Patterns of density‐dependent growth in juvenile stream‐dwelling salmonids

Literature data for 19 populations from 16 different studies of six species of stream‐dwelling salmonids were analysed to test the hypothesis that density‐dependent growth is stronger at low rather than at high population density. Fifteen of 19 populations showed evidence of a significant decrease in growth rate with increasing density. In 11 of these 15 populations, the pattern of density‐dependent growth was better described by a negative power curve than by a linear regression ( i.e . Akaike Information Criterion, AIC _linear −  AIC _power > 2), whereas only one population was better described by a linear regression than by a negative power curve; three populations were adequately described by both models ( AIC  < 2). In 10 of the 11 populations that were best described by a negative power curve, most of the decrease in growth rate occurred at population densities <1 fish m⁻², when space limitation is unlikely. This analysis provides broad support for the hypothesis that density‐dependent growth in stream salmonids occurs primarily at low population densities, probably due to exploitative competition.     

Elevated summer temperatures delay spawning and reduce redd construction for resident brook trout (Salvelinus fontinalis)

Redd (nest) surveys for resident brook trout (Salvelinus fontinalis) were conducted annually in a mountain lake in northern New York for 11 years with multiple surveys conducted during the spawning season in eight of those years. Repeated surveys throughout the spawning season allowed us to fit an individually based parametric model and estimate the day of year on which spawning was initiated, reached its midpoint, and ended during each year. Spawning phenology was then assessed relative to (1) mean of maximum daily air temperature and (2) mean of maximum daily water temperature at the lake bottom during summer in each year using a linear model. Elevated temperatures in summer were correlated with a delay in spawning and a reduction in the total number of redds constructed. Increasing the summer mean of maximum daily air temperatures by 1 °C delayed spawning by approximately 1 week and decreased the total number of redds constructed by nearly 65. Lake spawning brook trout select redd sites based on the presence of discharging groundwater that is relatively constant in temperature within and across years, leading to relatively consistent egg incubation times. Therefore, delayed spawning is likely to delay fry emergence, which could influence emergence synchrony with prey items. This work highlights non‐lethal and sub‐lethal effects of elevated summer temperatures on native resident salmonids in aquatic environments with limited thermal refugia.     

Density‐dependent growth in hatchery‐reared brown trout released into a natural stream

Hatchery‐reared brown trout Salmo trutta stocked in a natural stream in addition to resident wild brown trout grew more slowly than those stocked with an experimentally reduced density of brown wild trout. In both cases, hatchery‐reared brown trout grew more slowly than resident wild fish in control sections. Mortality and movements did not differ among the three categories of fish. The results showed that growth of stocked hatchery‐reared brown trout parr was density‐dependent, most likely as a consequence of increased competition. Thus, supplementary release of hatchery‐reared fish did not necessarily increase biomass.     

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 effects of introduced salmonids on two native stream‐dwelling salmonids through interspecific competition

Using an artificial stream, habitat use by two sympatric native salmonids in the presence and absence of introduced salmonid species was investigated experimentally. When only native white‐spotted charr Salvelinus leucomaenis and masu salmon Oncorhynchus masou were sympatric, they occupied different microhabitats. In the presence of introduced brown trout Salmo trutta or rainbow trout Oncorhynchus mykiss , however, white‐spotted charr and masu salmon were observed to use a similar habitat and interspecific competition between white‐spotted charr and masu salmon was initiated. The study suggested that the coexistence of native salmonids was negatively affected through interspecific competition between native and introduced salmonids.     

Seasonal temperature and precipitation regulate brook trout young‐of‐the‐year abundance and population dynamics

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A new method to evaluate egg‐to‐fry survival in salmonids, trials with Atlantic salmon

A new method to estimate the survival of salmonids from egg fertilization to fry emergence is described. Fine mesh screen cylindrical capsules, 12 cm³ in volume, filled with batches of 10 eggs of Atlantic salmon Salmo salar were implanted in the substratum using small removable guiding tubes inserted with a metal spike. The method was compared with two other commonly used techniques, capping redds with fry‐traps and fine mesh screen incubation‐emergence boxes buried into the gravel. Egg‐to‐fry survival was recorded for the three methods run in parallel in artificial redds created on three sites of the Nivelle River watershed (south‐west of France), two in the stream and one in an experimental spawning channel. In the channel, survival to the eyed stage and to hatching in capsules and incubation‐emergence boxes was also compared. The implant of capsules proved easier and faster to use than other methods, the structure of the surrounding substratum was less disturbed and the capsules were less vulnerable to spates. This technique provided survival values largely free of the bias induced by other methods and intermediate between that of incubators and of traps.     

