Sexual maturity increases mobility and heterogeneity in individual space use in Atlantic salmon (Salmo salar) parr

We assessed the effects of sexual maturity on space use in Atlantic salmon (Salmo salar) parr as facultative early maturation enables us to work on individuals belonging to the same cohort. We monitored the space use of 40 1-year-old males in natura throughout a breeding season. First, mature individuals covered longer distances (absolute and upstream) and located within broader home ranges than immature parr. Second, sexual maturity also generated a higher interindividual variability in space use. Finally, mature individuals exhibited a higher probability of association with likely breeding sites on average. However, some mature individuals experienced a lower probability than immature individuals, suggesting that the space use of some mature individuals may not be optimal. Moreover, mature parr exploiting a broader home range or covering longer upstream distances had a higher probability of association with likely breeding sites. Covering longer upstream distances may therefore increase the reproductive success of mature parr, while involving higher energetic costs and a greater risk of predation.     

Morphological plasticity in a native freshwater fish from semiarid Australia in response to variable water flows

In fishes, alterations to the natural flow regime are associated with divergence in body shape morphology compared with individuals from unaltered habitats. However, it is unclear whether this morphological divergence is attributable to evolutionary responses to modified flows, or is a result of phenotypic plasticity. Fishes inhabiting arid regions are ideal candidates for studying morphological plasticity as they are frequently exposed to extreme natural hydrological variability. We examined the effect of early exposure to flows on the development of body shape morphology in the western rainbowfish (Melanotaenia australis), a freshwater fish that is native to semiarid northwest Australia. Wild fish were collected from a region (the Hamersley Ranges) where fish in some habitats are subject to altered water flows due to mining activity. The offspring of wild‐caught fish were reared in replicated fast‐flow or slow‐flow channels, and geometric morphometric analyses were used to evaluate variation in fish body shape following 3, 6, 9, and 12 months of exposure. Water flows influenced fish morphology after 6 and 9 months of flow exposure, with fish in fast‐flow environments displaying a more robust body shape than those in slow‐flow habitats. No effect of flow exposure was observed at 3 and 12 months. Fishes also showed significant morphological variation within flow treatments, perhaps due to subtle differences in water flow among the replicate channels. Our findings suggest that early exposure to water flows can induce shifts in body shape morphology in arid zone freshwater fishes. Morphological plasticity may act to buffer arid zone populations from the impacts of anthropogenic activities, but further studies are required to link body shape plasticity with behavioral performance in habitats with modified flows.     

Effect of stripping on the length, condition and gonadal state of mature male Atlantic salmon, Salmo salar L., parr during autumn and winter

Hatchery-reared mature male Atlantic salmon parr were shorter than immature parr of the same age through the winter period and this difference increased prior to smolting in spring. Stripping had no effect on growth. In autumn mature male parr had a higher condition factor (CF) than immature parr, but from early January no difference was observed between unstripped mature males and immature parr. Stripping reduced the CF of male parr but the difference had disappeared by the time of smolting when the CFs of both mature groups were lower than that of immature fish. The quantity of expressible milt in stripped males decreased from c . 3% body weight in November and December to <0.05% by 2 March, after which no further milt expression was recorded. Milt expression in previously unstripped males was about 1% of body weight on 2 March, similar to that of stripped males 1 month earlier.     

A method for tracking the behaviour of mature and immature salmon parr around nests during spawning

