Movements and habitat use by PIT-tagged Atlantic salmon parr in early winter: the influence of anchor ice

1. Movements and habitat use by Atlantic salmon parr in Catamaran Brook, New Brunswick, were studied using Passive Integrated Transponder technology. The fish were tagged in the summer of 1999, and a portable reading system was used to collect data on individual positions within a riffle‐pool sequence in the early winter of 1999. Two major freezing events occurred on November 11–12 (Ice 1) and November 18–19 (Ice 2) that generated significant accumulations of anchor ice in the riffle. 2. Individually tagged parr (fork length 8.4–12.6 cm, n = 15) were tracked from 8 to 24 November 1999. Over this period, emigration (40%) was higher from the pool than from the riffle. Of the nine parr that were consistently located, seven parr moved <5 m up‐ or downstream, and two parr moved more than 10 m (maximum 23 m). Parr moved significantly more by night than by day, and diel habitat shifts were more pronounced in the pool with some of the fish moving closer to the bank at night. 3. During Ice 2, there was relatively little movement by most of the parr in the riffle beneath anchor ice up to 10 cm in thickness. Water temperature was 0.16 °C above the freezing point beneath anchor ice, suggesting the existence of suitable habitats where salmon parr can avoid supercooling conditions and where they can have access to low velocity shelters. To our knowledge, these are the first data on habitat use by Atlantic salmon parr under anchor ice.     

Habitat overlap between predatory benthic fish and their invertebrate prey in streams: the relative influence of spatial and temporal factors on predation risk

1. The spatial heterogeneity of ecosystems as well as temporal activity patterns of organisms can have far‐reaching effects on predator–prey relationships. We hypothesised that spatiotemporal constraints in mesohabitat use by benthic fish predators would reduce habitat overlap with benthic invertebrates and lead to mesohabitat‐specific predation risks. 2. We analysed the spatiotemporal activity patterns of two small‐bodied benthivorous fishes, gudgeon (Gobio gobio) and stone loach (Barbatula barbatula), and of benthic invertebrates in a small temperate stream during three 24‐h field experiments. By applying a novel method of field video observation, we monitored the spatiotemporal foraging behaviour of the fish in their natural environment. A parallel analysis of invertebrate mesohabitat use by means of small area Hess sampling allowed a direct estimation of habitat overlap at a pool–riffle scale. 3. Gudgeon showed a dominant spatial activity pattern preferring pools at all times of day, whereas stone loach used both mesohabitats but with a distinct temporal (nocturnal) activity pattern. The patterns of residence were not identical with those of active foraging. Invertebrate community composition differed significantly between mesohabitats but not between times of day. More than half of the total dissimilarity between pools and riffles was accounted for by six invertebrate taxa. Five of these were subject to higher fish predation in pools than in riffles. The total prey consumption of the two fish species together in pools was about three times as high as in riffles. Trophic niche breadth of stone loach and thus its predation range was broader than that of gudgeon. 4. These results indicate that the potential predation risk for stream invertebrates depends on the combination of spatial and temporal patterns of both predator and prey. Given the distinct differences in predation risk found between pools and riffles, we conclude that spatial heterogeneity at the mesohabitat scale can influence mechanisms and consequences of selective predation. We also suggest that the analysis of spatiotemporal predator–prey relationships should not be based on the premise that the main residence habitat and active foraging habitat of a predator are identical.     

Reconstructing riverine mesohabitat unit composition using fish community data and an autecology matrix

This research proposes a simplified method for estimating the mesohabitat composition that would favour members of a given set of aquatic species. The simulated composition of four types of mesohabitat units (deep pool, shallow pool, deep riffle and shallow riffle) could guide the design of in‐stream structures in creating pool‐riffle systems with ecological reference. Fish community data and an autecology matrix are used to support the development of a stream mesohabitat simulation based on regression models for reaches in mid to upper‐order streams. The fish community‐mesohabitat model results constitute a reference condition that can be used to guide stream restoration and ecological engineering decisions aimed at maintaining the natural ecological integrity and diversity of rivers.     