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

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

The evolution of redd site selection in brook charr in different environments: same cue,same benefit for fitness*

1. Despite the great interest in characterising fish reproductive habitat, the relationship between the selection of a given spawning site and individual fitness has not been experimentally tested. 2. In this study, we used an in situ experimental approach to determine (i) the relative contribution of substrate characteristics as well as hydrological and physicochemical variables to small‐scale redd site selection by brook charr, Salvelinus fontinalis (Mitchill) and (ii) if hatching and emergence success, used as a surrogate of fitness, are improved in selected compared to non‐selected sites in both lake and stream habitats. 3. Our results show that upwelling groundwater flow was always significantly higher in selected than in non‐selected sites in both lake and stream habitats. We found no significant difference in the mean geometric substrate diameter and no consistent differences in substrate composition between selected and non‐selected sites. Oxygen concentration was higher (significantly so in three of four comparisons) and conductivity tended to be lower in selected than in non‐selected sites, while temperature showed no significant or consistent variations. We found a significant positive relationship between the selection of a given spawning habitat and hatching and emergence success in these systems. 4. These results show that the main cue that brook charr use to select their spawning sites is upwelling groundwater in lake and stream habitats, and that active selection of these sites increases individual fitness. This suggests that natural selection acted on the same cues in lentic and lotic environments; this could have been highly adaptive in a species that used both habitats as colonisation routes after the last glaciation.     

Spatial patterns of hybridization between bull trout, <Emphasis Type="Italic">Salvelinus confluentus</Emphasis>, and brook trout, <Emphasis Type="Italic">Salvelinus fontinalis</Emphasis> in an Oregon stream network

Hybridization with introduced species represents a serious threat to the persistence of many native fish populations. Brook trout (Salvelinus fontinalis) have been introduced extensively throughout the native range of bull trout (S. confluentus) and hybridization has been documented in several systems where they co-exist and is seen as a significant threat to the persistence of bull trout populations. We identified a group of diagnostic microsatellite loci to differentiate bull trout and brook trout and then used these loci to examine the spatial distribution of hybrids in the Malheur River basin, Oregon USA. In random samples of approximately 100 fish from each of three creeks we identified 181 brook trout, 112 bull trout and 14 hybrids. Although bull trout, brook trout and hybrids were found in all three creeks, they were not evenly distributed; brook trout were primarily found in the lower sections of the creeks, bull trout further upstream, and hybrids in the areas of the greatest overlap. One creek with a population of brook trout in a headwater lake provided an exception to this pattern; brook trout were found distributed throughout the creek downstream of the lake. Several post-F1 hybrids were identified suggesting that hybrids are reproducing in the Malher River Basin. Mitochondrial DNA analysis indicated that both female bull trout and brook trout are involved in hybridization events. Analysis of population structure suggested that brook trout have established multiple spawning populations within the Malheur system. Data presented in this study suggest that relative abundance of brook trout and habitat quality are important factors to consider when evaluating the threat of hybridization to bull trout populations.     

Growth in length and in body depth in young‐of‐the‐year perch with different predation risk

Body shape of young‐of‐the‐year (YOY) perch Perca fluviatilis , and number and size of potential predators (perch and pike Esox lucius ) were compared across five lakes in northern Sweden. Body depth and dorsal fin ray length of YOY perch differed between lakes, with high relative body depth and long dorsal fin rays found in the lakes where the number of large piscivores was the highest. The most slender fish were found in the lake where the perch population had the smallest individuals and pike occurred in very low numbers. The average body mass in YOY perch from the two lakes with highest and lowest body depth, respectively, were the same, which indicates a difference between lakes in the relation between growth in length and in depth. Both body depth and fin ray length were correlated with predation risk by pike. Fin ray length was also correlated with number of piscivorous perch. Selection for different body shapes can be caused by different biotic and abiotic factors, singly or in combination, and the results from this study indicates that predation risk is one of these factors affecting body depth and fin ray length in perch.     

Food partitioning between coexisting Atlantic salmon and brook trout in the Sainte‐Marguerite River ecosystem, Quebec

Food resource partitioning between similar‐sized, sympatric Atlantic salmon Salmo salar and brook trout Salvelinus fontinalis was examined as a possible mechanism enabling their coexistence in a stream (Allaire) of the Sainte‐Marguerite River ecosystem, Quebec, Canada. Fish stomach contents and invertebrate drift were collected concurrently during three diel cycles in August to September 1996. The food and feeding habits of an allopatric brook trout population in a nearby stream (Epinette) were studied for comparison. The diel feeding rhythms of the two coexisting fish species were similar. The composition of their diet, however, showed significant differences. Atlantic salmon predominantly (60–90%) fed on aquatic insects, mainly Ephemeroptera (35–60% of the diet). The brook trout mostly (50–80%) fed upon the allochthonous terrestrial insects (mainly adults of Coleoptera, Hymenoptera and Diptera) which comprised 5–40% of the stream drift. The allopatric brook trout fed opportunistically on the more abundant aquatic insects and terrestrial insects rarely formed 25% of its diet. The allopatric trout fed nearly twice as much as the sympatric brook trout during a day. The results suggest that the differences in feeding by brook trout in the two streams (with and without Atlantic salmon) are the result of inter‐specific interaction with Atlantic salmon and are not related to the differences in food availability between the two streams. Food resource partitioning between Atlantic salmon and brook trout may be viewed as an adaptive response resulting in a greater exploitation of available resources and coexistence.     