A remote monitoring system was developed to provide information on the behaviour of mature and immature Atlantic salmon Salmo salar parr at nests during the spawning season. An octagonal passive integrated transponder (PIT) detector (0·865 m maximum diameter) designed to surround nests of Atlantic salmon was used to identify individual salmon parr present at 38 spawning events in three circular spawning channels. The range of the detector for PIT tags presented in the optimum orientation was 2·4 cm (range between tags 1·7–3·0 cm). Using a sub-sample of 20 spawnings, the mean efficiency of the detector (number of fish passes registered relative to number of passes observed on video) was 70·5% (range 32-100%). There were no significant effects of time from spawning, total number of registrations, body size or maturity status (mature or immature) on efficiency. However, fish were more likely to be detected entering nests than leaving, as departures were more rapid and higher in the water column. The PIT detector did not affect the numbers of parr at spawnings or between spawning intervals of females, and allowed for the individual identification of 65 of the 72 parr observed in nests during spawning. In all cases where certain identifications were not possible and the video was of satisfactory quality, this was due to obstruction of the camera view by anadromous fish. The remote monitoring system was thus effective in identifying behavioural differences, and only one of 20 immature parr was ever detected during the period encompassing 10 min before and after spawning compared with 30 or 40 mature parr.     

Use of a flooded salt marsh habitat by an endangered cyprinodontid fish (<Emphasis Type="Italic">Aphanius iberus</Emphasis>)

We report the first data on the use of occasionally inundated habitats in a Mediterranean coastal lagoon by the Spanish toothcarp (Aphanius iberus), a cyprinodontid fish in danger of extinction. During a flooding period, we sampled the fish population and the macroinvertebrate community in three contrasting habitats: mats of green algae, open water, and a habitat dominated by glasswort (Salicornia patula) that is occasionally inundated. The three habitats displayed strong variation in density and species composition of invertebrates in the water column and the benthos. In general, algal mats had higher invertebrate biomasses, but glasswort had higher diversity of organisms, in part of terrestrial origin. The density of toothcarp was very low in the open water. The habitat that is occasionally inundated (glasswort) significantly had the highest density of mature toothcarp, while immature fish were similarly abundant in the glasswort and algal mat habitats. Condition (weight–length relationship) and total food biomass in the gut contents of immature toothcarp was significantly higher in algal mats than in glasswort, whereas there were no such differences for mature fish. Therefore, the occasionally flooded habitat (glasswort) was positively selected by large mature toothcarp but seemed a disadvantageous habitat for immature individuals. Handling editor: L. Mauricio Bini.     

Effect of habitat types on survival, spatial distribution and production of an allopatric cohort of Atlantic salmon, Salmo salar L., under conditions of low competition

An allopatric cohort of Atlantic salmon, Salmo salar L., introduced to a small previously fishless stream was studied from parr to the smolt stages. In May 3900 0+ parr (mean total length 30mm) were planted at three different densities in habitats with slow, intermediate and fast water velocities. During the first year, high mortality occurred during the first 7 weeks after planting in May, and in September–October. Survival from May 1985 to April 1986, before the smolt emigration, was24.8%. The smolt yield 1 year after planting was 15.5%. It is suggested that the high survival was caused by low competition. Most of the redistribution of the fish took place during the first months. Type of planting habitat affected the timing of redistribution. The parr left slow-flowing, deep habitats with fine substrate soon after planting, while redistribution was slowest in the fastest flowing habitats with coarse substrate. The observed avoidance of slow, deep habitat types in the absence of interspecific competition, suggests that this may be a fixed behavioural response, and not due to competition. Long movements, up to 800 m, were recorded only within the first 7 weeks after planting. The effect of planting densities on population density was most pronounced immediately after planting in the fast and also intermediate habitats. Planting density effects declined and were not detectable after 1 year. The effect of habitat type on fish numbers and biomass was pronounced irrespective of planting densities. Growth was fastest in the intermediate habitat, and at the lower planting densities. Production was 7.2 g m⁻² the first summer-autumn. Due to smolt emigration, few fish remained in the stream the second summer-autumn, and the production was 1.0 g m⁻².     