Ice‐dependent winter survival of juvenile Atlantic salmon

Changes in snow and ice conditions are some of the most distinctive impacts of global warming in cold temperate and Arctic regions, altering the environment during a critical period for survival for most animals. Laboratories studies have suggested that reduced ice cover may reduce the survival of stream dwelling fishes in Northern environments. This, however, has not been empirically investigated in natural populations in large rivers. Here, we examine how the winter survival of juvenile Atlantic salmon in a large natural river, the River Alta (Norway, 70°N), is affected by the presence or absence of surface ice. Apparent survival rates for size classes corresponding to parr and presmolts were estimated using capture‐mark‐recapture and Cormack‐Jolly‐Seber models for an ice‐covered and an ice‐free site. Apparent survival (Φ) in the ice‐covered site was greater than in the ice‐free site, but did not depend on size class (0.64 for both parr and presmolt). In contrast, apparent survival in the ice‐free site was lower for larger individuals (0.33) than smaller individuals (0.45). The over‐winter decline in storage energy was greater for the ice‐free site than the ice‐covered site, suggesting that environmental conditions in the ice‐free site caused a strong depletion in energy reserves likely affecting survival. Our findings highlight the importance of surface ice for the winter survival of juvenile fish, thus, underpinning that climate change, by reducing ice cover, may have a negative effect on the survival of fish adapted to ice‐covered habitats during winter.     

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.     

Mid-winter activity and movement of Atlantic salmon parr during ice formation events in a Norwegian regulated river

A telemetry study in a Norwegian regulated river was conducted through a 12-day period in mid-winter 2003. The objective was to study activity (defined as number of movement per hour) and movement (defined as distance moved per hour) during different ice formation events. Twenty-four Atlantic salmon (Salmo salar L.) parr were radio tagged and continuously monitored by both manually tracking (N = 24) and by fixed recording stations (N = 15). Detailed data on climate, flow and ice formation and its spatial distribution were collected and used in the analyses. Fish activity was not found to be affected by their size (L _F). There was a significant difference in activity between diel periods with highest activity during dusk (5–6 p.m.). Between high and low flow (mean ± SD, 21.1 m³ s⁻¹ ± 1.7 SD and 11.1 m³ s⁻¹ ± 1.7 SD, respectively) no significant difference in activity was found. During the experiment extensive anchor ice growth occurred mainly in the riffle part with thickness up to 50 cm. Juveniles tend to avoid riffle section during anchor ice formation and exploited ice covered areas, indicating critical and preferable habitats respectively. Further, a significant difference in movement was found between five selected ice events with highest mean movement during an anchor ice event and lowest mean movement during an ice break up with no anchor ice formation. No significant difference in activity or movement between parr exposed to frazil ice and parr not exposed were found.     

Beaver dams maintain fish biodiversity by increasing habitat heterogeneity throughout a low‐gradient stream network

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Habitat effects on invertebrate drift in a small trout stream: implications for prey availability to drift-feeding fish

In this study, we focused on the drivers of micro- and mesohabitat variation of drift in a small trout stream with the goal of understanding the factors that influence the abundance of prey for drift-feeding fish. We hypothesized that there would be a positive relationship between velocity and drift abundance (biomass concentration, mg/m³) across multiple spatial scales, and compared seasonal variation in abundance of drifting terrestrial and aquatic invertebrates in habitats that represent the fundamental constituents of stream channels (pools, glides, runs, and riffles). We also examined how drift abundance varied spatially within the water column. We found no relationship between drift concentration and velocity at the microhabitat scale within individual pools or riffles, suggesting that turbulence and short distances between high- and low-velocity microhabitats minimize changes in drift concentration through settlement in slower velocity microhabitats. There were also minimal differences in summer low-flow drift abundance at the mesohabitat scale, although drift concentration was highest in riffle habitats. Similarly, there was no differentiation of drifting invertebrate community structure among summer samples collected from pools, glides, runs, and riffles. Drift concentration was significantly higher in winter than in summer, and variation in drift within individual mesohabitat types (e.g., pools or riffles) was lower during winter high flows. As expected, summer surface samples also had a significantly higher proportion of terrestrial invertebrates and higher overall biomass than samples collected from within the water column. Our results suggest that turbulence and the short length of different habitat types in small streams tend to homogenize drift concentration, and that spatial variation in drift concentrations may be affected as much by fish predation as by entrainment rates from the benthos. Handling editor: Robert Bailey     

Do pools impede drift dispersal by stream insects?

