitat use by two sympatric peccary species ( Tayassu pecari and Tayassu tajacu ) in an Atlantic forest fragment on the plateau region of São Paulo state, Brazil. Previous studies showed that range use by T. pecari was highly nonrandom, and that both species persisted at population densities typical of larger forest fragments. To explain this, we quantified the use and availability of habitats and riparian zones with compositional analysis. Use by T. pecari was nonrandom with respect to availability. Habitat preferences were different during dry and wet seasons and corresponded closely with seasonal movements, core range and fruit use. Although aquatic habitats made up a small proportion of the herd home range, they were most preferred by T. pecari during both seasons. Headwater palmito Euterpe edulis swamps were most favored in the dry season, whereas swamps and marshes near larger streams were preferred in the wet season. Tayassu pecari preferred riparian zones <50 m from streams over drier zones. These habitats were important sources of fruits, travel routes and corridors between forest patches in the agricultural matrix. The least-preferred habitat of T. pecari during both seasons was disturbed forest edge dominated by bamboo. Habitat and riparian zone use were herd-specific for T. tajacu and related to habitat quality and composition where stable home ranges had been established. The persistence of viable peccary populations after 75 years of fragmentation-associated pressures is related to preservation of rare habitats and overall habitat diversity. Thus, T. pecari is an indicator of high habitat diversity in forest fragments and will function as an umbrella species when targeted for conservation.     

Effect of fluctuating flow and temperature on cover type selection and behaviour by juvenile brown trout in artificial flumes

Brown trout Salmo trutta in a flume used cover more often and were more aggregated at low water temperatures in winter compared with summer. Cover combinations providing the most cover (overhead, visual and velocity cover) were preferred in winter whereas velocity refuges were preferred in late summer; and brown trout rarely used any of these cover structures in early summer. Flow affected cover use especially in winter, when most of the trout moved to shelters during the first high flow period. Movements by brown trout were most frequent in early summer. The much higher aggression rate in summer compared with winter decreased during the first period of high flow in both summer experiments. The results indicate seasonal changes in cover habitat and cover type preferences suggesting that habitat complexity is important, and availability of cover is particularly important during different seasons and fluctuating flow.     

Comparisons of habitat availability and habitat use by an allopatric cohort of juvenile Atlantic salmon Salmo salar under conditions of low competition in a Norwegian stream

Allopatric Atlantic salmon parr occupied a wider and lower range of mean water velocities than have been previously reported for parr in sympatry. With regard to depth, substrate and cover no substantial differences were found. Different fish sizes did not affect habitat use by parr significantly, neither did habitat use change during the season. Habitat availability strongly influenced observed habitat use thus, invalidating use of habitat suitability curves based on observations of habitat occupancy by fish only. The allopatric parr consistently selected habitats different from the available habitat, i.e. had preferences also in the absence of intraspecific interactions between age classes. Habitat conditions not used by the fish were more readily identifiable than habitats used. The parr rarely used shallow (≤ 10 cm) and deep (> 60 cm) stream areas, low mean water velocities (< 10 cm s^-1), fine substrate (sand and finer) and stream areas without cover. However, the calculated habitat preferences were also affected by habitat availability. Multivariate analysis did not identify any of the habitat variables as substantially more important than the others, but suggested that the importance of variables may vary with type of habitat studied.     

Habitat use by the fish assemblages of two chalk streams

Patterns of habitat use by fish assemblages in two chalk streams in southern England were examined to identify species preferences with respect to major habitat gradients. Both study sites, although differing in some physical habitat characteristics, mainly channel width, water temperature and instream cover, could be arranged similarly along a continuum extending from erosional to depositional habitats. Twelve fish species were collected from stream surveys conducted during July 1993. The habitat was partitioned into six fish species associations that could be assigned to three habitat guilds: depositional ( Barbatula barbatula, Gasterosteus aculeatus and Anguilla anguilla ), riffle ( Leuciscus leuciscus, Thymallus thymallus and Salmo salar ) and generalist ( Salmo trutta, Phoxinus phoxinus, Gobio gobio, Cottus gobio, Lampetra planeri and Pleuronectes flesus ). At low to moderate densities the different species associations were collected in the same habitat patch, but at higher densities there was a clear tendency to mutual avoidance. In particular, large trout appeared as strong interactors that tended to exclude other species from their territories. It is hypothesized that the fish assemblage of the Mill Stream and Bere Stream have partitioned the habitat in such a way as to minimize potential Competition.     