Relative ventricular weight of wild Atlantic salmon parr in relation to sex, gonad maturation and migratory activity

Relative ventricular weight (RVW, ventricular weight/body weight · 100) was measured for five groups of Atlantic salmon parr from the Girnock Burn in Scotland. The mean RVW was: mature males, 0.087; immature resident males, 0.054; immature resident females, 0.056; immature migrant males, 0.056; immature migrant females, 0.053. For a group of mature male parr from the R. Nith, Galloway, Scotland, the mean RVW was 0.108. There were no significant differences in the RVW between groups of immature fish. The RVW of each group of mature male parr was significantly greater than those of each, group of immature parr. Condition indices (weight/length³) were closely similar between all groups of parr. The functional significance of plasticity in ventricular weight is discussed in relation to life histories of Atlantic salmon.     

Comparative proximate body composition of Atlantic salmon with emphasis on parr from fluvial and lacustrine habitats

The nutritional status of Atlantic salmon Salmo salar parr was assessed by examining how proximate body composition and energy content (kJ g⁻¹) were associated with the habitat rearing origin of parr utilizing fluvial or lacustrine areas. Comparisons were made among sampling sites within the two habitats, as well as between the two major habitat types applying both fixed effect and random effect models. Mean protein concentration varied from 13·87 to 15·67% among fluvial sites, and from 14·96 to 16·38% among lacustrine sampling locations. Mean fat concentration ranged from 1·71 to 3·32% and from 2·22 to 4·29% among fluvial and lacustrine locations, respectively, with energy concentration, on average, c . 11% higher in lacustrine parr. Interpretation of results of comparisons between habitats was dependent upon the statistical model used. Benefits associated with lacustrine rearing, such as greater body size and nutritional status, have implications for overwintering survival in fresh water as well as subsequent survival during initial sea entrance as smolts.     

Morphological differences between an artificial lentic and adjacent lotic environments in a characid species

Dam constructions cause fundamental changes in the natural landscape, creating new ecological and evolutionary challenges for aquatic organisms. In some cases, such water impoundments have been related with morphological changes in organisms. Understanding how populations respond to rapid environmental changes imposed by dams is the first step to elucidate the consequences that disturbed habitats may have on species evolution. In this work, we analyzed shape and size variation in Bryconamericus iheringii Boulenger 1887 from the Chasqueiro stream basin, south of Brazil, which was recently dammed. We used linear measurements and geometric morphometrics to identify morphological differences among specimens from the reservoir (lentic habitat) compared to the habitat upstream and downstream of the dam (lotic habitats). We also tested for size- and shape-related sexual dimorphism to determine whether variations observed were the same for both sexes. We found that B. iheringii from the artificial reservoir were distinct in shape and size to those from their natural habitat in the stream. The size variation between environments was the same for both sexes, but the shape variation differed between males and females. Regarding the linear measurements, lotic populations were larger (greater body length, width, pectoral fin base length and caudal peduncle length), probably in response to increased swimming activity. Regarding body shape, we found that both sexes have a more fusiform body in lotic habitats than in the reservoir. In addition, females showed an altered mouth position that was distinct between these environments. This work indicates that the water reservoir seems to be an important factor influencing morphological variation in B. iheringii, a species with sexual shape dimorphism.

     

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.     

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.     

Linking stream ecology with morphological variability in a native freshwater fish from semi‐arid Australia

Environmental variation is a potent force affecting phenotypic expression. While freshwater fishes have provided a compelling example of the link between the environment and phenotypic diversity, few studies have been conducted with arid‐zone fishes, particularly those that occur in geographically isolated regions where species typically inhabit intermittent and ephemeral creeks. We investigated morphological variation of a freshwater fish (the western rainbowfish, Melanotaenia australis) inhabiting creeks in the Pilbara region of northwest Australia to determine whether body shape variation correlated with local environmental characteristics, including water velocity, habitat complexity, predator presence, and food availability. We expected that the geographic isolation of creeks within this arid region would result in habitat‐specific morphological specializations. We used landmark‐based geometric morphometrics to quantify the level of morphological variability in fish captured from 14 locations within three distinct subcatchments of a major river system. Western rainbowfish exhibited a range of morphologies, with variation in body depth accounting for a significant proportion (>42%) of the total variance in shape. Sexual dimorphism was also apparent, with males displaying deeper bodies than females. While the measured local habitat characteristics explained little of the observed morphological variation, fish displayed significant morphological differentiation at the level of the subcatchment. Local adaptation may partly explain the geographic patterns of body shape variation, but fine‐scale genetic studies are required to disentangle the effects of genetic differentiation from environmentally determined phenotypic plasticity in body shape. Developing a better understanding of environment–phenotype relationships in species from arid regions will provide important insights into ecological and evolutionary processes in these unique and understudied habitats.     

