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

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

Swimming performance of various freshwater Newfoundland salmonids relative to habitat selection and fishway design

Swimming ability of wild brook trout Salvelinus fontinalis , brown trout Salmo trutta , anadromous Atlantic salmon Salmo salar , and landlocked Atlantic salmon was examined using fixed and increasing velocity tests. Although brook trout and salmon parr were collected from the same site, brook trout were found generally in slow-moving pools whereas salmon were more common in faster riffle areas. Salmon parr could hold station indefinitely in currents in which brook trout could only maintain themselves briefly. Therefore, selection of fast-water areas by salmon parr may impose a velocity barrier to sympatric juvenile brook trout, reducing competition between the species. Performance comparisons also indicate that anadromous Atlantic salmon possess slightly greater sustained ability than landlocked salmon, possibly due to altered selective pressure associated with their different life histories. Finally, fishways and culverts in Newfoundland can now be designed using models generated from performance data collected from native salmonid species.     

Habitat-specific estimates of competition in stream salmonids: A field test of the isodar model of habitat selection

Summary The population densities of sympatric Atlantic salmon,Salmo salar and brook charr,Salvelinus fontinalis, were measured in riffle and pool stream habitats to test whether non-linear isodars, a multispecific model of habitat selection based on ideal distribution assumptions, could (1) predict the distribution of densities between habitats and (2) reproduce the processes postulated to underlie spatial segregation and species interactions in previous laboratory and field studies. The model provided a good fit to observed density patterns and indicated that habitat suitability declined non-linearly with increased heterospecific competitor densities. Competitive effects in riffles appeared to be due to exploitative resource use, with salmon always emerging as the superior competitor. No evidence was found for interference competition in riffles. In contrast, interspecific competition in pools seemed to occur through exploitation and interference. The specific identity of the superior competitor in pools depended on the density of both species; pools provided the charr with refuge from competition with the salmon, presumably through the adoption by the charr of density-dependent behaviours, such as schooling and group foraging, that mitigated the negative impact of the salmon. Charr were displaced from the riffles toward the pools as the total salmon density increased. The isodar analysis, based on limited density data, successfully reproduced the processes suggested to underlie spatial segregation in previous field and laboratory studies and provided new insights into how changes in competitor densities modify habitat suitability in this system.     

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.     

Does life-history variability in salmonids affect habitat use by juveniles? A comparison among streams open and closed to anadromy

1. Migratory and resident forms of salmonids coexist in many river systems. Although such coexistence is widespread, little is known about its ecological basis and no studies have compared the habitat use of premigratory juveniles and residents. 2. We employed a comparative approach to explore the differential habitat use of juvenile anadromous and resident brook trout. This required the investigation of habitat use in streams closed to anadromy, containing only resident brook trout Salvelinus fontinalis ('resident-only' streams) and streams open to anadromy, containing coexisting Atlantic salmon Salmo salar and anadromous and resident brook trout ('migrant-resident' streams). 3. We demonstrate that fast habitats (riffles) are occupied more frequently in streams with migratory brook trout relative to riffle habitats of streams with only resident brook trout. In contrast, occupation of slow current velocities (pools) was observed in both migrant-resident and resident-only streams as both stream types contain resident brook trout. The net effect is a wider distribution of occupied habitats (pool and riffles) in migrant-resident streams relative to resident-only streams, resulting in few, if any, unused habitats. 4. These results are consistent with previously reported bioenergetic, morphological and stable isotope differences observed between anadromous and resident brook trout. 5. Our findings suggest that a link exists between juvenile habitat use, metabolic costs and life-history strategies.     

Comparison of allopatric cutthroat trout stocks with those sympatric with coho salmon and sculpins in small streams

Synopsis Juvenile stocks of allopatric (upstream of barrier falls) cutthroat troutSalmo clarki and those sympatric (downstream of barrier falls) with coho salmonOncorhynchus kisutch and sculpinsCottus spp., were sampled during the late summer period of low flows in six small coastal streams in British Columbia. The objective was to obtain and compare information on pattern of habitat use and fish size distribution of these two trout types. In most instances, density (n m^–2; g m^–2) of cutthroat trout was considerably greater in pools and glides in the allopatric than in the sympatric stocks. The sympatric salmonids were dominated by juvenile coho salmon in pools and cutthroat trout in riffles. Sympatric cutthroat trout constituted from 7 to 45 % of the salmonids present in pools and from 50 to 90% in riffles. Glides were areas of intermediate densities for both salmonids, although coho salmon was the more abundant species in most instances. The density of sculpins was high in all three habitat types, and frequently it exceeded that of coho salmon and cutthroat trout combined. Sympatric cutthroat trout consisted primarily of underyearling fish, whereas allopatric cutthroat trout consisted primarily of two or more age classes with a large proportion of them living in pools. When tested in a laboratory stream both types of cutthroat trout had similar habitat preferences and agonistic behaviours, with the exception that allopatric trout made greater use of cover and defended pools more intensely than sympatric trout when the flow was increased. The results of this study provide insight of potential impact of coho salmon juvenile transplants into stream segments supporting allopatric cutthroat trout.     