Habitat shifts in rainbow trout: competitive influences of brown trout

Summary We compared habitat use by rainbow trout sympatric (three streams) and allopatric (two streams) with brown trout to determine whether competition occurred between these two species in the southern Appalachian Mountains. We measured water depth, water velocity, substrate, distance to overhead vegetation, sunlight, and surface turbulence both where we collected trout and for the streams in general. This enabled us to separate the effects of habitat availability from possible competitive effects. The results provided strong evidence for asymmetrical interspecific competition. Habitat use varied significantly between allopatric and sympatric rainbow trout in 68% of the comparisons made. Portions of some differences refelected differences in habitats available in the several streams. However, for all habitat variables measured except sunlight, rainbow trout used their preferred habitats less in sympatry with brown trout than in allopatry if brown trout also preferred the same habitats. Multivariate analysis indicated that water velocity and its correlates (substrate particle size and surface turbulence) were the most critical habitat variables in the interaction between the species, cover in the form of shade and close overhead vegetation was second most important, and water depth was least important.     

Nonideal nest box selection by tree swallows breeding in farmlands: Evidence for an ecological trap?

Animals are expected to select a breeding habitat using cues that should reflect, directly or not, the fitness outcome of the different habitat options. However, human‐induced environmental changes can alter the relationships between habitat characteristics and their fitness consequences, leading to maladaptive habitat choices. The most severe case of such nonideal habitat selection is the ecological trap, which occurs when individuals prefer to settle in poor‐quality habitats while better ones are available. Here, we studied the adaptiveness of nest box selection in a tree swallow (Tachycineta bicolor) population breeding over a 10‐year period in a network of 400 nest boxes distributed along a gradient of agricultural intensification in southern Québec, Canada. We first examined the effects of multiple environmental and social habitat characteristics on nest box preference to identify potential settlement cues. We then assessed the links between those cues and habitat quality as defined by the reproductive performance of individuals that settled early or late in nest boxes. We found that tree swallows preferred nesting in open habitats with high cover of perennial forage crops, high spring insect biomass, and high density of house sparrows (Passer domesticus), their main competitors for nest sites. They also preferred nesting where the density of breeders and their mean number of fledglings during the previous year were high. However, we detected mismatches between preference and habitat quality for several environmental variables. The density of competitors and conspecific social information showed severe mismatches, as their relationships to preference and breeding success went in opposite direction under certain circumstances. Spring food availability and agricultural landscape context, while related to preferences, were not related to breeding success. Overall, our study emphasizes the complexity of habitat selection behavior and provides evidence that multiple mechanisms may potentially lead to an ecological trap in farmlands.     

Behavioural and physiological responses of brook trout Salvelinus fontinalis to midwinter flow reduction in a small ice-free mountain stream

This study presents an experimental analysis of the effects of midwinter flow reduction (50-75%, reduction in discharge in 4 h daily pulses) on the physical habitat and on behaviour and physiology of overwintering brook trout Salvelinus fontinalis in a small mountain stream. Flow reduction did not result in significant lowering of temperature or formation of surface or subsurface ice. The main findings were (1) daily movement by S. fontinalis increased (c. 2·5-fold) during flow reduction, but was limited to small-scale relocations (<10 m). (2) Undercut banks were the preferred habitat and availability of these habitats was reduced during flow reduction. (3) Although both experimental and reference fish did lose mass and condition during the experiment, no effects of flow reduction on stress indicators (blood cortisol or glucose) or bioenergetics (total body fat, water content or mass loss) were detected, probably because access to the preferred type of cover remained available. Like other salmonids, S. fontinalis moves little and seeks physical cover during winter. Unlike many of the more studied salmonids, however, this species overwinters successfully in small groundwater-rich streams that often remain ice-free, and this study identifies undercut banks as the critical winter habitat rather than substratum cover.     