Habitat selection by juvenile Atlantic salmon: the interaction between physical habitat and abundance

The effect of physical river habitat variables on the distribution of juvenile Atlantic salmon Salmo salar L. in the Rivière de la Trinité, Québec, Canada, was examined using generalized additive modelling. A survey of Atlantic salmon fry and parr densities and habitat variables (flow velocity, water column depth and substratum size) was conducted in the summer months from 1984 to 1992. Clear patterns of habitat use existed: specific ranges of habitat variables were selected, with parr preferring greater velocities, depths and substratum sizes than fry. There was a large variation, however, in juvenile densities for given velocities, depths or substratum sizes, with this variation being greatest in optimal habitats. On examination of an individual year, interaction between the variables was found to explain some of the variation. On a year‐to‐year basis the juvenile Atlantic salmon population was found to exhibit an 'Ideal Free Distribution', which resulted in greatest variation in optimal habitats with year‐to‐year changes in population abundance.     

Effects of habitat on individual reproductive success of mature male parr of Atlantic salmon Salmo salar

The effect of the presence of stone blocks in the spawning habitat on the reproductive success of mature male parr of Atlantic salmon Salmo salar of various sizes and ages was tested in an artificial channel. Shelters allowed smaller individuals to contribute to egg fertilization as much as large parr, suggesting that the size-based dominance observed in a shelterless habitat was not maintained in a more complex habitat.     

Morphological divergence and flow‐induced phenotypic plasticity in a native fish from anthropogenically altered stream habitats

Understanding population‐level responses to human‐induced changes to habitats can elucidate the evolutionary consequences of rapid habitat alteration. Reservoirs constructed on streams expose stream fishes to novel selective pressures in these habitats. Assessing the drivers of trait divergence facilitated by these habitats will help identify evolutionary and ecological consequences of reservoir habitats. We tested for morphological divergence in a stream fish that occupies both stream and reservoir habitats. To assess contributions of genetic‐level differences and phenotypic plasticity induced by flow variation, we spawned and reared individuals from both habitats types in flow and no flow conditions. Body shape significantly and consistently diverged in reservoir habitats compared with streams; individuals from reservoirs were shallower bodied with smaller heads compared with individuals from streams. Significant population‐level differences in morphology persisted in offspring but morphological variation compared with field‐collected individuals was limited to the head region. Populations demonstrated dissimilar flow‐induced phenotypic plasticity when reared under flow, but phenotypic plasticity in response to flow variation was an unlikely explanation for observed phenotypic divergence in the field. Our results, together with previous investigations, suggest the environmental conditions currently thought to drive morphological change in reservoirs (i.e., predation and flow regimes) may not be the sole drivers of phenotypic change.     

The effects of age,sex, and habitat on body size and shape of the blackstripe topminnow,Fundulus notatus (Cyprinodontiformes: Fundulidae) (Rafinesque 1820)