Summer stream habitat partitioning by sympatric Arctic charr, Atlantic salmon and brown trout in two sub-arctic rivers

Summer habitat use by sympatric Arctic charr Salvelinus alpinus, young Atlantic salmon Salmo salar and brown trout Salmo trutta was studied by two methods, direct underwater observation and electrofishing, across a range of habitats in two sub-arctic rivers. More Arctic charr and fewer Atlantic salmon parr were observed by electrofishing in comparison to direct underwater observation, perhaps suggesting a more cryptic behaviour by Arctic charr. The three species segregated in habitat use. Arctic charr, as found by direct underwater observation, most frequently used slow (mean ±s .d . water velocity 7·2 ± 16·6 cm s⁻¹) or often stillwater and deep habitats (mean ±s .d . depth 170·1 ± 72·1 cm). The most frequently used mesohabitat type was a pool. Young Atlantic salmon favoured the faster flowing areas (mean ±s .d . water velocity 44·0 ± 16·8 cm s⁻¹ and depth 57·1 ± 19·0 cm), while brown trout occupied intermediate habitats (mean ±s .d . water velocity 33·1 ± 18·6 cm s⁻¹ and depth 50·2 ± 18·0 cm). Niche overlap was considerable. The Arctic charr observed were on average larger (total length) than Atlantic salmon and brown trout (mean ±s .d . 21·9 ± 8·0, 10·2 ± 3·1 and 13·4 ± 4·5 cm). Similar habitat segregation between Atlantic salmon and brown trout was found by electrofishing, but more fishes were observed in shallower habitats. Electrofishing suggested that Arctic charr occupied habitats similar to brown trout. These results, however, are biased because electrofishing was inefficient in the slow-deep habitat favoured by Arctic charr. Habitat use changed between day and night in a similar way for all three species. At night, fishes held positions closer to the bottom than in the day and were more often observed in shallower stream areas mostly with lower water velocities and finer substrata. The observed habitat segregation is probably the result of interference competition, but the influence of innate selective differences needs more study.     

Growth differences between the Atlantic salmon parr,Salmo salar,of nursery brooks and natal rivers in the River Teno watercourse in northern Finland

Synopsis A comparison was done between growth in Atlantic salmon,Salmo salar, parr from natal rivers and small tributaries in the subarctic River Teno watercourse in northern Finland (70° N, 28° E). Salmon do not spawn in these tributaries but juveniles enter the brooks from their spawning rivers. Parr from these brooks (age groups 1 + and 2 +) were larger, so were their recorded annual growth increments, than those caught simultaneously in the River Teno. First year growth was also better in brook parr, although both groups had spent their first year in the main river. The brook water temperature was lower than that in the main rivers through-out the growing season. On the other hand, drifting food resources were more abundant in brooks. It has been suggested that better growth and a longer residence in fresh water result in the larger smolts being produced in the brooks relative to the main river. Differences in habitat selection and subsequent differences in the smolt age and size of fish from the same place of origin reflect the distinct life histories of the salmon in the River Teno.     

Habitat use and fish activity of landlocked Atlantic salmon and brook charr in a newly developed habitat compensation facility

Degradation and destruction of valuable spawning and rearing habitat due to anthropogenic changes (e.g., flow modification and channelisation) is known to have dramatic impacts on fish populations. To compensate for habitat losses due to hydropower development, an artificial fluvial habitat channel (‘Compensation Creek’) was constructed in south-central Newfoundland, Canada. The creek was designed to include appropriate habitat features for the two dominant salmonid fish species, landlocked Atlantic salmon (Salmo salar L.) and brook charr (Salvenius fontinalis Mitchell). The study examines the habitat use of landlocked Atlantic salmon and brook charr in the Compensation Creek using electromyogram (EMG) radio telemetry. Ten landlocked Atlantic salmon and eight brook charr were captured and tagged with EMG transmitters. In laboratory swimming experiments, the EMG values were calibrated against swimming speed. Fish were then released in the Compensation Creek and tracked on a daily basis. The results show that (1) during residence in the creek, both species used preferentially the habitat features designed to match their rearing habitat preferences, and (2) swimming speed did not vary among habitat types for either species.     