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Conflicting effects of woody debris on stream fish populations: implications for management

1. Coarse woody debris (CWD) in stream channels causes changes in flow, sedimentation and ratios of pool to riffle areas. There is a consensus among fishery managers and scientists that CWD is beneficial to stream fish communities because of its enhancement of habitat diversity, invertebrate production and cover. Our hypothesis was that CWD accumulation or introduction would not increase in‐stream habitat capacity for all species and their ontogenic stages at reach and stream scales. 2. The study used a system of gravel‐bed streams with naturally dynamic CWD accumulations and a fish community consisting of Salmo trutta, Cotttus gobio, Phoxinus phoxinus, Lampetra cf planeri, Nemacheilus barbatulus and Anguilla anguilla. Cotttus gobio and L. cf planeri are protected by an EU Directive and S. trutta is exploited for angling. Riffles, pools and CWD matrices, considered as the basic habitat/spatial units of channel structure, were sampled separately and abundance of each fish species quantified seasonally at each spatial scale. 3. Multiple‐pass electric fishing techniques were used. Capture efficiencies were calculated for species, habitat and season. Areal densities (number m^?2) were compared for habitat types and season using nonparametric anova . Canonical analysis and stepwise multiple regression were used to show the most influential physical variables on fish density. Densities were also compared by unit volume (numbers m^?3) for pools and CWD matrices to investigate direct three‐dimensional use for cover. Reach‐scale densities for each fish species in relation to habitat composition were made using Spearman rank correlation of habitat‐scale densities with proportionate areas of the different habitat units in the reach. 4. Habitat‐scale densities of bullheads and age 0+ trout were negatively correlated with depth and CWD areas for some seasons. Densities of lampreys, older trout, eels and minnows were positively correlated with depth in some seasons. Water depth had the most consistent influence on fish abundance at the habitat unit scale. Three‐dimensional comparisons of pools and CWD matrices indicated that only trout older than 1+ may use CWD habitats as cover. 5. Reach‐scale densities of 0+ trout and bullheads were significantly correlated with proportion of riffle area and negatively with CWD and combined CWD‐pool habitat area in the reach. Densities of older trout, large eels and lampreys were positively correlated with CWD area and combined CWD‐pool area in some seasons. Inundation of riffles caused by impoundment upstream of CWD accumulations reduced spawning habitat for trout, bullheads, brook lampreys, minnows and stone loach. A trade‐off was an increase in refugia for older trout, minnows and eels. 6. Coarse woody debris accumulation in streams is not beneficial to all species or ontogenic stages in a mixed species population and could severely limit essential habitat areas for some species. Thus, physical manipulation of channels should be implemented only after a thorough study of the habitat relationships of all species present, especially where protected species coexist with target species. The relative importance of in‐stream morphological changes depends on the spatial and temporal scale of the species life histories.     

Comparative diets and foraging strategies of subyearling Atlantic salmon,brown trout,and rainbow trout during winter

Over‐winter survival of salmonids in streams is thought to be an important population regulation mechanism. Yet because of the difficulty of conducting field studies due to adverse weather or ice conditions, compared to other seasons, salmonid ecology during winter is least understood. Consequently, we sought to examine interspecific feeding associations of an important salmonid stream assemblage in the Lake Ontario watershed during winter. The diets of Atlantic salmon (Salmo salar) parr, brown trout (S. trutta) parr, and rainbow trout (Oncorhynchus mykiss) parr were significantly different in February but not in March. Salmonid diets differed from the benthos and the drift during both months. Dipterans (chironomids, simuliids, and tipulids) and ephemerellids were the major prey taxa consumed. All three species fed more heavily on prey items from the benthos than from the drift. The diet of Atlantic salmon had the highest similarity to the benthos whereas the diet of brown trout had the lowest similarity to the drift. All three salmonid species generally selected ephemerellids, limnephilids, and chironomids and avoided elmids. These winter feeding observations are the first reported for this specific salmonid assemblage and will help managers better understand interspecific associations during this critical period.     

Maternal behaviours may explain riffle‐scale variations in some stream insect populations

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Identifying keystone habitats with a mosaic approach can improve biodiversity conservation in disturbed ecosystems

Conserving native biodiversity in the face of human‐ and climate‐related impacts is a challenging and globally important ecological problem that requires an understanding of spatially connected, organismal‐habitat relationships. Globally, a suite of disturbances (e.g., agriculture, urbanization, climate change) degrades habitats and threatens biodiversity. A mosaic approach (in which connected, interacting collections of juxtaposed habitat patches are examined) provides a scientific foundation for addressing many disturbance‐related, ecologically based conservation problems. For example, if specific habitat types disproportionately increase biodiversity, these keystones should be incorporated into research and management plans. Our sampling of fish biodiversity and aquatic habitat along ten 3‐km sites within the Upper Neosho River subdrainage, KS, from June‐August 2013 yielded three generalizable ecological insights. First, specific types of mesohabitat patches (i.e., pool, riffle, run, and glide) were physically distinct and created unique mosaics of mesohabitats that varied across sites. Second, species richness was higher in riffle mesohabitats when mesohabitat size reflected field availability. Furthermore, habitat mosaics that included more riffles had greater habitat diversity and more fish species. Thus, riffles (<5% of sampled area) acted as keystone habitats. Third, additional conceptual development, which we initiate here, can broaden the identification of keystone habitats across ecosystems and further operationalize this concept for research and conservation. Thus, adopting a mosaic approach can increase scientific understanding of organismal‐habitat relationships, maintain natural biodiversity, advance spatial ecology, and facilitate effective conservation of native biodiversity in human‐altered ecosystems.     