Lake and stream habitats pose a variety of challenges to fishes due to differences in variables such as water velocity, habitat structure, prey community, and predator community. These differences can cause divergent selection on body size and/or shape. Here, we measured sex, age, length, and eight different morphological traits of the blackstripe topminnow, Fundulus notatus, from 19 lake and stream populations across four river drainages in central Illinois. Our goal was to determine whether size and shape differed consistently between lake and stream habitats across drainages. We also considered the effects of age and sex as they may affect size and morphology. We found large differences in body size of age 1 topminnows where stream fish were generally larger than lake fish. Body shape mainly varied as a function of sex. Adult male topminnows had larger morphological traits (with the exception of body width) than females, in particular longer dorsal and anal base lengths. Subtle effects of habitat were present. Stream fish had a longer dorsal fin base than lake fish. These phenotypic patterns may be the result of genetic and/or environmental variation. As these lakes are human‐made, the observed differences, if genetic, would have had to occur relatively rapidly (within about 100 years). © 2013 The Linnean Society of London     

Inclusion of water temperature in a fuzzy logic Atlantic salmon (Salmo salar) parr habitat model

As water temperature is projected to increase in the next decades and its rise is clearly identified as a threat for cold water fish species, it is necessary to adapt and optimize the tools allowing to assess the quantity and quality of habitats with the inclusion of temperature. In this paper, a fuzzy logic habitat model was improved by adding water temperature as a key determinant of juvenile Atlantic salmon parr habitat quality. First, salmon experts were consulted to gather their knowledge of salmon parr habitat, then the model was validated with juvenile salmon electrofishing data collected on the Sainte-Marguerite, Matapedia and Petite-Cascapedia rivers (Québec, Canada). The model indicates that when thermal contrasts exist at a site, cooler temperature offered better quality of habitat. Our field data show that when offered the choice, salmon parr significantly preferred to avoid both cold areas (<15 °C) and warm areas (>20.5 °C). Because such thermal contrasts were not consistently present among the sites sampled, the model was only validated for less than 60% of the sites. The results nevertheless indicate a significant correlation between median Habitat Quality Index and parr density for the Sainte-Marguerite River (R² = 0.38). A less important, albeit significant (F-test; p = 0.036) relationship was observed for the Petite-Cascapedia river (R² = 0.14). In all instances, the four-variable (depth, velocity, substrate size and temperature) model provided a better explanation of parr density than a similar model excluding water temperature.     

Ecomorphological correlates in tropical stream fishes of southern India

Synopsis I studied ecomorphological correlations in 18 stream fish species (belonging to five families) commonly occurring in diversity rich habitats of rivers in the central Western Ghats (Southern India). Univariate and multivariate analyses indicated a clear segregation of species in morphological and ecological (habitat) space. Cluster analyses using morphological features segregated species based mainly on their body shape and feeding parameters while habitat parameters segregated species according to their position along the water column and preference to runs or pool habitats. Principal component analyses showed that body size, head dimensions, mouth position and eye diameter were important characters for species segregation in morphological space while run or pool habitat type and the position of the fish in the water column best explained segregation of species in habitat space. Consistencies in some of the species clusters obtained from analyses on morphological characteristics and habitat preferences warrant closer examination of the relationships between them. These were studied employing multivariate methods like canonical correspondence analysis and the Mantel test. The tests revealed weak correlations indicating that in addition to morphology, other factors like behavioural, physiological and evolutionary constraints together are likely to play an important role in ecosystem structuring of these complex tropical assemblages.     

Foraging modes of stream benthic fishes in relation to their predation effects on local prey density

Habitat use and foraging behavior of two benthic insectivorous gobies, Rhinogobius sp. CO (cobalt type) and Rhinogobius sp. DA (dark type), were examined in relation to their predation effects on local prey density in a small coastal stream in southwestern Shikoku, Japan. Correlations among the foraging range, frequency of foraging attempts and current velocity indicated that individuals using fast-current habitats had small foraging ranges and infrequently made foraging attempts while those in slow currents frequently foraged over large areas. The former and the latter were recognized as ambush and wandering foragers, respectively. Interspecific comparisons of habitat use, foraging behavior and prey preference suggested that Rhinogobius sp. CO selectively forage mobile prey by ambushing in fast currents, whereas Rhinogobius sp. DA randomly forage available prey by wandering in slow-current habitats. A cage experiment was conducted to assess prey immigration rate and the degree of predation effects on local prey density in relation to current velocity. The results of the experiment support, at least in part, our initial predictions: (1) prey immigration rates increase with current velocity and (2) the effects of fish predation on local prey density are reduced as current velocity increases. Overall results illustrated a link between the foraging modes of the stream gobies and their predation effects on local prey density: fish adopt ambush foraging in fast currents, where the decrease in prey density tends to be less, whereas fish actively forage over large areas in slow currents, where the decrease in prey is relatively large.     