Habitat selection of breeding brown eared-pheasants (Crossoptilon mantchuricum) in Xiaowutaishan National Nature Reserve, Hebei Province, China

From April to July of 2003–2005, we investigated habitat selection of breeding brown eared-pheasants (Crossoptilon mantchuricum) in the Xiaowutaishan National Nature Reserve, Hebei Province. Our results show that brown eared-pheasants preferred deciduous-coniferous mixed or deciduous forests with altitudes ranging from 1600 to 2200 m, steep gradients of slopes and medium or high positions on slopes. Moreover, they preferred habitats with more tree species, larger diameters at the breast height, more density and cover of trees and habitats with more types of grasses, high density and cover of grasses, but with fewer types of shrubs and lower density and cover of shrubs. They had larger territories in the early breeding stage than in the late breeding stage. There were significant differences in diameters at the breast height of trees, average height of trees, cover of trees, density of shrubs, cover of shrubs, aspect, distance to path, distance to water, distance to forest edge (P < 0.01, respectively), density of trees, type of shrubs, height of shrubs, type of grasses, density of grasses and proportion of bareness (P < 0.05, respectively). The result of the principal component analysis suggested that the principal components among the 23 habitat factors were the average height of trees, density of trees, diameters at the breast height of trees, density of shrubs, cover of shrubs and density of grasses. The differences in habitat selection of brown eared-pheasants between early breeding and late breeding stages may correlate with the dynamics of ambient environmental conditions, the reproductive behavior and energy requirements in their sub-stages of breeding.     

Habitat selection of breeding brown eared-pheasants (Crossoptilon mantchuricum) in Xiaowutaishan National Nature Reserve, Hebei Province, China

From April to July of 2003-2005,we investigated habitat selection of breeding brown eared-pheasants (Crossoptilon mantchuricum) in the Xiaowutaishan National Nature Reserve,Hebei Province.Our results show that brown eared-pheasants preferred deciduousconiferous mixed or deciduous forests with altitudes ranging from 1600 to 2200 m,steep gradients of slopes and medium or high positions on slopes.Moreover,they preferred habitats with more tree species,larger diameters at the breast height,more density and cover of trees and habitats with more types of grasses,high density and cover of grasses,but with fewer types of shrubs and lower density and cover of shrubs.They had larger territories in the early breeding stage than in the late breeding stage.There were significant differences in diameters at the breast height of trees,average height of trees,cover of trees,density of shrubs,cover of shrubs,aspect,distance to path,distance to water,distance to forest edge (P ＜ 0.01,respectively),density of trees,type of shrubs,height of shrubs,type of grasses,density of grasses and proportion of bareness (P ＜ 0.05,respectively).The result of the principal component analysis suggested that the principal components among the 23 habitat factors were the average height of trees,density of trees,diameters at the breast height of trees,density of shrubs,cover of shrubs and density of grasses.The differences in habitat selection of brown eared-pheasants between early breeding and late breeding stages may correlate with the dynamics of ambient environmental conditions,the reproductive behavior and energy requirements in their sub-stages of breeding.     