Spatial variation in population dynamics of juvenile Atlantic salmon: implications for conservation and management

Application of habitat models for predicting expected local densities of Atlantic Salmon Salmo salar in healthy populations has been hampered by a lack of generality in their fit to data from different systems. It is believed that this problem results at least in part from difficulties of effectively integrating factors that act across a range of spatial and temporal scales. Here, as an aid to developing more robust modelling and sampling methodologies, a simple process‐based model for local‐scale dynamics of Atlantic salmon juveniles is developed from first principles by integrating contemporary understanding of self‐thinning, density‐dependent growth and dispersal. The aim is to present a readily understood structure to illustrate the links between spawning and stocking strategies, habitat, migration and fish production. Based on this structure, contemporary understanding of the more complex biological processes that affect density, growth and habitat are discussed in relation to some of the key requirements of managers, including stocking for rehabilitation, assessment of predation impact and development of strategies for sampling populations effectively when deriving habitat‐production models. A major conclusion is that more structured, integrated research is required to provide the basic variables needed to model links between local and global scale habitat and fish production effectively. Nevertheless, application of the current understanding of the biology of Atlantic salmon should be of great benefit to managers in extracting key information from field surveys.     

The effects of population density and lake characteristics on growth and size structure of brook trout Salvelinus fontinalis (Mitchill, 1815) in boreal forest lakes in Canada

This paper examines the effect of lake characteristics on population density and how this variation affects growth, mortality and population size structure of brook trout, Salvelinus fontinalis. The study was conducted on 17 recreationally fished, reproductively isolated boreal forest lakes in Newfoundland, Canada from 1993 to 2000. A standardized sampling program, the Fyke Littoral Index Netting program (FLIN) was used to collected data that describes brook trout population parameters and life history attributes. Regression analyses showed significant relationships between fish density and biomass and characteristics of the lakes. Variation in fish density and biomass was explained by lake surface area and littoral habitat area. Significant relationships were found when growth, mortality and size structure were regressed against density. The proportional stock distribution and theoretical maximum size of brook trout were negatively related to density, and natural mortality was positively related to density. The largest maximum length and highest proportional stock densities occurred at brook trout densities of less than 30 fish/ha. In general, the higher the proportion of littoral habitat area the higher the densities of brook trout, which correspondingly had important effects on growth, natural mortality and size structure of the brook trout populations. This information is critical to the development of management strategies aimed at altering size distribution to produce specific fisheries management outcomes.     

Integration of acoustic telemetry and GIS to identify potential spawning areas for lake trout (Salvelinus namaycush)

Locations of potential spawning areas for lake trout (Salvelinus namaycush) were predicted in Lake Opeongo, Ontario, Canada using information gained via acoustic telemetry and geographic information system (GIS) technologies. From 1998 to 2000, 18 adult lake trout (mean fork length 553 mm) implanted with acoustic transmitters (battery life 2 years) were manually tracked. For evening fall locations within the erosive zone of the lake (determined using an existing sedimentation model), habitat variables (slope, depth, and effective fetch) were summarised using GIS. Sites selected by lake trout during the spawning window were in areas of mean fetch equal to 1.5 km and mean slope of 10.6% (n = 50 fixes). We used GIS to identify areas that matched the mean habitat criteria and thus locate potential spawning areas. This model correctly identified 19 of 21 known spawning sites, as well as additional sites used by spawning females in an earlier telemetry study. Depths of traditional fall netting sites are shallow compared to areas in which telemetered lake trout were found during evenings of the spawning period (means 3.1 vs. 5.1 m, respectively). Through the use of information on spawning habitat selection gained through telemetry and knowledge of the physical characteristics of the lake, we provide an alternative means of identifying potential spawning habitat for lake trout.     