The Atlantic salmon in fresh water: spawning,rearing and production

Summary Fluvial salmonids have evolved to use the diversity of habitats in natural streams for different life history stages and at different seasons. Required freshwater habitat of Atlantic salmon can be classified generally as that suitable (i) for spawning, (ii) for feeding during the major growing period, and (iii) for overwintering.Spawning habitat of salmon is usually in rapid water at the tail of pools on the upstream edge of a gravel bar, ideally with depths about 25 cm, in mean water velocities of about 30–45 cm s^-1, with maximum velocities about 2 body lengths s^-1, and with a substrate of irregularly shaped stones of cobble, pebble, and gravel.Underyearling salmon (<7 cm TL) are most common in shallow (<15 cm) pebbly riffles, whereas older and larger parr (>7 cm TL) are usually in riffles deeper than 20 cm with a coarse substrate. Depth preference increases with size. Multiple linear regression models quantifying parr habitat have identified substrate as an important variable, with a positive relationship to an index of coarseness. Negative relationships were found with mean stream width, range of discharge, and overhanging cover. Water chemistry, especially alkalinity, nitrates, and phosphates, are important regulators of production. Although similar variables had importance, coefficients among rivers differed. Interactions occur among variables. Further studies are required to quantify productive capacity of habitat for parr. Results suggest that useful models can be derived and if a river system is mapped, and stratified by habitat, then smolt yield could be predicted and the required egg deposition could be estimated.In winter, young salmon shelter among coarse substrate or move to pools, but continue feeding, with larger parr being more active.Feeding is in general opportunistic. Food consists mainly of insects, taken primarily in the water column, but also from the surface and at the bottom. Young salmon in flowing water are highly territorial but are less so in slow or still waters. In fast water, parr use their large pectoral fins to apply themselves to the substrate, allowing them to occupy this type of habitat with little expenditure of energy. Height above the substrate decreases with water velocity, but increases with temperature and social status. Although riffles are preferred habitat, and are relatively more productive, lentic waters can be occupied where there are few predators or severe competitors and may provide significant smolt yield in some systems. Selective segregation minimizes competition between salmon and brook charr or brown trout, but brook charr and brown trout may have negative effects on underyearling salmon, and on parr in pools, whereas salmon have negative effects on small brook charr and brown trout in riffles and flats. Competition by both interference and exploitation results in interactive segregation when the resource, mainly food, becomes limiting.Limited downstream movement of underyearling salmon may occur during the summer. Older juveniles may make upstream movements, but generally migrate downstream, with most movements in the spring, and a lesser peak of activity in the autumn. Dispersal tends to be mainly downstream, indicating that for full distribution, spawning areas are best located upstream. High densities of yearling parr may have negative effects on growth and survival of underyearlings in some river systems, but apparently not in others, so that future research is required in this regard. Density-dependent growth is evident where food is limiting, and can provide an indicator of densities of cohorts so that if a quantitative relationship has been derived, mean size from a sample can give an estimate of the density at that station, with minimum size occurring at carrying capacity. Such regressions vary between habitats with differing productive capabilities, so that future research could provide useful models for assessing productive capacity of a habitat, and optimum densities. Life history strategies can change with changes in density-dependent growth rates. Present stock-recruitment functions do not take environmental variables into consideration, and have limited applicability. Further research is required to determine optimum spawning requirements for salmon in different types of river systems in different geographical areas.