Comparison of brook trout reproductive success and recruitment in an acidic adirondack lake following whole lake liming and watershed liming

Limestone applications to the catchment of one tributary to Woods Lake were highly effective in reducing stream acidity and stabilizing seasonal fluctuations in pH. The resulting improvement in stream water quality also led to a dramatic shift in reproductive strategy of the Woods Lake brook trout population. Prior to catchment liming, brook trout in Woods Lake were restricted to spawning on poor quality near shore substrate with limited ground water seepage. Reproductive success was limited by high mortality of eggs and larvae and recruitment from in lake spawning was not successful. Spawning brook trout did not utilize the tributary for spawning prior to watershed liming. Mitigation of acidity in the tributary, by catchment liming, effectively extended the spawning habitat available to the Woods Lake brook trout population and one year following treatment brook trout spawned successfully in the tributary for the first time in 6 years of observation. Significant recruitment of young trout into the lake population occurred from 1991 through 1993, although the absolute number of fish captured was relatively small. In the fall of 1993, four year classes of naturally spawned brook trout were present in the lake. Although reproductive success was enhanced by improving tributary spawning habitat in the Woods Lake basin, self maintenance of the population may be limited by low recruitment rates of young trout, due to high levels of summer mortality resulting from predation. Mitigation of this constraint would require substantially higher levels of fry production than were observed in Woods Lake and/or enhanced refugia for young trout. The results of this experiment suggest that re-establishment of tributary spawning populations of brook trout may be possible, with future reductions in acidic deposition, in acidic Adirondack lakes with limited in-lake spawning habitat.     

The evolution of cooperation and advanced social behaviour

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.     

Is there a threshold size regulating seaward migration of brown trout and Atlantic salmon?

We investigated the influence of variation in body size and growth rate on age of smolting in Atlantic salmon and brown trout in four different Norwegian rivers. In Atlantic salmon smolt ages varied between 2 and 6 years, and in brown trout between 2 and 7 years. Smolt age was negatively correlated with parr growth, and positively correlated with smolt size. Age at smolting was more variable in the two northern than the two southern rivers. Smolt sizes and ages were also more variable in brown trout than in Atlantic salmon. Based on the observed variation in smolt size and age, we reject the hypothesis that a threshold size alone regulates age at smolting. Within populations smolt age depends on growth rate so that fast-growing parr smolted younger and smaller than slow-growing parr. We hypothesize that smolt size and age is a trade-off between expected benefits and costs imposed by differences in individual growth rate.     

Local Movement as a Measure of Habitat Quality in Stream Salmonids

Habitat assessments are often based on the premise that spatial variation in population density arises from, and accurately reflects, underlying differences in quality among habitats. Nonetheless, this premise has been criticized on both theoretical and empirical grounds. Habitat quality perhaps is best evaluated by examining behavioural processes which directly influence habitat use. We present an approach based on the assumption that measures of local movement, such as habitat-specific immigration and loss rates, provide useful indicators of habitat quality. A dynamic turnover model was used in conjunction with mark-recapture techniques to estimate movement parameters for brook charr, Salvelinus fontinalis, and Atlantic salmon, Salmo salar, in different stream habitats during the summer. Immigration and loss rates were derived from mark-recapture experiments covering short periods of time (6 days). Movement-based rankings of habitat quality for both charr (pools glides > riffles) and salmon (riffles > glides > pools) were in agreement with results from earlier studies. Over evaluation periods of up to 65 days, observed abundances were highly variable in time and fluctuated about the equilibrium abundances calculated from movement parameters in the short-term experiments, suggesting that movement-based parameters may be more stable than measures of abundance for evaluating salmonid habitat. Because the approach based on movement behaviour focuses on immigration and loss, two processes that directly generate density differentials between habitats, it provides a more reliable mechanistic basis for understanding habitat selection than do traditional approaches based on density–quality relationships.     