Sympatric relationship between redband trout and non-native brook trout in the Southeastern Oregon Great Basin

Brook trout (Salvelinus fontinalis) and rainbow trout (Oncorhynchus mykiss) have been widely introduced outside their respective ranges within North America causing declines and displacement of native trout. Yet, successful coexistence of native and non-native trout has received little attention. Here we evaluated the effect of introduced brook trout on the size and density of native redband trout in two invaded sub-basins in southeastern Oregon. In a multi-year study, we investigated whether habitat and fish communities differed between streams and stream reaches where redband trout were allopatric versus where redband trout were sympatric with brook trout. We hypothesized that redband trout would be less dense and have smaller total length in sympatry with brook trout than in allopatry, but that total trout density would not differ. We investigated whether differences in habitat existed between sympatric and allopatric locations that would indicate differentiation in site level habitat preferences for each trout species. We found that sympatric locations had more wood but similar fish community structure. Mean length and densities of redband trout were higher at allopatric locations. However, in most years at sympatric locations total trout density was twice that of allopatric redband trout sites. Using comparable data from an eastern United States system where brook trout are native, sympatric sites had lower densities of brook trout; however, total trout density did not differ. We conclude that invading trout negatively impact native trout densities; but in southeastern Oregon system the negative impact is minimized.     

Telemetry and genetics reveal asymmetric dispersal of a lake‐feeding salmonid between inflow and outflow spawning streams at a microgeographic scale

The degree of natal philopatry relative to natal dispersal in animal populations has important demographic and genetic consequences and often varies substantially within species. In salmonid fishes, lakes have been shown to have a strong influence on dispersal and gene flow within catchments; for example, populations spawning in inflow streams are often reproductively isolated and genetically distinct from those spawning in relatively distant outflow streams. Less is known, however, regarding the level of philopatry and genetic differentiation occurring at microgeographic scales, for example, where inflow and outflow streams are separated by very small expanses of lake habitat. Here, we investigated the interplay between genetic differentiation and fine‐scale spawning movements of brown trout between their lake‐feeding habitat and two spawning streams (one inflow, one outflow, separated by <100 m of lake habitat). Most (69.2%) of the lake‐tagged trout subsequently detected during the spawning period were recorded in just one of the two streams, consistent with natal fidelity, while the remainder were detected in both streams, creating an opportunity for these individuals to spawn in both natal and non‐natal streams. The latter behavior was supported by genetic sibship analysis, which revealed several half‐sibling dyads containing one individual that was sampled as a fry in the outflow and another that was sampled as fry in the inflow. Genetic clustering analyses in conjunction with telemetry data suggested that asymmetrical dispersal patterns were occurring, with natal fidelity being more common among individuals originating from the outflow than the inflow stream. This was corroborated by Bayesian analysis of gene flow, which indicated significantly higher rates of gene flow from the inflow into the outflow than vice versa. Collectively, these results reveal how a combination of telemetry and genetics can identify distinct reproductive behaviors and associated asymmetries in natal dispersal that produce subtle, but nonetheless biologically relevant, population structuring at microgeographic scales.     

Native brook trout and invasive rainbow trout respond differently to seasonal weather variation: Spawning timing matters

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Inter‐individual differences in rates of routine energy loss and growth in young‐of‐the‐year juvenile Atlantic cod

Inter‐individual differences in rates of routine (non‐feeding) metabolism and growth were evaluated in young‐of‐the‐year (YOY) juvenile Atlantic cod Gadus morhua . Rates of O₂ consumption, CO₂ production and ammonia (TAN) excretion were measured in 64, 25–43 mm standard length ( L _S) YOY growing at different rates (0·27–0·47 mm day⁻¹) in a common rearing tank. Parameter rates ( y ) increased allometrically ( y = a·M^b ) with increasing body mass ( M ) with b ‐values for O₂ production, CO₂ consumption and TAN excretion equal to 0·81, 0·89 and 0·56, respectively. In some cases, residuals from these regressions were significantly negatively correlated to fish growth rate. In no cases did residuals of parameter rates increase with increasing growth rate. These data suggest that, during unfed periods, relatively fast‐growing fish were more metabolically efficient than slower‐growing fish from the same cohort. The fish condition factor, derived from     , also significantly decreased with increasing growth rate. Results indicated differences in both the rates of routine energy loss and the patterns of growth allocation among YOY Atlantic cod. Since these physiological attributes were positively correlated with growth rate, they may be indicative of 'survivors' in field populations.     

Assessing the reproductive contributions of sympatric anadromous and freshwater‐resident brook trout

The δ¹³C, δ¹⁵N and δ³⁴S isotope values of newly emerged, brook trout Salvelinus fontinalis alevins (free‐swimming individuals beginning exogenous feeding) were examined to determine if progeny of anadromous female spawners could be detected and their contributions to reproduction assessed in river systems with mixed migration strategies. Separation of anadromous and freshwater resident and immigrant sources of progeny could be detected primarily using δ¹³C values before alevins reached a size of 28 mm fork length and c . 20 days after exogenous feeding began. Among 10 populations, the contributions of anadromous females ranged from 0 to 42% of newly emerged alevins. Correlations between anadromous contributions and estimates of total brook trout density and alevin production were unexpectedly negative.