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.     

Macrozoobenthos response to environmental degradation in a heavily modified stream: Case study the Upper Elbe River,Czech Republic

Benthic macroinvertebrates are an important indicator of river health. However, their response upon water quality development downstream the pollution outlets considerably depends on the environmental habitat characteristics. Three successive stretches, each of them providing three different mesohabitats in stillwater (S), torrential (T) and riparian (R) zones were selected for evaluation of the impact of altered metapotamal river bed morphology (channelization) and chemical determinants of water quality on the Upper Elbe River. In downstream direction, the stretches are separated by weirs and characterized as a low polluted low modified natural stream (N), a low polluted channelized stream (C) and a channelized polluted stream (CP). Altogether, 111 benthic macroinvertebrate taxa were recorded in the Pardubice hotspot between Němčice and Přelouč. Despite different levels of stream bed and water quality degradation, micro- and mesohabitat characteristics appeared to be the most important factors determining the diversity of macrozoobenthos in riffle (substrate size structure) and in shoreline (macrophyte community composition and structure) mesohabitats. The diversity of macroinvertebrate communities was highest in riparian mesohabitats compared to stillwater and torrential ones. Saprobic indices increased in downstream direction, thus indicating the decline of water quality.     

Environmental factors influencing the regional distribution and local density of a small benthic fish: the stoneloach (<Emphasis Type="Italic">Barbatula barbatula</Emphasis>)

We investigated the relationships between different environmental variables and the spatial distribution patterns of the stoneloach (Barbatula barbatula) at the stream system, the stream site, and the mesohabitat (riffle/pool) scales in south-western France. Stoneloach occurred at 240 sites (out of 554 sampling sites), chiefly close to the source, in areas at low elevation and with weak slopes. Population density at a site was primarily influenced by physical conditions. Stream width was positively related to the probability of presence of stoneloach within the stream system, but negatively related to local density. These results indicate that stoneloaches can occur in a wide range of streams, but they are less abundant in wide rivers, probably because of lower habitat heterogeneity. Slope was negatively correlated to both fish presence at the regional scale and local density, suggesting that stoneloach’s swimming performance were weak under greater erosive forces. These results suggested that the distribution of populations and the density of stoneloach were governed by the suitability of physical habitat. Multi-scale studies of factors influencing a species’distribution allow to integrate patterns observed at different scales, and enhance our understanding of interactions between animals and their environment. The use of few pertinent variables in successful final models could reduce the effort and cost of data collection for water management applications.     

Behavioural strategies for species co‐existence during environmental extremes: competition between trout and salmon parr during dewatering

Competition between species of animals can be predicted to be intense under extreme environmental conditions during which niche overlap increases. Fluvial aquatic systems fluctuate naturally across a broad range of time‐scales. Land management activities impose further extreme fluctuations, particularly when water is abstracted for irrigation during summer. This study focused on the interaction between Atlantic salmon and brown trout parr during acute dewatering events. Brown trout are known to compete strongly for pool habitat whereas salmon can coexist by using riffle areas during normal flows. It is not known, however, how competition between the species affects their behavioural responses to extreme low flows when riffle areas decrease. Replicated groups of salmon were held in allopatry and sympatry with trout in sections of a large indoor stream, each of which was landscaped into riffles and pools. Space use and behaviours of the fish were recorded by direct visual observations and a network of Passive Integrated Transponder detectors. Here we report the response of the fish to dewatering and consider the system as a model for natural and forced responses of communities to environmental extremes.     