Spawning ecology of the American brook lamprey, <Emphasis Type="Italic">Lampetra appendix</Emphasis>

Synopsis We observed spawning American brook lamprey, Lampetra appendix, in coldwater streams in Minnesota to assess various aspects of their spawning behavior and spawning habitat requirements. Spawning occurred during April and May, at water temperatures ranging from 8.7 to 15.5°C. Average adult lamprey length and mass differed significantly among streams, but there were no significant differences in length or mass between males and females. Overall sex ratio was 1:1, although one stream had significantly more males than females and one stream held significantly more females. Lampreys spawned in groups of 2–14 individuals, averaging 4.2 adults per nest across all streams. Nests were constructed in gravel and cobble substrate just upstream of riffles, spaced at an average density of three nests m⁻². The typical nest was 16 cm in diameter in water 31 cm deep with a bottom current velocity of 14 cm s⁻¹, and was excavated to a depth of 4 cm below the stream bottom; however, some nest characteristics varied significantly in a few streams. Nests were larger in streams with larger spawning groups, deeper water, and slower current velocities. American brook lamprey exhibited spawning behaviors and spawning habitat requirements similar to those of other species of lamprey in North America.     

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.     

Relative production of Atlantic salmon from fluvial and lacustrine habitats estimated from analyses of scale characteristics

Empirical and back-calculated growth of Atlantic salmon parr were compared between fish reared in fluvial and lacustrine habitats of Conne River, Newfoundland. Length at age was significantly higher for lacustrine parr. Various classification and maximum likelihood estimators indicated that 75% or more of the fish used lakes for rearing. Lacustrine use is another aspect of the inherent variability and plasticity of Atlantic salmon life-history traits. As most Newfoundland river systems include lakes, estimates of regional spawning targets and potential smolt production will need to take lake habitat into account.     

Effect of mink, Mustela vison Schreber, predation on cohorts of juvenile Atlantic salmon, Salmo salar L., and brown trout, S. trutta L., in three small streams

Two cohorts of Atlantic salmon parr and one of brown trout were studied in periods with and without the presence of mink, Mustela vison . In all localities a marked increase in mortality rate was observed during periods when mink were present. Mink were observed catching salmon parr, and approximately 10% of the parr had bite marks, especially on the tail fins. In the smallest stream with brown trout, the mortality rate was 0.80 during a few days with mink present; remnants of trout were found along the stream. The present study suggests that mink predation may be a major cause of mortality of salmonids in small streams. The results indicate that predation efficiency may vary with characteristics of the habitat, especially stream width and discharge, and fish density.     

Spatial and temporal variability of macroinvertebrates in spawning and non-spawning habitats during a salmon run in Southeast Alaska

Spawning salmon create patches of disturbance through redd digging which can reduce macroinvertebrate abundance and biomass in spawning habitat. We asked whether displaced invertebrates use non-spawning habitats as refugia in streams. Our study explored how the spatial and temporal distribution of macroinvertebrates changed during a pink salmon (Oncorhynchus gorbuscha) spawning run and compared macroinvertebrates in spawning (riffle) and non-spawning (refugia) habitats in an Alaskan stream. Potential refugia included: pools, stream margins and the hyporheic zone, and we also sampled invertebrate drift. We predicted that macroinvertebrates would decline in riffles and increase in drift and refugia habitats during salmon spawning. We observed a reduction in the density, biomass and taxonomic richness of macroinvertebrates in riffles during spawning. There was no change in pool and margin invertebrate communities, except insect biomass declined in pools during the spawning period. Macroinvertebrate density was greater in the hyporheic zone and macroinvertebrate density and richness increased in the drift during spawning. We observed significant invertebrate declines within spawning habitat; however in non-spawning habitat, there were less pronounced changes in invertebrate density and richness. The results observed may be due to spawning-related disturbances, insect phenology, or other variables. We propose that certain in-stream habitats could be important for the persistence of macroinvertebrates during salmon spawning in a Southeast Alaskan stream.     

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

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

Is Atlantic salmon production limited by number of territories?

Thirty Atlantic salmon Salmo salar parr (mean 135 mm total length, L _T) in the River Alta, Norway, were radio‐tagged and tracked during a 11–13 day period. The parr stayed within defined home ranges with a 95% probability of localization within an average area of 1286 m²(241–3484 m²). On average, each parr had overlapping home ranges with 7·7 other parr, and the overlap between pairs of fish covered on average 24% of their home range areas. Mean length of the river stretch used was 90 m (22–383 m). On average, during 38%(16–77%) of the tracking surveys, the fish had moved >10 m since the previous survey. Mean total distance moved during the whole study was 402 m (208–862 m). The Atlantic salmon were most often recorded in riffles, but 27% of the parr alternated between riffles and pools. The extensive movements, flexible habitat utilization and relatively large home ranges, together with the fact that all the parr had home ranges partly overlapping with other radio‐tagged parr, indicate a flexible and variable behaviour and, thereby, a more complex social structure than a rigid defence of a fixed territory. Thus, Atlantic salmon production seems not to be limited by the highest potential number of territories for large parr in a given area.