Habitat selection by wintering male and female Snowy Owls on the Canadian prairies in relation to prey abundance and a competitor,the Great Horned Owl

Birds overwintering at high latitudes may find it challenging to meet their energy budgets when thermoregulatory costs are high and food availability is low. Snowy Owls (Bubo scandiacus), like most raptors, exhibit reversed sexual size dimorphism, so , if availability of high‐quality (food‐rich) habitats is limited, we predicted that larger and dominant females would use better ‐ quality habitat than males. During the winters of 2014–2015 and 2015–2016 in Saskatchewan , where many Snowy Owls overwinter annually, we measured prey (small mammal) abundance in fields with four types of cover, including cut stalks (stubble) of canola, grain and legume crops, and pasture, and related this estimate of quality to habitat selection by males and females. Small mammal abundance varied annually , but not among the three types of crop stubble. However, prey were less abundant in pastures than in the three types of crop cover in one of three years. Biweekly surveys of owls conducted during the two winters along a 60 ‐ km transect revealed weak selection for legume fields, especially by males. The home ranges of nine females with transmitters included proportionally less canola stubble than those of eight males with transmitters. Within home ranges, males avoided canola stubble and tended to use legume fields more, whereas females used all four habitat types in proportion to availability. Fewer Snowy Owls than expected were observed at locations along the transect within 800 m of Great Horned Owls (Bubo virginianus) and their associated habitats , suggesting that Snowy Owls also avoided these potential competitors on the landscape. Our results suggest that larger females outcompete smaller male Snowy Owls for home ranges in preferred habitat with less canola stubble because stubble‐free legume fields provide easier access to prey than canola fields with numerous rigid stalks.     

Seasonal changes in habitat use by Houbara Bustards Chlamydotis [undulata] macqueenii in northern Saudi Arabia

Habitat preferences by Houbara Bustards Chlamydotis [undulata] macqueenii in Harrat al-Harrah reserve, in northern Saudi Arabia were determined from sightings of birds in all seasons over three years. Vegetation and crawling invertebrate abundance were sampled in each habitat. Houbara Bustards showed seasonally changing habitat preferences that appeared to be influenced primarily by vegetation phenology, abundance and cover. More densely vegetated areas (10–17% cover) were preferred. Seasonal and inter-habitat variations in invertebrate numbers were not reflected in differential habitat use by Houbara Bustards. The highest selection ratio for a single habitat (dry lakes) occurred in summer, coinciding with the fruiting of Shafallah Capparis spinosa. Selectivity of habitats was least in spring, when green vegetation was most widespread. Changes in Houbara Bustard habitat preferences in response to marked seasonal changes in habitats brought about by well-defined patterns of rainfall indicate that studies of habitat selection should consider the entire annual cycle. The importance of vegetative cover and the sensitivity of Houbara Bustards to human disturbance suggest that reserves set aside for Houbara Bustards should be extensive, diverse and largely free of livestock, human occupation and its associated disturbances.     

Differential Habitat Selection by Pygmy Grasshopper Color Morphs; Interactive Effects of Temperature and Predator Avoidance

Habitat selection behavior is affected by complex interplays between competing requirements. Here we combine field observations with laboratory experiments to examine how thermal benefits and predator avoidance influences habitat selection by different color morphs of the pygmy grasshopper Tetrix undulata. The composition of substrate types and surface temperatures in areas selected by free-ranging individuals did not reflect relative availability, and varied among morphs and sexes. Surface temperatures of selected habitats deviated less from the range of preferred body temperatures than would result from a random utilization of surface temperatures, suggesting that grasshoppers selected habitats with thermal properties which were suitable for maintaining preferred body temperatures. The thermal property of habitats occupied by different color morphs suggests that darker morphs (which absorb more solar radiation) selected cooler habitats to avoid overheating. Dissimilarities in substrate use among color morphs in the field and laboratory emphasize a role also of predator avoidance by background matching for habitat choice. The degree of habitat selectivity was lowest in the striped morph, supporting the notion that a disruptive color pattern may constitute a solution to the trade-off between relative crypsis in different visual backgrounds. Finally, individuals modified their habitat use when subjected to elevated risk of predation, showing that habitat choice is governed by conflicting priorities. Collectively, our findings suggest that, as a result of direct and indirect effects of coloration on performance, alternative color morphs use different solutions to the trade-off between competing requirements. Our results also lend support to the notion that relative fitness of alternative color morphs and sexes may be dependent on microhabitat selection, as predicted by the theory of multiple niche polymorphisms.