Spatiotemporal variability of stream habitat and movement of three species of fish

Relationships between environmental variability and movement are poorly understood, due to both their complexity and the limited ecological scope of most movement studies. We studied movements of fantail (Etheostoma flabellare), riverweed (E. podostemone), and Roanoke darters (Percina roanoka) through two stream systems during two summers. We then related movement to variability in measured habitat attributes using logistic regression and exploratory data plots. We indexed habitat conditions at both microhabitat (i.e., patches of uniform depth, velocity, and substrate) and mesohabitat (i.e., riffle and pool channel units) spatial scales, and determined how local habitat conditions were affected by landscape spatial (i.e., longitudinal position, land use) and temporal contexts. Most spatial variability in habitat conditions and fish movement was unexplained by a site’s location on the landscape. Exceptions were microhabitat diversity, which was greater in the less-disturbed watershed, and riffle isolation and predator density in pools, which were greater at more-downstream sites. Habitat conditions and movement also exhibited only minor temporal variability, but the relative influences of habitat attributes on movement were quite variable over time. During the first year, movements of fantail and riverweed darters were triggered predominantly by loss of shallow microhabitats; whereas, during the second year, microhabitat diversity was more strongly related (though in opposite directions) to movement of these two species. Roanoke darters did not move in response to microhabitat-scale variables, presumably because of the species’ preference for deeper microhabitats that changed little over time. Conversely, movement of all species appeared to be constrained by riffle isolation and predator density in pools, two mesohabitat-scale attributes. Relationships between environmental variability and movement depended on both the spatiotemporal scale of consideration and the ecology of the species. Future studies that integrate across scales, taxa, and life-histories are likely to provide greater insight into movement ecology than will traditional, single-season, single-species approaches. The Virginia Cooperative Fish and Wildlife Research Unit is jointly sponsored by the US Geological Survey, Virginia Polytechnic Institute and State University, Virginia Department of Game and Inland Fisheries, and Wildlife Management Institute.     

Relationships between fishes and habitat in rainforest streams in Sabah, Malaysia

Distinct fish assemblages were found at the mesohabitat scale in 14 streams in eastern Sabah, Malaysia. Sites were designated a priori as pool, run or riffle on the basis of physical habitat structure and properties. Principal components analysis of physical habitat data confirmed the validity of the a priori designation with a major axis of three correlated variables: water velocity, depth and substratum type. Canonical discriminant analysis on fish abundance and biomass data confirmed the existence of a specialized assemblage of fishes from riffle areas of all streams. Overall, pool and run assemblages were highly variable, dependent on stream size, but also variable between streams of the same size. Multiple regression of species richness, diversity, abundance and biomass data on principal components revealed significant but low correlations with measured habitat variables. Riffle habitats showed lower species richness and diversity but high abundance. The fish assemblage in riffles was dominated by balitorid species, specialized for fast-water conditions. Pool assemblages had the highest species diversity and were dominated by cyprinid species of a number of morphological and ecological guilds. Run assemblages were intermediate in assemblage characteristics between riffle and pool assemblages. Between-stream variation in assemblage composition was less than within-stream variation. Of 38 species collected, seven could be designated as riffle specialists, 18 as pool specialists and 13 as ubiquitous, although most of the latter showed size-specific habitat use with larger size classes found in slower, deeper water.     

Role of habitat in determining macroinvertebrate production in an intermittent-stream system

1. The effect of channel drying on macroinvertebrate production was studied at the habitat and reach scale in a catchment drained by intermittent streams in Maine, U.S.A. The catchment includes two first‐order streams and their second‐order confluence. Six reaches were selected for study based on differences in channel slope and habitat cover (bedrock, riffle/run, debris dam and pool). Stream water in each reach was acidic and oligotrophic. 2. The study reaches had different degrees of channel drying. In the first‐order reaches, surface flow ceased earlier in the season and for longer periods than second‐order reaches. Regardless of reach, pool and debris dam habitats retained water longer than riffle/runs and bedrock. Unlike other habitats, debris dams retained moisture for relatively long periods following cessation of surface flow. 3. Reach‐specific macroinvertebrate production ranged from approximately 1.7 to 2.9 g AFDM m⁻² year⁻¹ which are among the lowest values ever reported. Habitat‐specific production ranged from approximately 0.5 to 5.0 g AFDM m⁻² year⁻¹ (bedrock and debris dams, respectively). 4. At the reach scale, quantities of stored benthic organic matter (range approximately 200–600 g AFDM⁻²) decreased in a downstream direction. 5. A combination of differences in the timing and duration of channel drying, habitat structure and detritus standing stocks appeared to influence levels of invertebrate production among the study reaches. 6. Our interpretation of a canonical correspondence analysis indicates that drying is more important than habitat in affecting macroinvertebrate production in this intermittent stream system.